Acta Bot. Croat. 63 (2), 171–202, 2004
Review paper
CODEN: ABCRA 25
ISSN 0365–0588
The genus Campanula L. (Campanulaceae) in Croatia,
circum-Adriatic and west Balkan region
SANJA KOVA^I]*
University of Zagreb, Faculty of Science, Department of Botany and Botanical Garden,
Maruli}ev trg 9a, HR-10000 Zagreb, Croatia
The status of the genus Campanula L. (Campanulaceae) in southeast-European, circum-Adriatic and west Balkan countries (Italy, Slovenia, Croatia, Bosnia and Herzegovina,
Serbia and Montenegro, FYR Macedonia, and Albania) is discussed, according to the local checklists, recent nomenclature and research. The flora of the region comprises at least
84 Campanula species and subspecies, out of which 75% are endemic, with a considerable number of incipient taxa. Accent is placed on the Croatian flora, which contains 30
species and 5 subspecies (42% of the regional taxa), while some older references are
found to be inaccurate or recently unconfirmed. The predominant chromosome number is
diploid, 2n = 34, while the most prevailing life form is hemichryptophytic (97% of the
taxa). More than 30% of the Croatian campanulas are endemic, particularly of the
Isophylla, Heterophylla (Rotundifolia), Pyramidalis and Waldsteiniana lineages, the unsolved relations among which are considered to be the most interesting in the region. The
genus Campanula, in its current circumscription, needs fundamental revision.
Key words: Campanula, Croatia, Adriatic coast, Balkan
Introduction
Members of the family Campanulaceae Juss. s.l. are widespread on most continents,
with up to 90 genera and 2200 species (JUDD et al. 2002). Although the family is found to be
monophyletic (COSNER et al. 1994, EDDIE et al. 2003), there is no single living genus that
could be regarded as the ancestor of the others. Three subfamilies – Campanuloideae,
Cyphoideae and Lobelioideae (SCHÖNLAND 1894) – may as well be recognized at the family level (LAMMERS 1992). There is still dispute even as to the number of recognized genera
within the family, while the criteria for delimiting taxa are problematic due to the complexity of characters within and among genera (EDDIE 1984, 1998).
Ancestors of the recent »campanuloids«, i.e. Campanula taxa and their relatives, probably started to develop in the early Tertiary (55 to 60 million years ago), in the warm climate of Gondwanaland. In Europe, the evolution of this group was additionally facilitated
in the Quaternary, by the Alpine Orogenesis and the Ice Ages. Today, Campanulaceae in
* Corresponding address: sanja@botanic.hr
ACTA BOT. CROAT. 63 (2), 2004
171
KOVA^I] S.
Eurasia are most frequently represented by the related genera Asyneuma Griseb. et
Schenk., Campanula, Phyteuma L. and Symphyandra DC., which are exceptionally rich
and uniquely developed in the higher mountains of Western and Central Europe, in the
Mediterranean area, the Middle East and the Caucasus. Apparently, the most primitive
campanuloids today are gathered around the Mediterranean Sea, e.g. Trachelium L. (including the eastern groups Tracheliopsis and Diosphaera), Legousia Durande, Edraianthus DC. and Jasione L., followed by archaic »true« campanulas of the subsection
Annuae (Boiss.) Fed. and subgenus Roucela (Dumort.) Damboldt (CONTANDRIOPOULOUS
1984, EDDIE 1998). It seems that the whole campanuloid lineage spread over the northern
hemisphere from this major evolutionary centre in the Mediterranean region (EDDIE et al.
2003).
The numerous Campanula taxa – 350 to 450 species of bellflowers, bluebells, harebells
and starbells, with many taxa lower than species – mostly inhabit steppe and mountainous
habitats in temperate and subtropical zones of the northern hemisphere. Many of them are
orographically, edaphically and microclimatically highly specialized and characterized by
extensive polymorphism. The first approaches to the Campanula taxonomy were often
geographically limited and based exclusively on (very variable) morphological characteristics, which led to the circumscription of a large number of species, subspecies, varieties,
subvarieties, forms, and even subforms (WITASEK 1906, HRUBY 1930, 1934). Many authors
over the centuries also tried to develop a suitable sub-classification of this large genus, in
order to help in systematization (DE CANDOLLE 1830, BOISSIER 1875, FIORI 1927, FEDOROV
1957, GADELLA 1964, PODLECH 1965, DAMBOLDT 1978, KOVANDA 1970b, 1977, GESLOT
1984, KOLAKOVSKY 1992). These positive efforts resulted though in many assemblages of
frequently heterogeneous taxa, causing more and more confusion. Such serious problems
in delimiting taxa have recently initiated partial investigations in Campanula, using classical taxonomic methods in combination with molecular and statistical analyses, which have
resolved some of the taxonomic problems of evolutionary-related and geographically-closer groups (CARLSTRÖM 1986, KOVANDA and AN^EV 1989, RUNEMARK and PHITOS
1996, EDDIE and INGROUILLE 1999, OGANESIAN 2001, SAEZ and ALDASORO 2003). In spite
of all this, no generally accepted criteria for the subgeneric delimitation of Campanula exist: phylogeny and relationships among the »campanuloids« in general are still poorly
known.
It is traditionally considered that at least two major Campanula lineages (»sections«)
have separate evolutionary patterns (BOISSIER 1875, FEDOROV 1957, CHARADZE 1949, FEDOROV and KOVANDA 1976), a conclusion based mainly on calyx morphology (presence/absence of appendages between the lobes) and on the mode of capsule dehiscence (apical or
lateral, valvate or porate). The section Rapunculus Dumort. is most probably older, while it
is widely distributed (also in North America), although outnumbered by the taxa within the
Section Campanula s.str. However, the most recent investigations indicate that not even
those two lineages are entirely natural.
The latest molecular phylogenetic research (EDDIE et al. 2003) reveals some very interesting details: according to the analysis of ITS-sequences of nuclear ribosomal DNA, the
genera Azorina, Campanulastrum, Edraianthus and Phyteuma actually nest within Campanula. The various taxa of that assemblage further group with related genera in three main
clades: the Campanula s.str.-clade, which includes Azorina, Edraianthus, Symphyandra,
Trachelium and some others; the Rapunculus-clade, with the addition of Adenophora,
172
ACTA BOT. CROAT. 63 (2), 2004
THE GENUS CAMPANULA IN CROATIA
Asyneuma, Legousia, Phyteuma and others; and a small clade called the »transitional-taxa«, composed of Jasione and several other genera. Although the family is monophyletic, the genus Campanula is therefore polyphyletic.
It appears that in its current circumscription Campanula is a classic »waste bin« taxon
that includes a number of dubious taxa left over after morphologically well-characterized
groups of campanuloids were removed to separate genera. Many taxa during the years have
been placed in Campanula for convenience, while even some of the satellite sister-genera
could well be restored to Campanula. Taking all of this into consideration, it is clear that
this heterogeneous genus (collective genus acc. to GADELLA 1964), as it now stands, is in
need of fundamental revision.
The genus Campanula in the west Balkans and circum-Adriatic region
According to the still valid literature (compilation of the Flora Europaea and the
MedCheck-list, EURO+MED, in preparation, as well as the Atlas Florae Europeae with
Campanulaceae included), approximately 200 to 250 Campanula species and subspecies
are listed for Europe (FEDOROV and KOVANDA 1976) and the Mediterranean region (GESLOT
1984), respectively. The actual number of taxa may be greater considering the fact that so
few have been adequately investigated.
In south-eastern Europe research on the genus Campanula began with the floristic investigations in 18th century – and yet even 300 years later the quest for new taxa is not over
(e.g. LUCCHESE 1993, RAN\ELOVI] and ZLATKOVI] 1998, LAKU[I] and CONTI 2004). Nevertheless, older botanists (e.g. BORBAS 1883, POSCHARSKY 1896, BECK-MANAGETTA 1901,
ADAMOVI] 1909, 1911, 1929, HEGI 1908–1931, JAVORKA 1924–1925, FIORI 1927, HAYEK
1925–1933) noticed a tremendous diversity of campanulas in the west Balkans and both
Adriatic coasts, occupying different habitats, from the shoreline to the highest mountains.
The circum-Adriatic regions of the Balkans and Apennines (Fig. 1) form the northern section of the eastern Mediterranean biogeographical region (QUEZEL 1985). These two large
European peninsulas are generally highly related in floristic and vegetation composition
(PIGNATTI 1982, JUNIKKA and UOTILA 2002), as well in Campanula taxa (DAMBOLDT 1965a,
FRIZZI 1988, BERNINI et al. 2002), which comprise a large share of the genus’ diversity and
endemicity in both European and Mediterranean checklists. Circum-Adriatic campanulas
share morphological characteristics and the similar vegetation conditions of the karstic
Mediterranean-mountainous region, i.e. heliophytic chasmophyta of mountain rock crevices, scree and rubble. Cytological investigations have confirmed high similarity in number
and form of chromosomes (GUTERMANN 1961, MERXMÜLLER and DAMBOLDT 1962, BÖCHER
1963, GADELLA 1964, PODLECH and DAMBOLDT 1964): campanulas of this region are primary diploids, often endemics and/or relicts reduced to small geographic areas, sometimes
listed in the local Red Books.
The distribution of Campanula species and subspecies in the countries of the eastern
Adriatic and western Balkan region is given in table 1. Nomenclature is adjusted according
to the data of FEDOROV and KOVANDA (1976) and GESLOT (in GREUTER et al. 1984), and supplemented by local floras and research of Italy (PIGNATTI 1982, LUCCHESE 1993), Slovenia
(MARTIN^I^ 1999, JOGAN ed. 2001), Croatia (LOVA[EN-EBERHARDT 2000), Bosnia and
Herzegovina (BJEL^I] 1983, [ILI] 2000, [OLJAN 2002), Former Yugoslav Republic of
Macedonia (NIKOLOV 2004), Serbia and Montenegro (ROHLENA 1942, OBRADOVI] 1974,
ACTA BOT. CROAT. 63 (2), 2004
173
KOVA^I] S.
Fig. 1. Position of the circum-Adriatic and west Balkan countries in Europe.
DIKLI] 1977, GAJI] 1986, RAN\ELOVI] and ZLATKOVI] 1998) and Albania (HAYEK 1923,
DEMIRI 1981, QUOSIA 1996). Endemicity and polymorphism are noted to illustrate each
taxon’s evolutionary pattern and range of variability.
According to the data in table 1, the flora of the circum-Adriatic and western Balkan region contains 84 Campanula taxa: 65 species and 19 subspecies. This number is not final,
while the immense range of lower taxa of uncertain taxonomical status (hereafter called the
»incipient« species) is neglected. The flora of Serbia and Montenegro contains 48 Campanula species and subspecies, of north-eastern Italy 42, of Albania 37, of Croatia 35 (discussed separately), of Bosnia and Herzegovina 33, of FYR Macedonia 33, and of Slovenia
26 (Fig. 2), but these numbers are not final. Italian campanulas of northwest and southwest
distribution, absent from the other countries in the region, were not listed: these are (according to PIGNATTI 1982) C. bertolae Colla, C. cenisia L., C. elatines L., C. elatinoides
Moretti, C. fragilis Cyr., C. forsythii (Arcang.) Podlech, C. isophylla Moretti, C. macrorrhiza Gay ex DC., C. pollinensis Podlech, C. pseudostenocodon Lacaita, C. rhomboidalis
174
ACTA BOT. CROAT. 63 (2), 2004
THE GENUS CAMPANULA IN CROATIA
50
48
42
37
40
35
33
33
26
30
20
10
0
S&M
NE Italy Albania
Croatia
B&H
FYRM Slovenia
Fig. 2. Number of Campanula species and subspecies in countries of the circum-Adriatic and
western Balkan region.
L., C. sabatia De Not., and C. stenocodon Boiss. et Reuter. Taxa stated as cultivated or dubious were excluded. As many as 63 taxa (75%) listed in Table 1 are indigenous to the region, and many of them subendemics with tiny distributions. About 35% of the species are
variable, some of them tremendously so (e.g. C. rotundifolia, C. glomerata L., C. patula
L.). Comparative experimental investigations using modern methods have rarely been conducted among the southeast European campanulas (FRIZZI 1988, KOVA^I] et al. 2003),
which makes sub-generic systematization extremely difficult.
The genus Campanula in Croatia
According to the data of LOVA[EN-EBERHARDT (2000), the genus Campanula is represented in Croatia by 38 species and 8 subspecies, as shown in table 2. Lova{en-Eberhardt
marked taxa that she considered to be of dubious taxonomic status or controversial occurrence in Croatia with »?«. She also treated C. fenestrellata Feer subsp. istriaca (Feer)
Damboldt as species C. istriaca Feer, which is not considered valid by most of the recent
authors. Complex subgeneric delimitation is adjusted by the present author, following the
most prominent works of DAMBOLDT (1965a, 1968), FEDOROV (1957), KOVANDA (1970b,
1977), FEDOROV and KOVANDA (1976) and GESLOT (1984), in order to represent the variety
of different views concerning the inter-taxonomic relationships of the genus. Additional
literature on Campanula and Campanulaceae in Croatia should be found in NIKOLI]
(2000).
Based on my research, I suggest that 7 species and 3 subspecies of Campanula are of
highly dubious existence in Croatia and should be removed from the Croatian checklist
pending further investigation. In table 2 I compare data presented by LOVA[EN-EBERHARDT
(2000) to those of the most outstanding older researchers of Croatian flora: VISIANI (1847,
ACTA BOT. CROAT. 63 (2), 2004
175
ACTA BOT. CROAT. 63 (2), 2004
Magyar Bot. Lap. 5: C. linifolia ssp. albanica (Witasek)
246 (1906)
Hayek, C. rotundifolia ssp. hellenica
Hayek, C. hellenica Hayek
P
2
alpestris
All.
Auct. Syn. Stirp.
Horti Taur. 11
(1773)
P
3
alpina ssp.
alpina
Jacq.
Enum. Stirp. Vindob.
36: 210 (1762)
P
4
– ssp. orbelica
(Pan~i})
Urum.
Spis. Balg. Akad.
C. alpina Jacq. ssp. orbelica Pan~i}
Nauk, 28: 147 (1923)
P
5
barbata
Linne
Syst. Nat. ed 10:
926 (1759)
C. firmiana Vandelli
6
beckiana
Hayek
Fl. Steierm. 2(1):
455 (1912)
C. hostii Baumg., C. baumgartenii J.
Becker ssp. beckiana
7
bononiensis
Linne
Sp. Pl. 165 (1753)
?C. obliquifolia Ten., C. ruthenica Bieb.,
C. simplex Lam. non Stev., incl. var. cana
Simonkai, var. concolor Pan~i}
8
calycialata*
Ran|elovi}
et Zlatkovi}
Flora Medit. 8,
85-92 (1998)
C. allionii Vill.
P
P
P
P
P
?
P
P
P
P
?
P
P
P
P
P
P
P
Albania
Witasek
FYR Macedonia
albanica
Montenegro
P
1
Serbia &
pt
Herzegovina
E
Bosnia &
Synonyms, included taxa
Croatia
Source
Slovenia
Author
NE Italy
Nr Campanula L.
P
P
?
P
?
P
P
P
P
P
P
P
P
P
P
P
KOVA^I] S.
176
Tab. 1. Campanula species and subspecies distribution in the circum-Adriatic and west Balkan countries: northeastern Italy, Slovenia, Croatia, Bosnia &
Herzegovina, Serbia & Montenegro, FYR Macedonia, and Albania. E = endemic taxon, pt = polymorphic taxon, grey cells = unreliable reference,
cult. = taxon stated as cultivated,? = taxon stated as dubious.
C. linifolia Scop. non L., C. scheuchzeri
Vill. var. carnica. Posp., C. rotundifolia
Bertol p.p. nec L., C. rotundifolia L. var.
forsythii Arcangeli, C. macrorrhiza Parl.
p.p. nec Gay
P
P
P
P
P
10 cervicaria
Linne
Sp. Pl. 167 (1753)
var. cervicaria, var. micrantha
P
P
P
11 cespitosa
Scop.
Fl. Carniol. 2(1):
143 (1772)
incl. C. caespitosa
P
P
P
12 cochleariifolia
Lam.
Encycl. Meth. Bot.
1:578 (1785)
C. pusilla Haenke, C. pusilla ssp. croatica P
Hruby, C. hochstetteri Schott et al., C. notata
Schott et al., C. tenella Jordan, C. caespitosa
Vill. non Scop., C. bellardii Allioni
P
P
P
13 crassipes
Heuffel
Österr. Bot. Zeitsch.
8: 27 (1858)
14 dichotoma
Linne
Cent. Pl. 2: 10
(1756)
15 erinus
Linne
Sp. Pl 169 (1753)
Roucela erinus (L.) Dumort, C. nanella
P. A. Smirn.
J. Bot. 28: 272
(1890)
C. garganica Vis. non Ten., C. garganica
Ten. ssp. fenestrellata (Feer) Hayek,
C. lepida Feer
16 fenestrellata ssp. Feer
fenestrellata
P
P
Albania
Croatia
Deutschl. Fl. 3(2):
158 (1826)
carnica
FYR Macedonia
Slovenia
Schiede ex
Merth. et
Koch
9
Montenegro
pt
Serbia &
E
Herzegovina
Synonyms, included taxa
Bosnia &
Source
Nr Campanula L.
P
P
P
?
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
177
THE GENUS CAMPANULA IN CROATIA
Author
NE Italy
ACTA BOT. CROAT. 63 (2), 2004
Tab. 1. – continued
KOVA^I] S.
ACTA BOT. CROAT. 63 (2), 2004
(Feer)
Damboldt
Jour. Bot. 28: 271
(1890)
P
19 foliosa
Ten.
Fl. Nap. 1, Prodr. 16
(1811)
P
20 formanekiana
Degen et
Dörfl.
Denkschr. Akad.
Wiss. Math.-Nat.
Kl. (Wien) 54: 728
(1899)
P
21 garganica
Ten.
Cat. Sem. Horti
Nap. (1827)
C. barbeyi Feer
22 glomerata ssp.
glomerata
Linne
Sp. Pl. 166 (1753)
C. aggregata Willd., C. speciosa Hoenem
non Pourr. C. viridis Rchb. (=var.
mediterranea Borb.), C. cephalotes
Fischer; div. var. et f.
23 – ssp.
(Schult.)
cervicarioides P. Fourn.
Quatre Fl. Fr. 914
(1939)
C. cervicarioides Schult.
P
P
24 – ssp. elliptica
Mitt. Thür. Bot.
Ges. 1(1): 118
(1949)
C. elliptica Schult.
P
P
Albania
18 – ssp. istriaca
FYR Macedonia
P
Montenegro
Bot. Jahrb. Syst. 84: C. debarensis Rech.f.
358 (1965)
Serbia &
(Rech.f.)
Damboldt
P
pt
Herzegovina
17 – ssp.
debarensis
C. istriaca Feer, C. garganica Ten. ssp.
istriaca (Feer) Hayek, C. fenestrellata
Feer ssp. istriaca (Feer) Fed.
E
Bosnia &
Source
Croatia
Author
Slovenia
Nr Campanula L.
(Kit. ex
Schult.)
O. Schwarz
Synonyms, included taxa
NE Italy
178
Tab. 1. – continued
P
P
P
P
P
P
?
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
26 – ssp. hispida
(Witasek)
Hayek
Prodr. Fl. Penins.
C. glomerata f. hispida Witasek, C.
Balcan. 2: 532 (1930) lamioides Witasek
27 – ssp. serotina
(Wettst.)
O. Schwarz
Mitt. Thür. Bot. Ges. C. serotina Wettst.
1(1): 118 (1949)
P
28 grossekii
Heuffel
Flora (Regensburg)
16: 353 (1833)
P
29 hawkinsiana
Hausskn.
et Heldr.
Mitt. Georg. Ges.
C. halacsyana Bald.
(Jena) 5(2): 87 (1887)
P
30 hercegovina
Degen et
Fiala
Österr. Bot. Zeitsch. ?C. tarana K. Maly, div. var, subvar, f.,
44: 303 (1894)
subf.
P
31 jordanovi
An~ev et
Kovanda
Preslia 61: 200
(1989)
P
32 justiniana
Witasek
Magyar Bot. Lapok C. linifolia ssp. justiniana (Witasek)
5:245 (1906)
Hayek
P
33 latifolia
Linne
Sp. Pl. 165 (1753)
C. eriocarpa Bieb., incl. C. latifolia var.
macrantha (A.DC.) Hornemann
34 lingulata
Waldst. et
Kit.
Descr. Icon. Pl.
Hung. 65 (1801)
C. cichoriacea Sm., C. capitata Sims.,
incl. var. farinosa Roch., elliptica Kit.,
mediterranea Borb., div. f. et subvar.
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
Albania
C. farinosa (Rochel) Andr., C. glaucophylla
Schlosser et Vuk., C. desertorum Weinm.
FYR Macedonia
Fl. Alsace 1: 378
(1852)
Montenegro
(Rochel)
Kirschl.
Serbia &
25 – ssp. farinosa
pt
Herzegovina
E
Bosnia &
Synonyms, included taxa
Croatia
Source
Slovenia
Author
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
THE GENUS CAMPANULA IN CROATIA
179
Nr Campanula L.
NE Italy
ACTA BOT. CROAT. 63 (2), 2004
Tab. 1. – continued
KOVA^I] S.
Sp. Pl. 167 (1753)
38 micrantha
Bertol.
Fl. Ital. 7: 623
(1851)
39 moesiaca
Velen.
Sitz. Königl. Böhm. C. hirsuta Pant.
Ges. Wiss. Prag,
Math.-Nat. Cl.
1892: 385 (1893)
P
40 moravica ssp.
moravica
(Spitzner)
Kovanda
Folia Geobot.
Phytotax. Bohem.
3: 409 (1968)
C. rotundifolia L. var. moravica Spitzn.
P
C. rotundifolia L. ssp. xylorrhiza O.
Schwartz
P
C. marchesettii auct. Fl. It. Centr., C.
apennina (Podlech) Podlech
P
P
P
P
P
P
P
P
P
P
P
P
P
Albania
Linne
P
P
cult.
37 medium
P
FYR Macedonia
Abh. K. K. Zool.Bot. Ges. Wien
1(3): 32 (1902)
Montenegro
Witasek
Herzegovina
36 marchesettii
Bosnia &
in Willd., Enum. Pl. C. multiflora Waldst. et Kit.
Hort. Berol.: 213
(1809)
pt
Croatia
Waldst. et
Kit.
C. scheuchzeri ssp. marchesettii non Vill.
E
Slovenia
35 macrostachya
41 – ssp. xylorrhiza (O. Schwartz) Folia Geobot.
Kovanda
Phytotax. Bohem.
3: 409 (1968)
Synonyms, included taxa
cult.
Source
NE Italy
Author
Serbia &
ACTA BOT. CROAT. 63 (2), 2004
Nr Campanula L.
(NW Italy)
180
Tab. 1. – continued
P
P
P
P
?
P
Linne
Sp. Pl. 163 (1753)
44 – ssp. abietina
(Griseb. et
Schenk.)
Simonkai
Enum. Fl. Transsilv. C. abietina Griseb. et Schenk.,
383 (1887)
C. stefanoffi F. Hermann, C. vajdae
Penzes
P
45 – ssp. costae
(Willk.)
Fedorov
Bot. J. Linn. Soc.
67: 281 (1973)
C. costae Wullk.
P
46 – ssp. epigaea
(Degen)
Hayek
Repert. Spec. Nov.
Regni Veg. Beih.
30(2): 546 (1930)
C. epigaea Degen,
C. velenovskyi Adamovi}
P
47 – ssp. jahorinae
(K. Maly)
Greuter et
Burdet
Willdenovia 11: 40
(1981)
C. patula f. jahorinae K. Maly
P
48 persicifolia ssp.
persicifolia
Linne
Sp. Pl. 164 (1753)
C. cristallocalyx Adamovi},
C. hispida Lej., C. pumila F. W. Schmidt,
incl. div. var. et f.
C. monanthos Pant., C. flaccida (Wallr.)
Dalla Torre et Sarnth., C. neglecta
Schult.; incl. C. patula ssp. costae
(Willkomm.) Fed., C.p. ssp. epigaea
(Janka) Hayek, div. f. et subf.
Albania
43 patula ssp.
patula
P
FYR Macedonia
Pl. Crit. 4: 18
(1826)
Montenegro
Rchb.
Serbia &
42 morettiana
pt
Herzegovina
E
Bosnia &
Synonyms, included taxa
Croatia
Source
Slovenia
Author
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
THE GENUS CAMPANULA IN CROATIA
181
Nr Campanula L.
NE Italy
ACTA BOT. CROAT. 63 (2), 2004
Tab. 1. – continued
KOVA^I] S.
Syst. Nat. ed 10:
926 (1759)
51 phygria
Jaub. et
Spach
Ill. Pl. Orient. 3:
233 (1848)
P
C. ramosissima Griseb. non Sibth. et
Sm.; div. f.
?
P
P
P
P
P
ACTA BOT. CROAT. 63 (2), 2004
52 portenschlagiana Schultes
in Römer et
C. muralis Portenschl., C. hederaefolia
Schultes, Syst. Veg. Portenschl, C. morettiana Vis. non Rchb.
5:93 (1820)
P
P
53 poscharskyana
Degen et
Fiala
Magyar Bot. Lapok C. elatines auct., non L. Reichenb.
7: 103 (1908)
P
P
54 pyramidalis
Linne
Sp. Pl. 164 (1753)
P
55 raineri
Perpenti
Bibliot. Ital. 5: 134
(1817)
P
P
56 ramosissima
Sibth. et Sm. Fl. Graecae Prodr.
1: 137 (1806)
C. loreyi Pollini, C. baldensis Balb.
P
P
57 rapunculoides
Linne
C. trachelioides M.Bieb., C. cordifolia
C. Koch, C. rhomboidea Falk,
C. rhomboidalis Gorter, C. lunariaefolia
Reich., C. setosa Fischer; div. f.
Sp. Pl. 165 (1753)
P
Albania
Linne
P
FYR Macedonia
50 petraea
pt
Montenegro
P
Serbia &
E
C. sessiliflora K. Koch, C. persicifolia L.
ssp. eriocarpa (K.Koch) U. Dettmann et
Rothm.
Herzegovina
Synonyms, included taxa
Willdenovia 12: 36
(1982)
Bosnia &
Source
(C. Koch)
Greut. et
Burd.
Croatia
Author
49 – ssp.
sessiliflora
Slovenia
Nr Campanula L.
NE Italy
182
Tab. 1. – continued
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
?
P
P
P
P
NE Italy
Slovenia
Croatia
Bosnia &
Herzegovina
Serbia &
Montenegro
FYR Macedonia
Albania
ACTA BOT. CROAT. 63 (2), 2004
Tab. 1. – continued
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
?
P
P
Nr Campanula L.
Author
Source
Synonyms, included taxa
E
pt
58 rapunculus
Linne
Sp. Pl. 164 (1753)
C. elatior Hoffmans. et Link, C. verruculosa
Hoffmans. et Link, incl. C. rapunculus
ssp. lambertiana Boissier, C.r. var.
spiciformis Boissier, div. var, f. et subf.
P
59 reatina
Lucchese
Fl. Medit. 3: 266
(1993)
60 rotundifolia
(s.l.)
Linne
Sp. Pl. 163 (1753)
? C. polymorpha (Witasek) Tacik, ? C.
tarana K. Maly, C. scheuchzeriformis
Hayek, C. lobata Schloss. et Vuk., C.
racemosa (Kra{an) Witasek, C. pinifolia
Uechtr.; div. ssp., var. et f.
P
61 scheuchzeri
Villars
Prosp. Hist. pl.
Dauph. 22 (1779)
C. linifolia Haenke non Scop., C. rotundifolia
L. ssp. balcanica (Adamovi}) Stoj. et
Stefanov, C. uniflora Vill. non L.
P
62 scutellata
Griseb.
Spic. Fl. Rumel.2:
282 (1846)
63 secundiflora
Visiani et
Pan~i}
Mem. Imp. Reale
Inst. Veneto Sci. 10:
442 (1861)
64 serrata
(Kit.)
Hendrych
Taxon 11: 123
(1962)
P
P
P
P
P
P
P
P
THE GENUS CAMPANULA IN CROATIA
183
C. pseudolanceolata Pant., C. napuligera
Schurr., Thesium serratum Schultes
(basion.)
P
KOVA^I] S.
ACTA BOT. CROAT. 63 (2), 2004
66 – ssp.
(Jav.)
divergentiformis Domin
Preslia 13-15: 222
(1936)
C. divergens W. et Kit. in Willd.,
C. sibirica var. divergentiformis Jav.,
C. spathulata W. et K.
P
67 sparsa ssp.
sparsa
Friv.
Magyar Tud. Tars.
Evk. 4(2): 201
(1840)
C. expansa Friv., C. sparsa ssp.
frivaldsky (Steudel) Hayek
P
68 – ssp.
sphaerothrix
(Griseb.)
Hayek
Repert. Spec. Nov.
Regni Veg. Beih.
30(2): 547 (1930)
C. sphaerothrix Gris., C. expansa var.
sphaerothryx Boiss.
P
69 spatulata ssp.
spatulata
Sibth. et Sm. Fl. Graecae Prodr.
1: 137 (1806)
C. sibthorpiana Halacsy
P
70 – ssp.
spruneriana
(Hampe)
Hayek
Repert. Spec. Nov.
Regni Veg. Beih.
30(2): 545 (1930)
C. spruneriana Hampe
P
71 spicata
Linne
Sp. Pl. 166 (1753)
72 tanfanii
Podlech
73 thyrsoides ssp.
thyrsoides
Linne
Sp. Pl. 167 (1753)
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
Albania
C. hohenackeri Fisch., C. parviflora
Lam., C. sibirica ssp. taurica Juz.
FYR Macedonia
Sp. Pl. 167 (1753)
Montenegro
Linne
pt
Serbia &
65 sibirica ssp.
sibirica
C. macrorrhiza Gay ex DC. var.
angustiflora Tanfani
E
Herzegovina
Synonyms, included taxa
Bosnia &
Source
Croatia
Author
Slovenia
Nr Campanula L.
NE Italy
184
Tab. 1. – continued
P
P
P
P
P
P
P
C. waldsteiniana R. et S. var. tubulosa DC., P
C. waldsteiniana R. et S. var. freyeri
Reichenb., C. tommasiniana Reuter,
C. waldsteiniana R. et S. var.
tommasiniana (Reuter) Arcangeli
76 trachelium ssp.
trachelium
Linne
Sp. Pl. 166 (1753)
C. urticifolia F. W. Schmidt, C. trachelioides
Bieb., C. athoa Boiss. et Heldr., incl.
C. trachelium var orientalis Boiss., div. f.
77 – ssp. athoa
(Boiss. et
Heldr.)
Hayek
Prodr. Fl. Penins.
Balcan. 2: 541
(1930)
C. athoa Boiss. et Heldr.
185
P
79 tymphea
Mitt. Georg. Ges.
(Jena) 5(2): 87
(1887)
P
Hausskn.
P
P
Cat. Pl. Hort. Neap. Asyneuma trich. (Ten.) K. Maly,
App. 1: 35 (1815)
Podanthum trich. (Ten.) Boiss., Phyteuma
trich. (Ten.) Tanfani, C. minae Strobl
P
P
P
?
P
P
P
P
P
P
P
P
P
P
P
P
P
P
78 trichocalycina** Ten.
P
P
Albania
Arch. Fl. France
Allemagne 229
(1852)
P
FYR Macedonia
C. Koch
?
Montenegro
75 tommasiniana
P
Serbia &
Ber. Bayer. Bot.
C. th. var. carniolica Sünd.
Ges. 37: 111 (1964)
P
P
P
THE GENUS CAMPANULA IN CROATIA
74 – ssp. carniolica (Sünd.)
Podlech
pt
Herzegovina
E
Bosnia &
Synonyms, included taxa
Croatia
Source
Slovenia
Author
NE Italy
Nr Campanula L.
(SW Italy)
ACTA BOT. CROAT. 63 (2), 2004
Tab. 1. – continued
KOVA^I] S.
ACTA BOT. CROAT. 63 (2), 2004
Andrews
Bot. Repos. 6: 396
(1804)
C. tenorii Moretti, C. mrkvickiana Velen.
P
82 waldsteiniana
Schultes
Syst. Veg. 5:99
(1820)
C. flexuosa Waldst. et Kit. non Michx.
P
83 witasekiana
Vierh.
Mitt. Naturwiss.
?C. inconcessa Schott et al. nom ambig.,
Vereins Univ. Wien, C. scheuchzeri Vill. ssp. witasekiana
2(4): 72 (1906)
(Vierh.) Hayek, div. var. et f.
84 zoysii
Wulf.
Collect. Bot. 2: 122
(1789)
P
P
P
P
P
P
P
P
P
P
P
P
P
Albania
81 versicolor
P
FYR Macedonia
Math. Termeszettud. C. farinulenta A. Kern. et Wettst.,
Ertesitö 1: 81 (1883) C. bulgarica Witasek, C. balcanica Hruby
p.p., C. rotundifolia L. ssp. velebitica
(Borbas) Hayek, C.r. ssp. bulgarica
(Witasek) Hayek, div. var. et f.
Montenegro
Borbas
pt
Serbia &
E
Herzegovina
80 velebitica
* extinct in the wild (Ran|elovi}, pers. cont.)
** Asyneuma pichleri acc. to LAKU[I] and CONTI (2004)
Synonyms, included taxa
Bosnia &
Source
Croatia
Author
Slovenia
Nr Campanula L.
NE Italy
186
Tab. 1. – continued
P
P
P
P
P
P
P
P
P
P
P
THE GENUS CAMPANULA IN CROATIA
1872), NEILREICH (1868, 1869), SCHLOSSER and VUKOTINOVI] (1869), HIRC (1908, 1912),
ROSSI (1924), JAVORKA (1925), ROSSI (1930), DEGEN (1938) and DOMAC (1950, 1994). Herbaria specimens of the Croatian Natural History Museum (CNHM), the Vienna Natural
History Museum (W), Vienna University (WU, including Herbarium Halaczy), Zagreb
University (ZA) and the Professor Ivo and Marija Horvat Herbarium (ZAHO) were examined. As campanulas of common occurrence in Croatian flora are present in all of the
named Herbaria, and dubious taxa just in ZA and WU, these two collections are listed as
exemplary in table 2. Just species and subspecies were taken into consideration.
In his »Flora Dalmatica« VISIANI (1847, suppl. 1872) listed all taxa of the Campanulaceae family as various sections of the genus Campanula. He registered 24 Campanula
species and subspecies, while some were included in the range of other taxa (e.g. C.
portenschlagiana Schult. as C. morettiana Reichenb.). NEILREICH (1868, suppl. 1869)
again assigned 24 taxa to the Croatian flora, with C. pulla L. (according to the localities, it
could have been C. justiniana Witasek), C. beckiana Hayek (probably some other incipient
taxon of the Rotundifolia group) and C. barbata L. (most likely cultivated). SCHLOSSER and
VUKOTINOVI] (1869) in their »Flora Croatica« quoted 23 species and subspecies known in
recent times in the Croatian flora. There were again C. barbata, C. beckiana, C. pulla (perhaps misspelled C. pusilla Haenke, syn. for C. cochleariifolia Lam., acc. to HIRC 1908), C.
morettiana (= C. portenschlagiana) and C. macrostachya Willd. found around Zemun in
Vojvodina, western Serbia. It is important to emphasize the good reasons for including recently absent taxa into the Croatian flora of that time. Firstly, there was a lack of efficient
determination keys and scientifically recognized taxa in the region (i.e.? many heterophyllous taxa were referred as C. rotundifolia, and isophyllous taxa as C. garganica Ten.).
Secondly, Croatian borders changed considerably over the years, expanding to include
parts of recent Serbia and Montenegro, and Bosnia and Herzegovina, while ceding the
Istria peninsula, Dalmatian coast and islands to Italy.
HIRC (1908, 1912) in his revisions of older authors corrected most of the misinterpretations and quoted 32 Campanula species and subspecies known in recent times. In his »Materials for the flora of southern Croatia« (1924) ROSSI listed 21 Campanula taxa, and in the
»Flora of Hrvatsko Primorje« (1930) 26 taxa. He wrongly assigned C. albanica Witasek
(most probably C. velebitica Borbas) to the Croatian flora, and stated that some taxa were
dubious. JAVORKA (1925) investigated the Croatian flora as a part of his »Magyar Flora«, in
which the magnificent »Iconographia« is still valuable for this part of Europe. He regarded
34 Campanula species and subspecies as belonging to Croatia, including C. barbata, C.
spicata L., C. serrata (Kit.) Hendrych and C. alpina Jacq., as dubious. HAYEK (1931), exploring the flora of the Balkans, listed 29 Campanula species and subspecies (with a number of lower taxa) for the Croatian area of that time. DEGEN (1938) noted 25 campanulas for
the Velebit mountain region (C. spicata L. as dubious), with three taxa absent today (but
possible, and very interesting): C. trachelium L. subsp. athoa (Boiss. et Heldr.) Hayek, C. x
murrii Dalla Torre et Sarnth. (hybrid between C. cochleariifolia and C. scheuchzeri Vill.)
and C. »staubii« Uechtr. (a common monstrosity of C. pyramidalis L.). Campanula
barbata and C. medium were listed as cultivated. DOMAC quoted (1994) 29 Campanula
species of recent circumscription. The Herbaria of Zagreb (ZA) and Vienna (WU) Universities have most of the Campanula taxa listed by LOVA[EN-EBERHARDT (2000), but lack
those that are missing in the works of the aforementioned authors.
ACTA BOT. CROAT. 63 (2), 2004
187
Rossi 1924
Javorka 1925
Rossi 1930
Hayek 1931
16
Hp
P
P
P
P
P
P
P
P
rapunculus L., Sp. Pl. 164 (1753)
20
Hb
P
P
P
P
P
P
P
P
patula L., Sp. Pl. 163 (1753),
ssp. patula
20
(40)
Hb
P
P
P
P
P
P
P
ramosissima Sibth. et Sm., Fl.
Graecae Prodr. 1: 137 (1806)
20
T
P
P
P
P
member of Croatian
flora (Kova~i})
Hirc 1908 /1912/
persicifolia L., Sp. Pl. 164
(1753), ssp. persicifolia
Domac 1950 /1994/
life
form
Campanula species
and subspecies
HERBARIA
Degen 1938
2n
Schloss.-Vuk. 1869
Campanula – references by older authors
Neilreich 1868 /1869/
Lova{en-Eberhardt (2000)
Visiani 1847 /1872/
syn. Subsection, Series,
Group, Aggregate
Subsection
(Fedorov 1957)
Campanulastrum
(Fourr.) Fedorov
Section
(Fedorov 1957)
Rapunculus (Fourr.) Boiss.
ACTA BOT. CROAT. 63 (2), 2004
Rapunculus Dumort.
Section (Fedorov and
Kovanda 1976)
Subgeneric delimitations
ZA
WU
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
KOVA^I] S.
188
Tab. 2. Campanula species and subspecies distributed in Croatia (acc. to LOVA[EN-EBERHARDT 2000) with subgeneric delimitations, diploid chromosome
number (2n) and life forms, compared to older literature and herbaria data. Hb = Hemichryptophyta-biennial, Hp = Hemichryptophyta-perennial, T
= Therophyta (annual); grey cells = unreliable reference or reference from a recently non-Croatian locality, cult. = taxon stated as cultivated,? =
taxon stated as dubious.
34
Hb(p)
P
P
lingulata Waldst. et Kit., Descr.
Icon. Pl. Hung. 65 (1805)
34
Hb
P
?
erinus L., Sp. Pl. 169 (1753)
28
T
P
P
189
Roucela (Dumort.)
Damboldt
Cervicaria Gris.,
Glomeratae Schur
member of Croatian
flora (Kova~i})
– ssp. divergentiformis (Jav.)
Domin, Preslia 13-15: 222 (1936)
P
Domac 1950 /1994/
P
cult.
HERBARIA
Degen 1938
P
Hayek 1931
Hb(p)
Rossi 1930
34
Javorka 1925
sibirica L., Sp. Pl. 167 (1753),
ssp. sibirica
Rossi 1924
Hb
Hirc 1908 /1912/
34
Schloss.-Vuk. 1869
Neilreich 1868 /1869/
Visiani 1847 /1872/
syn. Subsection, Series,
Group, Aggregate
Subsection
(Fedorov 1957)
Quinquelocula- medium L., Sp. Pl. 167 (1753)
r(es) Boiss.
Triloculares Boiss.
life
form
Annuae (Boiss.)
Fed.
Campanula species
and subspecies
Campanula – references by older authors
2n
Involucratae
(Fomin) Fedorov
Section
(Fedorov 1957)
Lova{en-Eberhardt (2000)
ZA
WU
P
P
P
P
P
P
cult. cult.
P
P
P
P
P
P
P
P
P
P
?
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
?
P
?
P
P
P
P
P
?
P
P
P
P
spicata L., Sp. Pl. 166 (1753)
34
Hb
P
P
P
thyrsoides L., Sp. Pl. 167 (1753),
ssp. thyrsoides
34
Hb
P
P
P
–?ssp. carniolica (Sünd.)
Podlech, Ber. Bayer. Bot. Ges.
37: 111 (1964)
34
Hb
P
P
P
?
THE GENUS CAMPANULA IN CROATIA
M e d i u m D C.
Section (Fedorov and
Kovanda 1976)
Subgeneric delimitations
Campanula
ACTA BOT. CROAT. 63 (2), 2004
Tab. 2. – continued
P
P
P
P
P
P
?foliosa Ten., Fl. Nap. 1, Prodr.
16 (1811)
34
Hp
glomerata L., Sp. Pl. 166 (1753),
ssp. glomerata
30
Hp
– ssp. cervicarioides (Schult.)
Arcang., Comp. Fl. Ital. 456 (1882)
30
Hp
– ssp. elliptica (Schult.) Kirschl.,
Fl. Alsace 1: 375 (1852)
30
Hp
– ssp. farinosa (Andr.) Kirschl.,
Fl. Alsace 1: 378 (1852)
30
Hp
– ssp. hispida (Witasek) Hayek,
Repert. Spec. Nov. Regni Veg.
Bieb. 30(2): 532 (1930)
30
Hp
–?ssp. serotina (Wettst.) O.
Schwarz, Mitt. Thür. Bot. Ges.
1(1): 118 (1949)
30
Hp
member of Croatian
flora (Kova~i})
P
Domac 1950 /1994/
P
Hb
Hb
Degen 1938
Javorka 1925
P
34
34
Hayek 1931
Rossi 1924
P
cervicaria L., Sp. Pl. 167 (1753)
?moesiaca Velen., Sitz.-Ber.
Böhm. Ges. Wiss. (Math.-Nat.
Kl.) 385 (1893)
HERBARIA
Rossi 1930
life
form
Hirc 1908 /1912/
2n
Schloss.-Vuk. 1869
Campanula species
and subspecies
Campanula – references by older authors
Neilreich 1868 /1869/
Lova{en-Eberhardt (2000)
Visiani 1847 /1872/
syn. Subsection, Series,
Group, Aggregate
Cervicaria Gris., Glomeratae Schur
Subsection
(Fedorov 1957)
Involucratae (Fomin) Fedorov
Section
(Fedorov 1957)
M e d i u m D C.
ACTA BOT. CROAT. 63 (2), 2004
Campanula
Section (Fedorov and
Kovanda 1976)
Subgeneric delimitations
ZA
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
WU
P
P
P
P
P
P
P
P
P
P
P
P
KOVA^I] S.
190
Tab. 2. – continued
member of Croatian
flora (Kova~i})
P
Domac 1950 /1994/
P
Degen 1938
P
Hayek 1931
P
Rossi 1930
P
Javorka 1925
Schloss.-Vuk. 1869
Hp
Rossi 1924
Neilreich 1868 /1869/
34
Hirc 1908 /1912/
life
form
Visiani 1847 /1872/
syn. Subsection, Series,
Group, Aggregate
Trachelium Jav.,
Rapunculoides Schur
trachelium L., Sp. Pl. 166
(1753), ssp. trachelium
2n
HERBARIA
ZA
P
P
P
P
P
P
WU
P
191
latifolia L., Sp. Pl. 165 (1753)
34
Hp
P
P
P
P
bononiensis L., Sp. Pl. 165 (1753)
34
Hp
P
P
P
P
P
P
P
P
P
P
P
P
P
rapunculoides L., Sp. Pl. 165
(1753), ssp. rapunculoides
68,
102
Hp
P
P
P
P
P
P
P
P
P
P
P
32
Hp
Pyramidalis (agg.
Geslot 1984)
Subsection
(Fedorov 1957)
Campanula species
and subspecies
Campanula – references by older authors
pyramidalis L., Sp. Pl. 164
(1753)
34
Hp(b)
P
P
P
P
P
P
P
P
P
P
P
P
P
Waldsteiniana
(agg. Geslot 1984)
D C.
Medium
Lova{en-Eberhardt (2000)
tommasiniana C. Koch in F.W.
Schultz, Arch. Fl. Fr. Allem. 229
(1852)
34
Hp
P
P
P
P
P
P
waldsteiniana Schult. in Roem.
et Schult., Syst. Veg. ed. nov.
15(5): 99 (1819)
34
Hp
P
P
P
P
P
P
(Asyne- trichocalycina Ten., Cat. Pl.
uma)
Hort. Neapol. App. 1: 35 (1815)
P
P
P
P
P
P
P
P
P
P
P
THE GENUS CAMPANULA IN CROATIA
Campanula
Eucodon Fedorov
Section
(Fedorov 1957)
Subgeneric delimitations
Section (Fedorov and
Kovanda 1976)
ACTA BOT. CROAT. 63 (2), 2004
Tab. 2. – continued
HERBARIA
poscharskyana Deg., Magyar
Bot. Lapok 7: 103 (1908)
34
Hp
fenestrellata Feer, J. Bot. 28: 272
(1890)
34
Hp
P
P
P
P
P
P
P
P
P
– (ssp.) istriaca Feer, Jour. Bot.
28: 271 (1890)
34
Hp
P
P
P
P
P
P
P
P
cespitosa Scop., Fl. Carniol. ed.
2(1): 143 (1771)
34
Hp
P
P
P
P
P
P
cochleariifolia Lam., Encycl.
Meth. Bot. 1: 578 (1785)
34
Hp
P
P
?
P
P
P
serrata (Kit.) Hendrych Taxon
11: 123 (1962)
34
Hp
witasekiana Vierh., Mitt. Naturwiss.
Vereins Univ. Wien, 2(4): 72 (1906)
34
Hp
scheuchzeri Vill., Prosp. Hist. Pl. 68
Dauph. 22 (1779)
(102)
Hp
WU
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
Degen 1938
P
ZA
Hayek 1931
P
Domac 1950 /1994/
P
Rossi 1930
Hp
Javorka 1925
34
Rossi 1924
portenschlagiana Schult. in Roem.
et Schult., Syst. Veg. 5: 93 (1819)
Hirc 1908 /1912/
life
form
Schloss.-Vuk. 1869
2n
Campanula species
and subspecies
P
P
P
?
P
P
P
P
P
P
P
P
P
P
P
member of Croatian
flora (Kova~i})
Campanula – references by older authors
Neilreich 1868 /1869/
Lova{en-Eberhardt (2000)
Visiani 1847 /1872/
syn. Subsection, Series,
Group, Aggregate
Scheuchzer
-ianae (Kovanda 1977)
Lanceolatae
(Kovanda
1970b)
Alpicolae
(Kovanda
1970b)
Subsect. Isophylla Damboldt
(series Garganicae Trinajsti})
Subsection
(Fedorov 1957)
Heterophylla
Fedorov
Section
(Fedorov 1957)
M e d i u m D C.
ACTA BOT. CROAT. 63 (2), 2004
Campanula
Section (Fedorov and
Kovanda 1976)
Subgeneric delimitations
P
P
P
P
P
P
P
P
P
P
P
KOVA^I] S.
192
Tab. 2. – continued
193
?hercegovina Degen et Fiala, Österr.
Bot. Zeitsch. 44: 303 (1894)
34
Hp
marchesettii Witasek Abh. Zool.- (34)
Bot. Ges. Wien 1(3): 32 (1902)
68
Hp
velebitica Borbas Math. Termesz. (34)
Ertesitö 1: 81 (1883)
68
Hp
moravica (Spitzn.) Kovanda,
Folia Geobot. Phytotax. (Praha)
3: 409 (1968), ssp. moravica
68
Hp
– ssp. xylorrhiza (O. Schwartz)
Kovanda, Folia Geobot.
Phytotax. (Praha) 3: 409 (1968)
102
Hp
34,
68
(102)
Hp
rotundifolia L., Sp. Pl. 163
(1753)
P
P
P
P
P
P
member of Croatian
flora (Kova~i})
Hp
P
Domac 1950 /1994/
34
P
HERBARIA
Degen 1938
justiniana Witasek Magyar Bot.
Lapok 5: 245 (1906)
Hayek 1931
P
Rossi 1930
Hp
Javorka 1925
34
Rossi 1924
carnica Schiede ex Merth. et
Koch in Röhling, Deutschl. Fl.
ed. 3(2): 158 (1826), ssp. carnica
Hirc 1908 /1912/
life
form
Schloss.-Vuk. 1869
2n
Campanula species
and subspecies
Neilreich 1868 /1869/
Campanula – references by older authors
Visiani 1847 /1872/
syn. Subsection, Series,
Group, Aggregate
Saxicolae (Kovanda 1970b)
Vulgares
(Kovanda
1977)
Subsection
(Fedorov 1957)
Fedorov
Heterophylla
Section
(Fedorov 1957)
Lova{en-Eberhardt (2000)
ZA
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
P
WU
THE GENUS CAMPANULA IN CROATIA
M e d i u m D C.
Section (Fedorov and
Kovanda 1976)
Subgeneric delimitations
Campanula
ACTA BOT. CROAT. 63 (2), 2004
Tab. 2. – continued
KOVA^I] S.
Considering the data in table 2, it is obvious that several Campanula species are indeed
highly dubious for Croatian flora. Campanula ramosissima Sibth. et Sm. is an annual species distributed in the southern Balkans and northern Italy (where it might be introduced,
acc. to PIGNATTI 1982). This species was originally referred to by all authors for the Kotor
region, which today is part of Montenegro. Several ambiguous herbaria samples localized
to »Dalmatien« (ZA) or »the border of Dalmatia and Montenegro« (WU) are not enough to
keep this taxon as a member of the Croatian flora, while no data on recent Croatian localities exist. Campanula medium (native to central Italy and southern France) is regarded as
cultivated in Croatia, like C. barbata in the past. As HIRC (1908) removed C. barbata from
the Croatian Flora, I recommend that C. medium be removed too. This biennial is not often
cultivated in Croatia and hardly ever escapes from gardens. The tall C. thyrsoides L. subsp.
carniolica (Sünd.) Podlech is, at best, very rare in Croatia (only one dubious sample in
WU), while the typical subspecies seems completely to prevail in Croatia. Sessile-flowered
C. moesiaca Velen. and C. foliosa Ten. are taxa of the central Balkans and the central and
southern Apennines, but are highly dubious for Croatia. At least two subspecies of the
poorly investigated C. glomerata L. complex (subsp. hispida (Witasek) Hayek and subsp.
serotina (Wettst.) O. Schwarz) could be excluded from the Croatian checklist until further
notice, as well as C. trichocalycina Ten. This species of uncertain origin and taxonomic position, between Campanula, Asyneuma and Phyteuma (LAKU[I] and CONTI 2004) requires
further investigation for a possible Croatian occurrence. Campanula serrata, centred in the
Carpathians, is unknown in recent Croatian phytocenological relevés, although often referred to by older botanists for the whole region. Actually, the entire subsection Heterophylla needs fundamental research in the west Balkans. Campanula rotundifolia L. is, according to KOVANDA (1970a), in its typical form absent from Southeast Europe. However,
large amounts of Croatian material in all the investigated herbaria are (mis)labelled as C.
rotundifolia. There is a strong possibility that some recently evolved heterophyllous incipient taxa are the local equivalents of C. rotundifolia in the Croatian flora, i.e.? C. moravica
(Spitzner) Kovanda with its subspecies. Heterophyllous campanulas of all groups are
poorly investigated in Croatia, so it might be better for the C. rotundifolia aggregate to be
retained in the Croatian flora pending further research.
3% 3%
6%
11%
2n=16
2n=28
2n=20
2n=30
63%
14%
2n=68
2n=34
Fig. 3. Chromosome numbers (2n) of 35 Croatian Campanula species and subspecies.
194
ACTA BOT. CROAT. 63 (2), 2004
THE GENUS CAMPANULA IN CROATIA
After the expulsion of the aforementioned taxa from the recent checklist (LOVA[ENEBERHARDT 2000), the Croatian flora comprises 35 Campanula species and subspecies
(42% of the regional campanulas). The prevailing diploid number is 2n = 34 (63%, in Fig.
3), while some uninvestigated taxa in Croatia could be represented by polyploid populations. Hemicryptophytes are dominant at 97% (Fig. 4).
T (2,9%)
Hb (22,8%)
Hp (74,3%)
Fig. 4. Spectrum of the life forms among 35 Croatian Campanula species and subspecies: Hp =
Hemichryptophyta-perennial, Hb = Hemichryptophyta-biennial, T = Therophyta (annual).
More than 30% of Croatian native campanulas are endemic. Only 3 species (Montenegrin C. ramosissima excluded) belong to the Section Rapunculus (sensu Dumort.): the
polymorphous and broadly distributed C. rapunculus L., C. patula L. and C. persicifolia
L.. The Croatian checklist indicates that these taxa are present only in their typical forms,
which is highly unlikely: they are actually insufficiently investigated complexes of large
distribution, with many included incipient taxa.. About 90% of Croatian campanulas are
members of the Campanula Section (=Medium DC.), assembled in several more or less
natural groups. There are only 3 (with the now excluded C. trichocalycina and C. medium)
rather isolated taxa without closer relatives in the Croatian flora: C. sibirica L., C. lingulata
Waldst. et Kit. (probably not closely related, though placed together in Triloculares by
Boissier 1875) and C. erinus L. (part of the circum-Mediterranean subgenus Roucela
(Dumort.) Damboldt). The subsection Involucratae (Fomin) Fedorov include sessile-flowered campanulas with thyrsiform (C. thyrsoides), spicate (C. spicata) and capitate (C.
cervicaria L., C. glomerata, excluded C. moesiaca and C. foliosa) inflorescences. Also
rather sessile-flowered are members of the subsection Eucodon Fedorov, as well as of the
aggregate Pyramidalis (sensu GESLOT 1984). »Eucodons« are broadly distributed in European forests (C. trachelium L. and its relative C. latifolia L. of the higher altitudes) and
open habitats (C. rapunculoides L. and its xerotherm relative C. bononiensis L.). Pyramidalis-taxa are of much narrower, south-European distribution, in Croatia represented only
by the Illyrian-Adriatic/Balkan endemic C. pyramidalis L.. Together with its closest relatives, the southern Balkan/southern Italian C. versicolor Andrews and the Serbian subendemic C. secundiflora Visiani et Pan~i}, C. pyramidalis forms a unique group of
»isophylloid« campanulas of Balkan origin. The Pyramidalis aggregate demonstrates the
phytogeographic relations of the eastern Adriatic and west Balkan campanulas to southeast
and central European floras. DAMBOLDT (1965a) excluded Pyramidalis-relatives from his
isophyllous group of taxa (henceforth referred to as »subsection Isophylla« sensu DamACTA BOT. CROAT. 63 (2), 2004
195
KOVA^I] S.
boldt), based on morphological differences. Then again, these two lineages must be related,
taking into consideration crossing-experiments by MUSCH and GADELLA (1972), and some
of the recent isoenzyme and molecular results (KOVA^I] et al. 2003, 2004). Close to this
group stands the small, relict and subendemic aggregate Waldsteiniana (GESLOT 1984),
whose relationships to all other campanulas remains rather controversial (FIORI 1927,
HAYEK 1931, GADELLA 1964, DAMBOLDT 1965b). Waldsteiniana consists of two »isophylloid« Dinaric (Adriatic) Alps subendemic diploids: the Mt Velebit C. waldsteiniana
Schult., and the Mt U~ka C. tommasiniana C. Koch. These two species share certain morphological characteristics with both isophyllous and heterophyllous campanulas, but are
well distinguished (DAMBOLDT 1965b). Waldstein’s and Tommasini’s campanulas do not
hybridise with any other isophyllous or heterophyllous taxa, although the data on such hybrids are extensive in horticultural literature (cp. CROOK 1951, LEWIS and LYNCH 1998).
There is a possibility, which requires a further research, that some isolated northeast Italian
subendemics could be the closest relatives to this group (DAMBOLDT 1965b), i.e. C. morettiana and C. raineri Perpenti.
»Starbells« of the endemic subsection Isophylla are the most recognizable and most
frequently cultivated campanulas of the western Balkan and circum-Adriatic region
(CROOK 1951, LEWIS and LYNCH 1998, BERNINI et al. 2002). After the classical research of
DAMBOLDT (1965a), the Isophylla are considered to be a natural group (this, however, was
never proved via modern methods, nor critically tested after 1968), and consists of about 12
mutually isolated taxa, mainly distributed in the sub-Mediterranean area of the Adriatic,
Ionian and Tyrrhenian coastal mountains. Cytological investigations (MERXMÜLLER and
DAMBOLDT 1962, PODLECH and DAMBOLDT 1964) clearly distinguished two groups according to their diploid chromosome numbers: 32 or 34. According to TRINAJSTI] (in LOVA[ENEBERHARDT and TRINAJSTI] 1978), there are three groups (»series«) of isophyllous taxa,
consistent with their distribution and morphology. The group Fragiles (2n = 32) includes
the Tyrrhenian taxa C. fragilis and C. isophylla. The north Italian C. elatines and C.
elatinoides are placed in the group Elatines (2n = 34). The Adriatic group Garganicae
(2n = 34) includes Italian C. garganica Tenore (with two Greek subspecies and possibly
the recently-discovered C. reatina Lucchese 1993), and the east Adriatic subendemics C.
portenschlagiana Schultes, C. poscharskyana Degen and C. fenestrellata (with its subspecies’). These relicts and Tertiary schizoendemics are, according to DAMBOLDT (1965a),
placed as a parallel lineage to the large and far more widely spread Subsection Heterophylla (Witasek) Fedorov and its group Rotundifolia.
KOVANDA (1970a, b, 1977) described five heterophyllous groups (»series«) gathered
around the extremely polymorphous C. rotundifolia (group Rotundifolia), described as
»too intricate to be solved« by BÖCHER (1963) and a »huge polyploid structure« by
KOVANDA and AN~EV (1989). Such a collective taxon is, by Kovanda’s opinion, in its typical
form of »true« C. rotundifolia of the group Vulgares with its several closest relatives, of
northern distribution. Kovanda assumed that in the mountains of the central Europe, western Balkan and circum-Adriatic region a number of neoendemic taxa of the groups
Saxicolae, Lanceolatae, Alpicolae and Scheuchzerianae (Tab. 2) are developing, slowly
replacing the typical C. rotundifolia of the North. Heterophyllous taxa (»harebells«) have
so far received little attention in the region, although some must have been isolated for a
long time and are undoubtedly determinable (e.g. the relict C. hercegovina or C. cespitosa
Scop.). Yet, there are many members of this group that are so difficult to precisely recog196
ACTA BOT. CROAT. 63 (2), 2004
THE GENUS CAMPANULA IN CROATIA
nize (BUZAS 1998) and are today disregarded and included in the range of the well-differentiated collective species (although in typical form quite divergent, e.g. the incipient taxa of
the polymorphous C. velebitica Borbas complex: C. balcanica Hruby, C. farinulenta A.
Kern. et Wettst., C. bulgarica Witasek etc.). Unlike the well isolated »garganicas«, different »rotundifolias« in Croatia often share wild habitats (e.g. C. scheuchzeri Vill., C.
marchesettii Witasek, C. velebitica s.l. and C. witasekiana Vierh.), and probably hybridise
(KOVANDA 1999). Also, unlike Garganicae, western Balkan Rotundifolia have no known
localities in the eastern Adriatic islands: they are exclusive members of the mainland
mountainous communities of karstic meadows, pastures, rock fissures and crevices.
Though several Campanula taxa should be excluded from the recent Croatian Flora until further research is done, it is also highly possible that some taxa are yet unrecognized,
have been disregarded, or included in other genera. Despite the isolated relicts, it seems
that the large part of this perplexing collective genus is still evolving in this part of Europe,
producing a number of local incipient species that are practically impossible to trace.
Maybe the best way to deal with taxonomy of the young, still developing taxa is to assemble collective species of the closest relatives, while taxonomic expediency is more important than the recognition of dubious taxa. On the other hand, even populations that are recognisable only at the molecular level may be of value when it comes to conservation
(EDDIE and INGROUILLE 1999). Many more biogeographical, morphological and molecular
investigations – in both Croatia / Southeast Europe and globally – are needed, to increase
our knowledge on the Campanula genus and its relatives’ taxonomy and evolution.
Acknowledgements
I am indebted to Prof. M. An~ev (Sofia, Bulgaria), Dr. A. Bernini (Stradella, Italy), Dr.
W. Gutermann (Vienna, Austria), Prof. T. Lammers (Oshkosh, WI, USA), Prof. F. Lucchese (Rome, Italy), Mr. H. Mylemans (Bouwel, Belgium), Mr. Z. Nikolov (Skopje, FYR
Macedonia), Dr. A. Oçak (Ess kisehir, Turkey), Dr. V. Ran|elovi} (Ni{, Serbia and Montenegro), Dr. G. Schneeweiss (Vienna, Austria), Prof. D. [oljan (Sarajevo, Bosnia and
Herzegovina), Prof. J. Topi} (Zagreb, Croatia) and Prof. I. Trinajsti} (Zagreb, Croatia), for
kindly sharing experience, literature and various unpublished data on the southeast European Campanula taxa. The assistance of Herbaria staff (CNHM, W, WU, ZA/ZAHO) is
greatly appreciated, as well as the technical support of Dr. T. Nikoli}. My special thanks to
Prof. W. M. M. Eddie (Edinburgh, Scotland, UK) for valuable improvements and most
helpful comments on an earlier version of the manuscript.
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