Carpathian bellflower Campanula carpatica, copyright Ghislain Chenais.

Belongs within: Campanuloideae.
Contains: Jasione, Edraianthus, Githopsis, Asyneuma, Phyteuma, Adenophora.

Campanula, harebells, is a diverse genus of herbs found in the Northern Hemisphere with tubular to bell-shaped flowers (Hickman 1993). Phylogenetic analysis indicates the genus to be extensively non-monophyletic with many taxa treated as distinct genera arising within it.

Characters (from Hickman 1993): Annual or perennial, from taproot, fibrous roots, or rhizome, glabrous to densely hairy. Stem reclining or erect, branched, 5–150 cm, four-angled. Leaves cauline, sometimes also basal, generally lanceolate to ovate, thin, fleshy, or leathery, entire to toothed, sessile or petioled. Inflorescence a raceme, panicle, or flowers solitary, terminal or axillary. Flower with corolla cylindric to funnel- or bell-shaped, white to deep blue, lobes linear to triangular; ovary inferior, hemispheric to obconic. Fruit dehiscing by two or three lateral pores. Seeds 2 mm, oblong.

Campanula [incl. C. sect. Medium subsect. Involucratae (Fomin) Fedorov 1957] H98
    |--+--+--‘Wahlenbergia’ hederacea CM16
    |  |  `--Jasione CM16
    |  `--+--+--C. lactiflora CM16
    |     |  `--Gadellia lactiflora CM16
    |     `--+--C. peregrina CM16
    |        `--+--C. primulifolia CM16
    |           `--Musschia CM16
    |                |--M. aurea CM16
    |                `--M. wollastonii CM16
    `--+--+--+--Trachelium caeruleum CM16
       |  |  `--+--C. fruticulosa CM16
       |  |     `--+--C. asperuloides CM16
       |  |        `--C. pubicalyx CM16
       |  `--+--+--+--C. pinatzii CM16
       |     |  |  `--+--C. erinus CM16
       |     |  |     `--+--C. creutzburgii CM16
       |     |  |        `--C. drabifolia CM16
       |     |  `--+--+--C. dimorphantha CM16
       |     |     |  `--+--C. balfourii CM16
       |     |     |     `--C. filicaulis [=C. spatulata ssp. filicaulis] CM16
       |     |     `--+--+--C. vidalii CM16
       |     |        |  `--Azorina vidalii CM16
       |     |        `--+--Feeria angustifolia CM16
       |     |           `--+--C. dichotoma CM16
       |     |              `--+--C. edulis CM16
       |     |                 `--C. mollis CM16
       |     `--+--+--+--C. jacquinii CM16
       |        |  |  `--+--C. rumeliana CM16
       |        |  |     `--Diosphaera rumeliana CM16
       |        |  `--+--C. semisecta CM16
       |        |     `--+--C. macrostyla CM16
       |        |        `--C. propinqua CM16
       |        `--+--+--C. barbata CM16
       |           |  `--C. hofmannii CM16
       |           `--+--+--+--C. alliariifolia CM16
       |              |  |  `--+--+--C. sibirica CM16
       |              |  |     |  `--+--C. grossheimii CM16
       |              |  |     |     `--+--C. bononiensis CM16
       |              |  |     |        `--C. rapunculoides CM16
       |              |  |     `--+--C. pendula CM16
       |              |  |        `--+--C. saxifraga [=C. bellidifolia ssp. saxifraga] CM16
       |              |  |           `--+--C. aucheri [=C. bellidifolia ssp. aucheri] CM16
       |              |  |              `--+--C. bellidifolia CM16
       |              |  |                 `--C. tridentata CM16
       |              |  `--+--C. mirabilis CM16
       |              |     `--+--+--C. chamissonis [=C. dasyantha ssp. chamissonis] CM16
       |              |        |  |--C. collina CM16
       |              |        |  |--C. dasyantha CM16
       |              |        |  `--C. sarmatica CM16
       |              |        `--+--+--C. punctata CM16
       |              |           |  `--+--C. ‘glomerata’ var. dahurica CM16
       |              |           |     `--C. takesimana CM16
       |              |           `--+--C. latifolia Linnaeus 1753 CM16, CD07
       |              |              `--+--C. sect. Campanula [incl. C. sect. Eucodon] H98
       |              |                 |    `--C. trachelium CM16
       |              |                 |         |--C. t. ssp. trachelium H98
       |              |                 |         `--C. t. ssp. athoa H98
       |              |                 `--C. sect. Elatae [incl. C. sect. Cordifoliae] H98
       |              |                      `--C. sclerotricha CM16
       |              `--+--+--C. (ser. Glomeratae) glomerata H98, CM16
       |                 |  `--C. subcapitata CM16
       |                 `--+--+--C. ptarmicifolia CM16
       |                    |  `--+--‘Edraianthus’ dalmaticus CM16
       |                    |     `--‘Edraianthus’ graminifolius CM16
       |                    `--+--+--+--Petkovia orphanidea CM16
       |                       |  |  `--C. petraea CM16
       |                       |  `--+--C. scoparia CM16
       |                       |     |--+--C. spicata CM16
       |                       |     |  `--C. thyrsoides CM16
       |                       |     `--+--C. stricta CM16 (see below for synonymy)
       |                       |        |    |--C. s. ssp. stricta P97
       |                       |        |    `--‘Wahlenbergia’ s. ssp. alternata P97
       |                       |        `--C. zangezura CM16
       |                       `--+--+--C. karakuschensis CM16
       |                          |  |--C. saxatilis CM16
       |                          |  |--+--C. conferta CM16
       |                          |  |  `--C. involucrata CM16
       |                          |  `--+--+--Michauxia CM16
       |                          |     |  |    |--M. campanuloides CM16
       |                          |     |  |    |--M. laevigata CM16
       |                          |     |  |    |--M. nuda CM16
       |                          |     |  |    `--M. tchihatcheffii CM16
       |                          |     |  `--+--C. hierapetrae CM16
       |                          |     |     `--+--C. foliosa CM16
       |                          |     |        `--C. incurva CM16
       |                          |     `--+--C. lanata CM16
       |                          |        `--+--C. carpatha CM16
       |                          |           `--+--+--C. laciniata CM16
       |                          |              |  `--C. tubulosa CM16
       |                          |              `--+--C. pelviformis CM16
       |                          |                 `--C. quercetorum CM16
       |                          `--+--+--‘Edraianthus’ hercegovinus CM16
       |                             |  `--+--‘Edraianthus’ dinaricus CM16
       |                             |     `--C. ser. Cervicariae H98
       |                             |          |--C. cervicaria CM16
       |                             |          `--C. macrostachya H98
       |                             `--+--+--+--C. betulifolia CM16
       |                                |  |  `--C. radula CM16
       |                                |  `--+--C. seraglio CM16
       |                                |     `--+--C. affinis CM16
       |                                |        `--C. speciosa CM16
       |                                `--+--Edraianthus CM16
       |                                   `--+--C. armena CM16
       |                                      `--+--C. radicosa CM16
       |                                         `--C. tymphaea CM16
       `--+--+--Peracarpa carnosa CM16
          |  |    |--P. c. var. carnosa CM16
          |  |    `--P. c. var. circaeoides CM16
          |  `--+--+--C. aparinoides CM16
          |     |  `--+--+--C. prenanthoides CM16
          |     |     |  `--C. wilkinsiana CM16
          |     |     `--+--C. exigua CM16
          |     |        `--+--Heterocodon rariflorum CM16
          |     |           `--Githopsis CM16
          |     `--+--+--C. pterocaula CM16
          |        |  `--+--C. persicifolia CM16
          |        |     `--C. stevenii CM16
          |        `--+--+--Legousia CM16
          |           |  |    |--L. pentagonia CM16
          |           |  |    `--+--L. hybrida CM16
          |           |  |       `--L. speculum-veneris CM16
          |           |  `--+--‘Legousia’ falcata CM16
          |           |     `--+--C. americana CM16
          |           |        `--+--C. reverchonii CM16
          |           |           `--Triodanis CM16
          |           |                |--T. biflora H93
          |           |                |--T. coloradoensis CM16
          |           |                |--T. leptocarpa CM16
          |           |                `--T. perfoliata H93
          |           `--+--+--C. cretica CM16
          |              |  `--Asyneuma CM16
          |              `--+--+--‘Asyneuma’ japonicum CM16
          |                 |  `--C. uniflora CM16
          |                 `--+--Petromarula pinnata CM16
          |                    `--+--Phyteuma CM16
          |                       `--Physoplexis comosa CM16
          `--+--Homocodon CM16
             |    |--H. brevipes CM16
             |    `--H. pedicellatus CM16
             `--+--+--C. fastigiata CM16
                |  `--Adenophora CM16
                `--+--C. aizoides CM16
                   `--+--+--C. elatines Linnaeus 1759 CM16, CD07
                      |  `--+--C. garganica CM16
                      |     `--C. poscharskyana CM16
                      `--+--+--C. lusitanica CM16
                         |  `--+--C. spatulata CM16
                         |     `--+--C. rapunculus CM16
                         |        `--+--C. olympica CM16
                         |           `--+--C. patula CM16
                         |              `--C. sparsa CM16
                         |                   |--C. s. ssp. sparsa CM16
                         |                   `--C. s. ssp. sphaerothrix CM16
                         `--+--+--C. hawkinsiana CM16
                            |  `--C. waldsteiniana CM16
                            `--+--C. decumbens CM16
                               `--+--+--C. carpatica CM16
                                  |  `--+--C. pyramidalis CM16
                                  |     `--C. versicolor CM16
                                  `--+--C. scouleri CM16
                                     `--+--+--C. piperi CM16
                                        |  `--+--C. lasiocarpa CM16
                                        |     `--+--C. divaricata CM16
                                        |        |--C. parryi CM16
                                        |        |    |--C. p. var. parryi CM16
                                        |        |    `--C. p. var. idahoensis CM16
                                        |        `--+--C. aurita CM16
                                        |           `--C. scabrella CM16
                                        `--+--+--C. cenisia CM16
                                           |  `--+--C. elatinoides CM16
                                           |     `--+--C. arvatica CM16
                                           |        `--C. herminii CM16
                                           `--+--+--C. fragilis CM16
                                              |  |    |--C. f. ssp. fragilis CM16
                                              |  |    `--C. f. ssp. cavolinii CM16
                                              |  `--+--C. caespitosa CM16
                                              |     `--C. cochleariifolia CM16
                                              `--+--C. excisa CM16
                                                 `--+--+--+--C. appenina CM16
                                                    |  |  `--C. rotundifolia CM16
                                                    |  `--+--+--C. moravica CM16
                                                    |     |  `--C. witasekiana CM16
                                                    |     `--+--C. rhomboidalis CM16
                                                    |        |--C. scheuchzeri CM16
                                                    |        `--C. stenocodon CM16
                                                    `--+--+--C. marchesetti CM16
                                                       |  `--+--C. forsythii CM16
                                                       |     `--C. macrorrhiza CM16
                                                       `--+--C. carnica CM16
                                                          |--C. bertolae CM16
                                                          |--C. brixiensis CM16
                                                          `--C. sabatia CM16

Campanula incertae sedis:
  C. abietina CM16 [=C. patula ssp. abietina H09]
  C. acarnanica CM16
  C. acutiloba CM16
  C. afra CM16
  C. aizoon CM16
  C. akgulii CM16
  C. alaskana CM16
  C. alata CM16
  C. algida C55b
  C. alpina CM16
  C. alsinoides CM16
  C. anchusiflora CM16
  C. andina CM16
  C. andrewsii CM16
  C. angustiflora H93
  C. aprica CM16
  C. ardonensis CM16
  C. argaea CM16
  C. argentea CM16
  C. argyrotricha O88
  C. ariena CM16
  C. aristata O88
  C. armazica CM16
  C. autraniana CM16
  C. axillaris CM16
  C. balansae CM16
  C. baumgartenii CM16
  C. bayerniana CM16
  C. beckiana CM16
  C. benthamii CM16
  C. betonicifolia CM16
  C. bluemelii CM16
  C. bohemica CM16
  C. bravensis CM16
  C. broussonetiana CM16
  C. buseri CM16
  C. cabezudoi CM16
  C. calamenthifolia CM16
  C. calcarata CM16
  C. calcicola CM16
  C. californica H93
  C. camptoclada CM16
  C. cana CM16
  C. candida CM16
  C. cashmeriana CM16
  C. caucasica CM16
  C. cecilii CM16
  C. celsii CM16
  C. cephallenica CM16
  C. choruhensis CM16
  C. ciliata CM16
  C. circassica CM16
  C. colorata CM16
  C. columnaris CM16
  C. constantini CM16
  C. coriacea CM16
  C. crassipes CM16
  C. crenulata CM16
  C. crispa CM16
  C. cymaea CM16
  C. cymbalaria CM16
  C. daghestanica CM16
  C. damascena CM16
  C. damboldtiana CM16
  C. darialica CM16
  C. davisii CM16
  C. debarensis CM16
  C. delavayi CM16
  C. delicatula PT98
  C. demirsoyii CM16
  C. dieckii CM16
  C. dolomitica CM16
  C. dzaaku CM16
  C. dzyschrica CM16
  C. ekimiana CM16
  C. euboica CM16
  C. evolvulacea CM16
  C. fedorovii CM16
  C. fenestrellata CM16
  C. flaccidula CM16
  C. floridana CM16
  C. formanekiana CM16
  C. gieseckiana CM16
  C. goulimyi CM16
  C. grandis CM16
  C. griffinii [incl. C. angustiflora var. exilis] H93
  C. hagielia CM16
  C. haradjanii CM16
  C. hedgei CM16
  C. hercegovina CM16
  C. heterophylla CM16
  C. hierosolyminata CM16
  C. hispanica CM16
  C. hohenackeri CM16
  C. humillima CM16
  C. hypopolia CM16
  C. iconia CM16
  C. incanescens CM16
  C. isaurica CM16
  C. isophylla [incl. C. floribunda] C55a
  C. jacobaea CM16
  C. justiniana CM16
  C. kadargavica CM16
  C. kenya CM16
  C. kermanica CM16
  C. khorasanica CM16
  C. kirikkaleensis CM16
  C. kolenatiana CM16
  C. komarovi CM16
  C. koyunceri CM16
  C. kremeri CM16
  C. kryophila CM16
  C. lambertiana CM16
  C. lamondiae CM16
  C. langsdorffiana CM16
  C. lavrensis CM16
  C. ledebouriana CM16
  C. legionensis CM16
  C. lehmaniana CM16
  C. leucantha CM16
  C. leucoclada CM16
  C. leucosiphon CM16
  C. lingulata CM16
  C. longistyla CM16
  C. lourica CM16
  C. luristanica CM16
  C. lyrata PT98
  C. macrochlamys CM16
  C. mardinensis CM16
  C. massakskyi CM16
  C. mekongensis CM16
  C. micrantha CM16
  C. modesta O88
  C. moesiaca CM16
  C. morettiana CM16
  C. myrtifolia CM16
  C. nakaoi O88
  C. nuristanica CM16
  C. odontosepala CM16
  C. oligosperma CM16
  C. oreadum CM16
  C. orphanidea CM16
  C. ossetica CM16
  C. pallida O88
  C. pamphylica CM16
  C. pangea Hartvig 1998 H98
  C. papillosa CM16
  C. paradoxa CM16
  C. perpusilla CM16
  C. peshmenii CM16
  C. petrophila CM16
  C. phryga CM16
  C. phyctidocalyx CM16
  C. pinnata CM16
  C. pinnatifida CM16
  C. podocarpa CM16
  C. polyclada CM16
  C. pontica CM16
  C. portenschlagiana CM16
  C. pseudostenocodon CM16
  C. psilostachya CM16
  C. pulla CM16
  C. pulvinaris CM16
  C. radchensis CM16
  C. raddeana CM16
  C. raineri CM16
  C. ramosissima CM16
  C. reatina CM16
  C. recta CM16
  C. reiseri CM16
  C. reuteriana CM16
  C. rhodensis CM16
  C. robinsiae CM16
  C. romanica CM16
  C. rupestris Sibthorp & Smith 1806 PL04
  C. rupicola CM16
  C. ruprechti CM16
  C. samothracica CM16
  C. sartorii CM16
  C. savalanica CM16
  C. saxanorum CM16
  C. saxifragoides CM16
  C. schimaniana CM16
  C. schitica CM16
  C. scopelia CM16
  C. scutellata CM16
  C. secundiflora CM16
  C. serrata CM16
  C. sharsmithiae H93
  C. shettleri CM16
  C. siegizmundii CM16
  C. similans CM16
  C. sosnowskyi CM16
  C. specularioides CM16
  C. stellaris CM16
  C. stenosiphon CM16
  C. strigillosa CM16
  C. strigosa CM16
  C. svanetica CM16
  C. takhtadzhianii CM16
  C. tanfanii CM16
  C. telephioides CM16
  C. telmesii CM16
  C. teucrioides CM16
  C. thessala CM16
  C. tokurii CM16
  C. tomentosa CM16
  C. tommasiniana CM16
  C. topaliana CM16
  C. trachyphylla CM16
  C. transtagana CM16
  C. trichocalycina CM16
  C. tristis CM16
  C. troegerae CM16
  C. turczaninovii CM16
  C. uyemurae CM16
  C. velebitica CM16
  C. veneris CM16
  C. wilkommii CM16
  C. xylocarpa CM16
  C. xylopoda CM16
  C. yaltirikii CM16
  C. yildirimlii CM16
  C. zoysii CM16

Campanula stricta CM16 [=C. gracilis var. stricta Brown 1810 P97, Wahlenbergia gracilis var. stricta P97; incl. C. erecta Sweet 1830 (n. n.) P97, C. vincaeflora Ventenat 1803 P97, C. gracilis var. vincaeflora P97, Wahlenbergia vincaeflora (Ventenat) Decaisne 1849 P97]

*Type species of generic name indicated


[C55a] Candolle, A. de. 1855a. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 1. Librairie de Victor Masson: Paris.

[C55b] Candolle, A. de. 1855b. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 2. Librairie de Victor Masson: Paris.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1–E44.

[CM16] Crowl, A. A., N. W. Miles, C. J. Visger, K. Hansen, T. Ayers, R. Haberle & N. Cellinese. 2016. A global perspective on Campanulaceae: biogeographic, genomic, and floral evolution. American Journal of Botany 103 (2): 233–245.

[H98] Hartvig, P. 1998. Campanula pangea, a new species of C. sect. Involucratae from Mt. Pangeon, NE Greece. Willdenowia 28: 65–68.

[H09] Heltmann, H. 2009. Der Königstein (Piatra Craiului), die Perle der Burzenländer Gebirge. Mauritiana 20 (3): 515–527.

[H93] Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[O88] Ohba, H. 1988. The alpine flora of the Nepal Himalayas: an introductory note. In: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 19–46.

[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi. Willdenowia 28: 95–116.

[P97] Petterson, J. A. 1997. Identity of the original Wahlenbergia gracilis (Campanulaceae) and allied species. An historical review of early collections. New Zealand Journal of Botany 35: 55–78.

[PL04] Pohl, G., & I. Lenski. 2004. Zur Verbreitung und Vergesellschaftung von Pennisetum orientale Rich. in Nordeuböa (Griechenland) (Poaceae, Paniceae). Senckenbergiana Biologica 83 (2): 209–223.

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