Campanula

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. Campanula angustiflora, C. exigua, C. griffinii and C. sharsmithiae are annual species localised to serpentine soils and talus slopes in California (Hickman 1993).

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’ hederaceaCM16
| | `--JasioneCM16
| `--+--+--C. lactifloraCM16
| | `--Gadellia lactifloraCM16
| `--+--C. peregrinaCM16
| `--+--C. primulifoliaCM16
| `--MusschiaCM16
| |--M. aureaCM16
| `--M. wollastoniiCM16
`--+--+--+--Trachelium caeruleumCM16
| | `--+--C. fruticulosaCM16
| | `--+--C. asperuloidesCM16
| | `--C. pubicalyxCM16
| `--+--+--+--C. pinatziiCM16
| | | `--+--C. erinusCM16
| | | `--+--C. creutzburgiiCM16
| | | `--C. drabifoliaCM16
| | `--+--+--C. dimorphanthaCM16
| | | `--+--C. balfouriiCM16
| | | `--C. filicaulis [=C. spatulata ssp. filicaulis]CM16
| | `--+--+--C. vidaliiCM16
| | | `--Azorina vidaliiCM16
| | `--+--Feeria angustifoliaCM16
| | `--+--C. dichotomaCM16
| | `--+--C. edulisCM16
| | `--C. mollisCM16
| `--+--+--+--C. jacquiniiCM16
| | | `--+--C. rumelianaCM16
| | | `--Diosphaera rumelianaCM16
| | `--+--C. semisectaCM16
| | `--+--C. macrostylaCM16
| | `--C. propinquaCM16
| `--+--+--C. barbataCM16
| | `--C. hofmanniiCM16
| `--+--+--+--C. alliariifoliaCM16
| | | `--+--+--C. sibiricaCM16
| | | | `--+--C. grossheimiiCM16
| | | | `--+--C. bononiensisCM16
| | | | `--C. rapunculoidesCM16
| | | `--+--C. pendulaCM16
| | | `--+--C. saxifraga [=C. bellidifolia ssp. saxifraga]CM16
| | | `--+--C. aucheri [=C. bellidifolia ssp. aucheri]CM16
| | | `--+--C. bellidifoliaCM16
| | | `--C. tridentataCM16
| | `--+--C. mirabilisCM16
| | `--+--+--C. chamissonis [=C. dasyantha ssp. chamissonis]CM16
| | | |--C. collinaCM16
| | | |--C. dasyanthaCM16
| | | `--C. sarmaticaCM16
| | `--+--+--C. punctataCM16
| | | `--+--C. ‘glomerata’ var. dahuricaCM16
| | | `--C. takesimanaCM16
| | `--+--C. latifolia Linnaeus 1753CM16, CD07
| | `--+--C. sect. Campanula [incl. C. sect. Eucodon]H98
| | | `--C. tracheliumCM16
| | | |--C. t. ssp. tracheliumH98
| | | `--C. t. ssp. athoaH98
| | `--C. sect. Elatae [incl. C. sect. Cordifoliae]H98
| | `--C. sclerotrichaCM16
| `--+--+--C. (ser. Glomeratae) glomerataH98, CM16
| | `--C. subcapitataCM16
| `--+--+--C. ptarmicifoliaCM16
| | `--+--‘Edraianthus’ dalmaticusCM16
| | `--‘Edraianthus’ graminifoliusCM16
| `--+--+--+--Petkovia orphanideaCM16
| | | `--C. petraeaCM16
| | `--+--C. scopariaCM16
| | |--+--C. spicataCM16
| | | `--C. thyrsoidesCM16
| | `--+--C. strictaCM16 (see below for synonymy)
| | | |--C. s. ssp. strictaP97
| | | `--‘Wahlenbergia’ s. ssp. alternaP97
| | `--C. zangezuraCM16
| `--+--+--C. karakuschensisCM16
| | |--C. saxatilisCM16
| | |--+--C. confertaCM16
| | | `--C. involucrataCM16
| | `--+--+--MichauxiaCM16
| | | | |--M. campanuloidesCM16
| | | | |--M. laevigataCM16
| | | | |--M. nudaCM16
| | | | `--M. tchihatcheffiiCM16
| | | `--+--C. hierapetraeCM16
| | | `--+--C. foliosaCM16
| | | `--C. incurvaCM16
| | `--+--C. lanataCM16
| | `--+--C. carpathaCM16
| | `--+--+--C. laciniataCM16
| | | `--C. tubulosaCM16
| | `--+--C. pelviformisCM16
| | `--C. quercetorumCM16
| `--+--+--‘Edraianthus’ hercegovinusCM16
| | `--+--‘Edraianthus’ dinaricusCM16
| | `--C. ser. CervicariaeH98
| | |--C. cervicariaCM16
| | `--C. macrostachyaH98
| `--+--+--+--C. betulifoliaCM16
| | | `--C. radulaCM16
| | `--+--C. seraglioCM16
| | `--+--C. affinisCM16
| | `--C. speciosaCM16
| `--+--EdraianthusCM16
| `--+--C. armenaCM16
| `--+--C. radicosaCM16
| `--C. tymphaeaCM16
`--+--+--Peracarpa carnosaCM16
| | |--P. c. var. carnosaCM16
| | `--P. c. var. circaeoidesCM16
| `--+--+--C. aparinoidesCM16
| | `--+--+--C. prenanthoidesCM16
| | | `--C. wilkinsianaCM16
| | `--+--C. exiguaCM16
| | `--+--Heterocodon rariflorumCM16
| | `--GithopsisCM16
| `--+--+--C. pterocaulaCM16
| | `--+--C. persicifoliaCM16
| | `--C. steveniiCM16
| `--+--+--LegousiaCM16
| | | |--L. pentagoniaCM16
| | | `--+--L. hybridaCM16 [=Campanula hybridaH64]
| | | `--L. speculum-venerisCM16
| | `--+--‘Legousia’ falcataCM16
| | `--+--C. americanaCM16
| | `--+--C. reverchoniiCM16
| | `--TriodanisCM16
| | |--T. bifloraH93
| | |--T. coloradoensisCM16
| | |--T. leptocarpaCM16
| | `--T. perfoliataH93
| `--+--+--C. creticaCM16
| | `--AsyneumaCM16
| `--+--+--‘Asyneuma’ japonicumCM16
| | `--C. unifloraCM16
| `--+--Petromarula pinnataCM16
| `--+--PhyteumaCM16
| `--Physoplexis comosaCM16
`--+--HomocodonCM16
| |--H. brevipesCM16
| `--H. pedicellatusCM16
`--+--+--C. fastigiataCM16
| `--AdenophoraCM16
`--+--C. aizoidesCM16
`--+--+--C. elatines Linnaeus 1759CM16, CD07
| `--+--C. garganicaCM16
| `--C. poscharskyanaCM16
`--+--+--C. lusitanicaCM16
| `--+--C. spatulataCM16
| `--+--C. rapunculusCM16
| `--+--C. olympicaCM16
| `--+--C. patulaCM16
| `--C. sparsaCM16
| |--C. s. ssp. sparsaCM16
| `--C. s. ssp. sphaerothrixCM16
`--+--+--C. hawkinsianaCM16
| `--C. waldsteinianaCM16
`--+--C. decumbensCM16
`--+--+--C. carpaticaCM16
| `--+--C. pyramidalisCM16
| `--C. versicolorCM16
`--+--C. scouleriCM16
`--+--+--C. piperiCM16
| `--+--C. lasiocarpaCM16
| `--+--C. divaricataCM16
| |--C. parryiCM16
| | |--C. p. var. parryiCM16
| | `--C. p. var. idahoensisCM16
| `--+--C. auritaCM16
| `--C. scabrellaCM16
`--+--+--C. cenisiaCM16
| `--+--C. elatinoidesCM16
| `--+--C. arvaticaCM16
| `--C. herminiiCM16
`--+--+--C. fragilisCM16
| | |--C. f. ssp. fragilisCM16
| | `--C. f. ssp. cavoliniiCM16
| `--+--C. caespitosaCM16
| `--C. cochleariifoliaCM16
`--+--C. excisaCM16
`--+--+--+--C. appeninaCM16
| | `--C. rotundifoliaCM16
| `--+--+--C. moravicaCM16
| | `--C. witasekianaCM16
| `--+--C. rhomboidalisCM16
| |--C. scheuchzeriCM16
| `--C. stenocodonCM16
`--+--+--C. marchesettiCM16
| `--+--C. forsythiiCM16
| `--C. macrorrhizaCM16
`--+--C. carnicaCM16
|--C. bertolaeCM16
|--C. brixiensisCM16
`--C. sabatiaCM16

Campanula incertae sedis:
C. abietinaCM16 [=C. patula ssp. abietinaH09]
C. acarnanicaCM16
C. acutilobaCM16
C. afraCM16
C. aizoonCM16
C. akguliiCM16
C. alaskanaCM16
C. alataCM16
C. algidaC55b
C. alpinaCM16
C. alsinoidesCM16
C. anchusifloraCM16
C. andinaCM16
C. andrewsiiCM16
C. angustifloraH93
C. apricaCM16
C. ardonensisCM16
C. argaeaCM16
C. argenteaCM16
C. argyrotrichaO88
C. arienaCM16
C. aristataO88
C. armazicaCM16
C. autranianaCM16
C. axillarisCM16
C. balansaeCM16
C. baumgarteniiCM16
C. bayernianaCM16
C. beckianaCM16
C. benthamii [incl. C. canescens]TS72
C. betonicifoliaCM16
C. bluemeliiCM16
C. bohemicaCM16
C. bravensisCM16
C. broussonetianaCM16
C. buseriCM16
C. cabezudoiCM16
C. calamenthifoliaCM16
C. calcarataCM16
C. calcicolaCM16
C. californicaH93
C. camptocladaCM16
C. canaCM16
C. candidaCM16
C. cashmerianaCM16
C. caucasicaCM16
C. ceciliiCM16
C. celsiiCM16
C. cephallenicaCM16
C. choruhensisCM16
C. ciliataCM16
C. circassicaCM16
C. colorataCM16
C. columnarisCM16
C. constantiniCM16
C. coriaceaCM16
C. crassipesCM16
C. crenulataCM16
C. crispaCM16
C. cymaeaCM16
C. cymbalariaCM16
C. daghestanicaCM16
C. damascenaCM16
C. damboldtianaCM16
C. darialicaCM16
C. davisiiCM16
C. debarensisCM16
C. delavayiCM16
C. delicatulaPT98
C. demirsoyiiCM16
C. dieckiiCM16
C. dolomiticaCM16
C. dzaakuCM16
C. dzyschricaCM16
C. ekimianaCM16
C. euboicaCM16
C. evolvulaceaCM16
C. fedoroviiCM16
C. fenestrellataCM16
C. flaccidulaCM16
C. floridanaCM16
C. formanekianaCM16
C. gieseckianaCM16
C. goulimyiCM16
C. grandisCM16
C. griffinii [incl. C. angustiflora var. exilis]H93
C. hagieliaCM16
C. haradjaniiCM16
C. hedgeiCM16
C. hercegovinaCM16
C. heterophyllaCM16
C. hierosolyminataCM16
C. hispanicaCM16
C. hohenackeriCM16
C. humillimaCM16
C. hypopoliaCM16
C. iconiaCM16
C. incanescensCM16
C. isauricaCM16
C. isophylla [incl. C. floribunda]C55a
C. jacobaeaCM16
C. justinianaCM16
C. kadargavicaCM16
C. kenyaCM16
C. kermanicaCM16
C. khorasanicaCM16
C. kirikkaleensisCM16
C. kolenatianaCM16
C. komaroviCM16
C. koyunceriCM16
C. kremeriCM16
C. kryophilaCM16
C. lambertianaCM16
C. lamondiaeCM16
C. langsdorffianaCM16
C. lavrensisCM16
C. ledebourianaCM16
C. legionensisCM16
C. lehmanianaCM16
C. leucanthaCM16
C. leucocladaCM16
C. leucosiphonCM16
C. lingulataCM16
C. longistylaCM16
C. louricaCM16
C. luristanicaCM16
C. lyrataPT98
C. macrochlamysCM16
C. mardinensisCM16
C. massakskyiCM16
C. mekongensisCM16
C. micranthaCM16
C. modestaO88
C. moesiacaCM16
C. morettianaCM16
C. myrtifoliaCM16
C. nakaoiO88
C. nuristanicaCM16
C. odontosepalaCM16
C. oligospermaCM16
C. oreadumCM16
C. orphanideaCM16
C. osseticaCM16
C. pallidaO88
C. pamphylicaCM16
C. pangea Hartvig 1998H98
C. papillosaCM16
C. paradoxaCM16
C. perpusillaCM16
C. peshmeniiCM16
C. petrophilaCM16
C. phrygaCM16
C. phyctidocalyxCM16
C. pinnataCM16
C. pinnatifidaCM16
C. podocarpaCM16
C. polycladaCM16
C. ponticaCM16
C. portenschlagianaCM16
C. pseudostenocodonCM16
C. psilostachyaCM16
C. pullaCM16
C. pulvinarisCM16
C. radchensisCM16
C. raddeanaCM16
C. raineriCM16
C. ramosissimaCM16
C. reatinaCM16
C. rectaCM16
C. reiseriCM16
C. reuterianaCM16
C. rhodensisCM16
C. robinsiaeCM16
C. romanicaCM16
C. rupestris Sibthorp & Smith 1806PL04
C. rupicolaCM16
C. ruprechtiCM16
C. samothracicaCM16
C. sartoriiCM16
C. savalanicaCM16
C. saxanorumCM16
C. saxifragoidesCM16
C. schimanianaCM16
C. schiticaCM16
C. scopeliaCM16
C. scutellataCM16
C. secundifloraCM16
C. serrataCM16
C. sharsmithiaeH93
C. shettleriCM16
C. siegizmundiiCM16
C. similansCM16
C. sosnowskyiCM16
C. specularioidesCM16
C. stellarisCM16
C. stenosiphonCM16
C. strigillosaCM16
C. strigosaCM16
C. svaneticaCM16
C. takhtadzhianiiCM16
C. tanfaniiCM16
C. telephioidesCM16
C. telmesiiCM16
C. teucrioidesCM16
C. thessalaCM16
C. tokuriiCM16
C. tomentosaCM16
C. tommasinianaCM16
C. topalianaCM16
C. trachyphyllaCM16
C. transtaganaCM16
C. trichocalycinaCM16
C. tristisCM16
C. troegeraeCM16
C. turczaninoviiCM16
C. uyemuraeCM16
C. velebiticaCM16
C. venerisCM16
C. wilkommiiCM16
C. xylocarpaCM16
C. xylopodaCM16
C. yaltirikiiCM16
C. yildirimliiCM16
C. zoysiiCM16

Campanula strictaCM16 [=C. gracilis var. stricta Brown 1810P97, Wahlenbergia gracilis var. strictaP97; incl. C. erecta Sweet 1830 (n. n.)P97, C. vincaeflora Ventenat 1803P97, C. gracilis var. vincaefloraP97, Wahlenbergia vincaeflora (Ventenat) Decaisne 1849P97]

*Type species of generic name indicated

References

[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).

[H64] Hogg, J. 1864. List of rarer phanerogamous plants discovered in the south-east of Durham since 1829. Report of the British Association for the Advancement of Science 33 (Notices and Abstracts): 96.

[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.

[TS72] Thaker, D. N., & S. D. Sabnis. 1972. Occurrence of Cochlearia cochlearioides (Roth) Sant. & Mah. & Campanula benthamii Wall. ex Kitamura in Gujarat State, India. Journal of the Bombay Natural History Society 69 (1): 227–229.

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