Campanulaceae Juss.
[NTK]
Pires de Godoy, S.A. (2009). Neotropical Campanulaceae.
- Morphology
-
Description
Herbs, shrubs, twining vines (Cyphioideae, some Centropogon C.Presl and Siphocampylus Pohl) and pachycaul rosettes (some Lobelia L.), annual or perennial plants frequently laticiferous (articulated laticifers); plants storing carbohydrate as inulin, accumulating polyacetylenes and sometimes pyridine alcaloids (some Lobelioideae); iridoids and tannins absent; vessel elements with simple perforation plates ( scalariform in the some Campanuloideae). Stipules absent. Leaves simple , frequently entire or toothed ended by hydathodes, sometimes profoundly lobed (Cyphocarpoideae), alternate , spiral (Campanula L. and Wahlenbergia Schrad. ex Roth.), whorled (about nine Siphocampylus species and Ostrowskia Regel.), rarely opposite (Codonopsis Wall., Cyananthus Wall. ex Benth.). Flowers hermaphrodite , rarely unisexual, isolated or in terminal inflorescence , diverse racemose or mixed or sometimes strictly cymose types, polysymmetric (Campanuloideae) or monosymmetric and resupinate by twisting of pedicel (Lobelioideae), (3-)5(-10)- merous ; sepals and petals connate , median sepal adaxial in Campanuloideae and abaxial in Lobelioideae; corolla -tube in Lobelioideae often fenestrate, upper lip 3- lobed and lower lip 2- lobed or 2- cleft , sometimes deeply cleft and corolla appearing unilabiate; stamens as many as and alternate with corolla lobes, filaments adnate to nectary -disk or corolla -tube, free or distinct at base and connate above (Lobelioideae), anthers free or tubelike, introrse, opening by longitudinal slits and totally or at least base connate ; pollen spheroid to oblate -spheroid, verrucate or with spicules in Nemacladoideae and Campanuloideae or prolate and usually reticulate in Lobelioideae, Cyphorcarpoideae and Cyphioideae; gynoecium syncarpous, (1)2-3(5-10), ovary inferior or seldom only half-inferior, rarely superior (Cyananthus), commonly with as many locules as carpels, exceptionally (some Lobelioideae) unilocular with 2 parietal placentas; stigmas wet or dry; ovules numerous on axile (rarely parietal ) placentas, anatropous. Fruit commonly capsules dehiscing through sides (apically) as in Lobelioideae and Cyphocarpoideae, with pores or slits (Campanula and Wahlenbergia) or circumscissile (some Lobelioideae and Parishella A.Gray); less often fruit is a berry (Centropogon and some Lobelia species). Seeds numerous, small, less often winged ; endosperm -development cellular, often oily with terminal haustoria.
- Distribution
-
Distribution in the Neotropics
- Burmeistera Karst. & Triana - 102 spp. Neotropics from Guatemala to Peru.
- Centropogon C.Presl - 212 spp. Neotropics from southern Mexico to Bolivia and Brazil - 2 spp. in the Lesser Antilles.
- Cyphocarpus Miers - 3 spp. Neotropics N Chile.
- Diastatea Scheidw - 5 spp. Neotropics, from central Mexico to Panama, with one species extending south to Bolivia.
- Downingia pusilla (G. Don ex A. DC.) Torr. is only spp. occurring in Neotropics, Chile and Argentina.
- Hippobroma longiflora (L.) G. Don - Jamaica, but now widely naturalized in tropics.
- Lobelia L. - pantropical, in the Neotropics occurring ca. 65 spp., from Mexico to Argentina, Bolivia and Brazil.
- Lysipomia Kunth - 30 spp. Neotropics, Andes to South America, from Venezuela to Bolivia.
- Siphocampylus Pohl. - 231 spp. Neotropics, from Costa Rica to Argentina and in the Greater Antilles .
- Wahlenbergia Schrad. ex Roth. - 6 spp. W. brasiliensis Cham. is endemic Brazil; W. calycina Schlecht. from Colombia to Northwest Argentina; W. globularis E.Wimm. - Peru; W. linarioides Lam. - from Equador to Southwest Brazil, Argentina and South Chile; W. perrottetii (A.DC.) Thulin - introduced; W. urcosensis E.Wimm. - Peru.
- Diagnostic
-
Notable genera and distinguishing features
- The "CBS clade" includes three shrubby neotropical genera, Centropogon, Burmeistera and Siphocampylus, which collectively comprise almost half of the species of Lobelioideae.
- Burmeistera and Centropogon have fleshy fruits and Siphocampylus is capsular type.
- Burmeistera is a monophyletic group primarily bat-pollinated, herbs or hemi-epiphytic herbs or subshrubs that climb nearby vegetation; tubular corollas with reproductive parts positioned above the opening.
- Siphocampylus is inferred to be a paraphyletic relative to fleshy -fruited Centropogon and Burmeistera, but fleshy fruits have evolved repeatedly, making Centropogon polyphyletic. Both genera are difficult to distinguish; apart from the fruit there are subtle differences regarding the lower anthers.
- The other representative genus in Neotropics is Lysipomia, a monophyletic group of small cushion-forming plants endemic to the high Andes. The genus is sister to the clade comprising the remaining three shrubby genera.
See above.
Key differences from similar families- Campanulaceae is easy to distinguish by combined presence of latex, simple leaves and inferior ovary.
- Material from other families with long and reddish or orange corolla -tube is often placed in Campanulaceae, mainly Acanthaceae, Lamiaceae and Rubiaceae. However, these families differing in leaves commonly opposite and free stamens and anthers.
- Allied families or others traditionally placed close to Campanulaceae such as Pentaphragmataceae, Stylidiaceae (including Donatiaceae), Sphenocleaceae and Goodeniaceae (including Brunoniaceae), can be distinguished from the Campanulaceae by the following features: they lack latex; Goodeniaceae have a style with apical hairy pollen -collecting indusium and stylar cup; Pentaphragmataceae and Stylidiaceae have extrorse anthers.
- Leaves simple, frequently entire or toothed ended by hydathodes, alternate.
- Flowers bisexual, isolate or in terminalinflorescence, polysymmetric (Campanuloideae) or monosymmetric and resupinate by twisting of the pedicel (Lobelioideae).
- The median sepaladaxial in Campanuloideae and abaxial in Lobelioideae.
- Corolla-tube in Lobelioideae often fenestrate, upper lip 3-lobed and lower lip 2-lobed or 2-cleft, sometimes deeply cleft and the corolla appears unilabiate.
- Filaments adnate to nectary-disk or corolla-tube, free or distinct at the base and connate above (Lobelioideae).
- Anthers free or tubelike.
- Ovary inferior or seldom only half-inferior, commonly with as many locules as carpels.
- Ovules numerous on axile placentas.
- Fruit commonly capsules dehiscing through sides (apically) as in Lobelioideae and Cyphocarpoideae, with pores or slits (Campanula and Wahlenbergia) or circumscissile (some Lobelioideae and Parishella).
- Endosperm often oily.
- Plants frequently laticiferous (articulated laticifers), storing carbohydrate as inulin, accumulating polyacetylenes; iridoids and tannins absent.
- Stipules absent.
- Leaves simple.
- Sepals and petals connate.
- Stamens as many as and alternate with corolla lobes.
- Anthers introrse, opening by longitudinal slits.
- Gynoecium syncarpous.
- Ovules numerous, anatropous.
- Seeds numerous, small.
- Endosperm-development cellular with terminal haustoria.
- General Description
-
General notes
- The Campanulaceae consists of about 84 genera, 2,319 species, 391 subspecies and 27 named hybrids, world-wide distribution.
- The family is divided into five subfamilies, two are well marked and widespread: Campanuloideae (N temperate Old World, very few species in the Australia-New Zealand area) and Lobelioideae (largely tropical, especially common in the New World, not Arctic and absent from the Near East and central Asia).
- Three other subfamilies are restricted in their distribution: Nemacladoideae (SW USA. and Mexico), Cyphocarpoideae (Chile) and Cyphioideae (Africa especially to south).
- Noteworthy is the presence of numerous endemic Lobelioideae on the Hawaiian Islands.
- The major center of diversity is in the Andes of South America for Lobelioideae and Eurasia from the Mediterranean to the Caucasus for Campanuloideae.
- Campanulaceae exhibits different mechanisms of secondary pollen presentation. Campanuloideae: stamens often sprawling at bottom of corolla tube after pollen is shed although bases conceal nectar; pollen is held among hairs along the style, and these hairs later retract. Cyphioideae: style hairs have bulbous bases, pollen is deposited in pollen box and may be transferred onto the pollinators in portions by pushing down the pollen box. Lobelioideae: "pump and piston" or as a "noodle squeezer" mechanism, as the style pushes the pollen out of the tip of a tube that is formed by the fused stamens.
Native:
- Burmeistera
- Centropogon
- Cyphocarpus (endemic to northern Chile)
- Hippobromalongiflora (Jamaican origin currently widespread by cultivation)
- Siphocampylus
- Lysipomia (endemic to high Andes)
Ten genera:
- Burmeistera.
- Centropogon.
- Cyphocarpus.
- Diastatea.
- Downingia pusilla.
- Hippobroma longiflora.
- Lobelia.
- Lysipomia
- Siphocampylus
- Wahlenbergia.
- The Campanulaceae family is currently placed within the order Asterales.
- The relationship of Campanulaceae to the rest of the Asterales is uncertain (Lundberg & Bremer 2003).
- The molecular data suggest a basal polychotomy with Rousseaceae, Pentaphragmataceae and Campanulaceae together as sisters to the other Asterales.
- Other possibilities indicate the Campanulaceae family is sister to the rest of Asterales.
- Literature
-
Important literature
Antonelli, A. 2008. Higher level phylogeny and evolutionary trends in Campanulaceae subfam. Lobelioideae: Molecular signal overshadows morphology. Molecular Phylogenetics and Evolution 46: 1-18.
Knox, E. B.; Muasya, A. M. & Muchhala, N. 2008. The predominantly South American clade of Lobeliaceae. Systematic Botany 33: 462-468.
Lammers, T.G. 2007. World checklist and bibliography of Campanulaceae. Royal Botanic Gardens, Kew. 675 p.
Lundberg, J. & Bremer, K. 2003. A phylogenetic study of the order Asterales using a morphological and three molecular data sets. International Journal of Plant Sciences 164: 553-578.
Wimmer, E.F. 1943. Campanulaceae-Lobelioideae, I. In R. Mansfeld (ed.) Das Pflanzenreich. Leipzig, Wilhelm Engelmann, IV-276b. p. i-viii + 1-260.
Wimmer, E.F. 1953. Campanulaceae-Lobelioideae, II. In R. Mansfeld (ed.) Das Pflanzenreich. Berlin, Akademie-Verlag, IV-276b, p. i-viii + 261-814.
Wimmer, E.F. 1968. Campanulaceae-Lobelioideae. Supplementum. In A. Engler (ed.) Das Pflanzenreich. Berlin, Akademie Verlag, IV-276c, p. 815-1024.
[FTEA]
Campanulaceae, Mats Thulin (University of Uppsala). Flora of Tropical East Africa. 1976
- Morphology General Habit
- Annual or perennial herbs, subshrubs, or rarely small shrubs, laticiferous
- Morphology Leaves
- Leaves alternate, rarely opposite, simple, entire, dentate to incised or rarely variously lobed, exstipulate
- Morphology Reproductive morphology Inflorescences
- Inflorescences generally cymose, panicle-, raceme-, spike- or head-like, or flowers solitary
- Morphology Reproductive morphology Flowers
- Flowers bisexual, usually protandrous, regular, (3–)5(–10)-merous, mostly with a bract and 2 bracteoles
- Morphology Reproductive morphology Flowers Calyx
- Calyx ± adnate to the ovary; lobes usually free, persistent, valvate
- Morphology Reproductive morphology Flowers Corolla
- Petals connate to various degrees, sometimes almost free, valvate in bud
- Morphology Reproductive morphology Flowers Androecium
- Stamens alternating with the corolla-lobes, free or rarely adnate to the corolla; anthers very rarely entirely or partly connate, introrse; filaments usually dilated at the base
- Morphology Reproductive morphology Flowers Gynoecium
- Ovary ± inferior, rarely superior, 2–10-locular; ovules few-many, anatropous, on axile placentas; style 1, furnished with pollen-collecting hairs on and usually below the style-lobes
- Morphology Reproductive morphology Fruits
- Fruit capsular, variously dehiscing by apical or lateral valves or pores, or ± baccate
- Morphology Reproductive morphology Seeds
- Seeds 1-many, albuminous; embryo straight, terete
[FWTA]
Campanulaceae, F.N. Hepper. Flora of West Tropical Africa 2. 1963
- Morphology General Habit
- Herbs to small trees, nearly always with milky juice
- Morphology Leaves
- Leaves alternate, rarely opposite, simple; stipules absent
- Morphology Reproductive morphology Flowers
- Flowers hermaphrodite, actinomorphic
- Morphology Reproductive morphology Flowers Calyx
- Calyx-tube adnate to the ovary, 3-10-lobed
- Morphology Reproductive morphology Flowers Corolla
- Corolla gamopetalous, tubular or campanulate, lobes valvate
- Morphology Reproductive morphology Flowers Androecium
- Stamens as many as the corolla-lobes and alternate with them, inserted towards the base of the corolla or the disk; anthers free, 2-celled, opening lengthwise
- Morphology Reproductive morphology Flowers Gynoecium
- Ovules mostly numerous Ovary inferior or rarely superior, 2-10-celled, with axile placentas
- Morphology Reproductive morphology Fruits
- Fruit capsular or baccate
[FZ]
Campanulaceae, M. Thulin. Flora Zambesiaca 7:1. 1983
- Morphology General Habit
- Annual or perennial herbs, subshrubs, or rarely small shrubs, laticiferous
- Morphology Leaves
- Leaves alternate, rarely opposite, simple, entire, dentate to incised or rarely variously lobed, exstipulate
- Morphology Reproductive morphology Inflorescences
- Inflorescences generally cymose, panicle–, raceme–, spike– or head–like, or flowers solitary
- Morphology Reproductive morphology Flowers
- Flowers hermaphrodite, usually protandrous, regular, (3–) 5(–10)–merous, mostly with a bract and 2 bracteoles
- Morphology Reproductive morphology Flowers Calyx
- Calyx –{_ adnate to the ovary; lobes usually free, persistent, valvate in bud
- Morphology Reproductive morphology Flowers Corolla
- Petals connate to various degrees, sometimes almost free, valvate in bud
- Morphology Reproductive morphology Flowers Androecium
- Stamens alternating with the corolla–lobes, free or rarely adnate to the corolla; anthers very rarely entirely or partly connate, introrse; filaments usually dilated at the base
- Morphology Reproductive morphology Flowers Gynoecium
- Ovary ± inferior, rarely superior, 2–10–locular; ovules few–many, anatropous, on axile placentas; style 1, furnished with pollen–collecting hairs on and usually below the style–lobes
- Morphology Reproductive morphology Fruits
- Fruit capsular, variously dehiscing by apical or lateral valves or pores, or 4; baccate
- Morphology Reproductive morphology Seeds
- Seeds 1–many, albuminous; embryo straight, terete
- Distribution
- About 35 genera and some 700 species, especially well represented in the Mediterranean region and S. Africa, but relatively sparsely developed in the tropics
[FZ]
Lobeliaceae, M. Thulin. Flora Zambesiaca 7:1. 1983
- Morphology General Habit
- Annual or perennial herbs, subshrubs, shrubs or trees, laticiferous
- Morphology Leaves
- Leaves alternate or more rarely opposite or verticillate, simple, entire to dentate or incised, more rarely lobed or dissected, exstipulate
- Morphology Reproductive morphology Inflorescences
- Inflorescences generally racemose, or flowers solitary in leaf–axils; pedicels mostly with 2 bracteoles
- Morphology Reproductive morphology Flowers
- Flowers bisexual or rarely unisexual, usually protandrous, zygomorphic, 5–merous, often resupinate
- Morphology Reproductive morphology Flowers Calyx
- Calyx regular or somewhat 2–lipped; lobes valvate in bud
- Morphology Reproductive morphology Flowers Corolla
- Petals usually united into a tube, which is often ± split on the back (or on the lower side in non–resupinate flowers), rarely 2 or all petals free or almost so, 2–lipped or rarely 1–lipped or subregular, valvate in bud
- Morphology Reproductive morphology Flowers Androecium
- Stamens alternating with petals, free or ± adnate to the corolla; filaments linear, rarely dilated at the base, ± connate, rarely free; anthers united into a tube or rarely free (Cyphia), introrse
- Morphology Reproductive morphology Flowers Gynoecium
- Ovary ± inferior, rarely ± superior, (1)2–locular; ovules few–many, anatropous, on axile placentas, style 1, with pollen–collecting hairs at or near the apex
- Morphology Reproductive morphology Fruits
- Fruit capsular, usually dehiscing by 2(4) apical valves, rarely by a lid, or ± baccate
- Morphology Reproductive morphology Seeds
- Seeds 1–many, albuminous, embryo straight
[FTEA]
Lobeliaceae, Mats Thulin (University of Uppsala). Flora of Tropical East Africa. 1984
- Morphology General Habit
- Annual or perennialherbs, subshrubs, shrubs or trees, laticiferous
- Morphology Leaves
- Leaves alternate or more rarely opposite or verticillate, simple, entire to dentate or incised, more rarely lobed or dissected, exstipulate
- Morphology Reproductive morphology Inflorescences
- Inflorescences generally racemose, or flowers solitary in leaf-axils; pedicels mostly with 2 bracteoles
- Morphology Reproductive morphology Flowers
- Flowers bisexual or rarely unisexual, usually protandrous, zygomorphic, 5-merous, often resupinate
- Morphology Reproductive morphology Flowers Calyx
- Calyx regular or somewhat 2-lipped; lobes valvate in bud
- Morphology Reproductive morphology Flowers Corolla
- Petals usually united into a tube, which is often ± split on the back (or on the lower side in non-resupinate flowers), rarely 2 or all petals free or almost so, 2-lipped or rarely 1-lipped or subregular, valvate in bud
- Morphology Reproductive morphology Flowers Androecium
- Stamens alternating with petals, free or ± adnate to the corolla; filaments linear, rarely dilated at the base, ± connate, rarely free; anthers united into a tube or rarely free (Cyphia), introrse
- Morphology Reproductive morphology Flowers Gynoecium
- Ovary ± inferior, rarely ± superior, (1–)2-locular; ovules few–many, anatropous, on axile placentas; style 1, with pollen-collecting hairs at or near the apex
- Morphology Reproductive morphology Fruits
- Fruit capsular, usually dehiscing by 2(–4) apical valves, rarely by a lid, or ± baccate
- Morphology Reproductive morphology Seeds
- Seeds 1–many, albuminous; embryo straight
[FWTA]
Lobeliaceae, E. Wimmer. Flora of West Tropical Africa 2. 1963
- Morphology General Habit
- Herbs, shrubs or small trees, always with milky juice; leaves alternate, rarely opposite or verticillate, simple; stipules absent
- Morphology Reproductive morphology Flowers
- Flowers hermaphrodite, zygomorphic
- Morphology Reproductive morphology Flowers Calyx
- Calyx-tube adnate to the ovary, 5-lobed
- Morphology Reproductive morphology Flowers Corolla
- Corolla gamopetalous, tubular, usually bilabiate, lobes 5, valvate
- Morphology Reproductive morphology Flowers Androecium
- Stamens 5, epigynous, alternate with the corolla-lobes, inserted on or free from the corolla; filaments partly free, anthers completely connate in a tube around the filiform style, 2-celled, opening lengthwise
- Morphology Reproductive morphology Flowers Gynoecium
- Ovary inferior, 2-celled with axile placentation, rarely 1-celled with parietal placentation; ovules numerous, very rarely few
- Morphology Reproductive morphology Fruits
- Fruit capsular or baccate; seeds minute, embryo straight in fleshy endosperm
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- Adenophora Fisch.
- Asyneuma Griseb. & Schenk
- Berenice Tul.
- Brighamia A.Gray
- Burmeistera H.Karst. & Triana
- Campanula L.
- Campanulastrum Small
- Canarina L.
- Centropogon C.Presl
- Clermontia Gaudich.
- Codonopsis Wall.
- Craterocapsa Hilliard & B.L.Burtt
- Cryptocodon Fed.
- Cyananthus Wall. ex Benth.
- Cyanea Gaudich.
- Cyclocodon Griff. ex Hook.f. & Thomson
- Cylindrocarpa Regel
- Cyphia P.J.Bergius
- Cyphocarpus Miers
- Delissea Gaudich.
- Dialypetalum Benth.
- Diastatea Scheidw.
- Dielsantha E.Wimm.
- Downingia Torr.
- Eastwoodiella Morin
- Echinocodon D.Y.Hong
- Edraianthus A.DC.
- Favratia Feer
- Feeria Buser
- × Fockeanthus H.R.Wehrh.
- Githopsis Nutt.
- Grammatotheca C.Presl
- Gunillaea Thulin
- Hanabusaya Nakai
- Hesperocodon Eddie & Cupido
- Heterochaenia A.DC.
- Heterocodon Nutt.
- Heterotoma Zucc.
- Himalacodon D.Y.Hong & Qiang Wang
- Hippobroma G.Don
- Homocodon D.Y.Hong
- Howellia A.Gray
- Isotoma (R.Br.) Lindl.
- Jasione L.
- Kericodon Cupido
- Legenere McVaugh
- Legousia Durande
- Lithotoma E.B.Knox
- Lobelia Plum. ex L.
- Lysipomia Kunth
- Melanocalyx (Fed.) Morin
- Merciera A.DC.
- Michauxia L'Hér.
- Microcodon A.DC.
- Monopsis Salisb.
- Muehlbergella Feer
- Musschia Dumort.
- Namacodon Thulin
- Nemacladus Nutt.
- Nesocodon Thulin
- Ostrowskia Regel
- Palmerella A.Gray
- Palustricodon Morin
- Pankycodon D.Y.Hong & X.T.Ma
- Peracarpa Hook.f. & Thomson
- Petromarula Vent. ex R.Hedw.
- Physoplexis Schur
- Phyteuma L.
- Platycodon A.DC.
- Poolea Morin
- Porterella Torr.
- Prismatocarpus L'Hér.
- Protocodon Morin
- Pseudocodon D.Y.Hong & H.Sun
- Pseudonemacladus McVaugh
- Ravenella Morin
- Rhigiophyllum Hochst.
- Roella L.
- Rotanthella Morin
- Ruthiella Steenis
- Sclerotheca A.DC.
- Sergia Fed.
- Siphocampylus Pohl
- Siphocodon Turcz.
- Smithiastrum Morin
- Solenopsis C.Presl
- Theilera E.Phillips
- Trachelium Tourn. ex L.
- Treichelia Vatke
- Trematolobelia Zahlbr. ex Rock
- Triodanis Raf.
- Unigenes E.Wimm.
- Wahlenbergia Schrad. ex Roth
- Wimmeranthus Rzed.
- Wimmerella Serra, M.B.Crespo & Lammers
- Zeugandra P.H.Davis
Campanulaceae Juss. appears in other Kew resources:
First published in Gen. Pl. [Jussieu] 163. 1789 [4 Aug 1789] (1789)
Accepted by
- APG IV (2016) http://dx.doi.org/10.1111/boj.12385
-
Flora Zambesiaca
Flora Zambesiaca
http://creativecommons.org/licenses/by-nc-sa/3.0
-
Flora of Tropical East Africa
Flora of Tropical East Africa
http://creativecommons.org/licenses/by-nc-sa/3.0
-
Flora of West Tropical Africa
Flora of West Tropical Africa
http://creativecommons.org/licenses/by-nc-sa/3.0
-
Kew Names and Taxonomic Backbone
The International Plant Names Index and World Checklist of Selected Plant Families 2022. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0
-
Neotropikey
Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics.
http://creativecommons.org/licenses/by/3.0