1. Campanulaceae Juss.

    1. This family is accepted.

[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.
Useful tips for generic identification

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.
Other important characters
  • 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.
Distinguishing characters (always present)
  • 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.
Status

Native:

  • Burmeistera
  • Centropogon
  • Cyphocarpus (endemic to northern Chile)
  • Hippobromalongiflora (Jamaican origin currently widespread by cultivation)
  • Siphocampylus 
  • Lysipomia (endemic to high Andes) 
Number of genera

Ten genera:

  • Burmeistera.
  • Centropogon.
  • Cyphocarpus.
  • Diastatea.
  • Downingia pusilla.
  • Hippobroma longiflora.
  • Lobelia. 
  • Lysipomia
  • Siphocampylus
  • Wahlenbergia.
Notes on delimitation
  • 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

Habit
Annual or perennial herbs, subshrubs, or rarely small shrubs, laticiferous
Leaves
Leaves alternate, rarely opposite, simple, entire, dentate to incised or rarely variously lobed, exstipulate
Inflorescences
Inflorescences generally cymose, panicle-, raceme-, spike- or head-like, or flowers solitary
Flowers
Flowers bisexual, usually protandrous, regular, (3–)5(–10)-merous, mostly with a bract and 2 bracteoles
Calyx
Calyx ± adnate to the ovary; lobes usually free, persistent, valvate
Corolla
Petals connate to various degrees, sometimes almost free, valvate in bud
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
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
Fruits
Fruit capsular, variously dehiscing by apical or lateral valves or pores, or ± baccate
Seeds
Seeds 1-many, albuminous; embryo straight, terete
[FWTA]

Campanulaceae, F.N. Hepper. Flora of West Tropical Africa 2. 1963

Habit
Herbs to small trees, nearly always with milky juice
Leaves
Leaves alternate, rarely opposite, simple; stipules absent
Flowers
Flowers hermaphrodite, actinomorphic
Calyx
Calyx-tube adnate to the ovary, 3-10-lobed
Corolla
Corolla gamopetalous, tubular or campanulate, lobes valvate
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
Gynoecium
Ovules mostly numerous Ovary inferior or rarely superior, 2-10-celled, with axile placentas
Fruits
Fruit capsular or baccate
[FZ]

Campanulaceae, M. Thulin. Flora Zambesiaca 7:1. 1983

Habit
Annual or perennial herbs, subshrubs, or rarely small shrubs, laticiferous
Leaves
Leaves alternate, rarely opposite, simple, entire, dentate to incised or rarely variously lobed, exstipulate
Inflorescences
Inflorescences generally cymose, panicle–, raceme–, spike– or head–like, or flowers solitary
Flowers
Flowers hermaphrodite, usually protandrous, regular, (3–) 5(–10)–merous, mostly with a bract and 2 bracteoles
Calyx
Calyx –{_ adnate to the ovary; lobes usually free, persistent, valvate in bud
Corolla
Petals connate to various degrees, sometimes almost free, valvate in bud
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
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
Fruits
Fruit capsular, variously dehiscing by apical or lateral valves or pores, or 4; baccate
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

Habit
Annual or perennial herbs, subshrubs, shrubs or trees, laticiferous
Leaves
Leaves alternate or more rarely opposite or verticillate, simple, entire to dentate or incised, more rarely lobed or dissected, exstipulate
Inflorescences
Inflorescences generally racemose, or flowers solitary in leaf–axils; pedicels mostly with 2 bracteoles
Flowers
Flowers bisexual or rarely unisexual, usually protandrous, zygomorphic, 5–merous, often resupinate
Calyx
Calyx regular or somewhat 2–lipped; lobes valvate in bud
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
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
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
Fruits
Fruit capsular, usually dehiscing by 2(4) apical valves, rarely by a lid, or ± baccate
Seeds
Seeds 1–many, albuminous, embryo straight
[FTEA]

Lobeliaceae, Mats Thulin (University of Uppsala). Flora of Tropical East Africa. 1984

Habit
Annual or perennialherbs, subshrubs, shrubs or trees, laticiferous
Leaves
Leaves alternate or more rarely opposite or verticillate, simple, entire to dentate or incised, more rarely lobed or dissected, exstipulate
Inflorescences
Inflorescences generally racemose, or flowers solitary in leaf-axils; pedicels mostly with 2 bracteoles
Flowers
Flowers bisexual or rarely unisexual, usually protandrous, zygomorphic, 5-merous, often resupinate
Calyx
Calyx regular or somewhat 2-lipped; lobes valvate in bud
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
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
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
Fruits
Fruit capsular, usually dehiscing by 2(–4) apical valves, rarely by a lid, or ± baccate
Seeds
Seeds 1–many, albuminous; embryo straight
[FWTA]

Lobeliaceae, E. Wimmer. Flora of West Tropical Africa 2. 1963

Habit
Herbs, shrubs or small trees, always with milky juice; leaves alternate, rarely opposite or verticillate, simple; stipules absent
Flowers
Flowers hermaphrodite, zygomorphic
Calyx
Calyx-tube adnate to the ovary, 5-lobed
Corolla
Corolla gamopetalous, tubular, usually bilabiate, lobes 5, valvate
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
Gynoecium
Ovary inferior, 2-celled with axile placentation, rarely 1-celled with parietal placentation; ovules numerous, very rarely few
Fruits
Fruit capsular or baccate; seeds minute, embryo straight in fleshy endosperm

Images

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

Sources

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 2018. 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