Neotropical Campanulaceae

Silvana Aparecida Pires de Godoy

Universidade de São Paulo (USP), São Paulo, SP., Brazil. 

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.

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.

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.

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.

Other important characters

  • Leaves simple, frequently entire or toothed ended by hydathodes, alternate.
  • Flowers bisexual, isolate or in terminal inflorescence, polysymmetric (Campanuloideae) or monosymmetric and resupinate by twisting of the pedicel (Lobelioideae).
  • The 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 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.

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.

Number of genera

Ten genera:

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

Useful tips for generic identification

See above.

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.

Status

Native:

  • Burmeistera
  • Centropogon
  • Cyphocarpus (endemic to northern Chile)
  • Hippobroma longiflora (Jamaican origin currently widespread by cultivation)
  • Siphocampylus 
  • Lysipomia (endemic to high Andes) 

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.

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.

How to cite

Pires de Godoy, S.A. (2009). Neotropical Campanulaceae. In: Milliken, W., Klitgård, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Campanulaceae.htm.

Click images to enlarge


Inflorescence, Centropogon cornutus © Herbier de Alan Jouy B1014Bis, Brazil.



Corolla with reproductive parts positioned above the opening, Centropogon cornutus © Herbier de Alan Jouy B1014Bis, Brazil.



Habit vine, Siphocampylus convolvulaceus © W. Saburi Jr., São Paulo, Brazil.



Lateral view of the flower: immediately above the obconical hipanthium is the isthmus formed by filaments adnate to corolla-tube, Siphocampylus convolvulaceus © W. Saburi Jr., São Paulo, Brazil.



Corolla resupinate: upper lip 3-lobed and lower lip 2-lobed inverted by twisting of the pedicel, Siphocampylus concolvulaceus © W. Saburi Jr., São Paulo, Brazil.



Habit shrubby, Siphocampylus corymbiferus © A.C. Bonifácio da Silva, São Paulo, Brazil.



Floral branch: terminal corymb and alternate leaves, Siphocampylus corymbiferus © L.R. Reato, São Paulo, Brazil.



Corolla ventricose and linear and narrow leaves, Siphocampylus lauroanus © W. Saburi Jr., São Paulo, Brazil.



Habit shrubby, Siphocampylus longipedunculatus © W. Saburi Jr., São Paulo, Brazil.



Habit shrubby, red corolla-tube with yellow lobes, Siphocampylus macropodus © W. Saburi Jr., São Paulo, Brazil.



Habit, shrubby with bright foliage, Siphocampylus nitidus © W. Saburi Jr., São Paulo, Brazil.



Corolla sulphur coloured with one of the lobes separate from the base, Siphocampylus sulfureus © Ivan Sazima, UNICAMP, Brazil.



Discolourous leaves, Siphocampylus umbellatus © Ivan Sazima, UNICAMP, Brazil.



Verticillate leaves, red corolla-tube, and yellow basal lobes, which become green toward their apices, Siphocampylus westinianus © Ivan Sazima, UNICAMP, Brazil.