Francoaceae A.Juss.

Melianthaceae, B. Verdcourt (East African Herbarium). Flora of Tropical East Africa. 1958

Morphology General Habit

Shrubs or trees

Morphology Leaves

Leaves alternate, pinnate; stipules present and often large

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or often functionally unisexual, more or less irregular, in conspicuous racemes

Morphology Reproductive morphology Flowers Calyx

Calyx-segments 4–5, somewhat unequal, imbricate

Morphology Reproductive morphology Flowers Corolla

Petals 5, free, unequal, clawed

Morphology Reproductive morphology Flowers Nectaries

Disc pentagonal-annular or variously unilateral

Morphology Reproductive morphology Flowers Androecium

Stamens 4–6, free or somewhat connate

Morphology Reproductive morphology Flowers Gynoecium

Style simple; stigma small or capitate Ovary superior, 4–5–locular Ovules axile, 1–4 per loculus

Morphology Reproductive morphology Fruits

Fruit a papery or woody capsule

Morphology Reproductive morphology Seeds

Seeds large with copious endosperm

Melianthaceae, F. White. Flora Zambesiaca 2:2. 1966

Morphology General Habit

Trees or shrubs

Morphology General Indumentum

Indumentum of simple hairs

Morphology Leaves

Leaves alternate, imparipinnate or 3-foliolate

Morphology Leaves Stipules

Stipules present, usually large, either in pairs at the base of the petiole or fused and intrapetiolar (Bersama)

Morphology Reproductive morphology Inflorescences

Inflorescence of conspicuous terminal or axillary racemes

Morphology Reproductive morphology Flowers

Flowers bisexual in appearance but often unisexual and then apparently dioecious or polygamous, slightly to markedly zygomorphic, usually 4–5-merous; sepals and petals dissimilar

Morphology Reproductive morphology Flowers Calyx

Sepals 4–5, shorter or longer than the petals, united at the base, with lobes imbricate

Morphology Reproductive morphology Flowers Corolla

Petals 4–5, free, imbricate, unequal, unguiculate

Morphology Reproductive morphology Flowers Nectaries

Disk extrastaminal, annular-pentagonal or variously unilateral

Morphology Reproductive morphology Flowers Androecium

Stamens 4–5(8), free or connate at the base; anthers 2-thecous, dehiscing longitudinally

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 4–5-locular, with basal or axile placentation; ovules 1–4 per loculus; style 1, stigma capitate

Morphology Reproductive morphology Fruits

Fruit a papery or woody loculicidal capsule

Morphology Reproductive morphology Seeds

Seeds large, with copious endosperm; aril present or absent

Weigend, M. (2009). Neotropical Vivianiaceae.

Morphology

Description

Shrubs, very rarely annual herbs, spine -tipped short-shoots (brachyblasts) often present. Leaves opposite throughout, rarely in whorls of three, simple , mostly ovate and coarsely crenate to pinnatisect to pinnatifid , sometimes tiny (ca. 1-2 mm, on conical short shoots forming little green cones), base cuneate to cordate ; estipulate. Inflorescences terminal , thyrsoids or pleiothyrsoids, sometimes apparently single and terminal ; inflorescence bracteose or frondose, bracts in Balbisia Cav. often deeply pinnatisect and situated diretly at base of calyx . Flowers hermaphrodite , (4-)5- merous , actinomorphic ; sepals well developed, sometimes larger than petals, entire with acute , usually aristate apex ; petals 0-4-5, (ob-) ovate to widely circular or obcordate, often apically emarginate , pink, white, or yellow; stamens (4-5)-10, usually obdiplostemonous and heterantherous with typically five long and five short stamens, filaments sometimes with pair of basal appendages; gynoecium of 3-5 carpels, syncarpous with 3-5 locules ; style very short, with 3-5 long, papillose stigmatic branches; ovary 3-5- lobed , with 1-20, pendulous, campylotropous ovules in each locule . Nectary disc absent, nectarines sometimes situated on filament appendages. Fruit septicidal or septifragal capsules with 1-8-seeded locules , rarely 5 schizocarps.

Distribution

Distribution in the Neotropics

Vivianiaceae comprise 3 genera and 17 species, all in Southern South America. Most of them are found in semi-arid areas on the western side of the Andes in South America; only Rhynchotheca is endemic to interandean valleys of Peru and Ecuador, where it is found in cloud-forest remnants (e.g. hedges).

• Rhynchotheca (1 species) -  Peru, Ecuador.
• Balbisia (11 species) - Chile, Peru, Bolivia, Argentina.
• Viviania (5 species) - Chile, 1 in Argentina, Uruguay, S Brazil.

Diagnostic

Key differences from similar families

• Similar to Geraniaceae, but always thin-stemmed and densely branched shrubs (1 herb!).
• Leaves ovate in outline and shortly petiolate to subsessile.
• Leaves never palmate.

Notable genera and distinguishing features

• Balbisia (11 species): flowers bowl-shaped, with yellow, sometimes yellowish-green or reddish-yellow corolla, fruit capsular,  flower subtended by pair of deeply divided bracts.
• Rhynchotheca (1 species): flowers apetalous, anthers pendulous, 5-partedschizocarp.
• Viviania (5 species): flowercampanulate, with white or pink petals, pedicellate without bracts directly at base, fruit capsular.

Distinguishing characters (always present)

• Predominantly shrubs, often with spines.
• Soft pubescence usually present.
• Flowers very similar to Geraniaceae.
• Sepals with aristate tip.
• Ovary superior and mostly 5-lobed (superficially identical to those of Geraniaceae).
• Fruit not falling into mericarpids, apart from Rhynchotheca spinosa Ruiz & Pav.
• R. spinosa with apetalous flowers.

General Description

Number of genera

• 3 genera.

Status

• All species are native, and some species are narrowly endemic.

General notes

• Hunziker & Ariza Espinar (1973) reduced Wendtia Meyen under Balbisia, this judgement is followed here.
• Viviania has been treated either in a wider sense (Knuth 1912) or segregated into a total of 4 genera (3 monotypic genera Cissabryon Meisn., Caesarea Cambess. and Araeoandra Lefor, Lefor 1975).
• The segregate genera of Lefor (1975) may or may not be natural entities, but Viviania s.l. including the segregates appears to be a natural unit and a single genus seems amply sufficient to accommodate its 6 species.
• Rhynchotheca may be the only anemophilous species of Geraniales; its schizocarpic fruits resemble those of Geraniaceae, but are apparently homoplasious.
• Viviania has nectar -flowers, whereas Balbisia is nectarless and has pollen flowers (Weigend 2005).

Notes on delimitation

• Members of the Vivianiaceae have often been placed in Geraniaceae or segregated into two families, Vivianiaceae (Viviania Cav.) and Ledocarpaceae (Balbisia, Rhynchotheca Ruiz & Pav.).
• The three genera are closely allied and are here considered as parts of a single family.
• Vivianiaceae together with Geraniaceae and Melianthaceae s.l. (incl. Francoaceae) make up the core (and possibly only) families of Geraniales.

Literature

Important literature

Ariza Espinar, L. 1995a. Flora Fanerogámica Argentina Fasciculo 8, 129b. Vivianiaceae. - Córdoba (Argentina): CONICET.

Ariza Espinar, L. 1995b. Flora Fanerogámica Argentina Fasciculo 18, 129a. Ledocarpaceae. - Córdoba (Argentina): CONICET.

Boelcke, O. 1989. Plantas Vasculares de la Argentina. Buenos Aires: Editorial Hemisferio Sur S.A. Brako, L., & Zarucchi, J. L.1993. Catalogue of the Flowering Plants and Gymnosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45.

Correa, N.M. 1988. Flora Patagonica V - Dicotyledones dialipétalas. Buenos Aires: INTA.

Hunziker, A.T., & Ariza Espinar, L.1973. Aporte a la rehabilitación de Ledocarpaceae, familia monotipica. Kurtziana 7: 233-240.

Jørgensen P.M., & Yanez León, S. 1999. Checklist of the Vascular Plants of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard. 75.

Lefor, M. W. M. 1975. 2012. A taxonomic revision of the Vivianiaceae. Univ. Conn. Occ. Papers, Biol. Sc. Ser. 2/15: 225-255.

Plazzesi, L., Gottschling, M., Barreda V. & Weigend, M. 2012. First Miocene fossils of Vivianiaceae shed new light on phylogeny, divergence times, and historical biogeography of Geraniales. Biological Journal of the Linnean Society 107: 67-85.

Price, R.A., & Palmer, J. D. 1993. Phylogenetic relationships of the Geraniaceae and Geraniales from rbcL sequence comparisons. Ann. Missouri Bot. Gard. 80: 661-671.

San Martín, J. 1983. Medicinal Plants in Central Chile. Economic Botany 37(2): 216-227.

Soltis, D. E.,  Soltis, P. S.,  Chase, M. W.,  Mort, M. E.,  Albach, D. C.,  Zanis, M.,  Savolainen, V.,  Hahn, W. H.,  Hoot, S. B.,  Fay, M. F.,  Axtell, M.,  Swensen, S. M.,  Prince, L. M.,  Kress, W. J.,  Nixon, K. C., & Farris, J. S. 2000. Angiosperm phylogeny inferred from 18S rDNA, rbcL, and atpB sequences. Bot. J. Linn. Soc. 133: 381-461.

Weigend, M. 2005. Floral morphology and Pollination in Vivianiaceae (Geraniales). Plant Systematics and Evolution 253: 125-131.

Weigend, M. 2006. Vivianiaceae. In: Kubitzki, K.: The Families and Genera of the Plants IX: 213-220. Springer-Verlag, Berlin, Heidelberg, New York.