Abstract
A diagnostic description of the genus is given with special emphasis on the occurrence of succulence amongst its species. The geographical distribution is outlined, together with a selection of important literature, and an explanation of the etymology of the name. This is followed by a short summary of its position in the phylogeny of the family and of the past and present classification in a phylogenetic context. The succulent features present amongst the species of the genus are shortly explained as to morphology and anatomy.
This is followed by a synoptical treatment of the succulent species of the genus, complete with typification details, full synonymy, geographical and ecological data, a diagnostic description, and, where applicable, notes on phylogenetic placement and relationships, as well as economic and/or horticultural importance.
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Vasconcellea A. St. Hilaire (Deux. Mém. Réséd., 12, 1837). Type: Vasconcellea quercifolia A. St. Hilaire [lectotype according to A. F. Carvalho, Molec. Phylog. Biogeogr. e-monograph Papaya family, 92, 2013]. – Lit: Badillo (1971: 60–159, monograph, as Carica p.p.); Badillo (1983: Flora Ecuador, as Carica p.p.); Badillo (2000: systematics); Kyndt & al. (2005: molecular & morphological phylogeny); Scheldeman & al. (2011: use, synopsis, distribution maps). Distr: Mexico, C & S America (to Uruguay, Ecuador, Peru, and C Chile). Etym: For Simão de Vasconcelos [also as Vasconcellos] (1597–1671), Portuguese jesuit and historian in Bahia and Rio de Janeiro, Brazil.
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Incl. Hemipapaya Missouri BG (Tropicos) (nom. inval., ICN Art. 29.1). Type: not typified.
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Incl. Vasconcella A. St.-Hilaire (1837) (nom. inval., ICN Art. 61.1). Type: Vasconcellea quercifolia A. St. Hilaire.
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Incl. Vasconella Walpers (1843) (nom. inval., ICN Art. 61.1). Type: Vasconcellea quercifolia A. St. Hilaire.
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Incl. Vasconcelia Bentham & Hooker (1867) (nom. inval., ICN Art. 61.1). Type: Vasconcellea quercifolia A. St. Hilaire.
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Incl. Vasconcellosia Caruel (1876). Type: Carica hastata Brignoli.
Usually dioecious (rarely monoecious or polygamous) mostly evergreen trees or shrubs, or rarely evergreen lianas (V. horovitziana (V. M. Badillo) V. M. Badillo), stem simple or generally few-branched, sometimes basally thickened and ± pachycaulous, bark smooth, interior tissue soft and pith-like; Br sometimes armed with spiny stipules; L large to very large, simple and entire or palmatilobed, pinnatilobed or palmatifid with up to 7 lobes, lobes simple or shallowly to deeply lobed; male Inf many-flowered and congested, or few-flowered; male Fl 5- or sometimes 4-merous; Sep alternating with the petals; Pet white, yellow or red, or greenish, forming a tube, glabrous or pubescent in the throat; St 5 + 5, shorter (sub-) sessile, longer with free filaments; Fil glabrous or hairy; female Inf few- to 1-flowered; Ov 5-locular in the lower part; Sty present or absent; Fr berries, variously shaped.
This is the largest genus of the family, comprising 20 species plus the naturally occurring hybrid V. ×pentagona (Heilborn) Mabberley (= V. ×heilbornii (V. M. Badillo) V. M. Badillo). It is most diverse in NW South America (esp. Ecuador, Colombia and Chile), with a centre of diversity in the Andean valleys of Ecuador, where 16 species occur and locally form fertile hybrids (Kyndt & al. 2005: 1034). The ecology of the species spans the continuum from wet to seasonally dry habitats. According to the molecular phylogeny of Carvalho & Renner (2012), Vasconcellea comprises 2 clades, a small clade of 4 species (incl. C. chilensis, C. quercifolia) and a larger clade with the remaining species (incl. V. parviflora).
For a long time, Vasconcellea was treated as synonym of Carica (as Sect. Vasconcellea (A. St. Hilaire) Hooker 1867), but Badillo (2000) accepted the taxon at generic rank, and subsequently also resolved the question of the correct spelling (Badillo 2001). Kubitzki (2002) supported this reclassification on morphological grounds, and the molecular studies of Kyndt & al. (2005), Carvalho & Renner (2012) and Sun & al. (2016: 380) confirmed a position completely separate from Carica, as sister to Jacaratia.
The majority of the species of Vasconcellea have comparatively massive trunks in relation to the crown, and at least minimal water storage capacity is probably universally present, esp. since flowering in the leafless state is reported for several species. A stem dry mass of 0.22 g/cm3 is reported for V. quercifolia by Zanne & al. (2009) (cited from Kempe & al. 2013). In the treatment below, only the most clearly water-storing species are covered by way of example.
The flowers of most species of Vasconcellea correspond to the sphingophilous syndrome. Cerino & al. (2015) found species from the hawkmoth families Arctiidae, Noctuidae and Pyralidae as flower visitors of the self-incompatible V. quercifolia. Only the male flowers offer nectar as a reward, while female flowers do not offer any reward and are pollinated by deceit. According to the results of Cerino & al. (2015), fruit set under natural conditions is low in this species, likely due to pollinator limitation, which is typical for deceit pollination systems.
The fruits of several species are used on a local level, and are eaten fresh, cooked or processsed as juices, jam or preserves. The only species that currently are of commercial importance are V. cundinamarcensis V. M. Badillo, V. pubescens A. De Candolle (“Highland Papaya”, “Mountain Papaya”, “Papayuelo”) and esp. V. ×pentagona (“Babacao”; usually still referred to as V. ×heilbornii, but the former name has nomenclatural priority; = V. pubescens × V. stipulata (V. M. Badillo) V. M. Badillo). The hybrid as well as V. pubescens and V. cundinamarcensis are cultivated throughout N South America. V. pubescens is esp. seen as extensive plantations in Chile, where its fruits are mostly marketed in the form of preserves or candies. Other species might also have potential for novel exotic fruit production (Scheldeman & al. 2011). Since at least V. quercifolia can be crossed with Carica papaya Linné, producing vigorous intergeneric hybrids, Vasconcellea could be used in Carica cultivar breeding (Drew & al. 2006, Scheldeman & al. 2011).
V. chilensis Planchon ex A. De Candolle (in De Candolle, Prodr. Syst. Regni Veg. 15(1): 416, 1864). Type [lecto]: Chile (Gay s.n. [P]). – Lit: Serra & al. (1986: conservation); Carrasco & al. (2014: conservation, genetics). Distr: C Chile (Región de Atacama, Región de Coquimbo, Región de Valparaíso: Huasco to Valparaíso); coastal cordillera, semi-arid stony valleys and hillsides, infrequent, to 700 (–1100) m. I: Muñoz Pizarro (1966: t. XLVI); Badillo (1971: t. 19); both as Carica; Scheldeman & al. (2011: 219).
≡ Carica chilensis (Planchon ex A. De Candolle) Solms (1889) ≡ Papaya chilensis (Planchon ex A. De Candolle) Kuntze (1891) (incorrect name, ICN Art. 11.4); incl. Carica microcarpa Gay (1846) (nom. illeg., ICN Art. 53.1).
Deciduous shrubs to 1.5 (–4) m, stem somewhat pachycaul, to 8 cm ⌀, with thickish branches with greyish to reddish smooth bark with conspicuous slightly raised leaf scars; L glabrous, petiole slender, 3–5 cm, lamina broadly triangular-ovate to suborbicular in outline, 2.5–8 cm ⌀, with 3–5 main lobes which are irregularly lobed giving a total of 10–18 uneven rounded lobes; Inf contracted shortly pedunculate few-flowered racemes (male), or 1-flowered (female); Fl green to reddish; male Fl tubular, 6–11 mm, corolla tube 3.5–6 mm, outside sparsely pubescent to subglabrous, lobes 3–6 mm; female Fl tubular, ± 11 mm, glabrous; Sty indistinct; Sti ± 2 mm; Fr 1.8–2.4 (–4) × 1.2–1.8 cm, shortly pedunculate, obovoid, pendent, dirty green to wine-red to violet; Se ellipsoid-fusiform, 5–6 (–7) × ± 2–3 mm, dirty yellowish with white blotches, smooth.
Individuals of V. chilensis occur highly scattered in coastal scrub, but it is unclear whether the low population densities are natural, or the result of grazing (Serra & al. 1986). According to Carrasco & al. (2014), only 3 populations remain due to relatively recent fragmentation.
V. parviflora A. De Candolle (Prodr. Syst. Regni Veg. 15(1): 417, 1864). Type [lecto]: Peru (Ruiz & Pavón s.n. [G, FI, MA]). – Distr: W & SW Ecuador (Manabí, Los Ríos, Guayas, El Oro, Lojas), NW Peru (Tumbes, Piura, Lambayeque, La Libertad, Cajamarca, Huánuco, San Martín); Pacific Andean slopes, semi-arid and arid places, 100–1500 (–2000) m. I: Badillo (1971: t. 13); Badillo (1983: 35); both as Carica; Scheldeman & al. (2011: 222). – Fig. 1.
≡ Carica parviflora (A. De Candolle) Solms (1889) ≡ Papaya parviflora (A. De Candolle) Kuntze (1891) (incorrect name, ICN Art. 11.4); incl. Carica paniculata Spruce (1869) ≡ Papaya paniculata (Spruce) Kuntze (1891) (incorrect name, ICN Art. 11.4); incl. Carica leptantha Harms (1922).
Deciduous shrubs or small trees to 3 m, frequently flowering when leafless, young plants with globose-elongate swelling at the stem base, bark greyish, smooth; L glabrous, petiole 8–20 cm, lamina 10–30 × 6–25 cm, narrowly ovate to suborbicular in outline, basally cordate, irregularly 3-, 5-, 7- or 9-lobed, the lower lobes sometimes overlapping, all lobes entire or again 3-lobed to pinnatilobed, acute to acuminate; male Inf lax or contracted, 1–12 cm pedunculate, often pinkish; male Fl rose-coloured, sessile to shortly pedicellate; Pet forming a tube 13 × 1 mm, lobes 4 × 1.7 mm; pistillode 3 mm; female Inf 2–4 cm pedunculate; female Fl rose-coloured, 4–7 mm pedicellate; Pet 20–25 mm, ascending-spreading; Ov narrowly ovoid, with 10 low ridges, 12 × 1–3.2 mm; Fr small, porrect, narrowly ovoid, 2–2.5 × 1–1.2 cm ⌀, yellowish to orange or reddish, edible but taste sweet-sour; Se ovoid, 5–6 mm.
V. quercifolia A. St. Hilaire (Deux. Mém. Réséd., 13, 1837). Type [neo]: Argentina, Jujuy (Horovitz 1111 [MY, B, BH, G, MO, SI, VEN, W]). – Lit: Carlquist (1998: wood anatomy, as Carica); Cerino & al. (2015: reproductive biology). Distr: SE Peru (Apurimac, Cusco), C Bolivia (La Paz, Potosí, Chuquisaca, Cochabamba, Santa Cruz, Tarija), Paraguay (Central, Alto Paraguay, Nueva Asunción, Paraguarí, San Pedro), E to S Brazil (Bahia, Minas Gerais, Goiás, Saõ Paulo, Paraná, Santa Catarina, Rio Grande do Sul), Uruguay (Cerro Largo), N Argentina (Misiones, Corrientes, Formosa, Chaco, Santa Fé, Jujuy, Salta, Tucumán, N Catamarca); humid to semi-arid forests and esp. forest margins, lowlands to 3000 (–3600) m. I: Badillo (1971: t. 16); Novara (1992: 8–10); both as Carica; Scheldeman & al. (2011: 223). – Figs. 2
and 3.
≡ Carica quercifolia (A. St. Hilaire) Hieronymus (1882) ≡ Papaya quercifolia (A. St. Hilaire) Kuntze (1891) (incorrect name, ICN Art. 11.4); incl. Carica hastata Brignoli (1862) ≡ Vasconcellosia hastata (Brignoli) Caruel (1876); incl. Vasconcellea lanceolata A. De Candolle (1864) ≡ Carica lanceolata (A. De Candolle) Bentham & Hooker ex Hieronymus (1882) ≡ Papaya lanceolata (A. De Candolle) Kuntze (1891) (incorrect name, ICN Art. 11.4); incl. Carica hastaefolia hort. ex Solms (1889); incl. Carica bonplandii Hort. Paris ex Solms (1889) (nom. inval., ICN Art. 36.1a); incl. Papaya tunariensis Kuntze (1898) (incorrect name, ICN Art. 11.4) ≡ Carica tunariensis (Kuntze) K. Schumann (1900); incl. Carica acuta Heilborn (1936).
Deciduous shrubs or little-branched trees, 3–8 (–15) m, trunk 10–35 (–60) cm ⌀, often conically tapering, rarely remaining short and lump-shaped, sometimes extending into thickened storage roots, bark smooth, peeling in small greenish-yellow, yellowish or ochre-coloured thin pieces; Br with well-visible closely-spaced leaf scars which become slightly raised; L very variable, glabrous, petiole 2–15 cm, lamina simple, to 30 (–40) × 4–15 (–22) cm, elongate-hastate, narrowly elliptic to variously rhombic, margin entire or lobed or serrate to dentate or variously incised, main vein well-developed, lateral veins inconspicuous; male Inf pendent, contracted-corymbiform, many-flowered, peduncle 5–15 cm; male Fl yellowish-green; Pet forming a tube ±12 mm long, lobes ±10 mm, recurved; pistillode subulate, 3–5 mm; female Inf 1- to 3- (to 4-) flowered, 3–6 cm; female Fl with rapidly caducous sepals; Pet ascending-spreading, to 15 × 3 mm, slightly thickish, greenish-yellow; Ov oblong, 5-costate, 7–8 × 3–4 mm; Sti lobes 2–5 mm; Fr obovoid, slightly apiculate, 2–5 (–8) × 2–3 cm, obtusely pentagonal, yellow to orange; Se ovoid, 4–5 × 2–3.5 mm.
This is an extremely variable taxon, esp. as to leaf form. Plants from lowland habitats have broad and lobed leaves reminiscent of oak leaves, hence the epithet. Plants from dry upland habitats, however, have leaves with much reduced lamina. The flowers appear towards the end of the dry season or early in the rainy season. They correspond to the sphingophilous syndrome, and hawkmoths from the families Arctiidae, Noctuidae and Pyralidae were found as flower visitors by Cerino & al. (2015). As for other taxa, only male flowers offer nectar as reward, and the female flowers are visited by deceit and do not offer any reward. The species can be successfully crossed with Carica papaya and the hybrids are vigorous and resistant against papaya ringspot virus (Drew & al. 2006).
The fruits are edible and are consumed fresh or made into preserves, or are used to produce a fermented beverage, at least in rural N Argentina (Novara 1992: 9).
References
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Eggli, U. (2023). Vasconcellea CARICACEAE. In: Eggli, U., Nyffeler, R. (eds) Dicotyledons: Rosids. Illustrated Handbook of Succulent Plants. Springer, Cham. https://doi.org/10.1007/978-3-030-93492-7_23
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