Solanum furcatum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 750. 1814.

Knapp, Sandra, Saerkinen, Tiina & Barboza, Gloria E., 2023, A revision of the South American species of the Morelloid clade (Solanum L., Solanaceae), PhytoKeys 231, pp. 1-342 : 1

19. Solanum furcatum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 750. 1814.

Figs 59 , 60

Solanum deltoideum Colla, Herb. Pedem. 4: 273. 1835. Type. Cultivated in Italy at "h. Ripul:" [ Hortus Ripulensis], the seeds originally sent by C. Bertero from Chile ["Chili Quillota"] (no specimens cited; lectotype, designated by Särkinen et al. 2018, pg. 73: TO [herb. Colla]).

Solanum furcatum Dunal var. glabrum G.Don, Gen. Hist. 4: 412. 1837. Type. Cultivated "Native of Peru" (no specimens cited; no original material located).

Solanum furcatum Dunal var. pilosum G.Don, Gen. Hist. 4: 412. 1837. Type. Cultivated "Native of Peru" (no specimens cited; no original material located).

Solanum furcatum Dunal var. acutidentatum Nees, Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 386. 1843, as " acutedentatum ". Type. "Chile ad Valparaiso, Februario; Peruvia in planitie circa Tacoram [ Volcán Tacora], alt. 14,000-17,000' [feet], Aprili" both syntypes collected by F.J.F. Meyen s.n. (no specimens cited; no original material located). Chile. Région V (Valparaiso): Prov. Valparaiso, Dunas de Concón, 22 Dec 2008, M. Gardner & S Knees 8356 (neotype, designated here: E [E00282600]; isoneotype: BM [BM001120031], CONC [?], SGO [?]).

Solanum furcatum Dunal var. obtusidentatum Nees, Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 386. 1843, as " obtusedentatum ". Type. "Chile. Prov. de San Fernando in Llano del Rio Tinguiririca, 3,000' [feet] alt., Martio"; Peruvia ad Arequipam, Aprili" both syntypes collected by F.J.F. Meyen s.n. (no specimens cited; no original material located). Chile. Région VI ( O’Higgins): Prov. Colchagua, San Fernando, s.d., R.A. Philippi s.n. (neotype, designated here: G [G00443353]).

Witheringia furcata (Dunal) J. Rémy, Fl. Chil. [Gay] 5: 67. 1849. Type. Based on Solanum furcatum Dunal.

Solanum pterocaulum Dunal var. dichotimiflorum Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, as ' opterocaulon '. Type. Cultivated in France at Montpellier " Solanum speciosum hort. botan" (no specimens cited, described from living plants "v.v. hort. Monsp."; neotype, designated by Särkinen et al. 2018, pg. 73: MPU [MPU310703]).

Solanum crenatodentatum Dunal, Prodr. [A. P. de Candolle] 13(1): 54. 1852. Type. Chile. Région VI ( O’Higgins): Colchagua, San Fernando, "in selibus chilensibus San Fernando", Mar 1831, C. Gay 2 (lectotype, designated by D’Arcy 1974a, pg. 738: P [P00337274]).

Solanum rancaguense Dunal, Prodr. [A. P. de Candolle] 13(1): 150. 1852. Type. Chile. Région VI ( O’Higgins): Rancagua, May-Oct 1828, C. Bertero 633 (lectotype, designated by Edmonds 1972, pg. 107 [as holotype], second step designated by Särkinen et al. 2018, pg. 73: P [P00384088]; isolectotypes: BM [BM000617677], G [G00144259], M [M-0171928], MO [MO-503700], NY [NY00743695], P [P00384089], P [P00384090], P [P00384091], P [P00384092], P [P00482266], W [acc. # 1889-0283789]).

Solanum bridgesii Phil., Linnaea 33: 203. 1864. Type. Chile. Región V ( Valparaíso): Panquegue, R.A. Philippi s.n. (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004549]).

Solanum coxii Phil., Linnaea 33: 200. 1864. Type. Chile. Región X (Los Lagos): Todos los Santos, 1862, G. Cox 38 (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004555]; isolectotype: W [acc. # 1903-0010246]).

Solanum rancaguinum Phil., Anales Univ. Chile 43: 523. 1873. Type. Chile. Región VI ( O’Higgins): Rancagua, Mar 1828, C. Bertero s.n. (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004594]).

Solanum caudiculatum Phil., Anales Univ. Chile 91: 12. 1895. Type. Chile. Región VIII ( Bío-Bío): Prov. Ñuble, Coigüeco, F. Puga s.n. (no original material located, not at SGO).

Solanum subandinum Phil., Anales Univ. Chile 91: 13. 1895, nom. illeg., not Solanum subandinum F.Meigen (1893). Type. Chile. Región XIII (Metropolitana): Santiago, Las Condes, R.A. Philippi s.n. (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004600 = F neg. 2745]).

Solanum ocellatum Phil., Anales Univ. Chile 91: 14. 1895. Type. Chile. Región XIII (Metropolitana): Prope Colina, F. Philippi s.n. (lectotype, designated by Särkinen et al. 2018, pg. 74: SGO [SGO000004582]; isotypes: SGO [SGO000004581], W [acc. # 1903-0010230]).

Solanum nigrum L. var. crentatodentatum (Dunal) O.E.Schulz, Symb. Antill. (Urban) 6: 160. 1909. Type. Based on Solanum crenatodentatum Dunal.

Solanum bridgesii Phil. var. ocellatum (Phil.) Witasek ex Reiche, Anales Univ. Chile 124: 460. 1909. Type. Based on Solanum ocellatum Phil.

Solanum andinum Reiche, Fl. Chile 5: 346. 1910. Type. Based on (replacement name for) Solanum subandinum Phil.

Solanum tredecimgranum Bitter, Repert. Spec. Nov. Regni Veg. 11: 6. 1912. Type. Chile. Región V ( Valparaíso): Valparaíso, 17 Aug 1895, O. Buchtien s.n. (lectotype, designated by Barboza et al. 2013, pg. 246: US [US00432692, acc. # 139293]; isolectotypes: HBG [HBG-511497], US [US00681745, acc. # 139294]).

Solanum robinsonianum Bitter, Repert. Spec. Nov. Regni Veg. 11: 7. 1912. Type. Chile. Región V ( Valparaíso): Juan Fernández Island, R.A. Philippi 742 (holotype: B, destroyed [F neg. 2743]; lectotype, designated by Särkinen et al. 2018, pg. 74: W [acc. # 0001347]).

Solanum masafueranum Bitter & Skottsb., Nat. Hist. Juan Fernandez & Easter Island 2: 167, pl. 14. 1922. Type. Chile. Región V ( Valparaíso): Juan Fernández Islands, Masafuera [Isla Alejandro Selkirk], Las Chozas, 715 m, 3 Mar 1917 [20 Feb 1917 on label], C. Skottsberg & I. Skottsberg 363 (lectotype, designated by Särkinen et al. 2018, pg. 74: S [acc. # 04-2947]; isolectotypes: BM [BM000617676], LD [1643307], K [K000585692], NY [00172084], GOET [GOET003548], GB [GB0048742], P [P00337092], UPS [acc. # 104031]).

Solanum spretum C.V.Morton & L.B.Sm., Revis. Argentine Sp. Solanum 132. 1976. Type. Argentina. Río Negro: Bariloche, 19 Mar 1939, A.L. Cabrera 5024 (holotype: GH [00077764]; isotypes F [v0073411F, acc. # 1007493], LP [LP006791]).

Type.

Peru? [more likely Chile]. "Cette plante croît au Perou", J. Dombey [343] (lectotype, first step designated by Edmonds 1972, pg. 107 [as holotype], second step designated by Barboza et al. 2013, pg. 246: P [P00335357]; isolectotypes: B, destroyed [F neg. 2729]; CORD [CORD00006928], F [v0043232F, acc. # 976864], G [G00359946], G-DC [G00144483], P [P00335358]) .

Description.

Annual or subwoody perennial herbs to 1 m high, erect to lax, sprawling to ca. 2 m across. Stems terete or ridged, green to purple tinged, not markedly hollow, sparsely pubescent with simple, uniseriate 1-5-celled eglandular trichomes 0.1-0.5 mm long; new growth sparsely to densely pubescent with similar simple, uniseriate 1-5-celled eglandular trichomes; older stems sparsely pubescent to glabrescent, pale yellowish brown. Sympodial units difoliate, the leaves not geminate. Leaves simple and shallowly sinuate, the blades (1.5-)4-8(-12) cm long, (0.6-)2.2-4.6(-6.5) cm wide, ovate to rhomboidal, widest in the lower half to third, membranous, discolorous; adaxial surface sparsely pubescent with simple, uniseriate trichomes like those on stem, these evenly spread along lamina and veins; abaxial surface more densely pubescent; major veins 4-6 pairs; base cuneate to acute, the two sides slightly unequal, decurrent on the petiole; margins entire or sinuate-dentate, this more pronounced in basal part of the leaf; apex acute; petioles 1-3.5 cm long, sparsely pubescent with simple uniseriate trichomes like those on stem. Inflorescences internodal, forked or more rarely unbranched, (1-)1.5-3(-4) cm long, with 6-14 flowers clustered at the tips (sub-umbelliform) or evenly spaced along the axis, sparsely pubescent with simple uniseriate trichomes like those on stem; peduncle (1-)1.5-2 cm long; pedicels 4-7.5 mm long, 0.2-0.3 mm in diameter at the base and 0.3-0.4 mm in diameter at the apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0.2-2.5 mm apart. Buds subglobose, the corolla exserted 1/3-1/2 from the calyx tube before anthesis. Flowers 5-merous, cosexual (hermaphroditic). Calyx tube 2-3 mm long, conical, the lobes 0.8-1.5 mm long, 0.6-1 mm wide, rectangular to narrowly obovate with obtuse to short-acute apices, pubescent with simple uniseriate trichomes like those on stem but shorter. Corolla 1.2-2 cm in diameter, white to lilac with a green or yellow-green central portion near the base, this sometimes purplish near the lobe midvein, stellate, lobed 1/3-1/2 of the way to the base, the lobes 5.5-7 mm long, 2.8-5.5 mm wide, strongly reflexed at anthesis, later spreading, densely pubescent abaxially with 1-4-celled simple uniseriate trichomes, especially along the margins and apex, these shorter than the trichomes of the stems and leaves. Stamens equal; filament tube minute; free portion of the filaments 0.9-1.6 (2) mm long, adaxially pubescent with tangled uniseriate 4-6-celled simple trichomes; anthers 2.3-3.3(-3.6) mm long, 0.8-1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style 6-6.5 mm long, straight or somewhat curved, long-exserted beyond the anther cone, densely pubescent with 2-3-celled simple uniseriate trichomes in the lower 1/2-2/3; stigma capitate, minutely papillate, yellow or green in live plants. Fruit a globose berry, 0.6-0.9 cm in diameter, dull green to purple at maturity, the pericarp matte, opaque, glabrous; fruiting pedicels 0.7-1.2 cm long, 0.2-0.4 mm in diameter at the base, 0.5-1 mm in diameter at the apex, strongly deflexed, not persistent; fruiting calyx not accrescent, the tube 1-2 mm long, the lobes 1.5-2.5 mm long, appressed against the berry. Seeds 30-40 per berry, 1.8-2 mm long, 1.4-1.5 mm wide, flattened and teardrop shaped with a subapical hilum, yellow-brown, the surface minutely pitted, the testal cells pentagonal in outline. Stone cells 6-14 per berry, 0.8-1 mm in diameter, scattered throughout the berry, cream-coloured. Chromosome number: 2n = 72 ( Stebbins and Paddock 1949, from Californian plants no voucher cited; Edmonds 1977, voucher from Chile Hjerting & Rahn 552, not found or verified).

Distribution

(Fig. 61 ). Solanum furcatum is native to temperate southern Chile (including the Juan Fernández Islands; Regions I [ Tarapacá], II [Antofagasta], IV [Coquimbo], V [Valparaiso], VI [ O’Higgins], VII [Maule], VIII [ Bío-Bío], IX [Araucania], X [Los Lagos], XIII [Metropolitana], XIV [Los Rios]) and adjacent southern Andean Argentina (Provs. Chubut, Neuquén, Río Negro). It is locally introduced and naturalised along the west coast of the United States of America, Australia and New Zealand (see Särkinen et al. 2018; Knapp et al. 2019).

Ecology and habitat.

In its native range S. furcatum is a plant of disturbed areas and forest edges in Nothofagus ( Nothofagaceae ) forests; from near sea level in the more southern part of its range to 2,300 m elevation.

Common names and uses.

Chile. Región V (Valparaiso): yerba mora (Philippi s.n.); Región VI ( O’Higgins): yerba mora (Bertero 633); Región VIII ( Bío-Bío): llaqui (Junge 2611). No uses recorded.

Preliminary conservation status

( IUCN 2022). Least Concern [LC]. EOO = 342,557 km2 [LC]; AOO = 168 km2 [EN]; calculated using South American distribution only and excluding the Juan Fernández Islands. Solanum furcatum has a relatively large range in Chile and adjacent Argentina and is a plant of disturbed areas. It occurs within several Chilean protected areas and in Parque Nacional Bariloche (Argentina). The populations on the Juan Fernández Islands are within the Juan Fernández Archipelago National Park (Chile).

Discussion.

Solanum furcatum is similar to S. arequipense , an endemic species found to the north in western Peru. The two taxa have long been confused (e.g., Nees von Esenbeck 1843) and in our treatment of S. furcatum in its introduced range outside of South America ( Särkinen et al. 2018), we considered them to be conspecific (e.g., Knapp et al. 2019: 67 illustrated S. furcatum with a drawing of S. arequipense ). We recognise these two species here based on the subtle morphological differences that correspond to geographically distinct distributions. They are supported as evolutionarily distinct based on a molecular phylogeny that shows samples of the two species in distinct well supported clades ( Gagnon et al. 2022). Both taxa exhibit the combination of a long-exserted style with the exserted portion equal to or longer than the length of the anthers, shallowly lobed corolla, calyx lobes separated by a paler sinus, forked to unbranched inflorescences (rarely several times branching), and usually large, lush leaves. The inflorescence branches of S. furcatum are less strongly divergent than those of S. arequipense , and the berries of S. furcatum always have more than six stone cells, while those of S. arequipense have only two or stone cells are absent. On an annotation on the now-destroyed duplicate of Dombey’s type gathering of S. furcatum in B (F neg. 2723) Georg Bitter noted the presence of eight stone cells in the berries. Material of S. furcatum from Juan Fernández Islands differs from that collected in mainland Chile in having flowers with styles barely exserted from the anther tube. This could indicate flowers that are autogamous, a reproductive adaptation commonly associated with island life ( Schueller 2004).

Solanum pentlandii also has similar globose buds and exserted styles but has much shorter anthers (less than 2 mm versus 2.5-3 mm in S. furcatum ) and shiny green berries that lack stone cells. Solanum pentlandii occurs at high elevations in disturbed, nitrogen-rich areas in Peru and Bolivia and is not sympatric with S. furcatum .

Details of typification for the synonyms of S. furcatum can be found in Särkinen et al. (2018). In earlier works ( Särkinen et al. 2018; Knapp et al. 2019) we did not lectotypify the four varieties of of S. furcatum described by Nees von Esenbeck (1843) from the collections of Franz Meyen’s trip around the world (1831-32). Franz Meyen’s herbarium from his South American travels was held in B and destroyed, and we have found no duplicates of these collections (see also S. arequipense ) nor were any specimens photographed by J.F. Macbride as is the case for many other species. Nees von Esenbeck’s four taxa were distinguished based on leaf shape differences, notoriously difficult in the Morelloid clade. He ( Nees von Esenbeck 1843) cited two collections each for three varieties, mixing plants from the distributions of S. arequipense and S. furcatum (see discussion of Solanum furcatum var. subdentatum and var. Solanum furcatum subintegerrimum under S. arequipense ). Var. Solanum furcatum acutedentatum was based on collections from Valparaiso and from "planitie circa Tacoram, alt. 14000-17000[ft]" [ Volcán Tacora on the Chile/Peru border]; Solanum furcatum var. obtusedentatum on collections from San Fernando in central Chile and Arequipa in Peru. We have chosen to neotypify these two varieties with specimens from areas near the cited collections from Chile, in order to link the varietal names with S. furcatum . Searches in SGO and CONC failed to reveal duplicates of either Philippi s.n. from San Fernando or Gardner & Knees 8356, so we have reluctantly used as neotypes for var. Solanum furcatum obtusedentatum a collection we have seen in G and for var. Solanum furcatum acutedentatum the E duplicate we have seen of Gardner & Knees 8356, rather than duplicates in Chilean herbaria.

Specimens in Paris used by J. Rémy (1849) to describe Witheringia rubra J. Rémy are of plants of S. furcatum (see P00335356). He cited S. rubrum Mill. in synonymy and was clearly making a new combination based on that name. The type of the name Witheringia rubra is the type of S. rubrum Mill., itself a synonym of S. villosum (BM000942563 see Särkinen et al. 2018) although Särkinen et al. (2018) failed to site it in the synonym of S. villosum . Witheringia rubra (Mill.) J. Rémy is not a synonym of S. furcatum .

Figures

Figure 59. Solanum furcatum A habit B flower (A, B Anonymous s. n., grown from seed sent by J. Edmonds, originally from California [ADW 42421]). Illustration by M. L. Szent-Ivany, first published in Symon (1981) and previously published in Saerkinen et al. (2018). Courtesy of the Board of the Botanic Gardens and State Herbarium (Adelaide, South Australia), reproduced with permission.

Figure 60. Solanum furcatum A flowering branch B inflorescence with flowers at full anthesis C developing fruits D mature fruits (A, B, D Knapp s. n. Golden Gate Park C Gardner & Knees 8322). Photos by S. Knapp and M. Gardner. A, B, D previously published in Saerkinen et al. (2018) and Knapp et al. (2019).

Figure 61. Distribution map of Solanum furcatum in South America. For adventive distribution in North America and Australia, see Knapp et al. (2019) and Saerkinen et al. (2018) respectively.