Solanum urbanum var. ovatifolium Chodat

3. LYCIANTHES GLANDULOSA (Ruiz & Pavon) Bitter, Abh. Naturwiss. Vereine View in CoL Bremen 24: 348. 1919. Solanum glandulosum Ruiz & Pavon, Fl. Peruv. 2: 35, t. 167, f. b. 1799. TYPE: PERU. Departamento Junın: Prov. Chanchamayo, “circa Vicum Vitoc”, H. Ruiz & J. Pavon s.n. (lectotype: MA 747073 [image!]; isolectotype F V0073277F fragment [image!]). Lectotypified by Knapp (2008).

Solanum vitocense Dunal, Prodr. 13(1): 176 0.1852. TYPE: PERU. Departamento Junın: Prov. Chanchamayo, J. J. Taffala s.n., ex Herb. J. Pavon 39 (holotype: G! barcode: G00357863 [image!]) ; isotype G! barcode G00357865 [image!]. Synonymized by Bitter (1919).

Lycianthes hypomalaca Bitter, Abh. Naturwiss. Vereine View in CoL Bremen 24: 344. 1919. Solanum hypomalacum (Bitter) C.V. Morton, Contr. U.S. Natl. Herb. View in CoL 29: 64. 1944. TYPE: BRAZIL. Amazonas: Acre-gebiet, Estella, Seringal Auristella, Abr 1911, E. Ule 9734 (holotype B† [probably destroyed], F Field Museum negative F2579 [photo!]; lectotype designated here: B 10–0248778 [image!]; isolectotypes; G!, K barcode K000585767 [image!], L 2881728 [image!], MG 14568!, S barcode S04–2918 [image!], US 1615226 barcode US 00027884 [image!]). Synonymized by Nee (2014).

Lycianthes poeppigii Bitter, Abh. Naturwiss. Vereine View in CoL Bremen 24: 345. 1919. Solanum poeppigii (Bitter) C.V. Morton. View in CoL TYPE: PERU. Departamento Loreto: Prov. Alto Amazonas, [“Maynas”], proximo a Yurimaguas , Apr. 1830, E. Poeppig 2406 lectotype designated here: W barcode W0003089 [image!], F Field Museum negative 23069 [photo!]; isolectotypes: F V0043190F fragment [image!], P! barcode P00430724 [image!], W barcode W111233 [image!]). Synonymized by Costa-Silva and Agra View in CoL (2018b).

Lycianthes australe (C.V. Morton) Hunz. & Barboza, Darwiniana View in CoL 31: 29–30. 1992. Solanum australe C.V. Morton, Contr. U.S. Natl. Herb. View in CoL 29: 62. 1944. Lycianthes pseudolycioides (Chodat & Hassl.) Bitter, Abh. Naturwiss. Vereine View in CoL Bremen 24: 352. 1919. Solanum pseudolycioides Chodat & Hassl., Bull. Herb. Boissier View in CoL , ser. 2, 4: 84. 1904, non Rusby (1899). TYPE: PARAGUAY. Departamento Canindeyu: ın silva pr. Arroyo Mocoy, Sierra de Maracayu, Oct 1898 –1899, E. Hassler 4912 (holotype: G! G364763/1 barcode G00229596; isotypes: BM barcode BM00074098 [image!]; G 364763/2 barcode G00229598, G 364763/4 barcode G00392299, K barcode K000585756 [image!], MO! 1574591 barcode MO-503606 [image!]; NY! barcode NY 00172142 [image!]; P! barcode P000368685 [image!], P barcode P00634448 [image!]). Synonymized by Nee (2014).

Solanum urbanum var. ovatifolium Chodat View in CoL , Syn. Nov. Bull. Soc. Bot. Geneve 8: 152. 1916. TYPE: PARAGUAY. Caaguazu, Balansa 2080 (holotype: G! barcode G00392294).

Scandent shrubs, 2.0–6.0 m tall. Stem and branches terete, ferruginous, pubescent, trichomes branched and stellate, short-stalked, with 3–5 lateral rays. Leaves solitary on older stems and geminate in the upper branches; petiole 0.4–1.0 cm long, terete, trichomes similar to the branches; lamina 3.6–8.7 3 1.9–4.6 cm, ovate, ovate-eliptic, or narrowly-ovate, dull, apex acute, base oblique, rounded or cuneate, margin entire to slightly crenate, adaxial surface pubescent to tomentose on the veins, trichomes stellate, short stalked, abaxial surface pubescent to tomentose, more compact than on the adaxial surface, the trichomes branched and stellate. Inflorescences 3–8 flowered. Pedicels terete, pendent, 1.0– 1.2 cm long, tomentose-ferruginous, the trichomes similar to the branches. Calyx cupuliform, 3.0–4.0 mm diam, tube 2.0– 3.3 cm long, appendages 10, unequal, rounded at the apex, emerging from the 1 = 4 of the calyx apex, arranged in three series (a smaller, 2.0– 2.4 mm long, one intermediate 3.0– 3.7 mm long, and one longer 5.0–8.0 mm long), glabrescent to pubescent, trichomes similar to the pedicel. Corolla white, campanulate, 2.0– 2.5 cm diam., lobes lanceolate to elliptic-lanceolate, 1.0– 1.3 cm long, puberulous externally, interpetalar tissue 5.6–6.0 mm, linking the corolla lobes 1.5 mm below the lobe apex; filaments unequal (1 1 4), the longer, 5.2–6.0 mm long, 4 shorter, 1.0– 1.3 mm long; anthers 5.7–6.0 mm long, attenuated to the apex, sagittate at the base; ovary subglobose, 1.0– 1.4 mm diam; style 10.0 mm long; stigma somewhat bilobed, glandular, ca. 1.0 3 0.5 mm diam, dark green. Fruit globose, orange when ripe, 0.8–1.0 cm diam, calyx accrescent, 1.5–1.7 cm diam, appendages perpendicular to the fruit, slightly recurved at the apex, 0.4–0.5 cm long; pedicel erect, terete, puberulous, trichomes stellate, sessile. Seeds reniform, compressed, 2.8–3.0 3 2.2–2.5, yellowish, partially reticulate.

Distribution, Habitat, and Phenology — Lycianthes glandulosa has a wide distribution in South America ( Argentina, Bolivia, Brazil, Paraguay, and Peru). It occurs in both northern and southern Brazil along humid forest edges, but may inhabit drylands and clayey soils, disturbed environments, and secondary forests ( Fig. 10 View FIG ), at elevations from 150 to 800 m. Plants in flower were observed from April to November and with fruits in May and December.

Notes — Lycianthes glandulosa is characterized by its scandent habit, with geminate leaves, stellate trichomes, inflorescences sessile to short stalked, 3–8-flowered, corolla rotate, and calyx patelliform-cupular on fruits, with 10 unequal appendages.

Some collections of Lycianthes glandulosa have been misidentified as L. pauciflora in Brazilian and international herbaria, mainly due to their sympatric distribution in the Brazilian Amazonia and their similar characteristics, such as a climbing habit, stellate trichomes, and white corollas. Lycianthes glandulosa can be clearly distinguished from L. pauciflora , however, by having broader leaves, more congested inflorescences, and unequal and erect calyx appendages in more than two series.

Several names have been proposed for Lycianthes glandulosa (which were synonymized by Bitter (1919), Morton (1944), Nee (2014), and Costa-Silva and Agra (2018b)), mainly due to its wide distribution, the absence of data concerning the types for some of those names, and the fact that specimens from the Berlin herbarium were destroyed.

Lycianthes hypomalaca was proposed by Bitter (1919) based on the collection E. Ule 9734 (which corresponds to Field photo negative F2579) that was deposited in the Berlin Herbarium and probably destroyed. There is another exsiccate of this same collection at the Berlin Herbarium under barcode B 10–0248778 that has flowers and fruits and is well-preserved and, for this reason, was chosen as the lectotype.

Bitter (1919) proposed Lycianthes poeppigii as a new species and prepared a detailed description based on the specimen E. Poeppig 2406 deposited at the Vienna Herbarium (W). Our analyses of the types revealed the presence of two exsiccates, both belonging to the E. Poeppig 2406 collection deposited at W, therefore indicating the need to designate a lectotype. There is also a duplicate at P, but the leaves are not well preserved, and it has only a fruiting calyx, no flowers or fruits. We therefore chose exsiccate number W0003089 as the lectotype, which has flowers and fruits and handwritten notes by Bitter.

Solanum australe View in CoL was a new name proposed by Morton (1944) to replace Lycianthes pseudolycioides (Chodat & Hassl.) Bitter View in CoL , whose homonym of Rusby (1899) has priority according to Article 53.1 of the International Code of Nomenclature for Algae, Fungi and Plants ( Turland et al. 2018). Lycianthes pseudolycioides View in CoL was based on the type of Solanum pseudolycioides, E. Hassler View in CoL 4912 collection, deposited at G Herbarium.

Solanum urbanum var. ovatifolium View in CoL was described by Chodat (1916) based on the collection Balansa 2080. It was considered to be synonymous with L. pseudolycioides View in CoL by Bitter (1919), but

was later synonymized under L. pauciflora View in CoL by Hokche (2008). We were able to analyze the holotype of this name deposited in Herbarium G, however, and identified a set of characters (calyx cupuliform, appendages slightly curved at the apex in the fruits) clearly different from the concept of L. pauciflora View in CoL , which led us to synonymize it here under L. glandulosa View in CoL .

Representative Specimens Examined — Brazil. — ACRE: Brasileia, Basin of Rio Acre, tribuary of Rio Purus , Colonia ^Santo Ant^onio , 21 Oct 1997, Daly 9674 (NY) ; M^ancio Lima , 8 Nov 1991, Cid Ferreira 10619 (NY) ; Porto Acre, 22 Jun 1994, Oliveira 274 (MO), Rio Branco , 5 Jun 1991, Daly 6850 (MO) ; 23 Oct 1980, Cid Ferreira 3020 (NY, RB); Tarauaca , 26 Jun 2006, Silveira 4076 (RB) . — AMAPÁ: Mazag ~ ao, 19 Sep 1985, Pires 631 (NY) ; Porto Terezinha , 20 Nov 1954, Cowan 38465 (NY) . — AMAZONAS: Embira , Jun 1933, Krukoff 4874 (F, MO, S, US) ; Humaita , 14 Sep–11 Oct 1934, Krukoff 6364 (MO, S, US) . — MARANHÃO: Vila Rondon, 09 Aug 1976, Pinheiro 46 (IAN). Mato Grosso do Sul : Chapada dos Guimar ~ aes, 23 Feb 1997, Nave et al. 1176 (RB) ; Porto Murtinho , 14 Nov 2002, Hatschbach 74208 (NY) . — PARÁ: Ananideua , 21 Nov 1957, Silva 469 (IAN) ; Rodovia Belem- Brasılia km 90, 21 set 1959, Kuhlman 266 (SP) . PARANÁ: Lobato , 18 Jul 1962, Gomes e Matos 1189 (RB) ; Londrina, Mata do Godoy, Favoretto s.n. (RB), Fazenda Santa Helena , 04 Sep 1986, Paro 14 (RB), 15 May 1986, Pizzaia 65 (RB) ; 20 Jun 1985, Rodas 10 (RB); Medianeira , 23 Feb 1971, Hatschbach 26482 (F) ; Parque Nacional do Iguaçu , 12 May 1949, Duarte 1758 (F) ; Sao ~ Pedro do Ivaı , 15 Oct 2003, Ribas 5616 (RB), 19 Dec 2003, Ribas 5272 (RB) . — RONDÔNIA: Chapada dos Parecis , 02 Nov 1979, Vieira 865 (INPA, MO) ; Vilhena , 02 Nov 1979, Vieira 865 (MO, NY) .

Additional Specimens Examined — Argentina. — MISIONES: Iguazu, Parque Nacional Iguazu , 16 Dec 1991, Vanni 2885 (F) . Bolıvia. — BENI: Vaca Diez , 1 Dec 1983, Boom 4059 (F) . — PANDO: Abuna, Rio Madeira , 19 Nov 1968, Prance 8638 (F) . — SANTA CRUZ: Sercado , 29 Aug 1925, Steinbach 7194 (F) . Paraguay. — ALTO PARANÁ: Estancia Rio Bonito , 26 Dec 1994, Zardini 41758 (RB) ; Reserva Biologica, cerca del rio Itabo , 16 Oct 1996, Schinini 31473 (F) . Peru. — AMAZONAS: Zungaro Cocha, near Iquitos , 17 Dec 1964, C. Dodson & J. Torres 2927 (F) . — HUÁNUCO: Divisoria , 14 Sep 1946, Woytkowski 34530 (F) . — JUNIN: between Puerto Bermudez and Cahuapanas , 28–19 Jul 1929, Killip 26709 (F) . — LORETO: Coronel Portillo, Iparia , 16 Sep 1968, Schunke 2785 (F) ; Gamitanacocha , 6 Feb 1935, Schunke 178 (F) ; Maynas, 20 Jul 1968, McDaniel 10718 (F), 20 Oct 1976, Revilla 1558 (F) ; Prov. Pachitea, Dtto. Honoria , 26 Apr 1967, Schunke 1893 (F) ; Quebrada Shanuce above Yurimaguas , 11 Jul 1972, T. Croat 18030 (F) ; Santa Rosa , 11 Nov 1929, Williams 4956 (F) . — SAN MARTÍN: Chazutu, Rio Huallaga , Mar 1935, Klug 3991 (F) .