Lycianthes coffeifolia, Bitter, Abh. Naturwiss. Vereins

Costa-Silva, Rafael & Agra, Maria de Fatima, 2022, Taxonomic Revision of Lycianthes (Capsiceae, Solanaceae) in Brazil, Systematic Botany (Basel, Switzerland) 47 (3), pp. 642-666 : 656-657

2. LYCIANTHES COFFEIFOLIA Bitter, Abh. Naturwiss. Vereins Bremen 24(2): 432. 1919. TYPE: BRAZIL. Amazonas: Acre-gebiet, Monte Mo, Nov 1911, E. Ule 9737 (holotype: B†, F Field Museum negative 2566 [photo!]); lectotype: US, US 01615264 barcode US 00027875 [image!], designated by Costa-Silva and Agra (2018a); isolectotypes: G! [2 sheets]; L 2881751 [image!], K 000585765 [image!]).

Shrubs, erect, 0.5–1.0 m tall, branched; stem and branches brown, striate, rugose, glabrescent, trichomes simple, eglandular, multicellular, uniseriate, 0.1–0.13 mm long, yellowish, hyaline. Leaves solitary or geminate; petiole short, 0.3–0.5 cm long, striate, glabrous; lamina chartaceous, 6.0–17.2 3 1.9–5.5 cm, concolorous, elliptical, oblong-elliptic, or obovate, glabrous on both surfaces, apex acuminate or acute, margin slightly wavy, base cuneate, puberulent on the midrib, trichomes simple, eglandular, uniseriate. Inflorescences 3–5-flowered; pedicels pendent in flower, 3.0–6.0 mm long, glabrescent, trichomes simple, eglandular, similar to those of the petiole. Calyx cupuliform, tube 1.4–1.5 cm long, appendages 10 in one series, 2.0– 2.3 mm long., lanceolate, attenuate toward to the apex, glabrous, emerging below the edge of the tube. Corolla white, stellate, 1.2–1.5 cm diam, lobes lanceolate, acute, 0.6–6.3 3 0.2–0.25 cm, interpetalar tissue connecting the petals on the basal 1 = 2; stamens equal, filaments 8–10 mm long, anthers oblong, connate, 3.6–3.7 mm long.; ovary ovoid to conical, 1.8–1.9 3 1.0– 1.2 mm, glabrous; style 3.8–4.0 mm long, erect; stigma clavate, bright green, glandular. Fruit long-ellipsoid, 1.5–1.8 cm diam, red, glabrous; fruiting pedicels pilose, trichomes setose; calyx accrescent, appendages 10, subulate, reflexed, 1.5–2.0 mm long. Seeds 3.5–4.0 3 3.0 mm, lenticular, flattened, reniform, pale yellow to beige. Figure 3 .

Distribution, Habitat, and Phenology — Lycianthes coffeifolia is a South American species distributed in Colombia, Bolivia, Peru, and Brazil. It is restricted to Acre State in Brazil ( Fig. 9 ) where it occurs in Open Ombrophilous Forests, Flood Terraces, Dense Ombrophilous Forests, and Deciduous Forests in areas near rivers and gallery forests, with a preference for humid soils. It may also occasionally occur in disturbed areas at forest edges and in wetlands at elevations from 50 to 300 m. Flowers and fruits were observed from May to November.

Taxonomic Notes — Lycianthes coffeifolia can be easily recognized by its elliptic-obovate leaves, pedicels pendent in flower and erect in fruit, calyces patelliform with 10 short and recurved appendages, and ellipsoid and red berries. Collections of Lycianthes coffeifolia in Brazilian herbaria are very infrequent, probably because its distribution in Brazil is restricted to Acre State, with only nine specimens being found during our searches.

Lycianthes coffeifolia was included in Lycianthes series Piliferae by Bitter (1919), although Dean et al. (2019) highlighted the probable misplacement of L. coffeifolia into this series based on habitat preferences, floral morphology, and seed characters. As we did not analyze the other species belonging to this series, no taxonomic decisions are being made on this matter.

Representative Specimens Examined — Brazil. — ACRE: Municipality of Cruzeiro do Sul , s.d., Daly 7638, 10907 (NY) ; s.d., Maas 12867 (NY); Municipality of Manoel Urbano, Parque Estadual Chandless , 26 Nov 2010, H. Medeiros 753 (RB) ; Municipality of Marechal Traumaturgo , 1 May 2001, Daly 10807 (NY) ; Municipality of Santa Rosa dos Purus , s.d., Silveira 1461 (NY) ; Municipality of Sena Madureira , 4 Oct 1968, Prance 7819 ( INPA,

NY); Municipality of Tarauca, s.d., C. Figueiredo 856 (NY); same locality, s.d., Eringhaus 401 (NY).

Additional Specimens Examined — Peru. — HUANUCO: Huamalies , 2 Nov 1938, Stork 9565 (F) . — JUNIN: Puerto Bermudez , 14–17 Jul 1929, Killip 26635 (F) . — LORETO: Lower Rio Huallaga, Oct-Nov 1929, Williams 5270 (F) . — SAN MARTÍN: Fundo Correviento , 28 May 1964, Schunke 6511 (F) ; Mariscal Caceres, Tocache Nuevo , 8 Jun 1970, Schunke 4040 (F) ; same locality, 17 Dec 1970, Schunke 4576 (F) .

3. LYCIANTHES INAEQUILATERA (Rusby) Bitter, Abh. Naturwiss. Vereins Bremen 24(2): 439. 1919. Brachistus inaequilaterus (Rusby) Rusby, Bull. New York Bot. Gard. 4: 470. 1907. Bassovia inaequilatera Rusby, Mem. Torrey Bot. Club 6(1): 90. 1896. TYPE: BOLÍVIA. Depto. La Paz: Prov. Larecaja, between Tipuani and Guanay, Dez 1892, M. Bang 1708 (lectotype: NY! barcode NY00138545 [image!], designated by Dean and Reyes (2018); isolectotypes: NY! barcode NY00138544 [image!]; NY! barcode NY00138546 A 76919 [image!], BM barcode BM00815915 [image!], E 00190709 [image!], F V0072756F!, GH 00077688 [image!], M! 0171840, MICH 1109881 [image!], MO02495247, PH 00030426 [image!], US 01175833 [image!], W 1893–0005612 [image!]).

Lycianthes fendleri (Rusby) Rusby, Bull. Torrey Bot. Club. 53: 210. 1926. Brachistus fendleri (Rusby) Rusby, Bull. New York Bot. Gard. 4: 470. 1907. Bull. New York Bot. Gard. 4: 470. 1907. Bassovia fendleri Rusby, Bull. Torrey Bot. Club 26: 197. 1899. TYPE: BOLIVIA. Depto. La Paz: Yungas, 6000 ft, 1885, H. Rusby 770 (lectotype designated here: NY barcode NY00138541[image!]; isolectotypes: NY barcode NY00138542 [image!], US barcode 1416211[image!]). Synonymized by Nee (2014).

Lycianthes subfalcata (Rusby) Rusby, Mem. New York Bot. Gard. 7: 349. 1927. Brachistus subfalcatus Rusby, Bull. New York Bot. Gard. 8(28): 117. 1912. TYPE: BOLIVIA. Depto. La Paz, Prov. Iturralde, San Buenaventura , 1400 ft, 14 Nov 1901, R. S. Williams 660 (holotype: NY! barcode NY00138563 [image!]; isotype US! barcode US 01175846 [image!]). Synonymized by Nee (2014).

Subshrub, erect, 1.0–2.0 m tall; stem and branches flexuous, terete, zigzag, green, striate, indument strigose, glabrescent on older branches; trichomes simple, uniseriate, 0.3–0.7 mm. Leaves geminate, anisophyllous; petiole terete, 0.5–0.8 cm long, strigose, trichomes similar to the branches; lamina entire, the longer 16.5–20 3 4–6.5 cm, elliptic, narrowly or broadly elliptic, 8–12 veins on each side, the smaller 2.3–3.6 3 1.3–1.6 cm, membranaceous to chartaceous, discolorous, opaque, acuminate at the apex, margin ciliate, base strongly oblique or obtuse, slightly decurrent to the petiole, strigose, the hairs denser on the midrib, trichomes simple. Inflorescences 1–4-flowered; pedicel terete, erect, 2.0–3.0 cm long, strigose, pubescence similar to the petiole. Calyx campanulate, 2.3–2.5 3 2.0– 2.1 mm, indument and trichomes similar to the petiole, appendages 10, subulate, emerging from below the calyx rim, sometimes inconspicuous, slightly unequal, 2.0–4.0 mm long. Corolla stellate, white, 0.8–0.9 3 0.9–1.0 cm, lobes lanceolate, 3.0–5.0 mm long, pubescent on the outer surface, trichomes simple, eglandular; stamens equal, filaments 3.0– 3.5 mm long; anthers yellow, narrowly elliptic, slightly attenuated to the apex, 2.5–3.0 mm long; ovary ovoid, 1.0–1.1 3 0.5–0.6 mm; style glabrous, 6–6.5 mm long, stigma capitate, 0.8–1.0 mm long. Fruit subglobose, 0.4–1.0 cm in diameter, red when ripe, fruiting pedicels erect, 0.6–1.5 cm long, pilose, trichomes simple, eglandular; calyx appendages erect, 2.0–3.0 mm long, slightly unequal, glabrous. Seeds numerous, 0.5–0.6 3 0.6 mm, somewhat triangular, yellowish to beige, reticulate. For illustration see Benıtez de Rojas and D’ Arcy (1997).

Distribution, Habitat, and Phenology — Lycianthes inaequilatera is a neotropical species with a South American distribution in Bolivia, Brazil, Colombia, and Venezuela. In Brazil, the species has been recorded exclusively for Acre State ( Fig. 9 ), in areas near the Purus River and its tributaries, at elevations from 140 to 300 m. It is common in wetlands near rivers, in forests on steep slopes, and in lowland or dryland forests, on poorly drained soils; it usually prefers shaded areas, but may also occur in open canopy vegetation with a shaded understory. It can also occur along the edges of disturbed areas. Flowers and fruits were collected from October to March.

Notes — Lycianthes inaequilatera is characterized by a subshrub habit, simple trichomes, geminate and anisophyllous leaves with asymmetric blades, and calyces with short and sometimes inconspicuous appendages.

Lycianthes inaequilatera is morphologically similar to L. amatitlanensis (and often misidentified as such). Brako and Zarucchi (1993), Martınez et al. (2002), and Nee (2014) considered L. amatilanensis a synonym of L. inaequilatera , although close examinations of the types of these taxa clearly indicated them as different species. Lycianthes inaequilatera can be recognized by having zigzag branches, dull leaves, and inconspicuous calyx appendages less than 1 mm long that are glabrescent at

anthesis. In contrast, L. amatitlanensis has erect branches, shiny leaves, and conspicuous calyx appendages (2.0–3.0 mm long) that are linear and pilose at anthesis. We consider L. amatilanensis to be a distinct taxon and not synonymous with L. inaequilatera , thus corroborating previous observations by Benıtez de Rojas and D’ Arcy (1997) and Bohs (2015) who also considered them distinct taxa.

Representative Specimens Examined — Brazil. — ACRE: Brasileia, Basin of Rio Purus, upper Rio Acre, Col^onia Santo Antonio ^, 29 Mar 1998, Daly 9875 (NY); Manoel Urbano, Rio Purus, Seringal Terra Nova, Colocaç ~ ao Terra Nova, 23 Nov 1996, Daly 9111 (NY); Marechal Taumaturgo, Reserva extrativista do Alto Jurua, Rio Jurua, right Bank, 6 Apr 1993, Daly 7802 (NY); Santa Rosa, Rio/Igarape Santa Rosa, tributary of Rio Purus, 15 Mar 2002, Daly 11374 (MO, NY); Sena Madureira, Basin of Rio Purus, Rio Iaco, right bank, Nova Olinda, 25 Oct 1993, Daly 7919 (NY); Basin of Rio Purus, Rio Macau ~ a, Colocaç ~ ao Apuı, 29 Mar 1994, Daly 8070 (MO, NY); same locality, Colocaç ~ ao Cala Boca, 29 Mar 1994, Lima 505 (NY).

Additional Specimens Examined — Peru. — MADRE DE DIOS: Manu, Parque Nacional Manu, 8 Oct 196, Foster 11688 (F) . — SAN MARTÍN: Mariscal Caceres, Tocache Nuevo , 23 Sep 1986, Knapp 8376 (MO) ; same locality, 14 Dec 1981, Plowman 11390 (F) . Venezuela. — SUCRE: Peninsula de Paria , 17 Oct 1984, Knapp 6766 (F) .

LYCIANTHES SECTION EUPOLYMERIS Bitter, Abh. Naturwiss. Vereine

Bremen 24: 423. 1919. TYPE SPECIES: Lycianthes rantonnetti

(Carriere) Bitter

Shrubs erect or scandent, perennial; trichomes simple, branched, bifurcate, and stellate. Lycianthes sect. Eupolymeris is represented in Brazil by six species: L. bitteri , L. cearaensis , L. glandulosa , L. stenoloba L. leptocaulis , L. rantonnetii .

Figures

FIG. 3. Lycianthes coffeifolia Bitter (Prance 7819). A. Flowering branch. B. Detail of the main vein of the leaf blade. C. Detail of eglandular trichome. D. Detail of a flower. E. Detail of calyx and gynoecium. F. Detail of the fruit and fruiting calyx. G. Detail of an isolated seed.

FIG. 9. Map of the geographic distribution of of L. amatitlanensis, L. asarifolia, L. coffeifolia, L. inaequilatera, and L. repens, based on herbarium specimen data.