Argentina. Salta: [Dtto. La Caldera], Pasaje del Río Juramento, Feb 1873, P.G. Lorentz & G. Hieronymus 364 (holotype: B [destroyed]; lectotype, designated by Barboza et al. 2013, pg. 262: GOET [GOET003574]; isolectotypes: CORD [CORD00004269], DR [DR054234], US [00027741, acc. # 282274]).

Bolivia and Argentina.

Type material for the varietal names coined by Georg Bitter in the original publication of S. physalidicalyx (Bitter 1912) may correspond to duplicates of the type collection of the species itself, making all three names homotypic. In describing var. integrascens Bitter (1912) states “var. integrifolia quod in descriptione specie pro typo habui” [var. integrifolia I had in the description of the type of the species] suggesting this name at least is based on Lorentz & Hieronymus 364. Bitter often used duplicates with minor leaf variations as material for describing infraspecific variation (Knapp 2013). None of the duplicates of these Lorentz and Hieronymus collections from Salta have annotations in Bitter’s hand, and we have found no other collections of S. physalidicalyx made by Lorentz and Hieronymus from “Pasaje de Juramento”.

Cultivated at the Glasgow Botanical Garden [protologue] from seeds sent by J. Tweedie from “near Buenos Ayres”, Anon. s.n. (lectotype, designated by Edmonds 1972, pg. 102 [as “holotype”], second step designated here: K [K000585739]; isolectotype: K [K000585738]).

Bolivia and Argentina.

Edmonds (1972) stated that the holotype of S. tweedieanum was held at K; Barboza et al. (2013) repeated this citation but added reference to a single sheet (K000585739). This is not effective lectotypification under Art. 9.23 of the Code (Turland et al. 2018). The specimen cited as holotype by Barboza et al. (2013) and selected as lectotype here (K000585739) has open flowers and several buds and is a better match for the illustration in the protologue than the other sheet at Kew (K000585738); both specimens are annotated “S. Tweedianum Hook./B^os. Ayres. Cult.” in W.J. Hooker’s handwriting and bear the herbarium stamp “Herbarium Hookerianum/1867” indicating they come from Hooker’s own herbarium. There is no evidence on the specimens themselves that they were taken from the plants cultivated in Glasgow mentioned in the protologue, nor that they were collected prior to 1835; but the top part of the stem mounted on the sheet we have selected as the lectotype at Kew (K000585739, https://plants.jstor.org/stable/pdf/10.5555/al.ap.specimen.k000585739) is an excellent (mirror-image) match for Tab. 3385 in the protologue (Hooker 1835; see https://www.biodiversitylibrary.org/item/14341#page/36/mode/1up) suggesting it does represent original material. The original, and all subsequent, spelling of the name was “Tweedianum” where John Tweedie’s name was implicitly latinized as “Tweedius” in which the terminal vowel was eliminated. This is not acceptable under Art. 60.9(a)(1) of the Code (Turland et al. 2018) and the name should be formed as “tweedieanum” (e.g., Art. 60.9, Ex. 31, Turland et al. 2018).

Like Solanum tweedieanum Hook. but differing in sessile leaves with broadly winged petioles, pedicels in flower longer than 1 cm, larger flowers and anthers more than 1 mm wide.

Figure 2. 

Solanum hunzikeri Chiarini & Cantero A flowering stem B inflorescence C flower D open flower E immature fruit showing the accrescent calyx not completely covering the berry F flower showing pubescent adaxial surface of the filaments G calyx H style with pubescence confined to the portion inside the anther cone I adaxial surface of the anther showing the pores elongating with age J abaxial surface of the anther K seed L stone cell (sclereid).

Argentina. Catamarca: Dtto. Ambato, Los Morteritos, Sierra de Ambato, falda E, subiendo desde El Rodeo hacia el Cerro Manchado [Cerro Manchao], 2300–2400 m, 13 Jan 1973, A.T. Hunziker & R. Subils 22205 (holotype: CORD [CORD00013086]).

Herb or subshrub from a woody base ca. 50 cm tall; stems terete or only slightly angled, densely glandular pubescent with glandular papillae and transparent spreading simple 3–8-celled uniseriate trichomes 0.5–1 mm long, some to 1.5 mm long; bark of older stems pale brown, glabrescent; new growth densely glandular pubescent with simple uniseriate trichomes to 1 mm long. Sympodial units plurifoliate, the leaves not geminate. Leaves simple to shallowly toothed, (2-)4.5–14 cm long, (1.1-)2–7 cm wide, elliptic in outline, membranous or somewhat thick and fleshy, concolorous; adaxial surface moderately and evenly glandular pubescent with transparent spreading, simple uniseriate trichomes ca. 0.5 mm long on the lamina, ca. 1 mm long on the veins; abaxial surface moderately and evenly glandular pubescent like the adaxial surface, but the trichomes denser and longer, to 1.5 mm long; principal veins 4–7 pairs, densely glandular pubescent; base attenuate and strongly decurrent onto the petiole; margins entire or shallowly toothed, the teeth if present 1–2 mm long, 2–3 mm wide, broadly deltate with somewhat rounded tips; apex acute; petioles absent and the leaves sessile or 0–0.1 mm long, the decurrent leaf bases running onto the stem, glandular pubescent like the stems and leaves. Inflorescences 2.5–4 cm long, opposite the leaves, unbranched but occasionally forked (Rodríguez 1421), with 10–20 flowers, densely glandular pubescent with transparent spreading simple uniseriate trichomes to 1.5 mm long; peduncle 1.2–2.5 cm long; pedicels 1.3–1.5 cm long, 0.5–0.7 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, spreading at anthesis, densely glandular pubescent, articulated at the base; pedicel scars irregularly spaced 1–2 mm apart. Buds ellipsoid, the corolla ca. halfway exserted from the calyx before anthesis. Flowers 5-merous, perfect. Calyx tube 2–3 mm long, conical, the lobes 2.5–4 mm long, long-triangular, densely glandular pubescent with simple uniseriate trichomes like the pedicels and rest of the inflorescence, the tips acuminate and somewhat recurved at anthesis. Corolla 1.6–2.5 cm in diameter, pale lilac to violet with a yellow-green central star, stellate, lobed ca. 1/2 way to the base, the lobes 5–5.5 mm long, 4–5.5 mm wide, deltate, reflexed or spreading at anthesis, adaxially glabrous, abaxially sparsely glandular papillate especially on the midvein, tips and margins; stamens equal; filament tube 0.35–0.5 mm; free portion of the filaments 1–1.5 mm, almost glabrous, but with a few tangled transparent eglandular simple uniseriate trichomes adaxially; anthers 4–5.5 mm long, 1.25–1.6 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary conical, glabrous; style 7–8 mm long, densely papillate with a few longer simple trichomes in the lower third; stigma large capitate to slightly bilobed, the surface minutely papillate. Fruit a globose berry, 1–1.2 cm in diameter, green (?) at maturity, opaque, the surface of the pericarp glabrous, thin, matte; fruiting pedicels 1.5–2 cm long, ca. 1.5 mm in diameter at the base, ca. 2 mm in diameter at the apex, somewhat woody, deflexed from the weight of the berry, glandular pubescent to somewhat glabrescent; fruiting calyx accrescent in young fruit tightly investing the berry, the tube 3–5 mm long, later tearing and the berry exposed, the lobes 3–5 mm long, ca. 3 mm wide, appressed to spreading. Seeds ca. 40 per berry, 1.5–2 mm long, 1–1.7 mm wide, flattened teardrop shaped with an apical hilum, reddish brown, the surfaces minutely pitted, testal morphology not clearly seen. Stone cells 10–11 per berry, 1–1.3 mm in diameter, globose, scattered throughout the berry. Chromosome number not known (but see comments on DNA content below).

(Figure 3). Solanum hunzikeri occurs in Argentina in the provinces of Catamarca and adjacent Salta and Tucumán and extends north to Bolivia in the departments of Tarija and Chuquisaca. The distribution is somewhat disjunct possibly due to loss of the wet high elevation foggy grassland habitat in the intervening areas.

Figure 3. 

Distribution of Solanum hunzikeri Chiarini & Cantero.

Solanum hunzikeri is confined to wet cloud forests and foggy grasslands above 1800 m elevation; it also grows in the ecotones between these vegetation types. These foggy grasslands are dominated by tall grasses (e.g., Festuca hieronymi Hack., Cinnagrostis polygama Griseb., Elionurus muticus (Spreng.) Kuntze [Poaceae]) and shrubs (e.g., Baccharis spp., Stevia spp. [Asteraceae]). Solanum hunzikeri can also be locally frequent on open grassy terraces with scattered palms, in narrow valleys with the lower slopes covered in seasonally moist forest dominated by Parajubaea torallyi (C.Mart.) Burret (Arecaceae) and with abundant Podocarpus spp. (Podocarpaceae) and can be found on steep, stony slopes in undisturbed grassland areas.

This species is named in honour of the late Ing. Armando T. Hunziker of IMBIV in Córdoba, whose life work on the Solanaceae inspired a generation of solanologists, in both Argentina and globally.

(IUCN 2019). AOO (80 km² – EN); EOO (97,627 km² – LC). Although the large extent of occurrence would suggest S. hunzikeri is not of conservation concern, the limited number of localities, the specialised habitat and the disjunct distribution suggest the species should be considered at risk. Solanum hunzikeri occurs in a very restricted habitat in which there are few officially protected areas. In these landscapes the main threat to the ecosystem is over-grazing; the introduction of alien forage species such as Pennisetum clandestinum Hochst. ex Chiov. (Poaceae) has severely altered the nature of the high elevation foggy grasslands and forest edges in which S. hunzikeri occurs. Although some populations are found in currently protected areas such as the Parque Nacional Aconquija, these areas are considered too small and isolated to provide long term conservation (Brown 1995). Based on the area of occupancy, the number of localities (ca. 8) in a disjunct distribution and threats to the habitat, we assign a preliminary threat status of Vulnerable (VU B2a,biii) for S. hunzikeri. The exploration of these relatively inaccessible habitats in the area between the currently known populations of S. hunzikeri is a priority.

Solanum hunzikeri had been recognized as distinct from other glandular-pubescent species in Argentina in the early 20^th century by the German botanist Georg Bitter as “Solanum catamarcae”, a name already occupied in Solanum (S. catamarcae Bitter ex Brücher, a synonym of S. boliviense Dunal, see Spooner et al. 2019). Morton (1976) in his treatment of Solanum for Argentina, cited Sleumer 2259 as part of his concept of S. atriplicifolium and stated “This last appears to be a local form that has the petioles broadly winged nearly throughout instead of at the apex only. According to determinations by Dr. Sleumer this plant was given the unpublished name of “Solanum catamarcae” by Bitter.” Morton annotated the sheet of Sleumer 2259 in US as “Solanum atriplicifolium var. sleumeri Morton HOLOTYPE” in 1971 but did not publish the infraspecific epithet; he also annotated Sleumer 2311 (US) as a paratype of the same.

The species is now known from a wider distribution, and additional specimens have clarified its differences from the widespread and highly variable S. tweedieanum. Solanum hunzikeri can be distinguished from S. tweedieanum populations in similar high elevation areas in its strongly attenuate and winged leaf bases, those of S. tweedieanum are more truncate. The single collection we have seen of S. hunzikeri with mature fruit (Rodríguez 1421 from Salta) has the calyx not covering any part of the mature berry; berries of S. tweedieanum are tightly covered by the accrescent calyx for at least 50% of their length. More collections of S. hunzikeri in fruit are needed to assess these differences. Preliminary data on DNA content for S. hunzikeri and S. tweedieanum (F. Chiarini unpubl.) show differences but suggest that, like S. tweedieanum (Moscone 1992), S. hunzikeri is diploid.

(paratypes). Argentina. Catamarca: Dtto. Ambato, camino desde El Rodeo rumbo al Cerro el Manchado [Manchao], Falda El Morro, 2593 m, 24 Feb 2016, Barboza et al. 4703 (CORD); Dtto. Ambato, Sierra de Ambato (falda E), subiendo desde El Rodeo hacia el Cerro Manchado [Manchao], 2300 m, 23 Feb 1967, Hunziker 19073 (CORD, US); Dtto. Pomán, Rumbo al Cerro Manchado [Manchao], Sierra de Ambato, falda E, subiendo El Rincon hacia Las Casitas, rumbo al Cerro Manchado [=Cerro del Manchao], 2300–2500 m, 18 Feb 1970, Hunziker & Ariza 20319 (CORD); Dtto. Pomán, Rumbo al Cerro Manchado, Sierra de Ambato, falda E, subiendo El Rincon hacia Las Casitas, rumbo al Cerro Manchado [= Cerro del Manchao], 2300–2500 m, 18 Feb 1970, Hunziker & Ariza 20329 (CORD); Dtto. Ambato, Los Morteritos, Sierra de Ambato, falda E, subiendo desde El Rodeo hacia el Cerro Manchado [Manchao], Los Morteritos, 2300–2400 m, 13 Jan 1973, Hunziker & Subils 22205 (CORD); Dtto. Ambato, Los Morteritos, Sierra del Ambato, falda E, subiendo desde El Rodeo hacia el Cerro Manchado [Manchao], 2300–2400 m, 13 Jan 1973, Hunziker & Subils 22206 (CORD); Dtto. Andalgalá, Río Potrero, 2600 m, 13 Feb 1942, Rohmeder s.n. (LIL); Dtto. Andalgalá, Río Lampacillo-Río Potrero, Entre Río Lampacillo y Río Potrero, 2700–2900 m, 26 Feb 1951, Sleumer 1834 (LIL, US); Dtto. Andalgalá, Mesada La Primera, Mesada La Primera, Las Estancias, 1900 m, 11 Feb 1952, Sleumer 2132 (LIL); Dtto. Andalgalá, Los Queñoales, arriba de la Mesada de Las Rosas, 2300–2400 m, 15 Jan 1952, Sleumer 2259 (G, LIL, US); Dtto. Andalgalá, Cuesta de la Negrilla, cerca de la Mina de Capillas, 3000–3100 m, 2 Mar 1952, Sleumer 2690 (CORD, G, US); Dtto. Andalgalá, Cuesta de la Negrilla cerca de la Mina de Capillitas, 3100 m, 2 Mar 1952, Sleumer 2691 (CORD, G, US, W); Dtto. Andalgalá, Mina de las Capillitas, cerca de los edificios, 2350 m, 2 Jan 1952, Sleumer 2692 (US, W). Salta: Dtto. Cafayate, Peñas Blancas, Cerros de Cajón [Sierras de Quilmes], 4040 m, 30 Mar 1914, Rodríguez 1421 (BR, CORD, SI). Tucumán: Dtto. Alberdi, Escaba, 2300 m, 27 Dec 1913, Monetti 1838 (LIL); Dtto. Alberdi, Estancia Yunka Suma, Valle del Río Las Chacras [as Catamarca, Dtto. Andalgalá on labels], 1800 m, 23 Feb 1951, Sleumer 1610 (LIL); Dtto. Alberdi, Cumbres de Suncho, Quebraditas del Portezuelo Sta. Anna [as Catamarca, Dtto. Andalgalá on labels], 2150 m, 8 Feb 1952, Sleumer 2311 (LIL, US).

Bolivia. Chuquisaca: prov. Zudañez, a 82 km de Sucre, entre Tarabuco y Sudanéz, paraje Lambayo, 2756 m, 25 Feb 2004, Cocucci et al. 3357 (CORD); prov. Azurday, Tarvita, ca. 3 km S of summit on road from Tarvita to Azurduy, 2800 m, 4 Dec 1999, Wood et al. 15303 (K); prov. Tomina, ca. 1 km W of summit of pass between Villa Tomina and Villa Serrano, 2700 m, 17 Mar 2002, Wood 17868 (K); prov. Azurduy. Bajando de la cumbre hacia Duraznal en el camino de Azurduy, 2459 m, 11 Dec 2004, Wood & Huaylla 21130 (K); prov. Tomina, entre Villa Serrano y Tomina, en la cumbre, 2580 m, 4 Mar 2006, Wood et al. 22394 (K); prov. Zudañez, AMNI El Palmar, AMNI El Palmar, along trail from Torotoro to El Palmar crossing Río Mission Waypu., 2800 m, 2 Feb 2007, Wood et al. 22612 (K). Tarija: Sama, between Tarija and Villazón, 3546 m, 27 Feb 1939, Balls 6111 (E, K, US); de Tarija a Narváez, 2000–2500 m, 19 Mar 1982, Kiesling et al. 3734 (SI); de Tarija a Iscayachi, 2000–3000 m, 20 Mar 1982, Kiesling et al. 3845 (SI); Mun. O’Connor, at the top of the first pass W of Entre Ríos on road to Narváez and Tarija, 1800 m, 21 Jan 2001, Wood & Goyder 16901 (K).

Argentina. Córdoba: IMBIV, Universidad Nacional de Córdoba [plant grown from Barboza et al. 4703], 450 m, 15 Feb 2017, Barboza 4763 (CORD).

Like Solanum nitidibaccatum Bitter but differing in eglandular, white pubescence, strongly winged stems, fleshy calyx lobes that are spreading in fruit and larger berries; also similar to S. americanum Mill. but differing in strongly winged stems and dark green mature berries marbled with white markings.

Figure 4. 

Solanum marmoratum Barboza & S. Knapp A habit (Barboza et al. 5099) B, C details of the winged stems (both at the same scale, B from Barboza et al. 5136, C from Barboza et al. 5073) D inflorescence (Barboza et al. 5136) E flowers, showing the included style and the filaments that elongate with flower age (Barboza et al. 5136) F mature fruits (Barboza et al. 5073) G Detail of berries showing the spreading fleshy calyx in fruit (Barboza et al. 5130). All photographs by S. Knapp.

Argentina. La Pampa: Dtto. Loventué, 10 km al W de Luan Toro, rumbo a Loventué, 297 m, 9 Feb 2020, G.E. Barboza, S. Knapp, F. Chiarini & R. Fortunato 5099 (holotype: CORD [CORD00007007]; isotypes: BAB, BM).

Figure 5. 

Solanum marmoratum Barboza & S. Knapp (Barboza et al. 3668, BM [BM001134643]).

Watery annual herb, 10–100 cm tall, sprawling and somewhat prostrate when very large. Stems strongly winged, the wing to 1 mm side, sometimes with spinose processes (old trichome bases), sparsely to moderately pubescent with spreading to appressed eglandular simple 5–8-celled uniseriate trichomes 0.5–1 mm long, these drying white; new growth densely pubescent with eglandular, white simple uniseriate trichomes 0.5–1 mm long; older stems greenish white, not woody. Sympodial units difoliate, the leaves not geminate, axillary shoots common. Leaves simple and shallowly toothed, 2–10 cm long, 1.5–6 cm wide, much larger in older plants, ovate, widest in the lower third, membranous, watery and somewhat succulent, concolorous, very bright green on live plants; adaxial and abaxial surfaces evenly white-pubescent with eglandular simple 5–8-celled uniseriate trichomes 0.5–1 mm long, these longer and denser on the veins; principal veins 5–6 pairs; base attenuate onto the petiole; margins shallowly and irregularly toothed, the teeth 2–4 mm long, 2.4- mm wide, broadly deltate, with blunt tips; apex acute; petioles 0.5–2.5 cm long, somewhat winged from the attenuate leaf base, pubescent with simple uniseriate trichomes like the stems and leaves. Inflorescences (1)2–3 cm long, internodal and extraaxillary, unbranched, with 5–7 flowers clustered at the tip, usually only 1–2 open at a time, sparsely and evenly pubescent with antrorse simple uniseriate trichomes 0.5–1 mm long like the stems and leaves; peduncle 1.4–2.5 cm long; pedicels 0.4 cm long, ca. 0.5 mm in diameter at the base, ca. 0.6 mm in diameter at the apex, slightly tapering, spreading, eglandular pubescent like the rest of the inflorescence, articulated at the base; pedicel scars tightly packed at the tip of the inflorescence, 0.5–1.5 mm apart. Buds broadly ellipsoid, the corolla included in the calyx tube until just before anthesis. Flowers 5-merous, perfect. Calyx tube 1.2–1.5 mm long, cup-shaped, the lobes 1–1.5 mm, narrowly deltate-triangular, fleshy and recurved in live plants, sparsely pubescent with eglandular white trichomes on both surfaces like the rest of the plant. Corolla 0.5–0.8 cm in diameter, white with a green central star, stellate, lobed ca. halfway to the base, the lobes ca. 2.5 mm long, ca. 2 mm wide, spreading to slightly reflexed at anthesis (flowers closing daily and lasting for several days), adaxially glabrous, abaxially densely pubescent with tiny simple uniseriate trichomes especially at the tips. Stamens equal or slightly unequal with one anther marginally longer than the rest; filament tube ca. 0.1 mm long; free portion of the filaments 0.5–1 mm long, elongating through anthesis, with a few tangled transparent simple uniseriate trichomes adaxially; anthers 1–1.5 mm long 0.6–1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores elongating with age. Ovary conical, glabrous; style 2–2.5 mm, included within the anther cone or the stigma just beyond, densely papillate in the lower 3/4; stigma large capitate, held at the level of the anthers when flowers first open, later included within the anther cone, bright green in life plants, the surfaces minutely papillate. Fruit a globose berry, 0.8–1.5 cm in diameter, dark green marbled with white at maturity, glabrous, translucent, the pericarp surface thin, shiny; fruiting pedicels 1.2–1.5 cm long, ca. 1 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, fleshy and watery, tapering to the spreading calyx, strongly deflexed at maturity, with a distinct bend at the pedicel base; fruiting calyx somewhat expanded, the tube 3–4 mm long, the lobes 4–5 mm long, ca. 3 mm wide, spreading and fleshy, the tips rounded. Seeds 50–70 per berry, ca. 2 mm long, ca. 1.7 mm wide, flattened teardrop shape with an apical hilum, pale tan to reddish brown, the surfaces minutely pitted, the testal cells mostly rectangular to pentagonal in outline, more sinuate towards the seed centre. Stone cells 1–2, 1–1.1 mm in diameter, found randomly positioned in the berry. Chromosome number: not known.

(Figure 6). Solanum marmoratum is endemic to Argentina and occurs in the provinces of Catamarca, La Pampa, La Rioja, San Juan and San Luis; we expect it also to be found in Mendoza, because several collections are known from Desaguadero (San Luis) a locality very close to the provincial border that crosses through uniform habitat.

Figure 6. 

Distribution of Solanum marmoratum Barboza & S. Knapp.

Solanum marmoratum is found in shady areas in Prosopis woodlands (Fig. 4A) and at the edges of arable fields; it usually grows under trees and shrubs with a number of other herbaceous plants such as S. tweedieanum, various species of Asteraceae and grasses. Specimens have been collected from 200 to 1400 m elevation.

The species is named for its distinctive marbled berries (Fig. 4F, G) that easily distinguish it from the similar tiny-flowered eglandular species S. americanum.

(IUCN 2019). AOO (84 km² – EN); EOO (239,336 km² – LC). Solanum marmoratum is a relatively widespread species, the extent of occurrence suggests is should be given a status of least concern. The small area of occupancy perhaps reflects a lack of collecting in the dry forest and partially degraded habitats where S. marmoratum occurs. The number of localities (ca. 9) is probably an underestimate due to the widespread perception that these habitats are not interesting; most collections are quite old and the species has not be collected recently (except by us). The large-scale conversion of land in the range of S. marmoratum to intensive monoculture of commercial crops such as maize, peanuts and sunflowers poses a risk for this and other species in these habitats; use of herbicides and elimination of patches of forest leave little room for even weedy species to persist. We suggest a preliminary threat status of Least Concern (LC) to S. marmoratum, but the widespread habitat conversion in central Argentina warrants further studies as to population status across the species’ historical range.

Solanum marmoratum has long confused botanists working with Argentinian solanums. In the herbarium at CORD specimens of S. marmoratum collected by P. Steibel in the province of San Luis were the subject of correspondence with A.T. Hunziker over their identification; they were tentatively identified as S. adventitium Polg., a synonym of S. americanum described from adventive material in Hungary (Särkinen et al. 2018). None of the pre-1970s specimens we have seen were cited in Morton (1976), but Semper s.n. at US (barcode 02837698) was annotated “Solanum dolichopteryx Morton, paratype” by C.V. Morton in 1971. We have not found specimens annotated as other types at US or elsewhere. A.T. Hunziker had kept specimens of this species aside with the herbarium name “Solanum alatocaule”, a reference to the strongly winged stems (Fig. 4B, C) on the folder, but never described it. We collected S. marmoratum in 2013 (Barboza et al. 3668) along with S. tweedieanum, and mistakenly noted the leaves of S. marmoratum as sticky (see Fig. 5); it was only examination of the dried specimens that alerted us to our error. Careful examination of all morelloid collections at CORD in early 2020 showed the distinctness of S. marmoratum, and its relatively widespread distribution.

The flowers of S. marmoratum are among the tiniest in the morelloid solanums (Fig. 4D, E) rivalled only by the globally distributed S. americanum and S. nitidibaccatum Bitter and the North American S. emulans Raf. (see Knapp et al. 2019). Solanum nitidibaccatum also has somewhat marbled berries but is always extremely sticky and covered with glandular trichomes, in contrast to the eglandular pubescence of S. marmoratum. Solanum americanum and S. emulans both have eglandular pubescence but have purplish black rather than green marbled berries. The fleshy spreading calyx lobes of S. marmoratum (Fig. 4G) are distinct from those of all of these taxa with tiny flowers.

Solanum marmoratum appears to be highly autogamous and is perhaps entirely self-fertilising. The style is completely included within the anther cone (Fig. 4D, E) and the filaments appear to elongate through anthesis (see. Fig. 4E) bringing the style further into the cone as the flower ages. Flowers stay open for several days (closing at night) and in cultivation the plant goes from bud to flower to fruit in 15–18 days with all flowers setting fruit. Over the course of anthesis the style becomes enclosed in the anther cone (Fig. 4E), with the anthers as they dehisce leaving pollen directly on the stigma. Ripe berries last more than two weeks after being gathered from desiccated plants, remaining unchanged as to colour or odour.

(paratypes). Argentina. Catamarca: Dtto. Santa María, Chiñucán, Sierra de la Aconquija, falda O, Chiñucán, 12 Apr 1948, Reales 1264 (CORD); Dtto. Belén, Yacutula, Mar 1879, Schickendantz 113 (CORD). La Pampa: Dtto. Toay, Reserva Provincial Parque Luro, pasando la laguna Luro, 233 m, 18 Jan 2013, Barboza et al. 3668 (BM, CORD, SI); Dtto. Utracán, Valle de Daza, rumbo a la Laguna El Loro, 10–12 km de la RP 18, 290 m, 8 Feb 2020, Barboza et al. 5073 (BM, CORD); Dtto. Toay, Reserva Parque Luro, ingreso S por ruta 35 desde General Acha, 116 m, 9 Feb 2020, Barboza et al. 5079 (BM, CORD); Bajo Lucero, cruce entre RP 11 y RP 10, 255 m, 9 Feb 2020, Barboza et al. 5094 (BM, CORD); Dtto. Atreucó, sin. loc., Mar 1960, Cano 960 (US); Dtto. Rancul, Chamaicó, 2 Mar 1984, Steibel & Troiani 7960 (CORD); Chamaicó, 2 Mar 1984, Steibel & Troiani 7963 (CORD); Dtto. Atreucó, Laguna Chillhué, 5 Apr 1984, Steibel et al. 8035 (CORD); Dtto. Capital, Barrancas Coloradas, 28 Feb 1991, Steibel 10111 (CORD); Dtto. Toay, Parque Luro, 8 Mar 1991, Steibel 10118 (CORD); Dtto. Capital, El Guanaco, 30 km al N de Santa Rosa, 13 Feb 1977, Troiani 4688 (CORD); Dtto. Atreucó, Laguna Chillhué, 13 Mar 1982, Troiani et al. 6820 (CORD); Dtto. Loventué, Luan Toro, 10 km al W, 10 Feb 1985, Troiani 8564 (CORD). La Rioja: Dtto. Famatina, Ruta 40 [now Ruta Prov. 11] (km 640/641), yendo de Famatina a Tinogasta, 20 Mar 1960, Hunziker et al. 15172 (CORD); Ruta 40 [now Ruta Prov. 11] (km 692), yendo de Famatina a Tinogasta, entre Santa Cruz y el límite con Catamarca, 20 Mar 1960, Hunziker et al. 15206 (CORD,US). San Juan: Dtto. Sarmiento, a 2 km de la estancia El Acequión, desde El Pedernal hacia el enpalme con Ruta provincial 412, por Ruta Prov. 312, 1400 m, 9 Apr 2004, Matesevach 10 C, (CORD). San Luis: Dtto. General Pedernera, Villa Mercedes, Estancia Agropecuaria INTA San Luis, lote 16, bajo, 510 m, 9 Jan 1969, Anderson & Galvani 1511 (CORD); Sierra El Morro, cuenca interior, querencia pisoteada, 1200 m, 27 Dec 1977, Anderson et al. 3427 (CORD); Dtto. Capital, Estancia Las Tres Marías, 30 km al sur de San Luis-Quemado, 700 m, 21 Mar 1979, Anderson et al. 3643 (CORD); Dtto. Chacabuco, Concarán, cerca de 2 km desde Concarán rumbo a Santa Rosa de Conlara, por el camino de tierra (RP 23), 660 m, 24 Feb 2020, Barboza et al. 5130 (BM, CORD); Concarán, cerca de 5.5 km desde Concarán rumbo a Santa Rosa de Conlara, por el camino de tierra (RP 23), 652 m, 24 Feb 2020, Barboza et al. 5136 (BM, CORD); Dtto. Capital, Potrero de los Funes, 2 Apr 1989, Del Vitto & Petenatti 3455 (CORD); Los Puquios, a 200 m del badén sobre el río Los Puquios en la ruta El Volcán-Cruz de Piedra, rumbo a Cruz de Piedra, 21 May 1972, Giordano & Guerreiro 23 (CORD); Dtto. Chacabuco, a ca. 3 km al N de Concarán, rumbo a Santa Rosa, por el viejo camino de tierra, 17 Feb 1989, Hunziker et al. 25335 (CORD); Dtto. Belgrano, Sierra del Gigante (falda O), Desaguadero, inmediaciones de Paso de Tropas, 7 Apr 1944, Ruiz Leal 9191 (CORD); Dtto. Capital, entre Estación Jarilla y Desaguadero., 500 m, 7 Apr 1944, Semper s.n. (BM, US).

Like Solanum aloysiifolium Dunal but differing in narrower leaves decurrent onto the stems, antrorse pubescence, ellipsoid buds and strongly deflexed fruiting pedicels.

Figure 7. 

Solanum tiinae Barboza & S. Knapp A habit (Barboza et al. 3491) B details of the attenuate leaf base and winged stems with antrorse trichomes (Barboza et al. 3491) C flower bud (Barboza et al. 3496) D inflorescence (Barboza et al. 3491) E flowers at anthesis, note the changing colour and size (Barboza et al. 3491) F mature fruits (Barboza et al. 3491). All photographs by S. Knapp.

Argentina. Tucumán: Dtto. Tafí del Valle, El Infiernillo, en el parador, 3042 m, 13 Feb 2012, G.E. Barboza, S. Knapp & T. Särkinen 3496 (holotype: CORD [CORD00013848]; isotypes: BM [BM001115408, BM001115409], others to be distributed).

Figure 8. 

Solanum tiinae Barboza & S. Knapp (isotype: Barboza et al. 3496, BM [BM001115409]).

Perennial herbs or subshrubs sprawling from a woody base, to 50 cm tall. Stems narrowly winged, the wing to 0.5 mm wide, often invested with spinose processes (enlarged trichome bases), sparsely pubescent with antrorse eglandular, simple uniseriate trichomes, 6–10-celled, ca. 0.5 mm long, these white when dry; new growth densely to moderately pubescent with antrorse eglandular, simple 2–8-celled uniseriate trichomes, ca. 0.5 mm long; bark of older stems pale greenish brown, glabrescent. Sympodial units plurifoliate, the leaves not geminate. Leaves simple, 2–5 cm long, 0.6–2 cm wide, narrowly elliptic to almost lanceolate in some individuals, membranous, concolorous; adaxial surfaces sparsely and evenly pubescent with antrorse eglandular simple 2–4-celled uniseriate trichomes to 0.5 mm long, the trichomes slightly longer on the veins, white when dry; abaxial surfaces with similar, but denser eglandular antrorse pubescence; principal veins 4–6 pairs, drying yellow, especially abaxially; base attenuate and decurrent onto the winged stem and the leaves sessile or nearly so; margins entire or with a few teeth ca. 2 mm long, ca. 2 mm wide with blunt tips in the lower third to half; apex acute to slightly blunt-tipped; petiole absent to 0.2 mm long, eglandular pubescent like the stems and leaves. Inflorescences 2.5–5 cm long, opposite the leaves or internodal, forked with 2 short branches, with 10–20 flowers clustered at the tips of the inflorescence branches, sparsely pubescent with antrorse eglandular simple uniseriate trichomes like those of the stems; peduncle 1.2–2.5 cm long; pedicels 0.8–1 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, strongly tapering, spreading to somewhat deflexed at anthesis, sparsely to moderately sparsely pubescent with antrorse eglandular simple uniseriate trichomes like the rest of the inflorescence, articulated at the base; pedicel scars clustered at the tips of the inflorescence branches, ca. 0.5 mm apart. Buds ellipsoid to somewhat turbinate (widest in lower third), the corolla strongly exserted from the calyx tube before anthesis, the style sometimes exserted from the bud before anthesis. Flowers 5-merous, perfect. Calyx tube 1.5–2 mm long, conical, the lobes (0.5)1–2 mm long, deltate with lanceolate tips, the sinuses rounded, sparsely pubescent with antrorse eglandular trichomes like the pedicels. Corolla 1.2–2.2 cm in diameter, white, pale violet or white tinged with violet, sometimes changing colour through anthesis, with a brownish yellow to yellow-green central star edged with brownish purple, stellate, lobed halfway to the base, the lobes 5–8 mm long, 4–5 mm wide, deltate to triangular, spreading or slightly reflexed at anthesis, adaxially glabrous, abaxially densely pubescent with eglandular papillae and simple uniseriate trichomes to 0.2 mm long. Stamens equal; filament tube minute; free portion of the filaments 0.5–1 mm long, adaxially densely pubescent with tangled transparent simple uniseriate trichomes; anthers 4–5 mm long, 1–1.25 mm wide, ellipsoid, yellow, the abaxial surfaces occasionally papillate, poricidal at the tips, the pores lengthening to slits with age. Ovary conical, glabrous; style 7–10 mm long, pubescent along almost the entire length, more densely in the lower half with tangled transparent simple trichomes to 0.5 mm long; stigma capitate to clavate, bright green in live plants, the surface minutely papillose. Fruit a globose berry, 0.8–0.9 cm in diameter, green with tiny white spots (immature?), opaque, the pericarp surface thin, matte, glabrous; fruiting pedicels 0.8–1 cm long, ca. 0.75 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, thickened but not woody, strongly deflexed with a distinct bend at the pedicel base; fruiting calyx not enlarged or accrescent, the lobes appressed to the surface of the berry. Seeds 10–30 per berry, 1.7–2 mm long, 1–1.5 mm wide, not markedly flattened, teardrop shaped with an apical hilum, pale tan, the surfaces minutely pitted, the testal cells sinuate in outline. Stone cells 4–9 per berry, 0.7–1.5 mm in diameter, 2 usually larger than the rest. Chromosome number: n=12 (Moscone 1992, as S. lorentzii Bitter var. montigenum C.V.Morton).

(Figure 9). Solanum tiinae is endemic to Argentina; it has been collected from the provinces of Jujuy, La Rioja, Salta and Tucumán, with most collections from the area around the type locality at El Infiernillo.

Solanum tiinae grows among rocks and in open areas in pre-puna habitats in the Andes (Fig. 7A), from 2400 to 4000 m elevation.

Solanum tiinae is named in honour of our long-term collaborator and colleague Dr. Tiina Särkinen of the Royal Botanic Garden Edinburgh; she was the first to notice the uniqueness of these plants, giving them the field name “Solanum misterioso” while in the field in 2012.

(IUCN 2019). AOO (76 km² – EN); EOO (41,143 km² – NT). Most collections of S. tiinae are from a very few commonly visited localities and the main road between Tafí del Valle and Amaicha del Valle in the province of Tucumán. Solanum tiinae is not found in protected areas, and based on the number of localities (ca. 5), the area of occupancy and the extent of occurrence, we assign a preliminary threat status of Vulnerable (VU B2a,biii). Where it occurs S. tiinae is not common or weedy, although it does grow in open areas.

It is surprising that S. tiinae has not been described previously, as the area from which the type and many other collections come is one of the most intensively collected Andean areas in Argentina. None of the collections we cite here were cited in Morton (1976). We have encountered specimens of S. tiinae identified as S. aloysiifolium (and its synonyms, see Barboza et al. 2013) and S. cochabambense. It is similar to those taxa in its forked inflorescence with a long peduncle, but differs from S. aloysiifolium in its larger, less deeply stellate purple or purplish cream (rather than white) corollas, and from S. cochabambense in its smaller habit and winged stems. The strongly antrorse pubescence of S. tiinae is distinctive and not found in either S. aloysiifolium or S. cochabambense.

Solanum tiinae also resembles the highly variable species S. salicifolium, from which it can be distinguished by its shorter (1–2 mm versus 2.5–3 mm long) calyx lobes, the strongly antrorse pubescence (Fig. 7B), the strictly furcate (versus only occasionally once branched) inflorescences with more flowers (10–20 versus 4–10) (Fig. 7B, C) and the calyx lobes (Fig. 7F) that are tightly appressed to the berry (versus spreading and slightly recurved). These two taxa have been collected in the same habitat (e.g., Barboza et al. 3491, S. tiinae and Barboza et al. 3494, S. salicifolium from km 92 on the Amaicha del Valle to Tafí del Valle road) and can be easily distinguished in the field using corolla shape – those of S. salicifolium are deeply stellate with relatively narrow lobes, while those of S. tiinae are less deeply and more broadly lobed (Fig. 7E).

(paratypes). Argentina. Jujuy: Dtto. Tilcara, Sierra de Zenta, 4000 m, Feb 1931, Budin 7471 (CORD). La Rioja: Dtto. Famatina, Rodeo de las Vacas, 3000–4000 m, Feb 1913, Flossdorf 55 (SI); Quebrada Encrucijada, 3500–5000 m, Mar 1913, Flossdorf 56 (SI). Salta: Dtto. San Carlos, Amblayo, 2371 m, 16 Mar 1943, Hunziker 2623 (CORD); Dtto. Cachi, Ruta Prov. 33, de Piedra del Molino a El Carril, La Herradura, 3110 m, 26 Feb 2009, Zuloaga et al. 11256 (CORD, SI). Tucumán: Dtto. Tafí del Valle, Pinar de los Ciervos, Km 70, 2400 m, 6 Mar 1998, Barboza et al. 139 (CORD); Dtto. Tafí del Valle, Pinar de Los Ciervos, km 70, 2400 m, 6 Mar 1998, Barboza et al. 140 (CORD); Dtto. Tafí del Valle, entre Tafì del Valle y Amaichá: Km 76, 6 Mar 1998, Barboza et al. 150 (CORD); Dtto. Tafí del Valle, entre Tafì del Valle y Amaichá: Km 76, 6 Mar 1998, Barboza et al. 151 (CORD); Dtto. Tafí del Valle, entre Tafí del Valle y Amaicha, 6 Mar 1998, Barboza et al. 152 (CORD); Dtto. Tafí del Valle, El Infiernillo, 2920 m, 19 Mar 2006, Barboza et al. 1705 (CORD); Dtto. Tafí del Valle, Carapunco, rumbo a Amaicha del Valle por RP307, 2864 m, 24 Feb 2009, Barboza et al. 2167 (CORD); El Infiernillo, 2960 m, 24 Feb 2009, Barboza et al. 2172 (CORD); Dtto. Tafí del Valle, La Quebradita, a unos pocos km de Tafí del valle rumbo a Amaicha del Valle, 2053 m, 21 Feb 2011, Barboza et al. 3014 (CORD); Dtto. Tafí del Valle, a 28 km de Tafí rumbo a Amaicha del Valle, 2857 m, 21 Feb 2011, Barboza et al. 3019 (CORD); Dtto. Tafí del Valle, desde Amaicha del Valle rumbo a Tafí del Valle, entre km 92–91, 3000 m, 13 Feb 2012, Barboza et al. 3491 (BM, CORD); Dtto. Tafí del Valle, entre Tafì del Valle y Amaichá, El Infiernillo, 19 Feb 1988, Cocucci et al. 293 (CORD); Tafí del Valle, 2500 m, 24 Feb 1998, Cocucci 989 (CORD); Dtto. Tafí del Valle, Km. 82, al N de Tafí del Valle, hacia la quebrada del Barón, 3100 m, 26 Feb 1959, Diers 285 (SI); Dtto. Tafí del Valle, El Molle, en el camino entre Tafí del Valle y Amaicha, km 91–92, 2800–2900 m, 12 Feb 1986, Hunziker et al. 24878 (BM, CORD [x2], E, MO); Dtto. Tafí del Valle, El Molle, en el camino entre Tafí del Valle y Amaicha, km 91–92, 2800–2900 m, 12 Feb 1986, Hunziker et al. 24879 (CORD); Dtto. Tafí del Valle, viniendo desde Tafí del Valle, rumbo a Amaicha del Valle, entre kms 75 y 76, 2600- 2700 m, 14 Dec 1995, Hunziker et al. 25546 (CORD); Dtto. Tafí del Valle, El Infiernillo, 22 km de Tafí del Valle, 2950 m, 18 Mar 1972, Krapovickas et al. 21885 (CTES); Tafí, 2000 m, 4 Dec 1908, Lillo 8691 (CORD, LIL, SI, US); Dtto. Tafí del Valle, La Ciénaga; Sierra de Tucuman, 10 Jan 1874, Lorentz & Niederlein 565 (CORD); Dtto. Tafí del Valle, 16 km N de Tafí del Valle, Mojon, K [km] 78, 16 Mar 1972, Maruñak et al.209 (CTES); Infiernillo, Tafí del Valle, 1850 m, 2 Mar 1972, Meyer s.n. (LIL); Dtto. Tafí del Valle, Ruta 40, ca. 10 km NW de Tafí del Valle, Cumbres Calchaquíes, 2985 m, 26 Jan 2007, Paula-Souza et al. 7912 (CTES); Dtto. Tafí del Valle, Infiernillo, 3040 m, 8 Mar 1955, de la Sota 236 (CORD); Dtto. Tafí del Valle, Cumbres Calchaquí, Quebrada Honda, 3100 m, 23 Jan 1952, Sparre 9232 (CORD); Dtto. Tafí del Valle, Cumbres Calchaquí, Quebrada Honda, 3100 m, 23 Jan 1952, Sparre 9233 (CORD); Dtto. Tafí del Valle, Tafi del Valle, 4 Dec 1960, Subils & Articó 285 (BM, CORD); Dtto. Tafí del Valle, Colalao del Valle, alrededores, 17 Feb 1979, Subils & Bernardello 2670 (CORD); Dtto. Tafí del Valle, pasando Tafí del Valle rumbo a Amaicha, Carapunco, 2942 m, 1 Apr 2012, Urdampilleta et al. 760 (CORD); Dtto. Tafí del Valle, Tafí del Valle, 2900 m, 29 Jul 1971, Without Collector s.n. (BAA); Dtto. Tafí del Valle, El Infiernillo, RP 307, 2995 m, 18 Mar 2018, Zuloaga et al. 16415 (SI).

Figure 9. 

Distribution of Solanum tiinae Barboza & S. Knapp.