Solanum cochabambense Bitter, Repert. Spec. Nov. Regni Veg. 10: 553. 1912.

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

12. Solanum cochabambense Bitter, Repert. Spec. Nov. Regni Veg. 10: 553. 1912.

Figs 3G, H , 4F , 38 , 39

Solanum extuspellitum Bitter, Repert. Spec. Nov. Regni Veg. 10: 555. 1912. Type. Bolivia. Tarija, 2,300 m, 30 Dec 1903, K. Fiebrig 2439 (holotype: B, destroyed [F neg. 2711]; lectotype, designated here: F [V0361919F, acc. # 621247]).

Solanum extuspellitum Bitter subsp. subcoeruleum Bitter, Repert. Spec. Nov. Regni Veg. 10: 556. 1912. Type. Bolivia. Tarija, 2,300 m, 30 Dec 1903, K. Fiebrig 2439 (holotype: B, destroyed [F neg. 2711]; lectotype, designated here: F [v0361919F, acc. # 621247]).

Solanum lorentzii Bitter var. tucumanicum Bitter, Repert. Spec. Nov. Regni Veg. 10: 556. 1912. Type. Argentina. Tucumán: sin. loc., P.G. Lorentz & G. Hieronymus 1155 (holotype: B, destroyed; lectotype, designated by Barboza et al. 2103, pg. 236: CORD [CORD00004238]; isotypes: CORD [CORD00004239, CORD00004240], F [v0073320F, acc. # 50929], K [K000585687], SI [SI003323]).

Solanum decachondrum Bitter, Repert. Spec. Nov. Regni Veg. 11: 228. 1912. Type. Bolivia. Cochabamba: Cercado, May 1909, O. Buchtien 2411 (lectotype, designated here: US [00027539, acc. # 700102]; isolectotypes: US [01014170, acc. # 1175973]).

Solanum decachondrum Bitter var. latiusculum Bitter, Repert. Spec. Nov. Regni Veg. 11: 229. 1912. Type. Bolivia. Cochabamba: Cercado, May 1909, O. Buchtien 2412 (lectotype, designated here: US [00027538, acc. # 1177823]; isolectotypes: GOET [GOET009219], NY [00139124]).

Solanum decachondrum Bitter var. longiusculum Bitter, Repert. Spec. Nov. Regni Veg. 11: 229. 1912. Type. Bolivia. Cochabamba: Cercado, May 1909, O. Buchtien 2411 (lectotype, designated here: US [01014170, acc. # 1175973]; isolectotype: US [00027539, acc. # 700102]).

Solanum probolospermum Bitter, Bot. Jahrb. Syst. 54, Beibl. 119: 10. 1916. Type. Peru. Huánuco: Valle del Río Pozuzo encima de Saria, 22 Jul 1913, A. Weberbauer 6789 (no herbaria cited; lectotype, designated here: MOL[MOL00005139]; isolectotypes: B, destroyed [F neg. 2682], F [v0043286F, acc. # 647965], GH [01011893], MOL [MOL00005138], US [00027756, acc. # 1444969]).

Solanum lorentzii Bitter var. montigenum C.V.Morton, Revis. Argentine Sp. Solanum 136. 1976. Type. Argentina. Tucumán: Dpto. Chicligasta: Estancia Santa Rosa, 8 Jan 1927, S. Venturi 4760 (holotype: US [03271889, acc. # 1548937]; isotypes: F [v0073318F, acc. # 695929; v0073319F, acc. # 637505], LP [LP010202, acc. # 010393], MO [MO-2127157, acc. # 960405] S [acc. # R-3117], SI [003322]).

Solanum montigenum (C.V.Morton) Cabrera, Fl. Prov. Jujuy 8: 435. 1983. Type. Based on Solanum lorentzii Bitter var. montigenum C.V.Morton.

Type.

Bolivia. Cochabamba: Vic. Cochabamba, 1891, M. Bang 1151 (lectotype, designated by Barboza et al. 2013, pg. 236: NY [00139097]; isolectotypes: BM [BM000617675], BR [BR0000005538553], CAL [acc.# 316673], E [E00190740], G [G00343347], GH [00077599], MO [MO-503629, acc. # 1815484], NY [00139096], PH [00030399], US [00610905, acc. # 92001; 00027515, acc. # 1324496], WIS [0256183WIS]).

Description.

Lax subwoody or woody shrubs, often vine-like with very long stems, to 5 m long, to 3 m if erect. Stems erect or sprawling, terete or slightly angled with tiny spinescent processes along the angles, moderately pubescent with eglandular white simple uniseriate 2-6-celled trichomes to 1 mm long, these soft and spreading; new growth densely white pubescent with eglandular simple uniseriate trichomes like those of the stems; bark of older stems pale brown, glabrescent. Sympodial units difoliate or plurifoliate, the leaves not geminate. Leaves simple or occasionally shallowly toothed, the blades 3.5-16 cm long, 1.5-8 cm wide, variable within an individual plant and always larger on lower stems, elliptic to narrowly elliptic, widest in the lower half, membranous, discolorous; adaxial surfaces sparsely pubescent with soft, spreading, eglandular simple uniseriate trichomes to 1 mm long, like those of the stems, these denser on the veins; abaxial surfaces more densely pubescent with simple uniseriate trichomes, the lamina still visible; principal veins 7-9 pairs, densely pubescent on abaxial surfaces; base acute, somewhat attenuate onto the petiole; margins entire or rarely shallowly toothed, the teeth if present in the basal part of the leaf, ca. 1 mm long, ca. 1.5 mm wide, with acute apices (see Brooke 5125, one duplicate entire, one toothed); apex acute to somewhat acuminate; petiole 0.5-2.8 cm long, slightly winged from the decurrent leaf bases in the distal part. Inflorescences internodal or terminating branches, several times branched, 3-13 cm long, with 10-80+ flowers clustered at the branch tips, moderately pubescent with soft, spreading eglandular simple uniseriate trichomes to 1 mm long like those of the stems; peduncle 1.7-10 cm long; pedicels 0.6-1 cm long, 0.5-0.75 mm in diameter at the base, 1-1.5 mm in diameter at the apex, tapering, spreading at anthesis, moderately pubescent like the inflorescence axes, articulated at the base; pedicel scars 0.5-1 mm part at the branched tips. Buds ellipsoid, occasionally somewhat inflated, the corolla strongly exserted from the calyx before anthesis. Flowers 5-merous, cosexual (hermaphroditic). Calyx tube 1-1.5 mm long, conical, the lobes 1.2-2 mm long, 1-1.5 mm wide, narrowly deltate, moderately pubescent with simple uniseriate trichomes like the rest of the plant. Corolla 2-3 cm in diameter, extremely variable through anthesis in size and colour, pale violet to whitish violet, with a pale greenish yellow eye, stellate, lobed 1/3 to 1/2 of the way to the base, the lobes 4-6 mm long, 4-5 mm wide, deltate or broadly deltate, spreading to slightly reflexed at anthesis, glabrous adaxially or with scattered uniseriate trichomes ca. 0.2 mm long at the tips and margins, abaxially densely papillate-puberulent with papillae and simple uniseriate 1-3-celled trichomes to 0.5 mm long along the lobe midveins, tips and margins, the interpetalar tissue glabrous. Stamens equal; filament tube to 0.25 mm long; free portion of the filaments 1-1.5 mm long, pubescent adaxially with densely tangled, transparent weak simple uniseriate trichomes; anthers 3.5-4.5 mm long, 1.2-1.5 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary conical, glabrous; style 9-10 mm long, straight at anthesis (curved in bud), exserted beyond the anther cone, densely pubescent with transparent simple uniseriate trichomes in the lower half; stigma clavate, somewhat bilobed or capitate, green or dark cream in live plants, the surface minutely papillate. Fruit a globose berry, (0.9)1-1.2 cm in diameter, green and usually maturing purplish black, the pericarp thin, matte, translucent when berry ripe, glabrous; fruiting pedicels 1-1.7 cm long, ca. 1.2 mm in diameter at the base, ca. 2 mm in diameter at the apex, slightly woody, deflexed or spreading, not persistent; fruiting calyx somewhat enlarged, the tube to 2 mm long, the lobes to 2 mm long, appressed to the berry. Seeds 30-50 per berry, 1.5-2 mm long, 1-1.5 mm wide, flattened and teardrop shaped, pale brown to golden tan, the surfaces minutely pitted, the testal cells elongate with sinuate walls. Stone cells 8-12 per berry, 0.5-1 mm in diameter, cream-coloured, distributed throughout the mesocarp. Chromosome number: n = 12 ( Edmonds 1972, voucher Steinbach 34, as S. polytrichostylum var. lorentzii ; Moscone 1992, voucher Subils et al. 3253, as S. aloysiifolium ; Moyetta et al. 2013, voucher Barboza et al. 2152, as S. aloysiifolium ).

Distribution

(Fig. 40 ). Solanum cochabambense occurs from the eastern Andean slopes from northern Peru (Depts. Amazonas, Ancash, Apurímac, Arequipa, Cajamarca, Cusco, Huancavelica, Huánuco, Junín, La Libertad, Puno, San Martín) throughout the Andean cordillera to Bolivia (Depts. Chuquisaca, Cochabamba, La Paz, Oruro, Potosí, Santa Cruz, Tarija) and northern Argentina (Provs. Jujuy, Salta, Tucumán).

Ecology and habitat.

Solanum cochabambense grows in a wide variety of middle to high elevation forest types, often at roadsides or in landslips and treefalls, from 150 to 4,120 m; most collections are from elevations above 1,000 m. The single collection from low elevation (Roque 295 from 150 m in Camaná, Arequipa, Peru) comes from an area where landslides ( ‘huaicos’) are common and perhaps represents seeds washed down from higher elevations.

Common names and uses.

Bolivia. La Paz: chinchi-chinchi (Beck 27781), cusmayo (Lewis 881659). Peru. Ancash: atoqpa papán (papa de zorro) (Gamarra 662); Cusco: ccaya-ccaya (Mexia 8079), chinchi-chinchi (Herrera 819); muya khaya (Franquemont et al. 297); qusmayllu (Franquemont et al. 348); Huánuco: shopta (Weberbauer 6789); Puno: chitinqoya ( Roersch 1994). In the southern Peruvian Quechua community of Chinchero (Cusco, Peru) leaves are used as cattle forage ( Franquemont et al. 1990, as S. aloysiifolium ) and as a wash for the head, especially for hangovers ( Franquemont et al. 1990, as. S. glandulosipilosum ). In southern Peru more generally leaves of S. cochabambense are used medicinally in a tea in the treatment of flu and colds ( resfrío) and to counter difficulty in urination, and as a macerated plaster to alleviate rheumatic pains ( Roersch 1994, as S. aloysiifolium ).

Preliminary conservation status

( IUCN 2022). Least Concern [LC]. EOO = 7,244,968 km2 [LC]; AOO = 1,132 km2 [VU]. Solanum cochabambense is a common plant of disturbed areas. Further study may reveal variation that warrants taxonomic distinction, and this preliminary assessment will need revisiting. Solanum cochabambense occurs within several protected areas across its range (see Supplementary materials).

Discussion.

Solanum cochabambense is one of the most variable and widespread morelloid species in South America. Barboza et al. (2013) placed S. cochabambense in synonymy with S. aloysiifolium , with which it is sympatric in northern Argentina. Further study throughout the range of S. cochabambense confirmed the distinctness of the two species, but individual specimens collected in sympatry can be difficult to identify. Solanum cochabambense differs from S. aloysiifolium in its more highly branched inflorescences (those of S. aloysiifolium are usually only forked), buds that are ellipsoid rather than narrowly ellipsoid and larger corollas and berries. The anthers of S. aloysiifolium are narrow relative to their length (3.9-5 mm long and 0.6-1 mm wide in S. aloysiifolium versus 3.5-4 mm long and 0.9-1.2 mm wide in S. cochabambense ) but this character can be difficult to see in the absence of comparative material. The berries of S. cochabambense are larger (1-1.2 cm in diameter) than those of S. aloysiifolium (0.5-0.6 cm in diameter), with similar numbers of stone cells.

In Bolivia S. cochabambense is partially sympatric with and morphologically very similar to S. pallidum . Solanum pallidum differs in its possession of dendritic trichomes, while S. cochabambense has only simple trichomes.

In the northern part of its range, S. cochabambense can be confused with S. arequipense , S. juninense and S. interandinum . Solanum juninense differs in its possession of glandular trichomes whereas S. cochabambense is always eglandular. Solanum arequipense has blunt-tipped calyx lobes, anthers 2.5-3 mm long and a strongly capitate stigma, while S. cochabambense has long-triangular calyx lobes with acute apices, anthers 3.5-4 mm long and a clavate to only somewhat capitate stigma. The calyx lobes of S. interandinum are longer and more pointed than those of S. cochabambense , and the flowers are smaller (0.8-1.4(1.8) cm in diameter versus 2-3 cm in diameter in S. cochabambense ).

The extreme variability seen across the range of S. cochabambense may indicate there are several distinct species contained within our rather broad circumscription. In some cases, duplicate collections from the same locality show that variation is present within a single population, which has helped us to recognise this group of specimens as a morphologically variable single species: an example of such variation is leaf margins varying from entire to toothed in duplicates of Brooke 5125. Similarly, variation in corolla shape and size was evident in the field in some populations, as well as inflorescence structure (e.g., Knapp et al. 10391, Knapp et al. 10392, Knapp et al. 10393, Knapp et al. 10669). Variation in other characters such as indumentum, calyx lobe shape and size, and other characters may represent fixed differences between populations, but based on our study of the specimens available across geographic space, we circumscribe this as a single highly variable species. Future studies at the population level throughout the range will be important to identify potential taxonomically recognisable segregates in this species.

Bitter (1912a) described S. extuspellitum and its variety subcoerulum using the same collection (Fiebrig 2439), citing "p. pt. herb. Berol.!" in each protologue. The sheet in B (now destroyed but photographed as F neg. 2711) has two stems, one with the label " Solanum extuspellitum n. sp." and the other with the label "Solanum (extuspellitum subsp.) subcoeruleum Bitt." - this latter suggesting he had originally considered naming the latter at the specific level. The stems differ only in pubescence density, and both are referrable to S. cochabambense . It is impossible to tell from which stem the fragment held in F came, so we are using it as the lectotype for both names. We have found no other duplicates of Fiebrig 2439.

Later that same year ( Bitter 1912c), he described S. decachondrum and its varieties Solanum decachondrum longiusculum and Solanum decachondrum latiusculum using two collections of Otto Buchtien (Buchtien 2411 and 2412), citing no herbarium but indicating with “!” that he had seen them. Otto Buchtien’s private herbarium that Bitter cited was donated to the Smithsonian (US) in the 1920s ( Morton and Stern 1966), so lectotypes for names based on Buchtien’s collections should be in US. Bitter cited both numbers in the protologue of the species, then used Buchtien 2411 for var. Solanum decachondrum longiusculum and Buchtien 2412 for var. Solanum decachondrum latiusculum . None of the duplicates of these collections we have seen has annotations in Bitter’s hand, but one duplicate in US (01014170, acc. # 1175973) is annotated as var. Solanum decachondrum longuisculum by Buchtien and is here designated the lectotype of var. Solanum decachondrum longuisculum . The other duplicate of Buchtien 2411 at US (00027539, acc. # 700102) is designated as the lectotype of S. decachondrum . We lectotypify var. Solanum decachondrum latiusculum with the US duplicate (000275538, acc. # 1177823) of Buchtien 2412, as it is the best preserved with both flowers and fruits and is annotated as var. Solanum decachondrum latiusculum in Bitter’s hand.

Bitter (1916) described S. probolospermum citing Weberbauer 6789 but without citing a herbarium. Many duplicates of this collection number have been preserved, and we select the better preserved of the two duplicates of Weberbauer 6789 in the herbarium of the Universidad Nacional Agraria La Molina (MOL00005139) as the lectotype. Weberbauer’s original personal herbarium is held in MOL.

Figures

Figure 3. Representative flowers of South American morelloids A campanulate purple corollas in forked internodal inflorescences in S. fiebrigii B pentagonal white corollas in S. annuum C broadly stellate purple corollas with long-exerted styles in S. pentlandii D broadly stellate white corollas with slightly exerted globose stigmas in S. radicans E deeply stellate pale yellow corollas in S. huayavillense F deeply stellate purple corollas with dark central eye colouration in S. salicifolium G bumblebee visiting and buzzing the anther cone in S. cochabambense H variation in corolla shape, size and colouration within a single individual of S. cochabambense (A Barboza et al. 3548 B Barboza et al. 3495 C Knapp et al. 10248 D Knapp et al. 10277 E Barboza et al. 3531 F Chiarini et al. 819 G Saerkinen et al. 4036 H Knapp et al. 10669). Photos by S. Knapp and T. Saerkinen.

Figure 4. Representative fruits, seeds and stone cells of South American morelloids A bilobed fruits in S. tripartitum with fully mature red berries at the base of the inflorescence and maturing green, yellow, and orange fruits more distally B orange-red fleshy berries of S. corymbosum in highly branched inflorescences C fully mature marbled fruits in S. physalifolium D immature ellipsoid fruits of S. antisuyo E fully mature fruits of S. antisuyo F immature green fruits amongst fully mature purple-black fruits in S. cochabambense G fully mature fruits of S. polytrichostylum H stone cells (also known as sclerotic granules or brachysclerids) found in the fruits of most species of the Morelloid clade (left side of photo) next to the teardrop shaped seeds (right side of photo; S. umalilaense Manoko) I stone cells visible in dried fruits of herbarium specimens (S. triflorum). (A Barboza 3563 B Saerkinen et al. 4604 B C Knapp et al. 10334 D Gonzales 10256 E Gonzales 10256 F Knapp et al. 10363 G Saerkinen et al. 5277 H Nijmegen accession A 24750133 I Podlech 8624 BM 000848286). Photos by P. Gonzales, S. Knapp, and T. Saerkinen.

Figure 38. Solanum cochabambense A flowering branch B detail of adaxial leaf surface C detail of abaxial leaf surface D flower bud E dissected flower F fruiting branch G maturing fruit (A-D Knapp et al. 10341; E Cardenas 5577 F, G Knapp et al. 10339). Illustration by R. Wise.

Figure 39. Solanum cochabambense A habit B inflorescence with purple corollas C infructescence with white corollas D mature fruits (A, C Knapp et al. 10363 B Knapp 10287 D Barboza et al. 3500). Photos by S. Knapp.

Figure 40. Distribution map of Solanum cochabambense.