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Regni Veg. Type species: Solanum decorticans Sendtn. Solanum section Lysiphellos Bitter Seithe, Bot. Solanum section Jasminosolanum Seithe, Bot. Solanum subsection Nitidum A. Child, Feddes Repert. Solanum section Californisolanum A. Vines or lax shrubs, woody at the base; stems weak and clambering or erect, sometimes winged, pubescent or glabrous, the trichomes unbranched or branched, never strictly stellate.

Sympodial units plurifoliate. Leaves simple or pinnatifid, sometimes deeply so such that the leaves appear pinnate, green, glabrous or variously pubescent with branched or unbranched trichomes, the branched trichomes sometimes very compact and echinoid multangulate ; petioles well developed, in vines often twining to aid in climbing. Inflorescences terminal, sometimes appearing lateral due to stem growth, simple or more usually many times branched, with up to flowers, not bracteate; peduncle present, often poorly distinguished from the leafy stem; pedicels inserted in a short sleeve of inflorescence tissue, articulated at the base inside the sleeve or just beneath the calyx Solanum boldoense.

Flowers 5-merous, actinomorphic or slightly zygomorphic, usually perfect, one species probably dioecious Solanum luculentum ; calyx 5-parted, glabrous or pubescent; corolla 5-parted, rotate to stellate, white to purple, sometimes with spots at the base of the lobes; stamens 5, the filaments equal or unequal, glabrous or pubescent; anthers yellow or occasionally purple, ellipsoid or tapering, separate to tightly connivent, dehiscing by terminal pores, these usually elongating to slits with age; ovary globose to conical, usually glabrous, occasionally pubescent; style straight or slightly curved; stigma capitate to clavate.

Fruit a globose berry, yellowish green to red, orange or black when ripe, glabrous, the pericarp thin, shiny or matte; calyx lobes in fruit not accrescent. Seeds flattened, often appearing hairy from outgrowths of the lateral cell walls. Most members of the clade possess a combination of other characters that, although occurring elsewhere in Solanum , together distinguish the group; these are, 1 vining habit, 2 twining petioles, 3 large, terminal, branched inflorescences, 4 brightly colored fruits, 5 pinnatifid leaves at least in juvenile plants.

In the absence of a well-sampled phylogenetic study see above , I have divided the Dulcamaroid clade into a series of morphologically and geographically delimited groups see Table 4. These should not be interpreted as monophyletic groups or taxa, and future phylogenetic analyses including more species will certainly reveal structure within this large and variable clade. Preliminary phylogenetic analyses with a number of plastid regions T. I have included them here on gross morphological grounds whilst realising their evolutionary relationships may be with other taxa.

Species are presented here in alphabetical order without regard to putative evolutionary relationships. Informal species groups in the Dulcamaroid clade. For country distribution see Table 2 and individual species descriptions. In the species treatments here I have cited all type specimens I have seen with barcodes or accession numbers if possible see Materials and Methods for citation style.

Those specimens I have not seen but whose existence is known from the literature or from others are indicated as n. Some of the infraspecific taxa, particularly in the widespread European species Solanum dulcamara were described from floristic accounts and did not have specimens cited in their original descriptions; I have not neotypified these.

Hispania. Volume 73, Number 2, May | Biblioteca Virtual Miguel de Cervantes

This synoptic set of observations is intended to simplify the task of identifying a member of this large and highly variable group. Sterile plants of members of the Dulcamaroid clade are often difficult to identify, and even plants with either flowers or fruit can be difficult. I have included many leaf and whole plant characters so that sterile plants can be in many cases identified to a choice of a few taxa. Once this step has been done the user can read the descriptions and match distributions considering other characters not used in this list to the plant in hand.

In the following list, character states are followed by a list of species epithets in alphabetical order. Epithets in parentheses indicate that the state is relatively uncommon in that species.

Species that vary in a particular character, for example in leaf trichome distribution, will be listed for each state present in that species. Not all relevant characters are considered; for example, I have listed tapering and globose anthers, but not ellipsoid anthers — common in most species in the group. In this way diagnostic states can be easily seen. A question mark? Many species of the group are rather narrow endemics, occurring in only one country or region. Table 2 lists species recorded for each country of the Neotropics; this list must be used with care, as new collecting is constantly revealing range extensions.

Shrubs or small trees: aligerum , angustifidum , coalitum , crispum , cutervanum , endoadenium , imbaburense , leiophyllum , macbridei , muenscheri , nitidum , pubigerum , ruizii , salicifolium , stenophyllum , storkii , umbelliferum , valdiviense , viscosissimum , wallacei. Stems winged or strongly angled: aligerum , amygdalifolium , valdiviense , salicifolium. Stems warty or knobbly from persistent leaf bases: coalitum , dichroandrum , endoadenium.

Stems and leaves with golden trichomes: aureum , ruizii , cutervanum , stenophyllum , storkii. Glandular trichomes present anywhere on the plant incl. Dendritic deer-antler-like trichomes present: agnoston , aligerum , aureum , calileguae , coalitum , crispum , dichroandrum , imbaburense , kulliwaita , leiophyllum , macbridei , muenscheri , nitidum , sanchez-vegae , stenophyllum , umbelliferum , uncinellum , valdiviense. Most leaves pinnatifid or lobed on mature reproductive stems: alphonsei , angustifidum , calileguae , lyratum , seaforthianum , septemlobum , salicifolium , triquetrum , uncinellum , umbelliferum , viscosissimum.

Leaves coriaceous and shiny: agnoston , coalitum , imbaburense , inodorum , leiophyllum , luculentum , macbridei , odoriferum , sanchez-vegae. Venation not visible on upper surface: agnoston , inodorum , luculentum. Lower surface of leaves with tufts of trichomes in vein axils: agnoston , aligerum , flaccidum , laxum. Both leaf surfaces densely pubescent with simple never branched trichomes: aspersum , endoadenium , umbelliferum , wallacei.

Inflorescences simple: umbelliferum , salicifolium , valdiviense. Flower buds inflated: boldoense , dulcamaroides , seaforthianum , laxum. Flowers with shiny green spots at the base of the corolla lobes: dulcamara , endoadenium , lyratum , pittosporifolium , septemlobum , umbelliferum , wallacei. Calyx lobes long-triangular or with long tips: aligerum , alphonsei , crispum , imbaburense , kulliwaita , leiophyllum , muenscheri , nitidum , ruizii , salicifolium , stenophyllum , triquetrum. Calyx lobes square quadrate : aligerum , pyrifolium , valdiviense.

Calyx lobes lacking mere enations of the calyx rim : boldoense , dulcamaroides , sousae. Filaments unequal: angustifidum , flaccidum, seaforthianum , sousae , uncinellum. Mature berries red: boldoense , crispum , dulcamara , dulcamaroides , inodorum , lyratum , nitidum , pittosporifolium , pubigerum , salicifolium , seaforthianum , septemlobum , triquetrum , uncinellum , valdiviense. Mature berries orange or yellowish green: aspersum? Mature berries black or purple: aligerum , amygdalifolium , angustifidum , aureum , coalitum , cutervanum , dichroandrum , flaccidum , imbaburense , kulliwaita , laxum , leiophyllum , macbridei , muenscheri , nitidum immature , odoriferum , pyrifolium , ruizii , sanchez-vegae , sousae?

North America and Mexico native or naturalised, not in cultivation : aligerum , dulcamara , dulcamaroides , pubigerum , sousae , triquetrum , umbelliferum , wallacei. Central America: aligerum , dulcamaroides , muenscheri , pubigerum , seaforthianum , storkii. SE Brazil: flaccidum , inodorum , laxum , odoriferum , viscosissimum.

Above treeline in Andes: aureum , coalitum , imbaburense , leiophyllum , macbridei , nitidum , stenophyllum. Eurasia: dulcamara , lyratum , pittosporifolium , septemlobum. Cultivated for ornament in both New and Old World often escaped : laxum , seaforthianum. Figure 8. Solanum agnoston S.

A, B drawn from Holm-Nielsen et al. Differs from Solanum sanchez-vegae S. Knapp in bearing tufts of dendritic trichomes in the vein axils of leaf undersides and elongate, densely pubescent buds; differs from Solanum laxum Spreng. Jaramillo et al. Bark of older stems pale greyish brown, glabrous. Leaves simple, 2—6. Buds narrowly ellipsoid, the corolla strongly exserted from from calyx before anthesis. Flowers all perfect, 5-merous. Calyx tube 1—1. Corolla known only from buds, ca.

Filaments with the tube ca. Ovary glabrous; style and stigma not seen. Fruit a globose berry, ca. Seeds ca. Chromosome number not known. Figure 9. Growing in dry scrub and high elevation forests in interAndean valleys at near m elevation. Data deficient DD ; known from only two specimens, assessment not possible. Solanum agnoston is known from only two localities, both along the main highway from Loja to Cuenca in southern Ecuador.

It superficially resembles Solanum laxum , a species of southern Brazil that is widely cultivated in tropical and subtropical environments, but differs from it in the dendritic trichomes of the lower leaf surfaces and the elliptic, pubescent buds. The leaves of the paratype specimen Holm-Nielson et al. Label data on both specimens indicate the flowers are violet, but both duplicates of Jaramillo et al. Holm-Nielson et al. Figure Type: Bolivia.

La Paz: Prov. Larecaja, Sorata, m, May , G. Solanum manicatum Bitter, Bot. Type: Peru. Ayacucho: Prov. Weberbauer holotype: B [F neg. Solanum dotanum C. Field Mus. Type: Costa Rica. Standley holotype: US [US]. Solanum grossum C. Morton, Revis. Argentine Sp. Solanum Type: Argentina.

Schiede s. Small tree or shrub, 1. Stems glabrous to densely pubescent with simple or dendritic uniseriate trichomes 0. Bark of older stems dark reddish brown, shiny. Leaves simple, 3. Inflorescences terminal or lateral, 4—15 cm long, many times branched, with 10—60 flowers, glabrous to pubescent, the trichomes simple or dendritic, ca. Buds pointed ovoid or turbinate with an apical nipple, the corolla strongly exserted from the calyx tube before anthesis.

Calyx tube 1.

Corolla 1. Filament tube minute, the free portion of the filaments 0. Ovary glabrous; style 6. Fruit a globose berry, 1—1. Seeds 20—40 per berry, 2. Chromosome number: not known. Common from Mexico to Argentina, from — m. A gap in the distribution of Solanum aligerum occurs from Panama to Peru see discussion.

Solanum aligerum has been collected most commonly in pine-oak forests in Mexico and adjacent Central America, and in cloud forests in the Andes. See Moat for explanation of measurements. Solanum aligerum has the widest latitudinal range of all the New World members of the Dulcamaroid clade, occurring all down the cordilleras of both continents, but with a distinct gap between western Panama and Peru see below.

In spite of this huge range, the species is remarkably uniform in general aspect, but varies considerably in pubescence. In Bolivia, several populations have been found with plants that are uniformly and densely pubescent with dendritic trichomes over all plant parts; at one time I thought these were a new species and annotated some material as such, but in studying the Dulcamaroid clade in detail it has become apparent that these plants are merely pubescence extremes in this widespread species.

They agree with other Bolivian material of Solanum aligerum in all other respects. It would be interesting to investigate if this pubescence variation is due to some habitat variation not apparent from herbarium labels. Solanum aligerum is extremely similar to Solanum pubigerum , with which it broadly overlaps in central Mexico and Central America. The two species can be very difficult to distinguish, but Solanum aligerum has dendritic trichomes concentrated in the vein axils or over the entire lamina, while Solanum pubigerum has simple trichomes found all along the midrib on the leaf undersides.

Flowers are larger in Solanum aligerum with the calyx lobes quadrate rather than deltate this can be difficult to see , and the berries of Solanum aligerum are also larger and usually black or blackish green, rather than red, when ripe. The stems of S. In general the two species appear to not occupy the same forest types where their ranges overlap. The absence of Solanum aligerum in Colombia and Ecuador could indicate the northern and southern populations are in the process of differentation.

This disjunction is unusual, but has also been found in the potato species Solanum morelliforme Bitter that is disjunct between Guatemala and Bolivia Simon et al. Solanum morelliforme populations from south of the disjunction were morphologically as here in Solanum aligerum and molecularly identical. Without the locality being revealed, it is impossible to distinguish a plant of Solanum aligerum as coming from any particular area, so I am confident that these all represent a single taxon. Molecular studies at the population and geographical level may reveal some differentiating characters.

The holotype of Solanum aligerum was destroyed in Berlin and no duplicates have been traced. I have selected that in G as a lectotype, as that is where Mueller worked. Costa Rica. San Isidro del General, 12 Aug , Spellman et al. El Salvador. Montecristo, plan de los helechos, m, 13 Mar , Carballo et al.

Montecristo, al final del atajo para el Trifinio, m, 24 Jan , Monterrosa et al. Hidalgo : Lolotla, carr. Pachuca-Huejutla, 3. Juxtlahuaca, a 2. Mixe, 28 Mar , Torres C. Cusco : Urubamba, Ollantaytambo, Dist. Ollantaytambo, Abra de Malaga, m, 2 Jun , Galiano et al. Lares, Mantto, m, 23 Jan , Valenzuela et al. Vilcabamba, m, 14 Oct , Suclli et al.

Puno : Carabaya, Ollachea to San Goban road, 2. Dunal, Prodr. Solanum alphonsei Dunal. A—H drawn from Anon. Solanum alphonsei Dunal var. Type: Chile. Bertero holotype: P [P, Morton neg. Solanum germainii Phil. Philippi s. Solanum tenuicaule Phil. Chile Woody vine or lax shrub to 1 m tall. Stems erect or spreading, glabrous or with simple uniseriate glandular trichomes to 0. Bark of older stems green to reddish brown, glabrous. Leaves simple or more often shallowly and irregularly pinnatifid, 0.

Inflorescences terminal or later lateral, 2—6. Buds globose when young, later ellipsoid, the corolla strongly exserted from the calyx before anthesis. Calyx tube ca. Corolla 1—1. Filament tube minute, the free portion of the filaments ca. Ovary glabrous; style 6—7 mm long, glabrous; stigma capitate, to ca.

Seeds 15—20 per berry, ca. Solanum alphonsei occurs in southern Chile and possibly also adjacent Argentina, from sea level to the summit of the Andes at ca. In Nothofagus Blume Nothofagaceae forest and forest margins. Solanum alphonsei is one of three species of the Dulcamaroid group found in the Nothofagus forests of southern Chile and probably also from adjacent Argentina ; the others are Solanum valdiviense and Solanum crispum. Solanum alphonsei differs from Solanum valdiviense in being a vine with non-angled stems and twining petioles, in having open and divaricately branched inflorescences and in its leaves that are rhombic or deltate in outline.

The corollas of Solanum alphonsei are in general smaller than those of Solanum valdiviense , and are less deeply lobed. Solanum crispum is much more common than either of the other two species, and can be distinguished from Solanum alphonsei by its larger flowers, denser pubescence and larger inflorescences.

Solanum alphonsei is much less commonly collected than is Solanum valdiviense. Dunal named Solanum alphonsei for Alphonse de Candolle, the editor of the Prodromus and son of his mentor from Montpellier, Agustin Pyramus de Candolle. Philippi named Solanum germainii after Philibert Germain, the Chilean botanist and collector. Both original spellings are correctable McNeill et al.

No specimens were cited in the original description of Solanum alphonsei , but the material was said to have been cultivated in Geneva in The specimen in G-DC [G] is dated , and is chosen here as the lectotype. They could, however, have been collected from the same individual plant, and could be considered topotypes. Curico, Cordillera de la Costa, Sep , Witasek s.

Solanum persicifolium Mart. Type: Brazil. Solanum angustifolium Lam. Buenos Aires: Buenos Aires, P. Commerson s. Bahia: sin. Guillot s. Solanum brittonianum Morong, Ann. New York Acad. Type: Paraguay. Pilcomayo River, 10 Jan , T. Solanum handelianum Morong, Ann. Type: Based on Solanum angustifolium Lam. Based on Solanum persicifolium Mart. Stems strongly ridged with 4 whitish green wings along the entire length, completely glabrous; new growth minutely papillose, occasionally pubescent with tangled simple uniseriate trichomes, these soon deciduous.

Bark of older stems green to pale yellowish green, the bark not markedly exfoliating. Sympodial units plurifoliate, not geminate. Leaves simple, 2—6 cm long, 0. Inflorescences terminal, becoming lateral and sometimes leaf-opposed, 4—13 cm long, usually 4—5 times branched, with 8—15 flowers, glabrous except for a few weak simple uniseriate trichomes at the tips of the branches; peduncle 0. Buds ellipsoid, the corolla ca. Corolla 2. Fruit a globose to ellipsoid berry, 1—1. Solanum amygdalifolium is also cultivated outside of its native range for its showy flowers Bolivia, Andean Argentina.

Occurs in chaco vegetation along streams and rivers, in thickets and in open vegetation. Solanum amygdalifolium occurs over a very broad geographical range in association with fresh non-brackish water, and has been characterised as semi-aquatic by some authors Mentz and Oliveira In chaco habitats in Paraguay it always grows in riverside thickets, and forms loose scrambling tangles.

With its very large, showy flowers, strongly angled stems and narrow, simple leaves, it is not easily confused with any other species of the Dulcamaroid clade; it is somewhat similar to other dulcamaroids from southern South America, particularly narrow-leaved specimens of Solanum flaccidum , but the smaller flowers, more pubescent leaves and unequal filaments of the latter species are distinguishing features. Solanum flaccidum grows in completely different types of habitats than does Solanum amygdalifolium , so confusion in the field is unlikely. Flower size may be related to water availability, as plants collected from near streams and wet places in the wet season all appear to have larger flowers than those from drier areas.

The specimens collected in the foothills of the Andes in the provinces of Jujuy and Salta, Argentina appear to have all been from cultivated plants Cabrera Morton annotated his photograph [Morton neg. Morong Morong and Britton clearly intended Solanum handelianum as a replacement name for Solanum angustifolium Lam. Tomas A. Malabrigo, F.

BM ; Rio de Janeiro, Macrae s. G ; Montevideo, Fruchard s. Bitter, Repert. Solanum angustifidum Bitter. A drawn from Tressens et al. Stuckert lectotype, designated by Morton , pg. Erect or lax, scandent shrubs, 1—2 m. Stems erect, glabrous; new growth glabrous. Bark of older stems reddish brown to grey.


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Leaves deeply pinnatifid pinnate with 1—4 pairs of leaflets, 2—6. Inflorescences terminal or occasionally lateral, 2—6 cm long, several times branched, with 20—50 flowers, glabrous but with a few simple uniseriate trichomes at the point of pedicel insertion; peduncle 1—3 cm; pedicels 0. Buds ellipsoid, the corolla exserted from the calyx tube before anthesis.

Filament tube minute, the free portion of the filaments variable, with one filament longer than the rest 2—2. Ovary glabrous; style 9—11 mm long, glabrous; stigma capitate, minutely papillose. Fruit a globose berry, 0. Seeds 10—20 per berry, ca. Central and northern Argentina, with a few collections from Bolivia; 45— m. Often cultivated see Figure 1 and found escaped; Bolivian specimens are thought to be escapes from cultivation Morton Widely distributed in many vegetation types, from chaco habitats to high elevation dry forests.

Solanum angustifidum is distinctive amongst members of the Dulcamaroid clade in having consistently pinnatifid leaves that are divided nearly to the midrib into usually three pairs of linear lobes; in general in the group pinnatifid and simple leaves both occur on individual plants.

Although normally a shrubby plant, the petioles of leaves on terminal branches occasionally are elongate and twine around other vegetation. This led Morton to suggest that this characteristic made Solanum angustifidum a linking species between the sections Dulcamara and Jasminosolanum both now recognised as part of the larger more inclusive Dulcamaroid clade, Weese and Bohs This species is not easily confused with any other in the region; Solanum salicifolium has simple or only shallowly pinnatifid leaves and simple inflorescences of usually white flowers, and Solanum endoadenium has densely glandular pubescent leaves and orange rather than black fruits.

Solanum viscosissimum of southeastern Brazil has deeply pinnatifid leaves, but is strictly vining and also always has simple leaves on the same stems. Solanum seaforthianum is the only other member of the Dulcamaroid clade to have leaves that are almost always pinnate and anthers on unequal filaments, but that species has much broader leaf divisions, rather than the narrow lobes of Solanum angustifidum , and is always a vine.

Cabrera states that Solanum angustifidum is commonly cultivated in Argentina for its abundant flowers see Figure 1 , and material from Bolivia appears to have been in association with habitations Morton ; M. Nee, pers. Solanum aspersum S. A—C drawn from MacDougal et al. Woody vine, of unspecified length or height. Stems densely and evenly pubescent with antrorsely curved simple uniseriate trichomes 0.

Leaves simple, 1- 3. Inflorescences terminal on leafy short shoots, 3—15 cm long, globose to ellipsoid in outline, branching many times, with 2 principal basal branches, with 12—60 flowers, densely pubescent with simple trichomes; peduncle 0. Buds narrowly ellipsoid, the corolla strongly exserted from the calyx tube. Ovary glabrous; style 5—6 mm long, pubescent with weak simple trichomes to 0. Solanum aspersum occurs in widely separated and isolated populations along the Andes from central Ecuador into Colombia, from to m.

The few specimens of Solanum aspersum have usually been annotated as the more common and widely distributed Solanum aureum , also from Andean Ecuador. Solanum aspersum differs from that species in its simple uniseriate, rather than congested-dendritic, pubescence and in the elongate buds that open to deeply stellate flowers. Specimens of Solanum aureum from Azuay province in Ecuador have shiny adaxial leaf surfaces like those of Solanum aspersum , but always have the characteristic golden dendritic pubescence of that species rather than the simple pubescence of Solanum aspersum.

The leaves of Solanum aspersum are usually more cordate than those of Solanum aureum , but some populations of Solanum aureum approach Solanum aspersum in overall leaf morphology at first glance. Solanum aspersum has a very scattered distribution all along the Andes from northern Colombia to central Ecuador and is likely to be found in more of the intervening parts of the cordilleras, but it is apparently rare and easily overlooked. A single collection from Cajamarca in northern Peru Diaz et al. Solanum loxense Dunal, Solan. Type: Ecuador. Loja: Loja, A. Bonpland s.

Solanum clematideum Bitter, Repert. Lehmann holotype: B [F neg. Solanum aureum Dunal var. Berlin-Dahlem Diels holotype: B, destroyed; no duplicates traced. Base of Chimborazo, June , A. Bonpland holotype: P [P, Morton neg. Woody vines or lax shrubs to 5 m tall, often in open areas.

Stems densely pubescent with stiff, golden dendritic trichomes to 1 mm long, the trichome branches numerous and very short; new growth densely golden-pubescent with congested dendritic trichomes. Bark of older stems pale brownish yellow, glabrescent. Leaves simple, 1.

Buds globose to broadly ellipsoid, the corolla strongly exserted from the calyx tube before anthesis. Calyx tube 2—2. Filament tube ca. Ovary glabrous; style 7—8 mm long, glabrous or densely and minutely papillate in the basal 1. Seeds 20—30 per berry, ca. In Andean Ecuador and Peru, from to m elevation, most commonly collected around m. Solanum aureum is widespread in Ecuador and has been considered to include several species here recognised as distinct: Solanum sanchez-vegae of northern Peru, Solanum dichroandrum of northern Colombia and Venezuela and Solanum aspersum of Ecuador and Colombia, with which Solanum aureum is sympatric.

Solanum aureum is almost always described as a vine, but occasionally as shrubby, a habit variously common in the Dulcamaroid clade. Solanum sanchez-vegae and Solanum dichroandrum differ from Solanum aureum in their looser, less dense leaf pubescence and somewhat larger flowers.

All three taxa have dark purplish black berries, but those of Solanum aureum are somewhat smaller. Solanum aureum and Solanum sanchez-vegae may overlap in distribution in northern Peru in the Huancabamba depression; Solanum dichroandrum is only known Colombia and Venezuela. Specimens now recognised as Solanum aspersum had long been considered as conspecific with Solanum aureum , but differ from it in their simple trichomes from bullate bases, elliptic buds and flowers with more lanceolate corolla lobes.

Hispania. Volume 73, Number 2, May 1990

These two species are sympatric in northern Ecuador. Solanum cutervanum is also sympatric with Solanum aureum through much of its range and is easily confused with it. The most obvious difference in the two taxa is habit, with Solanum aureum being a vine and Solanum cutervanum a shrub, but also the leaf and stem trichomes of Solanum cutervanum are beige, more elongate and the branches shorter than those of Solanum aureum , in which the trichomes are golden and more classically dendritic.

In Prov. Azuay, Ecuador, Solanum aureum has markedly shiny upper leaf surfaces and is superficially very like Solanum aspersum , but the trichomes from these plants are always dendritic with short branches and the flowers have deltate corolla lobes. The type of Solanum clematideum comes from among these populations.

Many specimens from the original Humboldt and Bonpland collection were apparently re-integrated into the general collections after use in the 19 th century rather than being returned to the special collection see Knapp and Spooner ; Knapp b. P ; Hac. Solanum cardiophyllum Dunal, Prodr. Type: Cuba. Boldo s. Based on Solanum cardiophyllum Dunal, non Solanum cardiophyllum Lindl. Woody vining shrub or liana. Stems flexuous, glabrous; new growth minutely pubescent, the trichomes ca. Bark of older stems dark reddish brown.

Leaves simple or occasionally very shallowly 3-lobed, 4.

Associated Data

Inflorescences leaf-opposed or terminal, 10—30 cm long, ovoid to ellipsoid in overall shape, branching to 5 times, with 50— flowers, glabrous; peduncle 1. Buds globose and somewhat inflated, the corolla strongly exserted from the calyx tube. Calyx tube 3. Corolla 2—2. Ovary conical, glabrous; style 0. Known only from Cuba and a single collection from Haiti, at low to middle elevations. Solanum boldoense is very similar to, and has been confused with see below , Solanum dulcamaroides of Mexico.

The pedicel articulation point serves to easily distinguish the two species: in Solanum boldoense it is the distal quarter of the length and long pegs are left when flowers fall, while in Solanum dulcamaroides the pedicel articulates at or very near the base. Dunal used the name Solanum cardiophyllum for this species, but before publication realised it had already been used by Lindley for a species of potato. His replacement name is based on the same type, collected by B. Boldo near Havana. MA is a specimen of Solanum boldoense , suggesting [confirming?

Another sheet of the same plant, MA, appears to be from the same gathering. The plate is only partly finished and could be either species, but the pedicel articulation near the rhachis suggests it is Solanum dulcamaroides ; epitype material is selected in the treatment of Solanum dulcamaroides to fix the usage this name. Havana : Havana, Delessert s. G ; Havana, , Karwinski s. Puerto Rico.

Solanum calileguae Cabrera. Jujuy: Dpto. Cabrera, N. Deginani, A. Giaioti, R. Kiesling, E. Zuloaga holotype: SI; isotype: LP [n. Woody vine. Stems scandent, glabrous to pubescent with a mixture of transparent simple and dendritic uniseriate trichomes to 1 mm long; new growth densely pubescent with simple and mostly dendritic trichomes. Bark of older stems yellowish brown. Leaves simple to deeply 5-lobed, the lower lobes complete and the leaves apparently pinnate, 5.

Buds broadly ellipsoid, the corolla about halfway exerted from the calyx before anthesis. Calyx tube 2—3 mm long, conical, irregularly splitting into lobes 2—4 mm long, densely pubescent with dendritic trichomes, these denser on the tube. Ovary glabrous; style ca. Seeds not seen from mature berries. Known only from the foothills of the Andes in the Provinces of Jujuy and Salta, Argentina, — m; possibly also occurring sterile voucher only in adjacent Bolivia. In premontane forests dominated by Juglans L. Juglandaceae and Podocarpus Pers.

Solanum calileguae is a distinctive species with its dense dendritic pubescence and large, white flowers. It has a restricted Andean distribution, and is easily distinguished from other members of the group potentially occurring in the area. It differs from Solanum uncinellum , which also has dendritic pubescence on the stems and leaves, in its ellipsoid rather than long pointed buds, its rotate-stellate rather than deeply stellate flowers and in its anthers borne on equal filaments.

Solanum uncinellum , although widespread over most of South America, has only once been collected in the Andean foothills of Argentina, but not from the same area where Solanum calileguae occurs. Solanum calileguae is most similar and perhaps most closely related to Solanum flaccidum , from which it differs in the dendritic pubescence and anthers borne on equal filaments. Solanum flaccidum occurs in southeastern Brazil, and the two species are unlikely to be found together in the field.

Solanum coalitum S. Lewis photographs of Lewis Loja: Yangana-Valladolid, km 1. Subshrubs to 1 m tall, sometimes scandent and trailing. Stems glabrous and shining, usually appearing warty from the prominent leaf scars; young stems and leaves completely glabrous or sometimes with a few scattered loose white branched trichomes to 0. Bark of older stems dark brown, shining. Leaves simple, 2. Inflorescences terminal, 2. Buds globose when very young, soon elliptic and strongly exerted from the calyx tube.

Calyx tube 2. Filament tube less than 0. Ovary conical, glabrous; style 1—1. Fruit a globose berry, 1. Chromosome number unknown. Knapp c gave a preliminary conservation status of EN endangered or critically endangered for this species. The fact that Solanum coalitum occurs only within the boundaries of the Parque Nacional Podocarpus is good news for its ultimate protection and conservation, but its very restricted distribution in an isolated habitat means it certainly is of concern.

Solanum coalitum is a striking species with its large, fleshy bright purple flowers and black fruits. Some specimens of Solanum coalitum have been identified as Solanum stenophyllum , with which it is very similar. Solanum coalitum differs from Solanum stenophyllum in its subshrubby, sometimes trailing habit, its glabrous stems and leaves except for the peculiar marginal trichome band , its cup-shaped rather than conical calyx and its slightly larger flowers that are glabrous abaxially. Trichomes of Solanum coalitum when they occur are looser and more openly dendritic than the almost echinoid trichomes that are distinctly yellowish of Solanum stenophyllum.

The fruiting pedicels of Solanum stenophyllum appear to be nodding when fruit are mature, while those of Solanum coalitum are erect. Specimens of Solanum stenophyllum have been collected from the province of Loja i. Solanum stenophyllum grows as a shrub or small treelet, usually in disturbed situations. Some individuals of Solanum stenophyllum in southern Ecuador are very sparsely pubescent, but the conical calyx and yellowish closely branched trichomes serve to distinguish these plants.

Solanum coalitum is distinguished from the very similar Solanum imbaburense by its broadly deltate, rather than long triangular calyx lobes, and its leaves with sparsely papillate undersides. Roads constructed by the military to allow access to radio towers have opened the area to others. Hofstede et al. Area next to antennas at summit of cerro, m, 27 Mar , Bohs et al. Solanum ligustrinum Lodd. Cuming in lectotype, designated here: Loddiges, Bot. Solanum concavum Lindl. Solanum laetum Kunze, Linnaea Type: Germany. Cultivated in Leipzig, , Anon. Bridges s. Witheringia berteroana J.

Tagua-Tagua, C. Bertero s. Witheringia gayana J. Gay s. Witheringia tomatillo J. Poeppig 74 [diar. Solanum congestiflorum Dunal, Prodr. Aviluco, C. Bertero lectotype, designated by Knapp , pg. Solanum congestiflorum Dunal var. Gay herb. Type: Based on Solanum ligustrinum Lodd. Solanum pyrrhocarpum Phil.

Chile 21 2 : Solanum sadae Phil. Solanum landbeckii Phil. Landbeck s. Solanum izquierdii Phil. Santiago, Aculeo ca. Izquierdo s. Solanum gayanum J. Type: Based on Witheringia gayana J. Solanum tomatillo J. Type: Based on Witheringia tomatillo J. Solanum pugae Phil. Chile 7. Puga s. Solanum pannosum Phil. Chile 9. Vidal s. Solanum berteroanum J. Chile 8. Type: Based on Witheringia berteroana J. Solanum tagua Kuntze, Revis. Solanum congestiflorum var.

Reiche, Anales Univ. Type: Based on Solanum pannosum Phil. Shrubs or small trees, often lax and scrambling, 0. Stems glabrous or pubescent with tiny dendritic trichomes; leaf scars somewhat prominent; new growth glabrous to densely pubescent with fine, dendritic trichomes. Bark of older stems pale brownish-yellow, glabrous, and shiny. Inflorescences terminal, later appearing lateral from overtopping of shoots, 2—10 cm long, flat-topped or pyramidal, branching 5—7 times, with 10—20 flowers, glabrous or sparsely pubescent with dendritic trichomes like those of the stems and leaves; peduncle 1—5 cm long; pedicels 1—1.

Buds globose when young, later elliptic, the corolla strongly exserted from the calyx tube. Ovary glabrous or with a few dendritic and simple trichomes at the apex, especially in otherwise pubescent plants; style 0. Fruit a globose or ellipsoid berry, 0. Ezcurra, pers. Solanum crispum grows in Nothofagus Nothofagaceae forest, often in second growth, and in a wide variety of moist microsites in otherwise dry habitats.

Chile: tomatillo, natre, natrien, natri, tomatilla see Knapp Solanum crispum is one of the most variable and is the most southerly species of the Solanum nitidum species group, occurring to 43 degrees S latitude. It also has a huge elevational range, occurring from sea level to nearly m in a wide variety of habitats. Two pubescence forms occur throughout the range of Solanum crispum : glabrous plants were traditionally called Solanum crispum and pubescent ones Solanum congestiflorum see Knapp Specimens of intermediate pubescence are rare, but the new growth of glabrous plants is always dendritic-pubescent.

In a cladistic analysis Knapp the two forms were treated as separate, but were strongly resolved as sister taxa. Pubescence may be related to habitat, but this effect has not been studied in any detail; polymorphism in pubescence is extremely common in members of the Dulcamaroid clade and elsewhere in Solanum. The pubescent form often has larger, more repand leaves than does the glabrous form. This raises the intriguing possibility that the pubescent form is paedomorphic, retaining the shape and indument of juvenile leaves.

Solanum crispum has been cultivated in the United Kingdom since the early part of the 19th century. Anderson Hooker and a variety still in cultivation today was developed at the Glasnevin Botanic Gardens in Dublin. Solanum crispum is usually classed as a climber, but this is due to its lanky habit in the British climate. It does not possess the twining stems, petioles or tendrils of a true climber, but it is not a robust, erect plant.

It is unlikely that any of these specimens are actually holotypes; the dispersal of specimens at the time of the expedition and subsequently through sale and loss means lectotypification is essential even if only a single sheet is present at MA. Knapp lectotypified Solanum crispum citing only a sheet in MA; this was rectified in by citation of the particular sheet Knapp c as the lectotype.

Cuming are cited. Lectotype of Solanum ligustrum Lodd. Loddiges : tab. BM ; Santiago, Oct , Grandjot s. G ; Santiago, , Philippi s. G ; Prov. G ; Coronel, , Oschenius s. G ; Valdivia, 22 Oct , Buchtien s. G ; Valdivia, 28 Sep , Buchtien s. Solanum cutervanum Zahlbr. Solanum pulverulentum Pers. Weberbauer holotype: B, destroyed; lectotype, designated by Knapp : F [F, F neg.

Cajamarca: Cutervo, C. Shrubs to small trees, 1—7 m tall. Stems and leaves densely covered with loosely branching golden tree-like trichomes; leaf scars somewhat raised, the stem not winged; new growth densely pubescent with golden tree-like trichomes above and below. Bark of older stems dark reddish-brown, sparsely pubescent with the tree-like trichomes of the young stems. Leaves simple, 6. Inflorescences terminal, later appearing lateral or in the fork of the branches, 4—10 cm long, pyramidal, branching ca.

Buds ellipsoid, the corolla strongly exserted from the calyx tube. Filament tube absent; free portion of the filaments 1—1. Ovary glabrous or with a few dendritic trichomes at the apex, glabrate in fruit; style 5—7 mm long, sparsely to densely pubescent at the base or along its entire length with golden dendritic trichomes; stigma bilobed, the surface minutely papillose.


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  6. Fruit a globose, purplish-black berry, with thin pericarp, 1—1. Andean Peru from Piura to Puno with a single collection known from Bolivia; — m. In rocky uplands, cloud forests and along trails in forest, usually growing in the open. Peru: rama de serrano Knapp Solanum cutervanum had long been confused with and placed in the synonymy of S.

    Specimens of both these species were previously annotated as Solanum pulverulentum Pers. The two species, however, are very different. Solanum cutervanum has golden or brownish tree-like trichomes on stems, leaves and inflorescences and black berries, while Solanum nitidum has more delicate, greyish, strictly dendritic trichomes and red ripe berries. Solanum cutervanum is closely related to Solanum ruizii , also from central Peru.

    It differs from that species in its often dichasial branching, rounded leaf bases, smaller flowers and deltate calyx lobes. These sister taxa are sympatric and flower size may contribute to reproductive isolation. Some populations of Solanum cutervanum from N Peru have broadly elliptical leaves superficially reminiscent of Solanum aureum , a vining species with golden strictly dendritic pubescence. Unfortunately the species name Solanum pulverulentum Pers. Linneaus only used the epithet pulverulentum in the edition of his Systemae Naturae , and in later publications reverted to the use of his original epithet tomentosum for the South African species.

    This is incorrect when several sheets are present in MA. Knapp lectotypified Solanum angustifolium citing only a sheet in MA; this was rectified in by citation of the particular sheet Knapp c. In describing Solanum aureum vars. He compared Weberbauer to a sheet at B collected by Humboldt an isotype of Solanum aureum and clearly stated Bitter that the Humboldt sheet represented the typical variety, and that various sheets of Weberbauer were his new taxa. I have selected the F duplicate as the lectotype of var. It is likely he was using material at Berlin now destroyed for his description. Lares, m, 18 Feb , Valenzuela et al.

    Solanum dichroandrum Dunal. A—F drawn from Riina et al. Solanum sanctaenevadae Dunal, Prodr. Type: Colombia. Solanum dichroandrum Dunal var. Solanum endotrichum Bitter, Repert. Solanum schlimii Bitter, Repert. Magdalena or Guajira: Prov. Linden holotype: G [G, F neg. Woody vine to 8 m long. Stems flexuous and appearing somewhat warty from prominent leaf scars, almost glabrous to densely pubescent with loose dendritic trichomes 1—1.

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