Monograph |
Corresponding author: Tiina Särkinen ( tsarkinen@rbge.org.uk ) Corresponding author: Sandra Knapp ( s.knapp@nhm.ac.uk ) Academic editor: Eric Tepe
© 2018 Tiina Särkinen, Peter Poczai, Gloria E. Barboza, Gerard M. van der Weerden, Maria Baden, Sandra Knapp.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Särkinen T, Poczai P, Barboza GE, van der Weerden GM, Baden M, Knapp S (2018) A revision of the Old World Black Nightshades (Morelloid clade of Solanum L., Solanaceae). PhytoKeys 106: 1-223. https://doi.org/10.3897/phytokeys.106.21991
|
The Morelloid clade, also known as the black nightshades or “Maurella” (Morella), is one of the 10 major clades within Solanum L. The pantropical clade consists of 75 currently recognised non-spiny herbaceous and suffrutescent species with simple or branched hairs with or without glandular tips, with a centre of distribution in the tropical Andes. A secondary centre of diversity is found in Africa, where a set of mainly polyploid taxa occur. A yet smaller set of species is found in Australasia and Europe, including Solanum nigrum L., the type of the genus Solanum. Due to the large number of published synonyms, combined with complex morphological variation, our understanding of species limits and diversity in the Morelloid clade has remained poor despite detailed morphological studies carried out in conjunction with breeding experiments. Here we provide the first taxonomic overview since the 19th century of the entire group in the Old World, including Africa, Asia, Australia, Europe and islands of the Pacific. Complete synonymy, morphological descriptions, distribution maps and common names and uses are provided for all 19 species occurring outside the Americas (i.e. Africa, Asia, Australia, Europe and islands of the Pacific). We treat 12 species native to the Old World, as well as 7 taxa that are putatively introduced and/or invasive in the region. The current knowledge of the origin of the polyploid species is summarised. A key to all of the species occurring in the Old World is provided, together with line drawings and colour figures to aid identification both in herbaria and in the field. Preliminary conservation assessments are provided for all species.
Africa, Asia, Australia, Eurasia, Europe, Madagascar, Pacific, polyploidy, Solanum nigrum complex, supervegetables, weeds
Solanum L., with approximately 1,400 species, is one of the largest genera of flowering plants (
The Morelloid clade of Solanum, also known as the black nightshades or “Maurella” (Morella), is amongst the 10 robustly supported major clades within Solanum (
“Solanum Hortense” from Joseph Banks' handcoloured copy of Dioscorides De Materia Medica; the flowers were originally painted white but have turned brown because a lead paint was used. This copy dates from the 15th century. Copyright The Trustees of the Natural History Museum, London. Reproduced with permission.
Morphology of the Old World species of the Morelloid clade of Solanum A Many species are cultivated for leaves and fruits (S. scabrum, NIJ994750012) B Most species show preference of disturbed, dry habitats in the wild (S. americanum, Knapp et al. 10210) C Strongly ridged and often purple coloured stems present in many species (S. tarderemotum, NIJ 2010-5) D Glandular hairs present in some species of the Morelloid clade (S. memphiticum, A34750473) E Typical stellate, white flowers with spreading or reflexed corolla lobes (S. tarderemotum, A14750151) F Fleshy round berries that are black, green, yellow, orange or red depending on the species (S. nigrum, A44750150) G Stone cells (also known as sclerotic granules or brachysclerids) are found in the fruits of most species of the Morelloid clade and are round in shape as compared to the tear-drop shaped or ellipsoid seeds (S. umalilaense, A24750133) H Stone cells are often easy to see in herbarium specimens (e.g. S. sarrachoides, Blom s.n. BM001207745). Photos by S. Knapp and G. van der Weerden.
Chromosome counts for species of the Morelloid clade of Solanum. For references, see individual species treatments.
Species | Haploid chromosome number |
Solanum alpinum Zoll. & Moritzi | – |
Solanum americanum Mill. | 12 (2n=2x=24) |
Solanum chenopodioides Lam. | 12 (2n=2x=24) |
Solanum furcatum Dunal | 36 (2n=6x=72) |
Solanum hirtulum A.Rich. | – |
Solanum memphiticum J.F.Gmel. | 24 (2n=4x=48) |
Solanum nigrum L. | 36 (2n=6x=72) |
Solanum nitidibaccatum Bitter | 12 (2n=2x=24) |
Solanum opacum A.Braun & C.D.Bouché | 36 (2n=6x=72) |
Solanum palitans C.V.Morton | 12 (2n=2x=24) |
Solanum pseudospinosum C.H.Wright | 24 (2n=4x=48) |
Solanum pygmaeum Cav. | 12 (2n=2x=24, some supernumerary anomalies, see species description) |
Solanum retroflexum Dunal | 24 (2n=4x=48) |
Solanum sarrachoides Sendtn. | 12 (2n=2x=24) |
Solanum scabrum Mill. | 36 (2n=6x=72) |
Solanum tarderemotum Bitter | 24 (2n=4x=48) |
Solanum triflorum Nutt. | 12 (2n=2x=24) |
Solanum umalilaense Manoko | 24 (2n=4x=48) |
Solanum villosum Mill. | 24 (2n=4x=48) |
General overviews of Black nightshade taxonomy and systematics have been published (
Within the Old World, Africa is a centre of species diversity with a total of 12 species (10 of them native), with fewer species found in Australasia (Figure
An overview of the entire section in the Old World, including a revision of the nomenclature and typification of the more than 370 names associated with these taxa, has never been done but is needed in order to provide identification tools for these difficult and morphologically very similar species. Here we provide a taxonomic revision of all 19 species of the Morelloid clade (black nightshades) occurring in the Old World based on a detailed morphological study; we include native and introduced taxa. This is part of our molecular systematic and taxonomic work focusing on producing a monographic treatment of the entire Morelloid clade which has to date focused on understanding species diversity and delimitation in the New World (e.g.
The European species of black nightshades were known to the ancient Greeks and Romans, who used them medicinally. In the 1st century AD, Pliny the Elder (
The first herbal written in English was by John Gerard. He (
By the mid-15th century, black nightshades were already becoming confused with the more toxic deadly nightshade (Atropa bella-donna). In his Complete herbal, first published in 1653, Nicholas Culpeper called his “Common nightshade” a “cold Saturnine plant” and warned “Have a care you mistake not the deadly nightshade for this [black nightshade]; if you know it not, then you may let them both alone” (
In the sixth edition of his Gardener’s dictionary, Philip Miller used Linnaean binomials for the first time (see
The name for the Morelloid clade is derived from
The very numerous treatments of these species in European floristic works (see species descriptions) usually did not specifically treat these taxa as belonging to infrageneric groups. Often these species were the only solanums treated in these floristic works, and they were often called “black nightshades” (e.g.
Within the Morelloids, four well-supported clades have been recognised based on a detailed molecular phylogenetic study (
The number of taxa included in this clade has not been clear, in part because it contains many widespread and morphologically variable species and has always been considered difficult. Although
Status and general distribution of Old World members of the Morelloid clade. Narrowly endemic taxa are shown in boldface Type.
Species | Status in the Old World | Distribution |
Solanum alpinum Zoll. & Moritzi | Native | Indonesia |
Solanum americanum Mill. | Native? | Circumtropical |
Solanum chenopodioides Lam. | Introduced | Southern South America; sporadically introduced elsewhere |
Solanum furcatum Dunal | Introduced | Southern South America; sporadically introduced elsewhere |
Solanum hirtulum A.Rich. | Native | Ethiopia |
Solanum memphiticum J.F.Gmel. | Native | Eastern Africa |
Solanum nigrum L. | Native | Europe, northern Africa, Eurasia and southeast Asia; sporadically introduced elsewhere |
Solanum nitidibaccatum Bitter | Introduced | Southern South America; widely introduced elsewhere |
Solanum opacum A.Braun & C.D.Bouché | Native | Australasia and the Pacific |
Solanum palitans C.V.Morton | Introduced | Southern South America; sporadically introduced elsewhere |
Solanum pseudospinosum C.H.Wright | Native | Cameroon, Equatorial Guinea (Bioko) |
Solanum pygmaeum Cav. | Introduced | Southern South America; sporadically introduced elsewhere |
Solanum retroflexum Dunal | Native | Southern Africa; sporadically introduced elsewhere from cultivation (see species discussion) |
Solanum sarrachoides Sendtn. | Introduced | Southern South America; sporadically introduced elsewhere |
Solanum scabrum Mill. | Native | Africa; introduced through cultivation elsewhere |
Solanum tarderemotum Bitter | Native | Africa |
Solanum triflorum Nutt. | Introduced | Southern South America and North America; introduced elsewhere |
Solanum umalilaense Manoko | Native | Tanzania |
Solanum villosum Mill. | Native | Europe, northern Africa, Eurasia and Africa; sporadically introduced elsewhere |
Numerical taxonomic studies have been undertaken in order to resolve species relationships, parental origin of polyploids and species delimitation in these species (
Modern regional taxonomic treatments of the Old World members of the Morelloid clade have been done for Europe (
The Old World species of the Morelloid clade do not form a monophyletic group. Phylogenetic analysis using plastid DNA sequence data (
Members of the Morelloid clade are either herbs or shrubs; species occurring in the Old World range from annual (e.g. S. triflorum) to short-lived perennials (e.g. S. retroflexum, S. tarderemotum) and most are herbaceous, although some species can develop woody bases and appear to be somewhat shrubby (e.g. S. villosum). Stems are usually weak and occasionally somewhat scrambling, but can reach 2+ m in height. Plants of all species usually have herbaceous upper stems, even if the base is woody. The stems can be hollow (drying flattened, e.g. S. tarderemotum) or solid (e.g. S. americanum, S. villosum); this can be a useful character for identification of herbarium specimens.
Sympodial growth is characteristic of Solanaceae, giving the stems a typical “zig-zag” appearance; details of sympodial structure have proved useful for infrageneric classification within Solanum (
“Spinose” processes are common on herbaceous stems in many species of black nightshades. They usually occur along the angles of upper parts of larger stems and are often decurrent from leaf bases. These are not true prickles, like those found in the “spiny” solanums (Leptostemonum clade, see
Species of the Morelloid clade have simple leaves that are generally elliptic or ovate in outline. Solanum retroflexum has a distinctive rhomboid leaf shape and S. scabrum leaves are usually broadly ovoid with long petioles. As with other vegetative characters in this group, leaf morphology can be extremely variable within a species or even in a single plant. Many of the infraspecific names in S. nigrum are based on variation in leaf morphology, particularly with respect to lobing of the margins.
Leaf margins vary from entire to quite deeply sinuate and lobed. Most populations of S. triflorum have deeply pinnatifid leaves, but a wing of leaf blade is always present along the midrib. Other species have variously entire or toothed margins and often the teeth occur only in the basal half to third of the leaf blade. The leaf blades are usually somewhat decurrent on to the petiole and the leaf base is cuneate to attenuate. Leaf apices are acute to attenuate, but vary considerably within species.
Petiole length to some extent is related to leaf size and, on individual plants, larger leaves always have longer petioles. As Solanum scabrum tends to have long petioles with little decurrent leaf blade tissue, this character can be helpful in distinguishing it from S. nigrum or from the sympatric S. tarderemotum.
Trichomes in species of the Morelloid clade are simple or branched (e.g. S. pallidum Rusby of the Andes), but never stellate (
The presence or absence of glandular trichomes has also been previously treated as taxonomically significant (see
As with all species of Solanum, the inflorescence of members of the Morelloid clade is developmentally terminal and later overtopped by the leading axillary shoot so that it appears lateral. The basic structure is an unbranched or variously branched scorpoid cyme. Most members of the black nightshades have unbranched (simple) or merely furcate (once-branched) inflorescences, but in populations cultivated for fruit (e.g. S. scabrum, S. umalilaense, S. villosum), complex branching can occur, presumably through human selection for higher fruit production. Populations of S. americanum from Hainan Island in southern China with highly branched inflorescences have been described as a distinct species (S. merrillianum), but we consider these to be variants of the common S. americanum (see description of S. americanum). The degree of inflorescence branching in species of the group may also depend upon plant or inflorescence age (e.g. S. tarderemotum). In all Solanum species, the inflorescence expands from the tip producing flowers in a proliferating manner (
All members of the group have distinct peduncles, usually somewhat longer than the distal flower-bearing portion, but inflorescence length and flower number vary both between and within species. Many species in the group have what are termed “sub-umbellate” inflorescences, where the flower-bearing rhachis is very short and the pedicels are all very closely spaced and congested at the very tip of the inflorescence. We use the term sub-umbelliform in the species descriptions for the extreme cases of this flower clustering. This inflorescence is not a true umbel, but is described as such in much previous literature, usually as an umbellate or subumbellate cyme (e.g.
Pedicels in flower are usually deflexed or spreading, but this can be very difficult to see in herbarium specimens. In fruit, pedicels are usually somewhat pendent from the weight of the berry, but are strongly (e.g. S. chenopodioides, S. tarderemotum) or weakly (e.g. S. nigrum) deflexed in some species. Solanum tarderemotum has the pedicels in fruit strongly bent at the base in an acute angle relative to the rhachis; this is a useful character for identification of herbarium specimens. Other species have markedly spreading pedicels in fruit (e.g. S. memphiticum). The abscission zone at the pedicel base in members of the Morelloid clade is at the very base and, if and when pedicels fall, the scars are generally flush with the rhachis. Pedicel persistence with fruit ripening is an important species character in this group. Ripe berries either fall or are taken from the plant with the pedicel still attached (e.g. S. opacum, S. tarderemotum) or the berry falls and the pedicel is left behind (e.g. S. americanum, S. nigrum, S. villosum). The presence of old pedicels can be useful for identification of non-flowering herbarium specimens.
The calyx in all members of the Morelloid clade is 5-merous and synsepalous. The calyx tube is generally conical or occasionally somewhat elongate (e.g. S. memphiticum) and the lobes are extremely variable in size and shape from minute and deltate (e.g. S. umalilaense) to long-triangular (e.g. S. pseudospinosum). The position of the calyx lobes in fruit is an important identification character; they can be strongly reflexed (e.g. S. americanum, S. villosum), spreading (e.g. S. nigrum, S. tarderemotum) or appressed to the berry (e.g. S. opacum). The calyces of the weedy introduced species S. nitidibaccatum and S. sarrachoides are accrescent in fruit with the calyx lobes expanding to envelop almost the entire berry (several other New World members of the group also have accrescent calyces, see
In common with most other species of Solanum, members of the Morelloid clade have 5-merous sympetalous corollas that are variously stellate. Floral mutants are often observed, where 4-6-merous corollas can occur on individual plants that are otherwise 5-merous (e.g. S. scabrum). Colour is generally white or pale violet-tinged, but anthocyanin pigmentation can vary depending on environmental growth conditions. In most species at least, some individuals (collections) with purple or violet flowers have been recorded. Solanum villosum often has distinctive dark purple coloration on the abaxial midvein of each corolla lobe. At the base of the corolla tube, there is a ring or irregular area of differently coloured tissue usually referred to as the “eye”. In the species of the Morelloid clade, this is usually yellow or greenish-yellow and, in some species, the eye has darker brown or blackish-purple margins (e.g. S. nitidibaccatum). The colours of the eye usually disappear in herbarium specimens and are rarely noted on labels. This eye is usually similar in texture to the rest of the corolla and not shiny as occurs in the Dulcamaroid clade (see
Corollas in the Morelloid clade are stellate to deeply stellate and corolla lobes are deltate to long-triangular. Solanum tarderemotum has deeply stellate corollas, with reflexed corolla lobes, while S. memphiticum has corollas with the lobes approximately the same length as the tubular portion and the lobes are not strongly reflexed at anthesis. These characters, particularly those of the degree to which corolla lobes are reflexed, can be very difficult to see in herbarium specimens. As is seen in many other groups of solanums (e.g. Dulcamaroid clade, ANS clade, see
Corollas of members of the Morelloid clade are usually very small, in fact representing the tiniest flowers of any Solanum. Corolla diameter varies from 4–20 mm; S. nitidibaccatum has the smallest corollas and S. hirtulum and S. furcatum the largest in the group of species occurring in the Old World. Adaxial lobe surfaces are usually glabrous, while abaxial corolla lobe surfaces are variously papillate, with longer simple uniseriate trichomes on the margins and tips.
The stamens of members of the Morelloid clade are ellipsoid and equal to very slightly unequal in size and length. The filament tube and filaments are variously pubescent adaxially. Most populations of S. memphiticum have completely glabrous filaments, but this is not completely consistent within the species; this was used as a species-specific character in
Anthers of members of the Morelloid clade conform to the poricidal morphology of all other species of Solanum (see
The gynoecium in members of the Morelloid clade is bicarpellate; the carpels are fused in a superior ovary with axile placentation. The ovary is glabrous and usually conical to globose. The flowers lack nectaries, as do all species of Solanum. The style is straight or slightly curved and usually sparsely to densely pubescent in the lower half to third where it is enclosed in the anther cone. It is usually exserted from the anther cone, but in some species (e.g. most populations of S. americanum, S. scabrum) only barely exceeds the length of the stamens. This may be related to self-fertilisation and thus self-compatibility, as has been observed in the tomatoes (
As with all species of Solanum, the fruit is a bicarpellate berry. Fruits of members of the Morelloid clade are usually brightly coloured and juicy. Most species have globose berries, but those of S. villosum are usually somewhat longer than wide. Berry colour can be green (S. nitidibaccatum, S. opacum, some populations of S. tarderemotum), greenish-yellow or yellow (some populations of S. nigrum and S. villosum, S. palitans), bright orange (S. villosum) or varying shades of purple (many species); immature berries are usually described as green on herbarium labels. Colour polymorphisms are common in species of this group; both S. nigrum and S. tarderemotum, for example, have individuals and populations with green or purple berries and S. villosum has either yellow or orange berries.
The pericarp (epicarp) of the berries is thin and either matte (e.g. S. chenopodioides, S. tarderemotum) or shiny (e.g. S. americanum, S. villosum). Surface characteristics are useful for species identification, especially when combined with other characters (see discussion of S. americanum). The mesocarp is always juicy and very liquid; these fruits are eaten by both birds and mammals (including people). In general, the mesocarp of fresh fruits is green or greenish-yellow, but in species with purple berries, it is sometimes purplish. In the cultivated S. scabrum, this may be related to human-mediated selection in Africa and, in China, where on Hainan Island people are beginning to cultivate S. americanum (S. Knapp, pers. obs.), coloured fruit pulp is selected for by local people. This character is rarely mentioned on herbarium labels.
Like some other groups of non-spiny solanum such as the Pachyphylla clade (
Members of the Morelloid clade have flattened seeds, like other solanums. Unlike other groups, however, they are usually tear-drop rather than kidney shaped, with the hilum and micropyle at one of the short ends of the seed. Seed size varies from 1–3 mm long, polyploid species usually have larger seeds than diploids (e.g. S. americanum seed size is 1–1.5 mm, while that of S. scabrum is 2–2.8 mm) and hence seed size is a good feature for distinguishing S. nigrum (hexaploid) from S. americanum (diploid). Seed number per berry in the Morelloid clade is generally quite high (
Seed coat morphology has been suggested as a useful character for species-level taxonomy in Solanum (
Chromosome numbers in the Morelloid clade are variations on the base number of 12 (Table
Many chromosome counts are reported for members of this group, often as unvouchered counts of “Solanum nigrum”. In the species treatments, we only record counts that are based on identifiable material or those that are vouchered and for which we have verified the specimen in question. Chromosome counts recorded in floras (e.g.
Members of the Morelloid clade are plants of disturbed habitats and occur in landslides, along roads and stream, and at the edges of cultivated fields. Many of the species have very broad elevational ranges (e.g. S. americanum) and extremely broad distributions, but a few of the Old World species are localised endemics (e.g. S. alpinum, S. hirtulum, S. pseudospinosum) primarily of montane areas.
These localised endemics are all from recent (ca. Miocene age), volcanic mountain ranges and are all probably of recent origin. Solanum pseudospinosum is a polyploid, while the ploidy levels of S. alpinum and S. hirtulum are not known. Molecular data suggests S. hirtulum to be a polyploid based on presence of multiple peaks in low copy nuclear gene sequences, similar to known polyploid species such as S. nigrum (
Several members of the group (e.g. S. nigrum, S. nitidibaccatum) are registered as noxious weeds of agriculture (see below) in both Europe and North America (
We list the status and general distribution of the species in the group in Table
Country distribution of members of the Morelloid clade in Africa. Introduced species are shown in parentheses. Countries where these species are expected to occur, but from which we have seen no specimens are included, but no species listed.
Country | Species |
---|---|
Algeria | S. nigrum, S. villosum |
Angola | S. americanum, S. scabrum, S. tarderemotum, S. villosum |
Benin | S. scabrum |
Botswana | S. retroflexum, S. scabrum, S. tarderemotum, S. villosum |
Burkina Faso | S. scabrum |
Burundi | S. memphiticum, S. tarderemotum, S. villosum |
Cape Verde | S. americanum, (S. nigrum), S. tarderemotum |
Cameroon | S. americanum, S. pseudospinosum, S. scabrum, S. tarderemotum |
Central African Republic (CAR) | S. scabrum |
Chad | S. tarderemotum, S. villosum |
Comoros | S. americanum, S. scabrum, S. tarderemotum |
Democratic Republic of the Congo | S. americanum, S. scabrum, S. tarderemotum |
Republic of the Congo | S. americanum, S. scabrum |
Cote d’Ivoire | S. americanum, S. scabrum |
Djibouti | – |
Egypt (incl. Hala’ib triangle) | S. memphiticum, S. nigrum, S. scabrum, S. tarderemotum, S. villosum |
Equatorial Guinea | S. americanum, S. pseudospinosum, S. scabrum |
Eritrea | S. americanum, S. memphiticum, S. scabrum, S. tarderemotum, S. villosum |
Ethiopia | S. americanum, S. hirtulum, S. memphiticum, S. scabrum, S. tarderemotum |
Gabon | S. americanum, S. scabrum |
Gambia | S. americanum |
Ghana | S. americanum, S. scabrum, S. tarderemotum |
Guinea | S. scabrum, S. tarderemotum |
Guinea-Bissau | S. americanum |
Kenya | S. americanum, S. memphiticum, S. scabrum, S. tarderemotum, S. villosum |
Lesotho | (S. chenopodioides), S. retroflexum, S. scabrum, S. tarderemotum |
Liberia | S. americanum, S. scabrum, S. villosum |
Libya | S. nigrum, S. villosum |
Madagascar | S. americanum, S. scabrum, S. tarderemotum |
Malawi | S. americanum, S. retroflexum, S. scabrum, S. tarderemotum, S. villosum |
Mali | S. tarderemotum |
Mauritania | (S. scabrum), S. villosum |
Mauritius | S. americanum, (S. chenopodioides), (S. tarderemotum) |
Morocco | S. nigrum, (S. triflorum), S. villosum |
Mozambique | S. americanum, S. scabrum, S. tarderemotum, S. villosum |
Namibia | S. retroflexum, S. scabrum, S. tarderemotum |
Niger | S. villosum |
Nigeria | S. americanum, S. scabrum, S. tarderemotum, S. villosum |
Rwanda | S. tarderemotum |
São Tome e Principe | S. americanum, S. scabrum |
Senegal | S. scabrum, S. tarderemotum |
Seychelles | S. americanum, S. scabrum |
Sierra Leone | S. americanum, S. scabrum, S. tarderemotum |
Somalia | S. americanum, S. memphiticum, S. tarderemotum, S. villosum |
Country distribution of members of the Morelloid clade in Asia. Introduced species are shown in parentheses. Countries where these species are expected to occur, but from which we have seen no specimens are included, but no species listed.
Country | Species |
Afghanistan | S. nigrum, S. villosum |
Armenia | S. nigrum, S. villosum |
Azerbaijan | S. nigrum, S. villosum |
Bahrain | S. nigrum, S. villosum |
Bangladesh | S. americanum, S. villosum |
Bhutan | S. americanum, S. nigrum, S. villosum |
Brunei | – |
Cambodia | S. americanum |
China | S. americanum, S. nigrum, (S. scabrum), S. villosum |
Cyprus | S. nigrum, S. villosum |
Georgia | S. nigrum, S. villosum |
India | S. americanum, S. nigrum, S. villosum |
Indonesia | S. alpinum, S. americanum, S. nigrum, S. opacum, (S. scabrum) |
Iran | S. nigrum, S. villosum |
Iraq | S. nigrum, S. villosum |
Israel | S. nigrum, S. villosum |
Japan | S. americanum, (S. chenopodioides), S. nigrum, S. opacum |
Jordan | S. memphiticum, S. nigrum, S. villosum |
Kazakhstan | S. nigrum, S. villosum |
Kuwait | S. nigrum, S. villosum |
Kyrgyzstan | S. nigrum, S. villosum |
Laos | S. americanum, S. nigrum |
Lebanon | S. nigrum, S. villosum |
Malaysia | S. americanum, S. nigrum, S. opacum |
Maldives | – |
Mongolia | S. nigrum |
Myanmar (Burma) | S. americanum, S. nigrum |
Nepal | S. americanum, S. nigrum, S. villosum |
North Korea | S. nigrum |
Oman | S. nigrum, S. villosum |
Pakistan | S. americanum, S. nigrum, S. villosum |
Palestine | S. nigrum, S. villosum |
Philippines | S. americanum, S. nigrum, S. opacum |
Qatar | S. villosum |
Russian Federation | S. nigrum, S. villosum |
Saudi Arabia | S. memphiticum, S. nigrum, S. villosum |
Singapore | (S. scabrum) |
South Korea | S. nigrum |
Sri Lanka | S. americanum, S. nigrum |
Syria | S. nigrum, S. villosum |
Taiwan | S. americanum, S. nigrum, S. opacum |
Tajikistan | – |
Thailand | S. americanum, S. nigrum |
Timor-Leste | – |
Turkey | S. nigrum, S. villosum |
Turkmenistan | S. nigrum, S. villosum |
United Arab Emirates (UAE) | S. nigrum, S. villosum |
Uzbekistan | S. nigrum |
Vietnam | S. americanum, S. nigrum |
Yemen | S. memphiticum, S. nigrum, S. villosum |
Country distribution of members of the Morelloid clade in Europe. Introduced species are shown in parentheses and species added from literature sources in boldface type (
Country | Species |
---|---|
Albania | S. nigrum, (S. nitidibaccatum), (S. triflorum), S. villosum |
Andorra | – |
Armenia | S. nigrum, S. villosum |
Austria | S. nigrum, (S. nitidibaccatum), S. villosum |
Azerbaijan | S. villosum |
Belarus | S. nigrum |
Belgium | S. nigrum, (S. nitidibaccatum), (S. triflorum), (S. villosum) |
Bosnia and Herzegovina | S. nigrum, S. villosum |
Bulgaria | S. nigrum, S. villosum |
Croatia | (S. chenopodioides), S. nigrum, (S. nitidibaccatum), S. villosum |
Cyprus | S. nigrum, S. villosum |
Czech Republic | S. nigrum, S. villosum |
Denmark | S. nigrum, (S. nitidibaccatum), S. villosum |
Estonia | S. nigrum |
Finland | (S. americanum), S. nigrum, (S. nitidibaccatum) |
France | (S. americanum), (S. chenopodioides), S. nigrum, (S. nitidibaccatum), S. villosum |
Georgia | S. nigrum, S. villosum |
Germany | (S. chenopodioides), S. nigrum, (S. sarrachoides), (S. triflorum), S. villosum |
Greece | (S. chenopodioides), S. nigrum, S. villosum |
Hungary | (S. americanum), (S. chenopodioides), S. nigrum, (S. scabrum), S. villosum |
Iceland | – |
Ireland | S. nigrum, (S. nitidibaccatum) |
Italy | (S. chenopodioides), S. nigrum, S. villosum |
Kazakhstan | S. nigrum, S. villosum |
Kosovo | – |
Latvia | S. nigrum |
Liechtenstein | – |
Lithuania | S. nigrum, (S. villosum) |
Luxembourg | – |
Macedonia (FYROM) | S. villosum |
Malta | S. nigrum, S. villosum |
Moldova | – |
Monaco | S. villosum |
Montenegro | – |
Netherlands | (S. chenopodioides), S. nigrum, (S. nitidibaccatum), (S. triflorum), (S. villosum) |
Norway | S. nigrum |
Poland | S. nigrum, S. villosum |
Portugal | (S. chenopodioides), S. nigrum, S. villosum |
Romania | S. nigrum, (S. triflorum), S. villosum |
Russian Federation | S. nigrum, S. villosum |
San Marino | S. villosum |
Serbia | S. villosum |
Slovakia | S. nigrum, S. villosum |
Slovenia | S. nigrum, S. villosum |
Spain | (S. americanum), (S. chenopodioides), S. nigrum, (S. sarrachoides), S. villosum |
Sweden | (S. americanum), (S. chenopodioides), S. nigrum, (S. nitidibaccatum), (S. sarrachoides), (S. triflorum), (S. villosum) |
Switzerland | (S. chenopodioides), S. nigrum, (S. pygmaeum), S. villosum |
Turkey | S. nigrum, S. villosum |
Ukraine | S. nigrum, S. villosum |
United Kingdom (UK) | (S. americanum), (S. chenopodioides), S. nigrum, (S. nitidibaccatum), (S. pygmaeum), (S. sarrachoides), (S. triflorum), (S. villosum) |
Country distribution of members of the Morelloid clade in the Pacific, including Australasia. introduced species in shown in parentheses. Countries where these species are expected to occur, but from which we have seen no specimens are included, but no species listed.
Country/Territory | Species |
American Samoa (USA) | S. americanum |
Australia | S. americanum, (S. chenopodioides), (S. furcatum), S. nigrum, (S. nitidibaccatum), S. opacum, (S. palitans), (S. retroflexum), (S. scabrum), (S. triflorum), (S. villosum) |
Cook Islands (New Zealand) | S. americanum, S. opacum |
Fiji | S. americanum, S. opacum |
French Polynesia (France) | S. americanum, S. opacum |
Guam (USA) | S. americanum |
Kiribati | – |
Marshall Islands | S. americanum, S. opacum |
Micronesia | S. americanum |
Nauru | – |
New Caledonia (France) | S. americanum, S. opacum |
New Zealand | S. americanum, (S. chenopodioides), (S. furcatum), S. nigrum, (S. nitidibaccatum), S. opacum, (S. villosum) |
Niue (New Zealand) | S. americanum |
Norfolk Island (Australia) | S. americanum |
Northern Mariana Islands (USA) | S. americanum |
Palau | S. americanum |
Papua New Guinea | S. americanum, S. nigrum, S. opacum |
Pitcairn Islands (UK) | S. opacum |
Rapa Nui (Chile, Easter Island) | S. americanum, S. opacum |
Samoa | S. americanum, S. opacum |
Solomon Islands | S. opacum |
Tokelau (New Zealand) | – |
Tonga | S. americanum, S. opacum |
Tuvalu | – |
Vanuatu | S. americanum, S. opacum |
Wake Island (USA) | – |
Wallis and Futuna (France) | – |
Like most solanums, flowers of members of the Morelloid clade are buzz-pollinated by bees (
Members of the Morelloid clade have juicy berries with thin pericarps (skins) that are typical for bird-dispersed fruits (
Glycoalkaloid concentrations are very low in ripe berries of S. americanum and other members of the Morelloid clade that have been tested (
Most Old World morelloid species are weedy and widely distributed; many are also cultivated (e.g. S. tarderemotum, S. scabrum, S. villosum) and are distributed widely via human migration. Many introductions of species to Europe have resulted from the use of wool shoddy as protection in market gardens (
The range-restricted endemic species of Old World morelloids suffer from taxonomic recognition issues that could seriously affect their conservation. In recent floras of many parts of the world, all taxa are treated as a single highly variable species (usually S. nigrum) and local endemic taxa are overlooked and equated with widespread invasive weeds. This means that these endemic taxa have possibly been placed at risk. Preliminary conservation assessments for the Old World members of the Morelloid clade (including introduced taxa) are presented in Table
Preliminary conservation assessments for the Old World members of the Morelloid clade. For details, see Materials and Methods and individual species treatments. Preliminary assessments are based on EOO (Extent of Occurrence) only (see Materials and Methods) and have been calculated for worldwide ranges for each species. The conservation status of species known to be introduced in the Old World has been assessed considering native range only and is discussed under each species treatments.
Species |
Preliminary conservation assessment ( |
EOO (km2) [worldwide range] |
Solanum alpinum | VU (Vulnerable) | 18,806 |
Solanum americanum. | LC (Least Concern) | 51,956,485 |
Solanum chenopodioides | LC (Least Concern) | 78,693,743 |
Solanum furcatum | LC (Least Concern) | 105,324,815 |
Solanum hirtulum | NT (Near-Threatened) | 26,035 |
Solanum memphiticum | LC (Least Concern) | 4,489,278 |
Solanum nigrum | LC (Least Concern) | 202,319,396 |
Solanum nitidibaccatum | LC (Least Concern) | 83,736,975 |
Solanum opacum | LC (Least Concern) | 78,180,634 |
Solanum palitans | LC(Least Concern) | 24,975,848 |
Solanum pseudospinosum | EN (Endangered) | 4,009 |
Solanum pygmaeum | LC (Least Concern) | 16,699,600 |
Solanum retroflexum | LC (Least Concern) | 19,067,920 |
Solanum sarrachoides | LC (Least Concern) | 114,305,743 |
Solanum scabrum | LC (Least Concern) | 49,116,048 |
Solanum tarderemotum | LC (Least Concern) | 22,131,524 |
Solanum triflorum | LC(Least Concern) | 91,711,479 |
Solanum umalilaense | EN [DD] (Endangered or Data Deficient) | 2,559 |
Solanum villosum | LC (Least Concern) | 364,611,726 |
The Morelloid clade is one of only a few cases of extensive ploidy level variation in Solanum. Potatoes and the Archaesolanum clade are the only other clades of Solanum in which ploidy level variation is extensive (
It is thought that most of the polyploids in the Morelloid clade are of alloploid, rather than autoploid origin, because bivalents show regular pairing at meiosis (
Summary of putative parental origins of the polyploid species in the Morelloid clade (see text for references).
Tetraploids 2n = 4× = 48 | Type | Putative parental species |
S. pseudospinosum | Unknown | Unknown |
S. retroflexum | autopolyploid | S. chenopodioides or S. nitidibaccatum |
S. umalilaense | Unknown | Unknown |
S. memphiticum | Unknown | Unknown |
S. tarderemotum | Unknown | Unknown |
S. villosum | autopolyploid | S. americanum |
Hexaploids 2n = 6× = 72 | ||
S. furcatum | Unknown | Unknown |
S. opacum | autoallopolyploid | S. retroflexum × S. chenopodioides |
S. scabrum | autoallopolyploid | S. americanum × S. villosum |
S. nigrum | autoallopolyploid | S. americanum × S. villosum |
The species of black nightshades are predominantly self-compatible (
There is considerable confusion regarding the toxicity of species of black nightshades (see
Most species of Solanum contain a wide range of steroidal alkaloids (also known as glycoalkaloids;
Species of black nightshades are used as leafy vegetables or fruits throughout the world, sometimes referred to as ‘supervegetables’ for their high protein and iron content (
In Africa, species are cultivated either for use as leaf vegetables (e.g. S. americanum, S. memphiticum, S. scabrum, S. tarderemotum, S. umalilaense, S. villosum) or for their fruits (e.g. S. americanum, S. retroflexum, S. scabrum, S. villosum). Leaves of African nightshades are high in flavonoids, vitamin C, folates, iron and antioxidants (
In Asia, including the Indian subcontinent, S. nigrum is also used as a leaf vegetable (
Disaggregating uses for these species is difficult because many authors treated all morelloids as a single, highly variable taxon;
Widely used common names for black nightshades in the Old World. Where names can be confidently assigned to individual species, these are listed in the species accounts. We only include names here from which the language is available and unambiguous; many common names in the literature are taken from herbarium labels and both the original and transcription are often very dubious. Sources include
Region | Name (language; alphabetically by language) | |
---|---|---|
Africa | North Africa | ‘anab el deeb (Arabic); morelle (French); yerba mora (Spanish) |
East Africa | ocogocuga, pot ocuga (Acholi); ociga/osiga (Alur [Jonam]); rinagu, amanagu (Gusii [Kisii]); ndolu (Kamba); encoratin (Karamojong); manago (Kikuyu); urusogo, isogo, ubusogosogo, ibwija, isiogwo (Kinyarwanda); isoyot, ol’momoit (Kipsigi); isogo, urusogo (Kirundi); manavu/mnavu (Kiswahili); kusochet (Kupsabiny); nswega (Kuria); lere (Kutuk na Kakwa); isusu (Loogoli); nsugga, nsugga enzirugavu (Luganda); asuaka (Lugbara); osuga, ocuga, ocokocok (Luo); mnavu tsaka, oromomoi/olomomoi (Maasai); isufa (Masaba); enswiga/enswiiga (Nkore/Kiga); cheporusion, kusuyo (Pökoot); okaka (Rutooro); esiga (Suvu); sochek (Tugen); | |
West Africa | fué (Abure); bogolo, fassa kuna (Bambara); efo-odu, sapota (Creole); shien (Dan); feibii (Edo); ebibirba (Fante); djagato-fórô (Fula); gautan kâdii (Hausa); ugunakpe (Igbo); àljáng (Kanuri); bundu kaemba (Kono); wosangu, wosangu njika (Kwpe, Ngomba); usiga (Luba); itofo, itotofu (Lombo); sibito jambo, bassa béné, suludjatô (Mandingo); mulunda, muzirhu (Mashi); kemba/kimba (Mende); wosangu (Mokpwe); rushoo (Nandi); kumbo (Nso); lin-sah, pe-noh (Samba Daka); suga (Swahili); nsusua (Twi); kholekodelen-na (Yalunka); odu/o’du, ogunmo, ogumo (Yoruba) | |
South Africa | nastergal, galbessie, nagskande (Afrikaans); umsobo, sheshoabohloko, umsobo-sobo (isiXhosa); umsobo, isihlalakuhe, udoye, umagqa, umgwaba, umsobo-sobo, umqunbabe (isiZulu); lethotho, seshoa-bohloko, momoli (Sotho); (u)msobo (siSwati); muxe, xaxadi (Venda); kophe (Xitsonga) | |
Madagascar | anamamy (Malagache); brède noir, brède morelle (French) | |
Asia | Middle East | ‘anab al deeb/’eneb el deeb (Arabic); mejaje, mesaeleha (Arabic); abriki (Arabic); ‘anamnam (Yemeni Arabic) |
Central Asia | taj rizi (Farsi) | |
Southeast Asia | peng pâhs khmôch, pum pou nam, to:ngx tè:ng, tumx to:yz (Laotian); ranti (Malay/Javanese); leuntja (Sundanese); toem tok, ya-tomtok (Thai); bóp bép, (cay) tăm bóp, (cay) hôt nut, tom bóp (Vietnamese) | |
Indian subcontinent | makŏg, gurkkamai (Hindi); kakarundi (Kannada); kakmachi (Marathi); manattakkâli, karintakâli, karimttakkâli (Malayam); kakamaci, kakamata (Sanskrit); manattakāli, milagutakkāli (Tamil); kâmanci, kacci, kaccipandu, gejjucettu (Telugu); ab makoh, inab-as-salab (Urdu) [also see http://envis.frlht.org/bot_search] | |
China and Japan | kong lui (Chinese, Mandarin); baihua cai, jia dengleng cao (Chinese, Cantonese); lung-kwei (Chinese, Taiwan); inuhozuki (Japanese) | |
Europe | belarri malaka, blets (Basque); pebre d’ase (Catalan); sort natskygge (Danish); zwarte nachtschade (Dutch); black nightshade (English); must maavits (Estonian); mustakoiso (Finnish); morelle noire (French); schwarzer nachtschatten (German); skilosaphilo (Greek); fuath dubh (Gaelic/Irish); erba morella, erba mora, solano nero (Italian); juodoji kiauliauogė (Lithuanian); svartsøtvier (Norwegian); psianka czarna (Polish); erva moira (Portuguese); paslen ĉernyj (Russian); pomocnica (Serbian); lulock čierny (Slovenian); yerba mora, hierba mora (Spanish); nattskatta (Swedish); kopek uuzimii (Turkish); codwarth du (Welsh) | |
Pacific | Australia | nightshade (English) |
New Guinea | – | |
Islands of the Pacific | popolo, huapopolo (Polynesian) | |
New Caledonia | morelle |
Solanum scabrum is by far the most commonly cultivated of the Old World species and is cultivated throughout Africa (
Solanum tarderemotum and S. villosum are more commonly cultivated in eastern Africa and many specimen labels note that the fruits of S. villosum are particularly prized by children (also see
Solanum “nigrum” has a long history of use in Ayurvedic medicine in India, but it is clear that the concept medicinally does not distinguish between S. nigrum s.s. and S. villosum (see
In Europe, seed remains of S. nigrum have been excavated from the Viking layers at York (
In the Pacific, uses for black nightshades (S. americanum and S. opacum, without distinction) are recorded from Hawaii (
Glycoalkaloids from S. americanum, S. nigrum and S. opacum were often used as a steroid base for human contraceptives until the industry moved to production of synthetic compounds in the 1980s (
Old World species of the Morelloid clade are also sources of resistance genes used in agriculture. Several species of black nightshades, including S. nigrum, S. americanum and S. villosum, are known to carry resistance genes against late blight (Phytophthora infestans (Mont.) de Bary), a major fungal pest in potato, tomato and eggplant (
Our goal for the treatment of the Old World species of the Morelloid clade has been to provide circumscriptions for the members of this morphologically variable group of species, while clearly highlighting areas, taxa and populations where further in-depth research is still needed to confirm current taxon concepts. Delimitation of species here basically follows what is known as the “morphological cluster” species concept (
Although infraspecific taxa have been recognised by others within the group, we do not recognise any here due to the complex morphological variation observed within each species, where the inspection of a larger number of specimens quickly reveals no apparent natural breaks in variation but rather a mixing between highly morphologically variable populations of widespread species. Results from seed protein studies (
Our taxonomic treatment is based on results from recent molecular systematic studies considering the taxonomy of the section, including focused morphological and AFLP studies of the African species by Jacoby (2003: South Africa),
Measurements were made from dried herbarium material supplemented by measurements from living material. Colours of corollas, fruits etc. are described from living material or from herbarium label data. Specimens with latitude and longitude data on the labels were mapped directly. Some species had few or no georeferenced collections; in these cases, we retrospectively georeferenced the collections using available locality data. Maps were constructed with the points in the centres of degree squares in a 1° square grid. Conservation threat status was assessed following the IUCN Red List Categories and Criteria (
Type specimens in this group have proved difficult to trace. Many taxa were described based on types housed in herbaria whose collections were lost during the Second World War, including the Berlin herbarium (B) (see
Many of the species and infraspecific names coined for European plants in the 19th and early 20th century floristic and other works have no specimens cited. In some cases, we have been able to trace potential original material, but in others, plants may have been described from living material. We have lectotypified these names where we have been able, but have not selected neotypes for those names that have rarely been used beyond their first description (mostly infraspecific taxa of S. nigrum and S. villosum). Where specific herbaria have not been cited in protologues, we have followed
The amateur Czech botanist, Paul M. Opiz, collected prolifically and described many taxa based on his and other collections, most of these specimens being housed in PR or PRC (
Georg Bitter (e.g.,
Type specimens are cited with their barcodes in square brackets after each herbarium code, written as they are spelled with barcode readers, either as a continuous string (i.e. [G00104280]) or with a dash (i.e. [MO–1781232]) depending on the style of barcode used in each herbarium. Sheet numbers are cited where barcodes are missing and these are indicated as distinct from barcodes (i.e. MO [acc. # 1037660]). For widespread species, we have not cited geographically representative specimens in the text under each species treatment, but instead listed the countries of occurrence by region (Africa, Asia, Europe, Pacific [incl. Australia]; also see Tables
Citation of literature follows BPH-2 (
Common names and uses given under each species treatment are recorded for verified specimens only. The aim here is to provide an authoritative summary; further details on uses and names can be checked from the literature cited. Similarly, we refer to chromosome counts based on voucher specimens that we have been able to verify or based on studies that describe the voucher material in adequate enough morphological detail that allows us to draw firm conclusions on the identity of the original material. This means that some studies (e.g.
Solanum grad. amig. Maurella Dunal, Hist. Solanum 119, 151. 1813. Lectotype species: S. nigrum L. (designated by
Solanum section Morella Dumort., Fl. Belg. 39. 1827. Lectotype species: S. nigrum L. (designated by
Solanum section Inermis G.Don, Gen. Syst. 4: 400. 1838. Lectotype species: S. nigrum L. (designated by
Solanum grad ambig. Morella G.Don, Gen. Syst. 4: 411. 1838. Lectotype species: S. nigrum L. (designated by
Solanum section Pachystemonum Dunal, Prodr. [A. P. de Candolle] 13(1): 28, 31. 1852. Lectotype species: S. nigrum L. (designated by
Solanum subsection Morella Dunal, Prodr. [A. P. de Candolle] 13(1): 28, 44. 1852. Lectotype species: S. nigrum L. (designated by
Solanum section Campanulisolanum Bitter, Repert. Spec. Nov. Regni Veg. 11: 234. 1912. Lectotype species: S. fiebrigii Bitter (designated by
Solanum section Episarcophyllum Bitter, Repert. Spec. Nov. Regni Veg. 11: 241. 1912. Lectotype species: S. sinuatirecurvum Bitter (designated by
Solanum section Morella (Dunal) Bitter, Bot. Jahrb. 54: 416, 493. 1917. Lectotype species: S. nigrum L. (designated by
Solanum section Chamaesarachidium Bitter, Repert. Spec. Nov. Regni Veg. 15: 93. 1919. Type species: S. chamaesarachidium Bitter (= S. weddellii Phil.).
Solanum series Transcaucasica Pojark., Bot. Mater. Gerb. Inst. Komarova Akad. Nauk S.S.S.R. 17: 332. 1955. Lectotype species: S. transcaucasicum Pojark. (= S. villosum Mill.) (designated by
Solanum series Alata Pojark., Bot. Mater. Gerb. Inst. Komarova Akad. Nauk S.S.S.R. 17: 336. 1955. Type species: S. alatum Moench [nom. et typ. cons. prop.] (= S. villosum Mill.) (designated by
Solanum series Pseudoflava Pojark., Bot. Mater. Gerb. Inst. Komarova Akad. Nauk S.S.S.R. 17: 338. 1955. Type species: S. pseudoflavum Pojark. (= S. villosum Mill.) (designated by
Solanum section Parasolanum A.Child, Feddes Repert. 95: 142. 1984. Type species. S. triflorum Nutt.
Solanum section Dulcamara (Moench) Dumort. subsect. 2 “herbaceous plants confined to the central Andes” of
Solanum section Solanum subsects. 1 “Solanum”, 2 “Glandular pubescent group”, 3 “Campanulisolanum”, 4 “Chamaesarachidium” and 6 “Episarcophyllum” of
Solanum series Lutea Pojark. ex Ivanina, Bot. Zhurn. (Moscow & Leningrad) 85(6): 144. 2000. Type species. S. villosum Mill.
Description. Herbs, occasionally woody at the base; unarmed. Stems terete or angled, sometimes hollow, lacking true prickles but sometimes with prickle-like processes along the angles, glabrous or pubescent with simple or branched (only in the Americas) uniseriate trichomes, these eglandular or glandular. Sympodial units difoliate or trifoliate, the leaves usually not geminate. Leaves simple with entirely or variously dentate or lobed margins or occasionally deeply pinnatifid, concolorous, glabrous to densely pubescent with eglandular and/or glandular simple or branched (only in the Americas) uniseriate trichomes; petioles well developed or not, the leaves never sessile. Inflorescences opposite the leaves or arising internodally, unbranched or many times branched, not bracteate (except in S. triflorum where a single bracteole sometimes present), with few to many (up to 100) flowers; peduncle various, usually not longer than the inflorescence branches; pedicels articulated at the base. Flowers 5-merous, actinomorphic to very slightly zygomorphic, all perfect. Calyx with the lobes deltate to spathulate to long-triangular. Corolla stellate or rotate-stellate, white or purplish-tinged to lavender, usually with an “eye” at the base of the lobes of a contrasting colour (yellow, green or dark purple-black), the lobes spreading or reflexed at anthesis. Stamens equal or very slightly unequal, the filaments equal, glabrous or more usually densely pubescent with tangled uniseriate weak-walled simple uniseriate trichomes, the anthers ellipsoid (sometimes slightly tapering in S. scabrum) and connivent, with distal pores that elongate to slits with drying and/or age. Ovary conical, glabrous or occasionally very minutely puberulent; style straight or curved and bent, usually pubescent with simple uniseriate trichomes in the lower half, only very slightly exserted from the anther cone; stigma minutely capitate to capitate or clavate. Fruit a globose or somewhat elongate juicy berry with thin pericarp, green, black, yellow or red-orange at maturity; fruiting pedicels spreading or deflexed; fruiting calyx lobes reflexed, appressed or accrescent at fruit maturity. Seeds flattened and tear-drop shaped, yellow or tan-brown. Chromosome number: n=12, 24, 36 (see section on Chromosomes, and individual species treatments).
Distribution. A worldwide species group occurring on all continents except Antarctica, but with highest species diversity in South America and Africa.
Discussion. In the synonymy here, we have included all groups that are members of the clade as we define it, not only those containing Old World species; for more detailed discussion of morphology and group definition see
Members of the Morelloid clade are amongst the most widely collected of solanums, in part because are they are herbaceous, widespread and weedy. They are also amongst the most difficult to identify, due to their extreme vegetative plasticity (see Morphology above) and their lack of striking distinguishing characters. Combinations of characters are most useful for identification and we have included these in the species treatments as well as in the keys. Species are often most usefully identified based on geography (i.e. where they are found), but the large number of potentially invasive and introduced species means one must exercise caution if a species is not readily identifiable (taking into account variation, of course). The many specimens grown in the 19th century in botanical gardens present special difficulties – these are not included here in species treatments or keys, except as type citations. Many morelloid species from the Americas were sent as seeds to European botanical gardens (e.g. S. americanum, S. chenopodioides, S. emulans Raf., S. nigrescens Martens & Galeotti) and, though cultivated, they did not escape and become part of the local floras of the regions. In order not to confuse users, we have limited our treatment here to include only native and a set of non-native introduced species that have become naturalised and persistent in the Old World.
The Morelloid clade suffers from two extreme sorts of taxonomic recognition issues. Firstly, in many parts of the world, especially in more recent floras, all taxa are treated as a single highly variable species (usually S. nigrum) and local endemic taxa are overlooked. Secondly and especially in Europe in the late 19th and early 20th century, many minor variants were described and then were transferred and recombined at different taxonomic levels, creating a confusing morass of names, many of which lack types. The latter is unfortunate because of the nomenclatural work entailed in sorting out the identities and types for these names, but the former is more serious, because endemic taxa have been overlooked and thus have possibly been placed at risk due to their being equated with widespread invasive weeds.
1 | Leaves 3-lobed nearly to the midrib; Australia (introduced and coastal) | S. palitans |
1' | Leaves entire, sinuate-dentate or pinnatifid, but never regularly 3-lobed | 2 |
2 | Leaves shallowly to deeply pinnatifid | S. triflorum |
2' | Leaves entire to sinuate-dentate | 3 |
3 | Glandular trichomes present (e.g. along stems, petioles and leaves), plants usually sticky to touch when fresh | 4 |
3' | Glandular trichomes absent (e.g. along stems, petioles and leaves), plants not sticky to touch when fresh | 17 |
4 | Anthers < 1.8 mm long | 5 |
4' | Anthers ≥ 1.8 mm long | 9 |
5 | Inflorescences with 10–40 flowers; pedicels spaced 1–2 mm apart, sharply bent at the base (near articulation point) in flower and fruit; Africa | S. tarderemotum |
5' | Inflorescences with 2–5(-10) flowers; pedicels spaced 0–1 mm apart, nodding, erect or spreading in flower and fruit, reflexed and slightly curved in some species in fruit but never in flower; pantropical | 6 |
6 | Calyx lobes 1–1.5 mm long in flower; fruiting calyces not accrescent, the tube remaining 1–1.7 mm long and lobes 1–1.5 mm long; fruit black when ripe, not markedly shiny | 7 |
6' | Calyx lobes 1.5–2.5 mm long in flower; fruiting calyces accrescent, the tube 3–4 mm long and lobes 2.5–8.0 mm long; fruit green when ripe, shiny | 8 |
7 | Leaves rhomboidal to lanceolate in shape; inflorescences unbranched, never furcate; buds globose; calyx lobes all equal in size, strongly reflexed in fruit; berries with a glaucous cast; southern Africa (also cultivated; introduced in Australia) | S. retroflexum |
7' | Leaves elliptic to slightly ovate; inflorescences unbranched or rarely furcate; buds ellipsoid; calyx lobes unequal with two larger lobes, appressed to somewhat spreading in fruit; berries matte, but not markedly glaucous; Australasia and the Pacific | S. opacum |
8 | Leaf bases attenuate to cuneate; inflorescences mostly internodal, with 4–8(-10) flowers; pedicels spaced 0.3–1 mm apart; calyx lobes 1.7–2.5 mm long; corollas with yellow-green central eye with black-purple “V”-shaped margins; anthers 1.0–1.4 mm long; berries dark green to green-brown marbled with white lines, becoming usually translucent and shiny, lower half of berries covered with enlarged calyces but berry mostly visible; seeds brown; stone cells (1-)2–3, these 0.5 mm in diameter | S. nitidibaccatum |
8' | Leaf bases truncate; inflorescences mostly leaf-opposed, with 2–5(-7) flowers; pedicels spaced 0(-1) mm apart; calyx lobes 1.5–2.0 mm long; corolla with yellow-green or translucent basal star without black-purple colouration; anthers 1.2–2.0 mm long; berries pale green, shiny becoming dull, opaque, usually completely enveloped by enlarged calyces; seeds pale yellow; stone cells 4–6, these (0.5-)0.8–1 mm in diameter | S. sarrachoides |
9 | Anthers ≥ 2.8 mm long | 10 |
9' | Anthers < 2.8 mm long | 12 |
10 | Inflorescences with bracteoles present in most individuals; buds narrowly ellipsoid; corolla lobes narrowly lanceolate; berries with >30 stone cells | S. triflorum |
10' | Inflorescences never with bracteoles; buds globose, ovoid or narrowly ellipsoid; corolla lobes variously triangular; berries with (0)2–4 stone cells | 11 |
11 | Flower buds globose to ovoid; calyx lobes 1–1.5(-2) mm long, long-triangular with rounded tips; corolla rotate-stellate; Arabian Peninsula to Egypt and south in eastern Africa | S. memphiticum |
11' | Flower buds narrowly ellipsoid; calyx lobes 0.5–1.0 mm long, broadly deltate to triangular with acute tips; corolla stellate; Java to Lesser Sunda Islands | S. alpinum |
12 | Calyx lobes appressed to spreading in fruit, never strongly reflexed | 13 |
12' | Calyx lobes strongly reflexed in fruit | 15 |
13 | Calyx accrescent in fruit, calyx tube 3–4 mm long and lobes 2.5–8 mm long | S. sarrachoides |
13' | Calyx not accrescent in fruit, calyx tube 1–2 mm long and lobes 1–1.5 mm long | 14 |
14 | Buds ellipsoid; calyx tube 1.5–2.0 mm long, lobes 1–1.5 mm long, elongate-deltate with rounded tips; fruiting pedicels persisting when fruits mature and fall off; Cameroon line (Cameroon and Equatorial Guinea), usually above 2,000 m elevation | S. pseudospinosum |
14' | Buds subglobose; calyx tube 0.8–1.0 mm long, lobes 0.5–0.8 mm long, triangular with rounded to acute tips; fruiting pedicels generally not persisting and falling off with mature fruits; common across Old World especially in temperate areas or below 1,000 m in the tropics and subtropics (Asia) | S. nigrum |
15 | Leaves rhomboidal to lanceolate; filaments 1.2–1.5 mm long, anthers 1.3–1.8(-2) mm long; seeds 1.6–1.8 mm long and 1.3–1.5 mm wide | S. retroflexum |
15' | Leaves broadly to narrowly ovate to elliptic; filaments 0.5–1.3 mm long; anthers 1.8–2.5 mm long; seeds 1.8–2.2 mm long and 1.5–1.7 mm wide | 16 |
16 | Calyx with broad and relatively transparent sinuses, lobes elliptic to triangular, rounded at tip; free part of the filaments 1.0–1.3 mm long; mature berries slightly ellipsoid, shiny yellow, orange or red; stone cells always absent | S. villosum |
16' | Calyx with narrow, sharp triangular sinuses, lobes deltate with acute or rounded tips; free part of the filaments 0.5–0.7 mm long; mature berries round, dull black or green; stone cells 0–4 | S. nigrum |
17 | Anthers < 1.8 mm long | 18 |
17' | Anthers ≥ 1.8 mm long | 21 |
18 | Pedicels spaced 1–2 mm apart, pedicels sharply bent at the base (near the articulation point) in flower and fruit; Africa | S. tarderemotum |
18' | Pedicels spaced 0–0.5 mm apart, pedicels nodding, erect or spreading in flower and fruit; pantropical | 19 |
19 | Leaves with entire margins, occasionally sinuate-dentate; calyx lobes 0.3–0.5 mm long in flower, 1(-2) mm in fruit; mature fruits black, the surface very shiny | S. americanum |
19' | Leaves shallowly toothed, occasionally entire; calyx lobes 1.0–1.5 mm long in flower, 1.5–2 mm in fruit; mature fruits purple-black or green, the surface matte | 20 |
20 | Leaves rhomboidal to lanceolate in shape; inflorescences unbranched, never furcate; buds globose; calyx lobes all equal in size, strongly reflexed in fruit; southern Africa (also cultivated; introduced in Australia) | S. retroflexum |
20' | Leaves elliptic to slightly ovate; inflorescences unbranched or rarely furcate; buds ellipsoid; calyx lobes unequal with two larger lobes, appressed to somewhat spreading in fruit; Australasia and the Pacific | S. opacum |
21 | Anthers < 2.8 mm long | 22 |
21' | Anthers ≥ 2.8 mm long | 31 |
22 | Berries without stone cells | 23 |
22' | Berries with 2–22 stone cells | 28 |
23 | Pedicels persisting and not dropping with mature fruits; calyx lobes in fruit mostly strongly reflexed | 24 |
23' | Pedicels dropping with mature fruits; calyx lobes in fruit appressed to slightly spreading, rarely strongly reflexed | 26 |
24 | Leaves rhomboidal to lanceolate; filaments 1.2–1.5 mm long, anthers 1.3–1.8(-2) mm long; seeds 1.6–1.8 mm long and 1.3–1.5 mm wide | S. retroflexum |
24' | Leaves broadly to narrowly ovate to elliptic; filaments 0.5–1.3 mm long; anthers 1.8–2.5 mm long; seeds 1.8–2.2 mm long and 1.5–1.7 mm wide | 25 |
25 | Calyx with broad and relatively transparent sinuses, lobes elliptic to triangular, rounded at tip; filaments 1.0–1.3 mm long; mature berries slightly ellipsoid, shiny yellow, orange or red; stone cells always absent | S. villosum |
25' | Calyx with narrow, sharp triangular sinuses, lobes deltate with acute tips; filaments 0.5–0.7 mm long; mature berries round, dull black or green; stone cells generally absent (2–4 stone cells common in Asian material) | S. nigrum |
26 | Buds elongate-oblong; fruiting peduncles strongly deflexed at the base (bent downwards at junction with the stem) | S. chenopodioides |
26' | Buds ellipsoid to subglobose; fruiting peduncles straight or ascending | 27 |
27 | Pedicels spaced 1–2 mm apart, sharply bent at the base (near the articulation point) in flower and fruit; seeds 1.5–2 mm long and 1–1.5 mm wide; Africa | S. tarderemotum |
27' | Pedicels spaced 0–0.7 mm apart, straight, spreading or reflexed in flower and fruit; pantropical; seeds 1.8–2 mm long and 1.5–1.6 mm wide; widespread | S. nigrum |
28 | Prostrate herb; leaves narrowly elliptic to lanceolate, base strongly attenuate; inflorescences with 1–5 flowers; pedicels stout and spreading; calyx lobes linear-oblong with rounded apices; mountains of Ethiopia | S. hirtulum |
28' | Upright or spreading herb; leaves broadly ovate to elliptic, base acuminate, acute, obtuse, truncate to abruptly attenuate; inflorescences with 2–40 flowers; pedicels spreading to strongly reflexed; calyx lobes triangular, broadly deltoid or ovate with acute to rounded apices | 29 |
29 | Pedicels strongly bent at the base (near articulation point) in flower and fruit | S. tarderemotum |
29' | Pedicels spreading, stout or pendent in flower, occasionally recurved in fruit but never strongly bent at the base | 30 |
30 | Inflorescences unbranched or more often branched, often with small leaves (bracteoles?); calyx lobes broadly deltate to mere enations of the rim; style exserted 1.0–1.5 mm beyond anther cone; mature berries 3–4(-5) mm in diameter, dull yellowish-brown | S. umalilaense |
30' | Inflorescences unbranched, never with small leaves; calyx lobes triangular; style exserted 0–1 mm beyond anther cone; mature berries 6–10 mm in diameter, dull black | S. nigrum |
31 | Inflorescences with bracteoles present in most individuals; buds narrowly elliptic; berries with >30 stone cells | S. triflorum |
31' | Inflorescences never with bracteoles; buds globose, ovoid or narrowly ellipsoid; berries with 0–14 stone cells | 32 |
32 | Berries without stone cells | 33 |
32' | Berries with 2–14 stone cells | 35 |
33 | Buds narrowly ellipsoid; pedicels 1.0–1.3 cm long; anthers 2.8–3.8(-4) mm long | S. alpinum |
33' | Buds elongate-oblong to globose; pedicels 0.5–1.0 cm long; anthers (2.0-) 2.3–3.0 mm long | 34 |
34 | Buds elongate-oblong; calyx lobes broadly deltate to triangular with acute tips; fruiting peduncles strongly bent at the base near junction with the stem; fruiting pedicels reflexed and slightly recurved; seeds 1.2–1.4 mm long and 1.0–1.2 mm wide | S. chenopodioides |
34' | Buds globose-subglobose; calyx lobes broadly deltate with rounded tips; fruiting peduncles straight; fruiting pedicels stout, erect and spreading; seeds 2–2.8 mm long and 1.5–1.8 mm wide | S. scabrum |
35 | Leaves narrowly elliptic to lanceolate, bases strongly attenuate | 36 |
35' | Leaves lanceolate, ovate or rhomboidal, bases cuneate-acute to occasionally attenuate | 37 |
36 | Calyx lobes 1.5–1.8 mm long, elliptic with long-acuminate to acute tips; anthers 3.0–3.8 mm long; Europe, occasional adventive associated with human activity, at or close to sea level | S. pygmaeum |
36' | Calyx lobes 0.8–1.2 mm long, linear-oblong with rounded tips; anthers 2.3–2.8 mm long; Ethiopia, usually above 2,000 m elevation | S. hirtulum |
37 | Leaves ovate to rhomboidal; inflorescences furcate, occasionally simple; calyx lobes rectangular to narrowly obovate with obtuse to shortly acute tips; Australia and New Zealand (introduced at or close to sea level) | S. furcatum |
37' | Leaves lanceolate to ovate, most commonly narrowly elliptic or broadly lanceolate; inflorescences always simple; calyx lobes broadly deltate to triangular with acute tips; Java and Lesser Sunda Islands, usually above 2,000 m elevation | S. alpinum |
Solanum viscidissimum Zoll. & Moritzi, Natuur- Geneesk. Arch. Ned.-Indië 2: 571. 1845.
Type. Indonesia. Java: Tengger [Tengger Range], above Gebok-Klacca [Klakah], 5500 ft, Oct 1844, H. Zollinger 2514 (lectotype, designated here: G-DC [G00357866]; isolectotypes: BM [BM000778312], G [G00144593, G00343069], P [P00379791, P00379790], W [1889-0052800]).
Solanum dichrophyllum Dunal, Prodr. [A. P. de Candolle] 13(1): 48. 1852, nom. illeg. superfl.
Type. Based on (renaming of) Solanum alpinum Zoll. & Moritzi
Solanum anacamptocarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 59. 1852.
Type. Indonesia. Java: Waliran [Gunung Welirang], 27 Aug 1844, H. Zollinger 2177 (holotype: G-DC [G00144590]; isotypes: A [A00248838], BM [BM000886120], G [G00343304], MPU [MPU012652], P [P00369059, P00369060 [mixed sheet with right-hand stem belonging to S. nigrum], P00368907, P00368908]).
Solanum bromoense Kuntze, Revis. Gen. Pl. 453. 1891.
Type. Indonesia. Java: Mount Tengger, Bromo [Gunung Bromo], 2000 m, 15 Sep 1875, O. Kuntze 5999 (lectotype, designated here: NY [0017228]; isolectotypes: GH [GH00077829], K [K000788117]).
Indonesia. Java: “in montibus Ardjune” [Gunung Arjuno], 2000–3000m, 14 Sep 1844, H. Zollinger 2255 (lectotype, designated here: P [P00368905] (original label on this sheet with date and locality – “an 2177 var?”); isolectotypes: BM [BM000886119], G [G00144225, G00301669], MPU [MPU014201], P [P00368906]).
Annual or short-lived erect to somewhat spreading perennial herbs, height not known, likely subwoody and branching at base. Stems spreading to decumbent, terete, sometimes recorded as greenish-violet (Afriastini 475), older stems yellowish-brown, with no prickle-like projections, not markedly hollow; new growth densely pubescent with simple, spreading, uniseriate, glandular or eglandular trichomes, the trichomes 6-8(-10)-celled, ca. 0.5 mm long, if glandular then with a terminal gland, sometimes drying with a yellowish-brown tinge. Sympodial units trifoliate to plurifoliate, the leaves not geminate. Leaves simple, (2-)5–12 cm long, (0.5-)1–3 cm wide, lanceolate to ovate, most commonly narrowly elliptic or broadly lanceolate, membranous, concolorous, smell not known; adaxial surface pubescent with simple, uniseriate, glandular or eglandular trichomes evenly and moderately spread along veins and lamina; abaxial surface similar but the pubescence denser along the veins; major veins 7–8 pairs; base cuneate to attenuate; margins entire or shallowly toothed, the teeth, if present, narrow and acute; apex acute to acuminate; petioles ca. 1 cm long, pubescent like the stems and leaves. Inflorescences 2–4 cm long, internodal, unbranched or furcate, with 5–10 flowers mostly clustered at the distal end of the rhachis, pubescent with simple uniseriate trichomes like those of the stems and leaves; peduncle 1.0–3.5 cm long, straight; pedicels 1.0–1.3 cm long, ca. 0.5 mm in diameter at base and apex, filiform, spreading, pubescent with simple uniseriate trichomes like the rest of the inflorescence, articulated at the base; pedicel scars mostly clustered at the distal end of the rhachis and overlapping, sometimes up to 1.5 mm apart in the distal half of the rhachis. Buds narrowly ellipsoid, the corolla soon exserted from the calyx tube. Flowers 5-merous, all perfect. Calyx tube 1–2 mm long, broadly conical, the lobes 0.5–1.0 mm long, ca. 1 mm wide, broadly deltate to triangular, tips acute, densely to moderately pubescent with simple uniseriate 6–8-celled trichomes. Corolla 9–12 mm in diameter, purple or violet, stellate, lobed 2/3 to 3/4 of the way to the base, the lobes 3–4.5 mm long, 1.5–2.0 mm wide, spreading or reflexed, densely papillate along the margins and on the tips. Stamens equal; filament tube < 0.2 mm long; free portion of the filaments ca. 1 mm long, densely pubescent adaxially with tangled simple uniseriate trichomes; anthers 2.8–4 mm long, ca. 1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary rounded, glabrous; style 5–6 mm long, densely pubescent with tangled simple uniseriate trichomes in the basal half, exserted up to 2 mm from the anther cone; stigma minutely bilobed, the surfaces minutely papillate. Fruit a globose berry, 6–9 mm in diameter, black, the pericarp thin and matte; fruiting pedicels 1.2–1.3 cm long, ca. 0.5 mm in diameter at the base and apex, spreading or strongly deflexed, spaced (0-)0.5–1.5 mm apart, not falling with the berry, persistent on older inflorescences; fruiting calyx not accrescent, the tube less than 1 mm long, the lobes ca. 1 mm long, loosely appressed to the berry, not markedly reflexed. Seeds 20–50 per berry, 2.5–3.0 mm long, ca. 1 mm wide, flattened reniform, golden brown to brown, the surfaces minutely pitted, the testal cells small, rectangular to pentagonal in shape. Stone cells (0)2–4 per berry. Chromosome number not known.
Grows in montane habitats, in grassy areas along forest margins or in clearings; between 1,700 and 3,100 m elevation.
Indonesia: ranti hutan.
Uses. No specific uses recorded, but labels (Afriastini 475) indicate young berries are “edible but rather bitter”.
(
Solanum alpinum is a plant of high elevations with both glandular and eglandular morphs. Glandular forms were named as S. viscidissimum, while in the same publication eglandular forms were named S. alpinum. As the two names have not been synonymised before (neither name was included in
The plant illustrated as “Solanum nigrum” in
A single collection of Zollinger was cited for each of S. alpinum (“Herb. N. 2255”) and S. viscidissimum (“Herb. N. 2514”), but no herbarium was cited; we have lectotypified S. alpinum with the P (P00368905) duplicate of Zollinger 2255, because it has a label with the protologue locality. We have selected the G-DC (G00357866) sheet of Zollinger 2514 as the lectotype for S. viscidissimum as the best preserved of the duplicates we have seen.
C.E.O. Kuntze named S. bromoense for Gunung Bromo in the Tenggar range, citing only “Java. Bromo 2000m” (
Indonesia. Bali: bei der Quelle Jaritie auf Weg zum Gunung Ajaung, 2 Jun 1912, Arens 19 (L); Kleine Soenda Eilanden, Bali, Z. helling Agoeng, 6 Apr 1936, van Steenis 7839 (K);. Java: Central Java, Blumbang, Mt. Lawu, Central Java, 26 Nov 1982, Afriastini 475 (A); West Java, Mt.Malabar, Anderson 369 (CAL); East Java, Ardjoeno, tjemarabosch boven Lalidjiwo, 17 Oct 1915, Arens s.n. (L); East Java, 12 Oct 1915, Arens 48 (L); East Java, Pasoeroean, G[unung] Tengge, boven Tosari, 4 Jun 1913, Backer 8380 (L); East Java, Te Pasoeroean, Ngadisari, Jan 1925, Backer 36563 (A); East Java, Pasoeroean, Tengge, boven Tosari, Backer 36564 (L); Central Java, Soerkarta, Top van de Lawoe [Mount Lawu], 16 Jul 1936, Brinkman 754 (NY); Sitiebondo, G[unung] Raneg [Raoeng] via Brembeinri, 15 May 1932, Clason-Laarman EHH-157 (L); East Java, SE Java, 1880, Forbes 1019 (BM); Central Java, Central Java, Slamet Mountain, between base camp Beraba-top of summit, 17 Mar 2004, Hoover et al. Deden-113 (A); Central Java, Mt. Prahu, Horsfield s.n. (BM); Central Java, Surakarta, Horsfield s.n. (BM); Central Java, Mt. Prahu, Horsfield s.n. (BM); Central Java, Blambangan & Mt. Prahu, Horsfield s.n. (BM); East Java, Pasoeroean G[unung] Smeroe, Jesweit s.n. (L); Central Java, Tenggu, Penandjaan, 31 Jul 1932, Kleinhoonte 371 (L); Central Java, Besoeki, G[unung] Merapi, 18 Jul 1916, Koorders & Koorders-Schumacher 43218 B (L); East Java, Res. Besoeki Kawah Idjen, 1 Nov 1916, Koorders & Koorders-Schumacher 43214B (L); East Java, Besoeki, Jang Plateau, 12 Aug 1916, Koorders & Koorders-Schumacher 43463 (L); East Java, Tdjan plateau, Mt. Raung, Nov 1938, Kostermans s.n. (L); Central Java, G[unung] Muria, Tjollo, N of Kudus, 25 Nov 1951, Kostermans 6278 (L); West Java, Malawar, Dec 1875, Kuntze 5410 (DUKE, NY); West Java, Gunong Gede, 6 Mar 1979, Murata et al. 2021 (L); East Java, Pasoeroean, 4 Jun 1935, Neth Ind Forest Service 7038 (L); East Java, Pasoeroean, 4 Jun 1935, Neth Ind Forest Service 7056 (A); East Java, Tosari 49, Proefstation voor de Javasuikerindustrie s.n. (L); East Java, Tosari 52, Proefstation voor de Javasuikerindustrie s.n. (L); Central Java, Central Java Merbaboe, Jul 1922, Roorda van Eysinga s.n. (MO); East Java, Tosari, Pasoeroean, Teysmann s.n. (CAL, K); East Java, Tengergebergte tusschen Ngadisari, Zandzee en Tosari, Went s.n. (L); Central Java, Meapi, Catu septr, Without collector s.n. (U); [iter Java secundum], Zollinger 1790 (LE, P). West Nusa Tenggara: Klein Soenda Eilanden, Plawangan, Segara anak, 16 Jun 1936, de Voogd 2654 (GH, NY); Lombok, Rindjani Vulkangebirge N Seite, 9 May 1909, Elbert 1327 (L).
Solanum nigrum var. patulum L., Sp. Pl. 186. 1753.
Type. “Solanum procerius patulum, vulgaris fructu”, cultivated in England, at James Sherard’s garden in Eltham (Hortus Elthamensis) (lectotype, designated here: Dillenius, Hortus Elthamensis 2: 367, t. 275, f. 355. 1732). “Solanum procerius patulum, vulgaris fructu”, Herb. Dillenius 441 (epitype, designated here [as lectotype by
Solanum nodiflorum Jacq., Collectanea [Jacquin] 2: 288. 1789.
Type. Cultivated in Austria at Vienna, said to be from Mauritius (“crescit insula Mauritii”), Herb. Jacquin s.n. (lectotype, designated by
Solanum patulum (L.) Roth, Catal. Bot. 2: 23. 1800.
Type. Based on Solanum nigrum var. patulum L.
Solanum papilionaceum Dum.Cours., Bot. Cult. 2: 135. 1802.
Type. Cultivated from seeds received from the Jardin Nat. [Paris] (no specimens cited, likely to have been described from living material). Cultivated in Paris (“H.P.”), 1825, Anon. [Herb. Maire] s.n. (neotype, designated here: P [P00582223]).
Solanum strictum Zuccagni, Cent. Observ. Bot. [p. 20] No. 49. 1806.
Type. Cultivated in Italy; “Semina nobis communicavit Cl. Thouin sub nomine S. nigri sp. nova” (no specimens cited; Zuccagni’s herbarium originally at FI but destroyed). Cultivated in Italy in Bologna “j. de Bologna”, 6 Jul 1808, Anon. s.n. (neotype, designated here: G-DC [G00144215]).
Solanum rumphii Dunal, Hist. Nat. Solanum 157. 1813.
Type. Indonesia. Malaku: Amboina (no specimens cited; lectotype, designated here: Rumphius, Herbarium Amboinenese 6: t. 26, fig. 2, 1750).
Solanum oleraceum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 750. 1814.
Type. “Antilles” Herb, Richard s.n. (lectotype, designated by
Solanum dillenii Schult., Öster. Fl., ed. 2, 1: 393. 1814, as “Dilleni”
Type. Hungary?. “In umbrosis Matra [Mátra]”, P. Kitaibel s.n. (lectotype, designated here: BP [Herb. Kit. fasc. IX, No. 102, small-flowered stem only]; isolectotype: B-W [B-W 04364-03]).
Solanum microspermum Dunal, Solan. Syn. 12. 1816.
Type. Origin unknown, 1815, Anon. (Herb. Thibaud) s.n. (lectotype, designated here: G-DC [G00144267]).
Solanum erythrocarpon G.Mey., Prim. Fl. Esseq. 109. 1818.
Type. Suriname. Saramacca: Hamburg (Essequibo), E.K. Rodschied 31 (lectotype, designated here: GOET [GOET003505]).
Solanum desvauxii Ham., Prod. Pl. Ind. Occ. 26. 1825, nom. illeg. superfl.
Type. Based on S. nodiflorum Jacq. (cited in synonymy).
Solanum nigrum Vell., Fl. Flumin. 85. 1829 [1825], nom. illeg., not Solanum nigrum L. (1753)
Type. Brazil. [Rio de Janeiro]: “undequaeque nascitur” (lectotype, designated by
Solanum tenuiflorum Steud., Nomencl. ed. 2, 2: 606. 1841.
Type. Based on (replacement name for) Solanum nigrum Vell.
Solanum indecorum A.Rich., Hist. Fls. Cuba, Fanerogamia 11: 121. 1841.
Type. Cuba. Sin loc., 1836, R. de la Sagra s.n. (lectotype, designated here: P [P00370899]).
Solanum nigrum subsp. nodiflorum (Jacq.) Sendtn., Fl. Bras. (Martius) 10: 16. 1846.
Type. Based on Solanum nodiflorum Jacq.
Solanum nigrum var. angulosum Sendtn., Fl. Bras. (Martius) 10: 16. 1846, as Solanum nigrum subsp. nodiflorum var. angulosum Sendtn.
Type. Based on Solanum tenuiflorum Steud. (=Solanum nigrum Vell.)
Solanum nigrum subsp. aguaraquiya Sendtn., Fl. Bras. (Martius) 10: 17. 1846.
Type. Brazil. Rio Grande do Sul: “Pat. Joan a St. Barbara”, C.F.P. Martius s.n. (lectotype, designated here: M [M-0171809]; isolectotype: M [M-0171810]).
Solanum nigrum var. minus Hook.f., Trans. Linn. Soc. London 20(2): 201. 1847, as “minor”
Type. Ecuador. Galápagos Islands: James Island [Santiago], C. Darwin s.n. (lectotype, designated here: CGE [CGE00297]; isolectotype: K [K000922162]).
Solanum amarantoides Dunal, Prodr. [A. P. de Candolle] 13(1): 55. 1852.
Type. Brazil. Rio de Janeiro, C. Gaudichaud 522 (lectotype, designated by
Solanum pterocaulum var. aguaraquiya (Sendtn.) Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, as ‘pterocaulon’.
Type. Based on Solanum nigrum subsp. aguaraquiya Sendtn.
Solanum ptychanthum Dunal, Prodr. [A. P. de Candolle] 13(1): 54. 1852.
Type. United States of America. Georgia: Chatham Co., Savannah, Anon. s.n. (holotype: G-DC [G00144485]).
Solanum nodiflorum var. macrophyllum Dunal, Prodr. [A. P. de Candolle] 13(1): 46. 1852.
Type. Brazil. Rio de Janeiro: Rio de Janeiro, C. Gaudichaud 521 (lectotype, designated by
Solanum nodiflorum var. acuminatum Dunal, Prodr. [A. P. de Candolle] 13(1): 46. 1852.
Type. Brazil. Minas Gerais: Sin loc., M. Vauthier 537 (lectotype, designated by
Solanum nodiflorum var. petiolastrum Dunal, Prodr. [A. P. de Candolle] 13(1): 46. 1852.
Type. Brazil. Rio de Janeiro: Novo Friburgo, 1842, P. Claussen 180 (holotype: P [P00319584]).
Solanum inops Dunal, Prodr. [A. P. de Candolle] 13(1): 55. 1852.
Type. Mexico. “sin. loc.” [Tamaulipas: Tampico, 4 Feb 1827], J.L. Berlandier 46 (holotype: G-DC [G00144469]; isotypes: BM [BM000775579], F [V0073104F, acc. # 680275], LE, P [P00336046, P00336047, P00336048], W [1889-0291394, 1889-0144848]).
Solanum nigrum forma nodiflorum (Jacq.) Miq., Fl. Ned. Ind. 2: 637. 1856.
Type. Based on Solanum nodiflorum Jacq.
Solanum nigrum forma rumphii (Dunal) Miq., Fl. Ned. Ind. 2: 637. 1856.
Type. Based on Solanum rumphii Dunal
Solanum nigrum forma nodiflorum (Jacq.) Miq., Fl. Ned. Ind. 2: 637. 1856.
Type. Based on Solanum nodiflorum Jacq.
Solanum nigrum forma uniflorum Miq., Fl. Ned. Ind. 2: 638. 1856.
Type. Indonesia. Java: “op den Diëng, 6000-8000 ft”, F.W. Junghuhn s.n. (lectotype, designated here: U [U0113977]).
Solanum
patulum
Kit. ex Kanitz, Linnaea 32: 440. 1863, nom illeg., not Solanum patulum (L.)
Type. Based on Solanum dillenii Schult. (cited in synonymy)
Solanum nigrum subsp. dillenii (Schult.) Schur, Enum. Pl. Transsilv. 478. 1866.
Type. Based on Solanum dillenii Schult.
Solanum nigrum var. dillenii (Schult.) A.Gray, Synopt. Fl. N. Amer. 2(1): 228. 1878.
Type. Based on Solanum dillenii Schult.
Solanum nigrum var. nodiflorum (Jacq.) A.Gray, Synopt. Fl. N. Amer 2(1): 228. 1878.
Type. Based on Solanum nodiflorum Jacq.
Solanum nigrum var. oleraceum (Dunal) Hitchc., Rep. Missouri Bot. Gard 4: 111. 1893.
Type. Based on Solanum oleraceum Dunal
Solanum
nigrum
var.
nodiflorum
(Jacq.) Hitchc., Rep. (Annual) Missouri Bot. Gard. 4: 111. 1893, nom. illeg., not Solanum nigrum var. nodiflorum (Jacq.) A.
Type. Based on Solanum nodiflorum Jacq.
Solanum nigrum var. americanum (Mill.) O.E.Schulz, Symb. Antill. (Urban) 6: 160. 1909.
Type. Based on Solanum americanum Mill.
Solanum nigrum forma grandifolium O.E.Schulz, Symb. Antill. (Urban) 6: 160. 1909, as Solanum nigrum var. americanum forma grandifolia O.E.Schulz
Type. Puerto Rico. “prope Cayey in sylvis ad rivulum superiorem m. Sept. fl. et. fr.”, P.E.E. Sintenis 2429 (probably type, no herbarium cited).
Solanum nigrum forma parvifolium O.E.Schulz, Symb. Antill. (Urban) 6: 160. 1909, as Solanum nigrum var. americanum forma parvifolia O.E.Schulz.
Type. Cuba. La Habana: Santiago de las Vegas, “Baker Herb. Cub. 3377” (probably type, no herbarium cited).
Solanum minutibaccatum Bitter, Repert. Spec. Nov. Regni Veg. 10: 549. 1912.
Type. Bolivia. La Paz: San Carlos, bei Mapiri, 750 m, Aug 1907, O. Buchtien 1443 (lectotype, designated here: US [00027684, acc. # 1175843]; isotypes: GOET [GOET003478], NY [00172089]).
Solanum inconspicuum Bitter, Repert. Spec. Nov. Regni Veg. 11: 204. 1912.
Type. Peru. Lima: Lima, 12 Jul 1910, C. Seler 222 (holotype: B, destroyed; no duplicates found).
Solanum tenellum Bitter, Repert. Spec. Nov. Regni Veg. 11: 219. 1912.
Type. Brasil. Minas Gerais: “Prope urbem Caldas florens fructibusque instructum”, 4 Oct 1869, A.F. Regnell III 970 (holotype: UPS; isotypes: US [00027821, acc. # 201069, 01931849, acc. # 201352]).
Solanum minutibaccatum subsp. curtipedunculatum Bitter, Repert. Spec. Nov. Regni Veg. 11: 205. 1912.
Type. Bolivia. La Paz: Guanai-Tipuani, Apr-Jun 1892, M. Bang 1462 (holotype: W; isotypes: BM [BM000617672], E [E00106087], M [M-0171808], MO [MO-503647], NDG [NDG42278], NY [00172090, 00172091, 00172092], PH [PH00030453], US [00027685, acc. # 1324656], WIS [0256198WIS]).
Solanum sciaphilum Bitter, Repert. Spec. Nov. Regni Veg. 11: 220. 1912.
Type. Brazil. Santa Catarina: Pedras Grandes, Aug 1890, E. Ule 1678 (holotype: B, destroyed, F neg. 2851; lectotype, designated here: HBG [HBG-511539]; isolectotype: HBG [HBG-511540]).
Solanum curtipes Bitter, Repert. Spec. Nov. Regni Veg. 11: 228. 1912.
Type. Paraguay. Cordillera: San Bernardino, Aug 1898-1899, É. Hassler 3104 (holotype: B, destroyed; lectotype, designated by
Solanum calvum Bitter, Repert. Spec. Nov. Regni Veg. 12: 81. 1913.
Type. Mexico. Baja California: Guadalupe Island, 1875, E. Palmer 60 [pro parte] (holotype: UPS; isotypes: BM [BM001017192], MO [MO-159620, MO-568722], NY [00138967, 00759880], YU [YU065319]).
Solanum depilatum Bitter, Repert. Spec. Nov. Regni Veg. 12: 88. 1913.
Type. Madagascar. Toliara: Fort Dauphin [anchorage], 1897, G. Paroisse 10 (holotype: P [P00338747]).
Solanum imerinense Bitter, Bot. Jahrb. Syst. 49: 566. 1913.
Type. Madagascar. Antananarivo: “Central Madagaskar, Imerina”, Dec 1880, J.M. Hildebrandt 3796 (lectotype, designated by
Solanum sancti-thomae Bitter, Bot. Jahrb. Syst. 49: 560. 1913, as “Sancti Thomae”
Type. São Tome and Principe: São Tome, F. Quintas & A. Moller 47 (syntypes: B, destroyed, COI, not located).
Solanum nodiflorum var. sapucayense Chodat, Bull. Soc. Bot. Genève, sér. 2, 8: 150. 1916.
Type. Paraguay. Paraguarí: Sapucaí [“Sapucay”], 1914, R. Chodat & W. Vischer 46 (holotype: G [G00306708]).
Solanum nigrum subsp. dillenii (Schult.) Probst, Mitteil. Naturfor. Gesellsch. Solothurn 9: 33. 1932.
Type. Based on Solanum dillenii Schult.
Solanum nigrum var. pauciflorum Liou, Contr. Inst. Bot. Natl. Acad. Peiping 3: 454. 1935.
Type. China. Hainan: Ngai District, Yeung Ling Shan, 5 Jun 1932, S.K. Lau 209 (lectotype, designated here: BM [BM000942311]; isolectotypes: A, LU?, K [K001152446]).
Solanum merrillianum Liou, Contr. Inst. Bot. Nat. Acad. Peiping 3: 455. 1935.
Type. China. Hainan: Thai Hang, Shek Kuet Ts’o, Lin Fa Shan and vicinity, Lam Ko District, W.T. Tsang 412 (holotype: LU [acc. no. 15911, not seen]; isotypes: A [A00077824, A00395157], E [E00718800], MO [acc. # 1037660], S [acc. # S-G-5703]).
Solanum photeinocarpum Nakam. & Odash., J. Soc. Trop. Agric., Taiwan 8: 54. 1936.
Type. Taiwan. “Taihoku” [Taipei?], 28 Feb 1936, K. Odashima 17720 (lectotype, designated here: TAI).
Solanum pachystylum Polg., Trans.& Proc. Roy. Soc. N. Z. 69: 280. 1940.
Type. New Zealand. North Island: Auckland, Mt. Wellington, near Auckland, Plant Research Station, H.H. Allan s.n. (holotype: CHR [8954]; isotypes: CHR [8954 A], CHR [8954 B]).
Solanum americanum var. nodiflorum (Jacq.) Edmonds, J. Arnold Arb. 52: 634. 1971.
Type. Based on Solanum nodiflorum Jacq.
Solanum suffruticosum var. merrillianum (Liou) C.Y.Wu & S.C.Huang, Fl. Hainan. 3: 586. 1974.
Type. Based on Solanum merrillianum Liou
Solanum nodiflorum subsp. nutans R.J.F.Hend., Contr. Queensland Herb. 16: 30. 1974.
Type. Australia. Queensland: Brisbane, Dept. Primary, Industrial grounds, 3 Jul 1969, R.J.F. Henderson 518 (holotype: BRI [AQ0023172]; isotypes K [K001080528], NSW [NSW568939], MEL [MEL2289999A]).
Solanum americanum var. patulum (L.) Edmonds, Bot. J. Linn. Soc. 75: 171. 1977.
Type. Based on Solanum nigrum var. patulum L.
Solanum americanum subsp. nodiflorum (Jacq.) R.J.F.Hend., Austrobaileya 2: 555. 1988.
Type. Based on Solanum nodiflorum Jacq.
Solanum pauciflorum (Liou) H.Y.Zhang, Bull. Bot. Res., Harbin 19(2): 131. 1999.
Type. Based on Solanum nigrum var. pauciflorum Liou
Cultivated at the Chelsea Physic Garden [in protologue said to “grow naturally in Virginia”], Herb. Miller s.n. (lectotype, designated by
Annual to short-lived erect or weakly scrambling perennial herbs up to 1.5 m tall, subwoody and branching at base. Stems spreading, terete or somewhat angled with ridges, green to somewhat purple tinged, older stems often appearing spinescent, not markedly hollow; new growth pubescent with simple, spreading, uniseriate, translucent, eglandular trichomes, these 2-8-celled, 0.2–0.8 mm long, often clustered along the stem angles; older stems glabrescent, with only the trichome bases persisting as pseudo-spines. Sympodial units difoliate, the leaves not geminate. Leaves simple, 3.5–10.5 cm long, 1.0–4.5 cm wide, ovate to elliptic, membraneous, concolorous, without odour; adaxial surface sparsely pubescent with simple, uniseriate trichomes like those on stem, these evenly spread along the lamina and the veins; abaxial surface similar but more densely pubescent; major veins 3–6 pairs; base attenuate, decurrent on the petiole; margins entire or occasionally sinuate-dentate; apex acute; petioles (0.3-)2.0–3.8(-4.0) cm long, sparsely pubescent with simple uniseriate trichomes like those on stems. Inflorescences 0.6–2.5 cm long, internodal, simple or very rarely furcate, umbelliform to sub-umbelliform, with (3-)4–6(-8) flowers (very rarely with more flowers in branched inflorescences), sparsely pubescent with simple uniseriate trichomes like those on stems; peduncle (0.5-)1.0–1.8 cm long, straight and stout; pedicels 3–9 mm long, 0.2–0.3 mm in diameter at the base and 0.4–0.5 mm at the apex, stout, straight and spreading, articulated at the base; pedicel scars spaced 0–0.5 mm apart, clustered at the tip of the inflorescence. Buds broadly ellipsoid, the corolla 1 exserted/3 beyond the calyx lobe tips before anthesis. Flowers 5-merous, all perfect. Calyx tube 0.8–1.3 mm long, conical, the lobes 0.3–0.5 mm long, 0.5–0.6 mm wide, broadly triangular, the tips obtuse, sparsely pubescent with simple uniseriate trichomes like those of the stem. Corolla 3–6 mm in diameter, white with a yellow-green central portion near the base, stellate, lobed 1/2-2/3 of the way to the base, the lobes 2.0–3.2 mm long, 1.0–2.5 mm wide, strongly reflexed at anthesis, later spreading, densely papillate abaxially with 1-4-celled simple uniseriate trichomes, these denser on the tips and margins. Stamens equal; filament tube minute; free portion of the filaments 0.5–0.8 mm long, adaxially pubescent with tangled uniseriate trichomes; anthers 0.8–1.5 mm long, 0.5–0.6 mm wide, ellipsoid to almost globose, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.2–2.6 mm long, densely pubescent with 2-3-celled simple uniseriate trichomes in the lower 2/3 where included in the anther cone, almost included to exserted 0.5(-1.0) mm beyond the anther cone; stigma minutely capitate, the surface minutely papillate, green in live plants. Fruit a globose berry, 4–9(-12) mm in diameter, purplish-black at maturity, the pericarp thin and markedly shiny; fruiting pedicels 13–18 mm long, 0.7–1.0 mm in diameter at the base and 0.8–1.0 mm at apex, stout, straight and spreading, spaced 0–0.5 mm apart, not falling with the fruit, remaining on the plant and persistent on older inflorescences; fruiting calyx not accrescent, the tube less than 1 mm long, the lobes 1(-2) mm long, strongly reflexed at fruit maturity. Seeds 30–50 per berry, 1.0–1.5 mm long, 0.8–1.3 mm wide, flattened and tear-drop shaped with a subapical hilum, pale yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells mostly absent (Australia, South Pacific and South America), but if present (North America, Mexico, Eurasia and Africa) 2–4(6) per berry, 2–4 larger ones >0.5 mm and two smaller ones <0.5 mm in diameter. Chromosome number: 2n=2x=24 (
Grows in disturbed habitats and associated with human activities in tropical moist to dry areas, in dry areas often found growing in full shade close to water sources; between sea level and 2,000 (-2,500) m elevation.
American Samoa: magalo; Australia: glossy nightshade (
In all parts of its range, the leaves of S. americanum are used as spinach and the ripe berries are eaten, either raw or cooked.
(
Solanum americanum is a diploid species that can be easily recognised by its shiny black fruits on spreading pedicels with strongly reflexed calyx lobes (to parallel with the pedicel) that are somewhat papillate abaxially. In fruit, the pedicels remain on the plant after fruits fully mature and drop off, leaving behind a distinct group of tightly clustered spreading pedicels with reflexed calyx lobes; this character is easily visible in many herbarium specimens. In flower, S. americanum has tiny, almost globose anthers 0.8–1.5 mm long borne on short filaments. It can be distinguished from S. opacum, which also has tiny anthers of the same size, in its shorter filaments relative to anther size and in its deltate calyx lobes with rounded tips. Solanum opacum has longer filaments relative to anther size, long-triangular calyx lobes and, in fruit, the calyx lobes are appressed to the base of the berry. Three other morelloids with such small anthers can be difficult to distinguish from S. americanum and are often confused with it in herbaria. Solanum emulans Raf. does not occur in the Old World (except sometimes in old botanical garden collections, see discussion of the typification of S. patulum and S. dillenii below; it is a species of the north-eastern United States and Canada) has matte berries and longer calyx lobes. Solanum nitidibaccatum also has extremely small anthers, but can be easily distinguished by its glandular pubescence and accrescent calyx in fruit. Solanum opacum is the third morelloid with tiny anthers and the most difficult to distinguish from S. americanum in the Old World. Solanum americanum and S. opacum co-occur across the Pacific and distinguishing individual specimens can be difficult, but the shiny fruits (versus matte in S. opacum) and persistent pedicels with strongly reflexed calyx lobes are good characters by which to recognise S. americanum.
In southeast Asia and China, S. americanum is at least partially sympatric with S. nigrum (see discussion of S. nigrum). The species can be distinguished by anther size (0.8–1.5 mm versus ca. 2–2.5 mm) and by inflorescence morphology; S. americanum usually has few flowers that are tightly congested in the distal part of the inflorescence, while S. nigrum usually has more flowers that are more spaced out along the inflorescence rhachis, although young inflorescences of S. nigrum can appear sub-umbellate. In fruit, the strongly reflexed calyx lobes of S. americanum are distinctive and the seeds are smaller than those of the hexaploid S. nigrum (ca. 1 mm versus ca. 2 mm long).
Solanum merrillianum was recognised as a distinct species in the Flora of China (
Solanum americanum exhibits the highest infraspecific genetic diversity compared to polyploids (
The taxonomic status and relationship of S. americanum to S. nodiflorum was studied by
The results described above, based on AFLP markers, should be tested with modern population genetic tools such as functional markers (
The identity of the species depicted in plate 355 (
Solanum papilionaceum was almost certainly described from living material only.
The specimens in Zuccagni’s herbarium in Florence were consumed by fire, making the designation of a neotype for S. strictum Zucc. necessary. The specimen we have selected is dated later than the description, but is labelled as “strictum Zucc.” and is from Italy in cultivation (G00144215); it was used by Dunal in his Prodromus treatment as S. strictum (
The identity of
In describing S. microspermum,
Although the protologue of S. erythrocarpon (
In describing S. minutibaccatum,
Solanum calvum was described using “Palmer 60 p. pte.” (
The protologue of S. nigrum var. pauciflorum (
A total of 1,074 specimens were examined from 73 countries during the study across Africa, Asia, Australia, Eurasia and the Pacific. Specimens from the New World were also studied in order to understand the full range of morphology within the species. All specimens examined from the Old World can be seen in Appendix 2 (csv format) and Appendix 3 (traditional Specimens Examined list in pdf format).
Solanum sublobatum Willd. ex Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 664. 1819.
Type. Argentina. Buenos Aires, Anon. s.n. [probably P. Commerson] (Herb. Willdenow 4336) (lectotype, designated by
Solanum besseri Weinm., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 593. 1819.
Type. “In America” [cultivated in Europe?], Anon. s.n. (no specimens cited; no original material located; neotype, designated here: G-DC [G00144198]).
Solanum subspatulatum Sendtn., Fl. Bras. (Martius) 10: 45, tab. 4, fig. 16–18. 1846.
Type. Brazil. Sin. loc., F. Sellow s.n. (holotype: B, destroyed, F neg. 3183; lectotype, designated by
Witheringia chenopodioides (Lam.) J.Rémy, Fl. Chil. [Gay] 5: 69. 1849.
Type. Based on Solanum chenopodioides Lam.
Solanum isabellei Dunal, Prodr. [A. P. de Candolle] 13(1): 153. 1852.
Type. Uruguay. Montevideo, Lat. aust. 34°45’08”, 1838, A. Isabelle s.n. (lectotype, designated here: G-DC (G00145645); isolectotypes: F [V0073298F, acc. # 680251; V0073299F, acc. # 680253)], K [K000585686], P [P00384071], W [1889-115034]).
Solanum chenopodifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 44. 1852.
Type. Argentina/Uruguay. “Buenos Aires et Montevideo”, P. Commerson s.n. (lectotype, designated
Solanum crenatodentatum var. ramosissimum Dunal, Prodr. [A. P. de Candolle] 13(1): 54. 1852.
Type. United States of America. Louisiana: “Basse Louisiane”, 1839, G.D. Barbe 82 (holotype: P [P00362535]).
Solanum gracile Dunal, Prodr. [A.P. de Candolle] 13(1): 54. 1852, nom. illeg., not Solanum gracile Sendtn. (1846).
Type. Brazil. Rio de Janeiro: “Rio de Janeiro”, 1831-1833, C. Gaudichaud 520 (lectotype, designated by
Solanum gracile var. microphyllum Dunal, Prodr. [A. P. de Candolle] 13(1): 54. 1852.
Type. Argentina/Uruguay. “ Circa Buenos Ayres et Montevideo”, P. Commerson s.n. (lectotype, designated by
Solanum nodiflorum var. microphyllum Hassl., Repert. Spec. Nov. Regni Veg. 9: 118. 1911.
Type. Paraguay. Estrella: Mar, É. Hassler 10271 (holotype: G?, Morton photo 8612).
Solanum vile Bitter, Repert. Spec. Nov. Regni Veg. 11: 221. 1912.
Type. Brazil. Rio de Janeiro: Restinga do Harpoador, E. Ule 4310 (lectotype, designated here: CORD [CORD00004277]; isolectotype: HBG [HBG511507]).
Solanum gracilius Herter, Rev. Sudamer. Bot. 7: 266. 1943.
Type. Based on (replacement name for) S. gracile Dunal
Solanum ottonis Hyl., Uppsala Univ. Årsskr. 7: 279. 1945.
Type. Based on (replacement name for) Solanum gracile Dunal
Solanum americanum var. baylisii D’Arcy, Ann. Missouri Bot. Gard. 61: 837. 1974.
Type. New Zealand. Sin. loc., cultivated, 1953, Momson s.n. (holotype: OTA [OTA-00419]).
Mauritius. “Ex ins. Mauritiana”, Herb. Lamarck s.n. (lectotype, designated by
Annual herbs to short-lived, erect to somewhat sprawling perennial herbs to 1.0 m tall, subwoody and branching at base. Stems spreading to decumbent, terete, green-grey to straw colour, older stems with no prickle-like projections, not markedly hollow; new growth pubescent with simple, appressed, uniseriate, eglandular trichomes, these 1-6-celled, 0.1–0.6 mm long; older stems more sparsely pubescent, glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.5–5.5(-7.0) cm long, 0.5–3.0(-3.5) cm wide, lanceolate to narrowly ovate, rarely ovate, discolorous; adaxial surface green, sparsely pubescent with appressed 1-4-celled translucent, simple, uniseriate trichomes like those on stem, these denser along the veins; abaxial surface pale grey, more densely pubescent with trichomes like those of the upper surface evenly across lamina and veins; major veins 3–6 pairs, not clearly evident abaxially; base attenuate, decurrent on the petiole; margins entire or sinuate; apex acute to obtuse; petioles (0.5-)1.0–1.5(-3.5) cm long, sparsely pubescent with simple uniseriate trichomes like those of the stems and leaves. Inflorescences 1.0–2.5(-4.0) cm long, generally internodal but appearing leaf-opposed on young shoots, simple or rarely furcate, sub-umbelliform, with 3–7(-10) flowers, sparsely pubescent with appressed 1-2-celled simple uniseriate trichomes; peduncle 1.0–2.3(-4.0) cm long, straight but becoming strongly deflexed downwards in fruit; pedicels 0.5–1 cm long, ca. 0.5 mm in diameter at the base and 1 mm at apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0–1 mm apart. Buds elongate-oblong, the corolla only slightly exserted from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 0.5–1.0 mm long, conical, the lobes 0.6–1.2 mm long, less than 1 mm wide, broadly deltate to triangular with acute to obtuse apices, sparsely pubescent with 1-4-celled appressed hairs like those on stem but shorter. Corolla 6–12 mm in diameter, white with a black or yellow-green central portion near the base, the black colour usually distal to the yellow-green, deeply stellate, lobed 4/5 of the way to the base, the lobes 3.5–4.0 mm long, 1.5–1.9 mm wide, strongly reflexed at anthesis, later spreading, densely puberulent-papillate abaxially with 1-4-celled simple uniseriate trichomes, these denser on the tips and margins. Stamens equal; filament tube minute; free portion of the filaments 0.6–1.0 mm long, adaxially pubescent with tangled uniseriate 4-6-celled simple trichomes; anthers (2.0-)2.3–2.8 mm long, 0.5–0.8 mm wide, narrowly ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying, the connective becoming darker brown with age in dry plants. Ovary globose, glabrous; style 3.7–4.5 mm long, densely pubescent with 2-3-celled simple uniseriate trichomes in the lower half where included in the anther cone, exserted up to 1.5 mm beyond the anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 4–9 mm in diameter, dull purplish-black at maturity, the pericarp thin and opaque, matte and somewhat glaucous; fruiting pedicels 6–13 mm long, 1.2–1.4 mm in diameter at the base and the apex, reflexed and slightly curving, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube less than 1 mm long, the lobes 1–1.5 mm long, appressed against the berry. Seeds (13-)20–35(-50) per berry, 1.2–1.4 mm long, 1.0–1.2 mm wide, flattened and tear-drop shaped with a subapical hilum, pale yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells absent or occasionally 1–2 diminutive apical stone cells present. Chromosome number: 2n=2x=24 (
Grows in humid, disturbed areas between rocks, along water sources and roads and in cultivated lands, common in areas with human disturbance; between sea level and 1,900 (-2,500) m elevation.
Australia: whitetip nightshade (
South Africa. leaves used as spinach (
(
Solanum chenopodioides can be distinguished from most of the other morelloids occurring in the Old World based on its narrowly lanceolate leaves with grey indumentum, inflorescences with ca. 3–7 flowers tightly congested near the tip of the peduncle, the stellate corollas that are deeply lobed to the base and usually with a dark purple or black central star and anthers that are usually more than 2 mm and up to 2.8 mm long. In fruit, the pedicels and proximal portion of the peduncle are strongly reflexed and the berries are not at all shiny. Solanum retroflexum has similar matte black berries, but has rhomboid leaves, less deeply divided corollas, shorter anthers and the calyx lobes are strongly reflexed in fruit. Solanum chenopodioides could also potentially be confused with the more common S. nigrum, especially in Europe, but differs from that species in its terete stems, matte black fruits on short strongly reflexed pedicels and its smaller seeds (1.2 mm long versus 2 mm long).
Solanum chenopodioides has a scattered distribution in the Old World, but it seems to be spreading, perhaps related to climate change and/or increased habitat alteration (
This diploid species possibly contributed its genome to the tetraploid S. retroflexum and hexaploid S. opacum.
In the protologue (
In the protologue of S. isabellei,
Solanum gracile was incorrectly typified by
Australia. New South Wales: N shore Lake Illawarra, Lake Hts, 5 Sep 2003, Andrew s.n. (WOLL); Nepean River, Douglas Park, 6 mi E of Picton, 11 Oct 1965, Constable 6161 (AD, K, NSW); Bega Valley, Twofold Bay, Nullica Bay Beach 5 km SSW of Eden, 8 Feb 1979, Haegi 1711 (AD, MEL, MO, NSW); Swamp 2 km N of Bodalla on Princes H[igh]w[a]y, just S of Tuross River crossing, 10 Feb 1979, Haegi 1756 (AD, NSW); Bega Valley, Tathra, 20 Mar 1995, Heyligers 95004 (PERTH); Bega Valley, Tathra, 20 Mar 1995, Heyligers 95006 (PERTH); Tamworth Regional, Hanging Rock, 13 Apr 2008, Hosking 3091 (CANB, MEL, NE, NSW); Murrumbidgee River at Cotter Bridge, Cotter Reserve, 1 Mar 1992, Lepschi 740 (AD, CANB, NSW); Hillview (3 mi SW of Liverpool), 11 Dec 1968, Rodd s.n. (NSW); Queensland: Brisbane, 20 Jan 1966, Henderson 128 (BRI, NSW); Brisbane, Clapham Junction, 20 Oct 1967, Henderson 301 (AD, BRI); South Australia: Tea Tree Gully, Gorge Rd, 15 Jan 2010, Brodie 1171 (AD); sin. loc, 1885, Lea s.n. (BM); Region 11, Southern Lofty, Lower Gorge Rd, 13 Feb 1997, Symon 15462 (AD, K, MEL); Victoria: East Gippsland, N of Mallacoota township, 4 Apr 1999, Clarke 2890 (AD, MEL); Melbourne, 70 m SW of Queens Bridge, 18 Feb 1983, Clarke 1544 (AD, CANB, MEL); Wellington, c. 13 km from Licola, 28 Jan 1989, Thompson 159 (AD).
France. Nouvelle Aquitaine: Gironde, Bordeaux, 28 Aug 1931, Bouchon 6703 (BM); Provence-Alpes-Côte d’Azur: Bouches-du-Rhône, La Valentine, banlieue E de Marseille, route vers Saint Menet, 19 Nov 1973, Martin 6848 (BM, H).
Germany. Baden-Württemberg: Karlsruhe, Carlsruhe, Oct 1834, Braun s.n. (K); Karlsruhe [?], Dec 1834, Braun s.n. (K).
Greece. Crete: sin. loc, de Tournefort s.n. (BM); East Macedonia and Thrace: Island of Thasos, SE of Potamia, 23 Nov 2016, Biel IM-16054 (N/A).
Italy. Friuli Venezia Giulia: Monfalcone, Friulia, 25 Sep 1953, Neumann s.n. (W).
Japan. Honshu: Tochigi, Utsunomiya Agricultural College, Japan, 1935, Kagawa s.n. (K)Kyoto, Koyama, Chitose, Chitose-cho, Kameoka-shi, 12 Sep 2010, Tsugaru et al. 7604 (MO).
Lesotho. Maseru airfield escarpment, Lesotho, 10 Dec 1969, Without collector 348 (K); Maseru Ekp Stn, 3 Mar 1970, Without collector 650 (K).
New Zealand. North Island: Auckland, Tawharanui Regional Park, 14 Jan 2012, Salter & Duff s.n. (AK); Northland, Te Arai Sanctuary, 27 Jan 1995, Wright 12532 (AK); South Island: Tasman, Collingwood, 1 km W on Cape Farewell rd, 13 Mar 2006, Brummitt 21567 (K).
Portugal. Azores: Faial, Horta, 27 Sep 1970, Brooke 11376 (BM); Faial, Capelo, 1 Dec 1971, Goncalves 374 (BM); Faial between Pedro Miguel and Espalhafatos, 15 Sep 2001, Henderson et al. 98 (AZU, BM); Centro: Vila Nova de Barquinha, Ribatejo, 16 Jul 1963, Rainha 6177 (W); Madeira: Between Monte and Funchal, 11 Sep 1984, Davis 70402 (BM); Norte: Amarante, Marcos de Canavezes, Douro Litoral, 6 Jul 1960, Pinto da Silva et al. 6754 (W).
South Africa. Eastern Cape: Old Town Quarry, Grahamstown, 20 Nov 1972, Bayliss 5292 (A, BH, MO); Belmont Valley, Albany Dist, 1 Sep 1974, Bayliss 6785 (K, MO); Humansdorp, Distr. Humansdorp, Tzitzikama Park, 2 Feb 1966, Liebenberg 7909 (K); Gauteng: Germiston District, Dowerglen, 22 Oct 1992, Balkwill 7163 (MO); Pretoria, Johannesburgh, Taamlik volop, 2627BB grid ref, Oct 1976, Liebenberg 8568 (K, MO); KwaZulu-Natal: Pretoria, Greytown, Natal, 11 Feb 1939, Galpin 14832 (K); Underberg, 2929CB Sani Pass, Undeberg Distr., 17 Feb 1982, Hilliard & Burtt 15537 (K); Nottingham rd, Natal, Mar 1939, McClean 869 (K, MO); Mpumalanga: Middelburgh District, just outside Middelburgh, 26 Jan 1995, Balkwill 9130 (MO); Western Cape: 3323 Willowmore DC, 28 Dec 1982, Goldblatt 6780 (MO).
Spain. Cantabria: Santander, rd to Pechon, 31 Jul 1972, Brenan 12271 (BM, K).
Sweden. Götaland: Västra Götaland, Göteborg, Backa prope Brunnsbo, Aug 1938, Blom s.n. (BM); Västra Götaland, Agnesbergs kvarn, 30 Sep 1938, Blom s.n. (K, W).
Switzerland. Ticino: Locarno, bank of Maggia river, 17 Sep 2000, Brummit 20476 (K).
United Kingdom. Channel Isles: Guernsey, nr entrance to Mont Cuet, Nov 2001, Dupree s.n. (BM); Guernsey, 6 Sep 1994, McClintock & Ryan s.n. (BM); Guernsey, St. Sampsons, 23 Jul 1968, Simpson 68010 (BM); England: Greater London, Victoria Park, 4 Sep 2008, Atchison 2 (BM); Worcestershire, Charlton, 25 Aug 1959, Pannister 983 (BM); Cornwall, Bude, Aug 1925, Thurston s.n. (K); Wales: Vale of Glamorgan, Barry Docks, 12 Sep 1935, Brenan & Sandwith 1449 (BM); Cardiff, Barry Docks, 12 Sep 1935, Sandwith & Brenan s.n. (K).
Solanum deltoideum Colla, Herb. Pedem. 4: 273. 1835.
Type. Cultivated in Italy at “h. Ripul:” [Hortus Ripulensis], the seeds originally sent by C. Bertero from Chile [“Chili Quillota”] (no specimens cited; lectotype, designated here: TO [herb. Colla]).
Solanum furcatum var. glabrum G.Don, Gen. Hist. 4: 412. 1837.
Type. “In Peruvia” (no specimens cited; no original material located).
Solanum furcatum var. pilosum G.Don, Gen. Hist. 4: 412. 1837.
Type. “In Peruvia” (no specimens cited; no original material located).
Solanum furcatum var. acutidentatum Nees, Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 386. 1843, as “acutedentatum”.
Type. “Chile ad Valparaiso, Februario; Peruvia in planitie circa Tacoram, alt. 14000-17000’, Aprili” both syntypes collected by F.J.F. Meyen s.n. (no specimens cited; no original material located).
Solanum furcatum var. obtusidentatum Nees, Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 386. 1843, as “obtusedentatum”.
Type. “Chile. Prov. de San Fernando in Llano del Rio Tinguiririca, 3000’ alf, martio”; Peruvia ad Arequipam, Aprili” both syntypes collected by F.J.F. Meyen s.n. (no specimens cited; no original material located).
Solanum furcatum var. subintegerrimum Nees, Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 386. 1843.
Type. “Chile: Copiapó, Aprili; Peruvia: circa Tacoram, Aprili” both syntypes collected by F.J.F. Meyen s.n. (no specimens cited; no original material located).
Witheringia furcata (Dunal) J.Rémy, Fl. Chil. [Gay] 5: 67. 1849.
Type. Based on Solanum furcatum Dunal
Solanum pterocaulum var. dichotimiflorum Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, as ‘pterocaulon’.
Type. Cultivated in France at Montpellier “Solanum speciosum hort. botan” (no specimens cited, described from living plants “v.v. hort. Monsp.”; neotype, designated here: MPU [MPU310703]).
Solanum crenatodentatum Dunal, Prodr. [A. P. de Candolle] 13(1): 54. 1852.
Type. Chile. Région VI (O’Higgins): Colchagua, San Fernando, “in selibus chilensibus San Fernando”, Mar 1831, C. Gay 2 (lectotype, designated by
Solanum rancaguense Dunal, Prodr. [A. P. de Candolle] 13(1): 150. 1852.
Type. Chile. Région VI (O’Higgins): Rancagua, May-Oct 1828, C. Bertero 633 (lectotype, designated by
Solanum bridgesii Phil., Linnaea 33: 203. 1864.
Type. Chile. Región V (Valparaíso): Panquegue, R.A. Philippi s.n. (lectotype, designated here: SGO [SGO000004549]).
Solanum coxii Phil., Linnaea 33: 200. 1864.
Type. Chile. Región X (Los Lagos): Todos los Santos, 1862, G. Cox 38 (lectotype, designated here: SGO [SGO000004555]; isolectotype: W [W1903-0010246]).
Solanum rancaguinum Phil., Anales Univ. Chile 43: 523. 1873.
Type. Chile. Región VI (O’Higgins): Rancagua, Mar 1828, C. Bertero s.n. (lectotype, designated here: SGO [SGO000004594]).
Solanum caudiculatum Phil., Anales Univ. Chile 91: 12. 1895.
Type. Chile. Región VIII (Bío-Bío): prov. Ñuble, Coigüeco, F. Puga s.n. (no original material located, not at SGO).
Solanum subandinum Phil., Anales Univ. Chile 91: 13. 1895, nom. illeg., not Solanum subandinum F.Meigen (1893).
Type. Chile. Región XIII (Metropolitana): Santiago, Las Condes, R.A. Philippi s.n. (lectotype, designated here: SGO [SGO000004600, F neg. 2745]).
Solanum ocellatum Phil., Anales Univ. Chile 91: 14. 1895.
Type. Chile. Región XIII (Metropolitana): Prope Colina, F. Philippi s.n. (lectotype, designated here: SGO [SGO000004582]; isotypes: SGO [SGO000004581], W [1903-0010230]).
Solanum nigrum var. crentatodentatum (Dunal) O.E.Schulz, Symb. Antill. (Urban) 6: 160. 1909.
Type. Based on Solanum crenatodentatum Dunal
Solanum bridgesii var. ocellatum (Phil.) Witasek ex Reiche, Anales Univ. Chile 124: 460. 1909.
Type. Based on Solanum ocellatum Phil.
Solanum andinum Reiche, Fl. Chile 5: 346. 1910.
Type. Based on (replacement name for) Solanum subandinum Phil.
Solanum tredecimgranum Bitter, Repert. Spec. Nov. Regni Veg. 11: 6. 1912.
Type. Chile. Región V (Valparaíso): Valparaíso, 17 Aug 1895, O. Buchtien s.n. (lectotype, designated by
Solanum robinsonianum Bitter, Repert. Spec. Nov. Regni Veg. 11: 7. 1912.
Type. Chile. Región V (Valparaíso): Juan Fernández Island, R.A. Philippi 742 (holotype: B, destroyed, F neg. 2743; lectotype, designated here: W [0001347]).
Solanum masafueranum Bitter & Skottsb., Nat. Hist. Juan Fernandez & Easter Island 2: 167, pl. 14. 1922.
Type. Chile. Región V (Valparaíso): Juan Fernández Islands, Masafuera [Isla Alejandro Selkirk], Las Chozas, 715 m, 3 Mar 1917 [20 Feb 1917 on label], C. Skottsberg & I. Skottsberg 363 (lectotype, designated here: S [acc. # 04-2947]; isolectotypes: BM [BM000617676], LD [1643307], K [K000585692], NY [00172084], GOET [GOET003548], GB [GB0048742], P [P00337092], UPS [acc. # 104031]).
Solanum spretum C.V.Morton & L.B.Sm., Revis. Argentine Sp. Solanum 132. 1976.
Type. Argentina. Río Negro: Bariloche, 19 Mar 1939, A.L. Cabrera 5024 (holotype: GH [GH00077764]; isotypes F [V0073411F, acc. # 1007493], LP [LP006791]).
Peru ?. “Cette plante croît au Perou”, J. Dombey [343] (lectotype, first step designated by
Annual or perennial erect to sprawling herbs or small shrubs to 1.0 m tall, subwoody and branching at base. Stems spreading to decumbent, terete or ridged, green to purple tinged, older stems pale yellowish-brown, not markedly hollow; new growth sparsely pubescent with simple, spreading, uniseriate, eglandular trichomes, these 1-5-celled, 0.1–0.5 mm long; older stems sparsely pubescent to glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, (1.5-)4.0–8.0(-12.0) cm long, (0.6-)2.2–4.6(-6.5) cm wide, ovate to rhomboidal, membranous, slightly paler beneath, without smell; adaxial surface sparsely pubescent with simple, uniseriate trichomes like those on stem, these evenly spread along lamina and veins; abaxial surface more densely pubescent; major veins 4–6 pairs; base cuneate to acute, slightly oblique, decurrent on the petiole; margins sinuate-dentate or entire; apex acute; petioles 1.0–3.5 cm long, sparsely pubescent with simple uniseriate trichomes like those on stem. Inflorescences (1.0-)1.5–3.0(-4.0) cm long, internodal, furcate or more rarely unbranched, sub-umbelliform or the flowers evenly spaced along the rhachis, with 6–14 flowers, sparsely pubescent with simple uniseriate trichomes like those on stems; peduncle (1.0-)1.5–2.0 cm long, straight; pedicels 4.0–7.5 mm long, 0.2–0.3 mm in diameter at the base and 0.3–0.4 mm at apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0.2–2.5 mm apart. Buds subglobose, the corolla exserted 1/3-1/2 from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 0.5–1.5 mm long, conical, the lobes 0.8–1.5 mm long, 0.6–1.0 mm wide, rectangular to narrowly obovate, tips obtuse to shortly acute, pubescent with simple uniseriate trichomes like those on stem but shorter. Corolla 12–20 mm in diameter, white to lilac with a green or yellow-green central portion near the base, this sometimes purplish near the lobe midvein, stellate, lobed 1/3–1/2 of the way to the base, the lobes 5.5–7.0 mm long, 2.8–5.5 mm wide, strongly reflexed at anthesis, later spreading, densely pubescent abaxially with 1-4-celled simple uniseriate trichomes, especially along the margins and apex, these shorter than the trichomes of the stems and leaves. Stamens equal; filament tube minute; free portion of the filaments 0.9–1.6(-2) mm long, adaxially pubescent with tangled uniseriate 4-6-celled simple trichomes; anthers 2.3–3.3(-3.6) mm long, 0.8–1.0 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary globose, glabrous; style 6.0–6.5 mm long, densely pubescent with 2-3-celled simple uniseriate trichomes in the lower 1/2–2/3 where included in the anther cone, exserted 2–3 mm beyond the anther cone and somewhat curved; stigma capitate, minutely papillate, yellow or green in live plants. Fruit a globose berry, 6–9 mm in diameter, dull green to purple at maturity, the pericarp thin and opaque; fruiting pedicels 7–12 mm long, 0.2–0.4 mm in diameter at the base and 0.5–1.0 mm at apex, strongly reflexed, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes 1.5–2.5 mm long, appressed against the berry. Seeds 30–40 per berry, 1.8–2.0 mm long, 1.4–1.5 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow-brown, the surface minutely pitted, the testal cells pentagonal in outline. Stone cells 2–13 per berry, largest 0.8–1.0 mm in diameter but varying in size. Chromosome number: 2n=6x=72 (
Solanum furcatum Dunal A Habit B Detail flower (A–BAnonymous s.n., grown from seed sent by J. Edmonds, originally from California [ADW 42421]). Drawing by M.L. Szent-Ivany, first published in
Solanum furcatum Dunal A Habit B Flowering branch C Infrutescence with developing fruits. D Fully mature fruit dropping off with pedicel (A–DKnapp s.n., United States of America, California, San Francisco, Golden Gate Park, in front of California Academy of Sciences, 11 June 2013). Photos by S. Knapp.
(Figure
Open places and disturbed areas, along roadsides and field margins; between sea level and 3,000 m elevation in its native range, in Australia collected in coastal habitats along the foreshore or along creeks near sea level.
Australia: broad nightshade (
None recorded for Old World collections.
(
Solanum furcatum can be distinguished from all other Old World species based on its branched inflorescences, anthers 2.3–3.3 mm long and flowers where styles are exserted up to 3.0 mm beyond anthers. It has been sparingly introduced into areas of Mediterranean habitat (e.g. Australia and California) but appears not to spread. It is potentially confusable with S. nigrum, but is distinguished from it by the characters mentioned above and by its 2–13 stone cells per berry. Its subglobose buds are also distinct from the more ellipsoid buds of S. nigrum and S. villosum, but this character can be hard to see in herbarium specimens.
The name S. douglasii Dunal has been misapplied to some specimens of S. furcatum in Australia (e.g.
Solanum furcatum was lectotypifed by
The infraspecific taxa of S. furcatum described by
Solanum rancaguense was lectotypified by
We have selected lectotypes in SGO for the taxa described by Philippi here considered synonyms of S. furcatum following the advice of
Australia. Tasmania: Circular Head, B22 between Irishtown and Edith Creek, 2 Nov 2004, Baker 981 (HO); Circular Head, Grooms Cross rd (B22), between Irishtown and Edith Creek, 2 Nov 2004, Baker 1010 (HO); Circular Head, Edith Creek, 2 Nov 2004, Baker 1019 (HO); Circular Head, Copper Creek nr Smithton, May 1947, Curtis s.n. (HO); Circular Head, Copper Creek, May 1948, Without collector s.n. (HO); Victoria: Greater Geelong, Edwards Point State Faunal Reserve, 19 May 1984, Albrecht 483 (MEL); Along Billy Creek in proposed extension to Morwell National Park, 15 Jul 1988, Harris 4 (K); South Gippsland, Boolarra to Mirboo North rd, 22 Nov 1987, Harris s.n. (MEL); East Gippsland, Gauging Station on the Genoa River nr the Wangarabell rd, 10 Mar 2011, Jeanes 2662 (MEL); Yarra Ranges, Tremont, 30 Jul 2006, Stajsic 4292 (HO); Yarra Ranges, Tremont, 30 Jul 2006, Stajsic 4292 (MEL)Greater Geelong, St Leonards, Bellarine Peninsula, 9 Nov 1947, Willis s.n. (MEL); Greater Geelong, St Leonards, Bellarine Peninsula, 9 Nov 1947, Willis s.n. (MEL); Mornington Peninsula, Main Creek, 24 Mar 1984, Willis s.n. (MEL); McCrae, between Dromana & Rosebud, 24 Feb 1963, Willis s.n. (MEL); McCrae, between Dromana & Rosebud, 24 Feb 1963, Willis s.n. (MEL).
New Zealand. North Island: Manawatu-Wanganui, Vinegar Hill, Rangitikei River near Rowa, 18 Nov 1964, Healy 64/425 (AK).
Solanum monactinanthum Dammer, Bot. Jahrb. Syst. 48: 236. 1912.
Type. Ethiopia. “Galla-Hochland: Arussi-Galla, Jidah”, 2600 m, Jul 1900, H. Ellenbeck 1452 (holotype: B, destroyed; no duplicates found).
Solanum hirtulum Steud. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 44. 1852, nom. illeg. (isonym), not S. hirtulum A.Rich. (1850).
Type. Based on same type as Solanum hirtulum A.Rich.
Ethiopia. Amhara: “Enschadcap” [Inch’et Kab Bota in Simien Mountains], 29 Jan 1838, G.H.W. Schimper 977 (lectotype, designated by
Annual to short-lived, mostly prostrate perennial herbs to 5–20(-150) cm high, branches ascending from woody tap-root. Stems decumbent to ascending, terete or very slightly winged from decurrent leaf bases, green or straw coloured, older stems not appearing spinescent, yellowish-brown, not markedly hollow; new growth moderately pubescent with simple, antrorse, uniseriate, eglandular trichomes, these 5-7-celled, 0.5–0.8 mm long, white. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.5–5.0(-6.0) cm long, 0.7–1.8 cm wide, narrowly elliptic to lanceolate, narrowing gradually to the base, concolorous, without smell; adaxial surface sparsely and evenly pubescent with simple, uniseriate trichomes like those on stem; abaxial surface with a few evenly scattered trichomes like those of the adaxial surface; major veins 5–7 pairs; base long-attenuate, decurrent on the petiole; margins entire to sinuate; apex acute to acuminate, the tip slightly rounded; petioles absent, the laminar tissue extending to the junction of leaf and stem. Inflorescences 0.5–2.2 cm long, opposite the leaves, simple, sub-umbelliform to shortly racemose, with 1–5 flowers clustered in the distal portion, sparsely pubescent with antrorse simple uniseriate trichomes like those of the stems; peduncle 0.4–1.8 cm long, straight; pedicels 0.7–1.6 cm long, ca. 0.5 mm in diameter at the base and apex, stout and spreading, articulated at the base; pedicel scars clustered at the tip of the inflorescence rhachis and overlapping, occasionally the basal scar 1–2.5 mm distant. Buds ellipsoid, the corolla halfway exserted from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 2.0–3.0 mm long, conical, the lobes 0.8–1.2 mm long, 0.8–1.2 mm wide, linear-oblong, tips rounded, densely pubescent with trichomes like those of the stems and pedicels. Corolla 12–18(-20) mm in diameter, deep purple to pale violet, stellate, lobed 3/4 of the way to the base, the lobes 4.5–8 mm long, 2–3 mm wide, spreading to reflexed, densely papillate abaxially, the papillae denser along margins and tips. Stamens equal; filament tube ca. 0.5 mm long; free portion of the filaments ca. 0.5–1.0 mm long, adaxially densely pubescent with tangled simple uniseriate trichomes; anthers 2.3–2.8 mm long, 0.6–0.9 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary rounded, glabrous; style 4.5–5.5 mm long, densely pubescent with simple trichomes in the basal 1/3, exserted ca. 1 mm beyond anther cone; stigma large-capitate, the surfaces minutely papillose. Fruit a globose berry, 5–6 mm in diameter (immature?), mature berry colour not known, the pericarp thin and matte or somewhat shiny; fruiting pedicels 0.7–1.6 cm long, 0.5 cm in diameter at the base and 1.0–1.2 mm at the apex, stout with slight curving at the base, reflexed, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes 1–1.5 mm long, appressed to the berry. Seeds (10-)20–35 per berry, ca. 1.8–2.0 mm long, ca. 1.5 mm wide, not markedly flattened, tear-drop shaped with a subapical hilum, pale yellow, the surfaces minutely pitted, the testal cells pentagonal to rectangular in outline. Stone cells 2 per berry, ca. 0.6 mm in diameter. Chromosome number: not known.
Grows in open grassy areas, arable land, along river banks, in forest and along roadsides; between 2,200 and 3,500 m elevation.
None recorded.
Uses. None recorded.
(
Solanum hirtulum is a distinctive prostrate, creeping herb with narrow lanceolate attenuate leaves with strigose, somewhat antrorse pubescence. The flowers are larger than others in the group in Africa and appear to always be purple or “blue”; this, however, must be taken with caution given the high degree of flower colour polymorphism in other species. The inflorescences are usually few-flowered with tightly spaced flowers at the very tip. Solanum hirtulum is similar to S. memphiticum in having black berries with a somewhat accrescent calyx, but the flowers of S. memphiticum are always smaller, usually white and more delicate (see description of S. memphiticum). The leaf bases in S. hirtulum are strongly attenuate and decurrent on to the stem.
Solanum hirtulum is a plant of Afromontane vegetation (sensu
The type number of S. hirtulum (Schimper 977) is also that for a plant collected in 1837 in Saudi Arabia (the type of Seddera intermedia Hochst., Convolvulaceae, see http://apps.kew.org/herbcat/getImage.do?imageBarcode=K000852498). He clearly reused numbers for his Ethiopian plants.
Solanum monactinanthum is here placed in synonymy based on the description and following
Ethiopia. sin. loc., 26 Oct 1862, Schimper 631 (BM, E, K, P, W); Addis Ababa: Addis Ababa, 30 Sep 1937, Piovano 511 (FT); Addis Ababa, 19 Oct 1937, Senni 1837 (FT); Amhara: Semien, Debarek Semien Gonder Region, 11 Jul 1909, Chiovenda 889 (FT); 10 km SE of Debre Markos along rd to Addis Ababa, between Debre Markos and Addis Ababa, 25 Oct 2004, Friis et al. 11912 (K); Bichena Awraja, c. 36 km NW of Debre Work (Gojjam region), 30 Oct 1981, Mesfin Tadese & Kagnew 1659 (K); in campis Debra Eski, 19 Oct 1850, Schimper 74 (P); Semien, Derasghie, 25 Dec 1952, Scott 292 (K); Dejen to Debra Marcos, 48 km NW of Dejen, 24 May 1980, Thulin 3914 (K, MO); mule track between Debarak and Geech, 16 Sep 1969, de Wilde & Gilbert 5 (EA, MO); Oromia: Semien Shewa region, Holetta, 4 May 1953, Mooney 4752 (K); Addis Alem, 20 Sep 1926, Omer Cooper s.n. (K); Tigray: Adua, Prope Adoam, Abyssinia, 1852, Schimper s.n. (P).
Solanum nigrum var. hirsutum Vahl, Symb. 2: 40. 1791.
Type. Based on Forsskål’s “S. aegyptiacum b) Fructu nigro; foliis integris villosissimus” (=Solanum memphiticum J.F.Gmel.)
Solanum hirsutum (Vahl) Dunal, Hist. Nat. Solanum 158. 1813.
Type. Based on Solanum nigrum var. hirsutum Vahl
Solanum grossidentatum A.Rich., Tent. Fl. Abyss. 2: 101. 1850, as “grossedentatum”.
Type. Ethiopia. “Tchélikote” [Chelicote], R. Quartin-Dillon & A. Petit s.n. (lectotype, designated by
Solanum nigrum var. rigidum Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852.
Type. Yemen. Sin. loc., 1837, P.E. Botta s.n. (lectotype, designated by
Solanum hirsutum var. abyssinicum Dunal, Prodr. [A. P. de Candolle] 13(1): 58. 1852.
Type. Ethiopia. Tigray: Adigrat, prope Adoam [Adoa] “nomen Abyssinicum: Alam tch’aguar”, G.H.W. Schimper 46 (lectotype, designated by
Solanum pruinosum var. pilosulum Dunal, Prodr. [A. P. de Candolle] 13(1): 59. 1852.
Type. Sudan. Blue Nile: Sennaar, 1831, G. Acerbi s.n. (holotype: G-DC [G00144592]).
Solanum subuniflorum Bitter, Repert. Spec. Nov. Regni Veg. 10: 546. 1912.
Type. Tanzania. Marangu near Mount Kilimanjaro, G. Volkens 2108 (neotype, designated by
Solanum plebeium var. grossidentatum (A.Rich.) Chiov., N. Giourn. Bot. Ital. 26: 159. 1919, as “grossedentatum”.
Type. Based on Solanum grossidentatum A.Rich.
Solanum nigrum var. grossidentatum (A.Rich.) De Wild., Pl. Bequaert. 1: 431. 1922, as “grossedentatum”.
Type. Based on Solanum grossidentatum A.Rich.
Solanum memphiticum var. abyssinicum (Dunal) Cufod., Bull. Jard. Bot. État Bruxelles 33(3): 872. 1963.
Type. Based on Solanum hirsutum var. abyssinicum Dunal
Locality unknown [Yemen or Saudi Arabia, but most likely Yemen], Herb. Forsskål 421 (lectotype, designated by
Annual or short-lived sprawling perennial herbs to 1.5 m tall, woody and branching at base. Stems spreading to decumbent, terete or occasionally very slightly angled, green, older stems green or straw colour, not markedly hollow; new growth densely viscid-pubescent with simple, spreading, uniseriate, mixed glandular and eglandular trichomes, these 3-10-celled, 0.5–2 mm long, with a terminal single-celled gland if glandular; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, (1.5-)2–9 cm long, (0.8-)1.2–5.5 cm wide, elliptic to ovate and widest in the basal third, membranous, concolorous, foul-smelling when crushed; adaxial surfaces moderately viscid-pubescent with a mixture of glandular and eglandular simple uniseriate trichomes 0.5–2 mm long like those of the stems, these denser along the veins; abaxial surfaces densely viscid-pubescent with similar glandular and eglandular simple uniseriate trichomes, these evenly distributed on veins and lamina; base acute, then attenuate and decurrent on to the petiole; margins entire or more often irregularly toothed, the teeth 2–4 mm long, acute; apex acute to acuminate, the tip often blunt and usually somewhat rounded; petioles 0.5–1.5 cm long, winged from the decurrent leaf base. Inflorescences 1–2.5(-3) cm long, internodal, simple, umbelliform to sub-umbelliform, with (2-)3–5(-8) flowers clustered at the tip, densely viscid-pubescent with mixed glandular and eglandular simple uniseriate trichomes like those of the stems; peduncle 0.9–2(-2.3) cm long, straight; pedicels 7–9 mm long, ca. 0.5 mm in diameter at the base, 0.5–0.8 mm in diameter at the apex, spreading, densely to moderately viscid-pubescent like the inflorescence axis, articulated at the base; pedicel scars clustered at the tip of the inflorescence, the scar from the basal flower spaced 1–2 mm from the rest. Buds globose to ovoid, the corolla exserted more than halfway from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 1–2.5 mm long, deeply conical, the lobes 1–1.5(-2) mm long, 0.5–0.8 mm wide, long-triangular, tips rounded, densely viscid-pubescent with mixed glandular and eglandular simple uniseriate trichomes to ca. 0.5 mm long. Corolla (8-)10–12 mm in diameter, white, rotate-stellate to stellate, lobed ca. 2/3 of the way to the base, the lobes 4–5 mm long, 3–4 mm wide, spreading or reflexed at anthesis, minutely pubescent-papillate abaxially with simple eglandular trichomes ca. 0.2 mm long, these white when dry. Stamens equal; filament tube minute; free portion of the filaments 0.5–1 mm long, glabrous or occasionally with a few tangled simple uniseriate trichomes adaxially; anthers (2-)2.5–3 mm long, 0.75–1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 4–6 mm long, minutely puberulent in the lower 1/4, merely papillate in plants with glabrous filaments, exserted ca. half the length of the anthers; stigma capitate-globose, bright green in live plants, the surface minutely papillate. Fruit a globose berry, 7–10 mm in diameter, black when ripe, the pericarp thin, matte and somewhat translucent; fruiting pedicels 0.8–0.9 cm long, ca. 0.75 mm in diameter at the base and to ca. 2 mm at the apex, somewhat woody, deflexed to spreading (often appearing like spokes of a wheel), dropping off with mature fruits, not persistent; fruiting calyx somewhat accrescent, the tube 2.5–3 mm long, the lobes 2.5–4 mm long, ca. 1.5 mm wide, somewhat spathulate, appressed to spreading, covering ca. 1/2 of the berry. Seeds 10–30 per berry, 1.5–2.5 mm long, 1.5–2 mm wide, flattened and tear-drop shaped with a subapical hilum, pale brown, the surfaces minutely pitted, the testal cells more or less pentagonal with some sinuate margins. Stone cells 2 per berry, 0.3–0.7 mm in diameter, usually borne near the base of the berry. Chromosome number: 2n=6x=72 (
Solanum memphiticum J.F.Gmel A Habit B Detail of stem with glandular indumentum C Detail of abaxial leaf surface D Glandular trichome E Eglandular trichome F Inflorescence G Flower at anthesis H Infructescence I Seed (A–E, Hde Wilde 8017; F–IGilbert 1387; G Nijmegen acc. A34570474). Scale bar: 3 cm (A), 3 mm (B), 4 mm (C, G), 0.8 mm (D–E), 1.5 cm (F, H) and 2 mm (I). Drawing by L. Smith.
Grows in open areas and along streams in forests and dry areas, also cultivated in eastern Africa; from 800 to 3,000 (-3,500) m elevation.
Ethiopia: alam tch’aguar; Kenya: ap-poinet, isusa, manage, nafu, ol’ momoit/olmomoi, soiyot, sonja sucha, sujet; Saudi Arabia: gebel soda; Uganda: ekyala ky’ente, kyalakyente, orushwiga, osiga.
In eastern Africa, leaves are eaten as spinach and berries are said to be edible.
(
Solanum memphiticum was long confused with S. villosum; both are similarly villous plants with toothed leaves that are sometimes pubescent with glandular trichomes.
In flower, S. memphiticum can be distinguished by its rotate-stellate corolla with small lobes that are strongly reflexed in flower and calyx lobes that extend beyond the corolla sinuses in open flowers. The flowers of S. memphiticum are delicate and only last a short time as compared to other species (the corolla bruises easily). Leaves of S. memphiticum are generally longer and thinner than those of S. villosum and the base is more decurrent on to the petiole so the free part of the petiole in S. memphiticum is much shorter than in S. villosum.
The parental origin of the tetraploid species has not been investigated in detail.
Solanum memphiticum was previously commonly known as S. grossidentatum or S. hirsutum (see
Selected specimens examined. Burundi. Ruyigi: Musongati, FTEA region: BUR, 10 May 1974, Reekmans 3419 (EA, MO).
Democratic Republic of the Congo. Nord-Kivu: Ruindi, Nov 1937, Lebrun 8380 (K).
Egypt. South Sinai: Wadi El Sheikh, Sinai, 15 Apr 1937, Shabetai s.n. (K).
Eritrea. Anseba: Geleb, Gheleb-Caropebir, 16 Jan 1893, Terracciano & Pappi 2027 (FT); Debub: Adi Ugri-Mai Tacala, 4 Sep 1909, Bellini 335 (FT); Saganeiti, gorge Gona pres Addingofon, 29 Mar 1892, Schweinfurth & Riva 1319 (FT, K); Gash Barka: Badum, lungo fiume Mareb, 10 Jan 1906, Pappi 6893 (FT); Get Arba, Gret-Arba, 7 Jan 1893, Terracciano & Pappi 1706 (FT); Maekel: Asmara, 12 Sep 1912, Baldrati 3536 (FT); Asmara, 2 Aug 1902, Pappi 2099 (EA, MO, P, SI); Semienawi Keyih Bahri: Nefasit-Maha-bar, 2 Feb 1909, Fiori 1593 (FT); Illalia-Scilliki; Assaorta, 28 Mar 1893, Pappi 3601 (FT).
Ethiopia. Addis Ababa: Addis Ababa, 18 Jul 1962, Mooney 9136 (FT, K); Amhara: c. 15 km S of Debre Sina on the main rd towards Debre Berhan, 22 Nov 2000, Friis et al. 10115 (K); Dessie, Wollo Prov., 18 Aug 1946, Hall 22 (BM); Wello Prov., Azewagedel mountain, 2 km E of Desse, 11 Apr 1969, Sutherland 167 (MO); Dessie, 26 May 1938, Vatova 2417 (FT); Oromia: Borena, Mega, presso fortino nuovo, 4 May 1937, Cufodontis 627 (FT, W); Agheremariam-Dilla rd, 3 Dec 1952, Gillett 14586 (EA, FT, K); lower slopes of Mt. Cilalo, nr Asella, 10 Sep 1965, de Wilde & de Wilde-Duyfjes 8017 (K, MO, P); Southern Nations (SNNP): Gurage Mountains, above Butajira towards the village of Ageta on the track towards Endibis, 24 Feb 2000, Friis et al. 10066 (K); Tigray: Amba Alaga pass, 9 Oct 1995, Friis et al. 6640 (K); Bellaka, 8 Nov 1854, Schimper 506 (E, FT, W); Arba Tensesa, 7 Oct 1862, Schimper 523 (BM).
Jordan. East Jordan, Pelit, 30 Apr 1963, Gillett 15973 (K); Aqaba: Wadi Rum, 13 Apr 1945, Davis 8987 (E); Ma’an: Petra, 28 Dec 1935, Dinsmore 12151 (E).
Kenya. Central: Nyeri, Aberdare National Park, The Ark, 7 Apr 1975, Hepper & Field 4912 (K); Kiambu, Makuyu, Fort Hall Dist, 26 Jun 1960, van Someren 11980 (EA, K); Eastern: Makueni, Chyulu North, 21 May 1938, Bally B-7788 (K); Makueni, Chyulu North, 28 Apr 1938, Bally B-8306 (K); Makueni, Chyulu Hills North, Apr 1938, van Someren 7644 (K); Nairobi: Mathare Valley, between Mathare Police Station and Eastleigh Section One, 11 Sep 1971, Mwangangi & Kasyoki 1797 (EA, K); Nairobi, FTEA region K4, 1942, Nattrass 341 b (EA, MO); Rift Valley: Kericho, Londiani to Elburgon, Dec 1905, Baker K-346 (EA, K); Nakuru, Lake Naivasha, West shore Mennel’s Farm, 14 Feb 1971, Gillett 19300 (EA, K); Glover et al. 181 (EA, K); Kajiado, Olekairtoror Escarpment, 20 mi from Narok on Nairobi rd, 14 Jul 1962, Glover & Samuel, 3110 (EA, K); K6 Rift Valley, Suswa volcano, 1 Jun 1997, Phillipson & Bytebier 4785 (MO); Western: Nandi, Kapsoret Forest, 15 Jun 1951, Williams 240 (EA, K).
Saudi Arabia. Al Mahmoud, 35 km N of Abha, 21 May 1980, Boulos & Ads 14150 (E, K); Abha Pass (nr. W. Abha), 25 Oct 1971, Popov 71 257 (BM); Asir: Jebel Sudah, c. 18 km N of Abba, 5 Apr 1979, Collenette 1269 (K).
Somalia. Upper Sheikh Kitchen Garden, 1 Dec 1919, Godman 67 (BM, MO); Saaxil: Ally Ully nr Sheikh, 11 May 1973, Wood S/73-65 (K).
Sudan. Darfur: Jebel Marra, Golel, c. 120 km E of Zalingei, 22 Jan 1965, de Wilde et al. 5504 (K, MO).
Tanzania. Arusha: Lake Manyara National Park, Mto ya Ukindu, 29 Nov 1963, Greenway & Kirrika 11096 (EA, K); Monduli Forest Reserve, T2. Nr Olchoropus Village, 25 Jan 2001, Simon et al. 720 (MO); Central: Kondoa, Great North rd, 11 Jan 1962, Polhill & Paulo 1130 (K); Kagera: Ngara, Murugwanza, Ibivyaza Bakobwa, Bugufi, 20 Jan 1961, Tanner 5617 (K); Kilimanjaro: Osirwa Farm, TBL Estates, 26 Jan 1994, Grimshaw 94-148 (K); Ulei, Kwa Sadala, 2 May 1994, Grimshaw 94-470 (K); Moshi, Sanya River, Mar 1928, Haarer 1209 (EA, K); Lake: Ngara, Kirushya, Bugufi, 23 Nov 1959, Tanner 4530 (K); Mbulu: Pienaars Heights, Great North rd, 120 mi S of Arusha, 5 May 1962, Polhill & Paulo 2342 (EA, K); Northern: Mbulu, Mbulumbul, Block D1, 24 Jun 1944, Greenway 6952 (EA, K).
Uganda. Central: Masaka, Kasambya, Kigezi, Feb 1948, Purseglove P-2595 (K); Eastern: Sironko, Budadiri, Bugishi, Jan 1932, Chandler 405 (K); Serere, at Tira, Jul 1926, Maitland 1290 (K); Northern: Zombo, Paidha, 28 Aug 1953, Chancellor 185 (K); Western: Kigezi D.F.I, 28 Aug 1972, Goode G-5 -72 (K); Kasese, Muhokya, Ruwenzori, 25 Dec 1925, Maitland 1790 (K); Kigezi, Kachwekano Farm, Sep 1949, Purseglove 3121 (EA, K); Kisoro, Virunga-Kette, Muhavura, Nkanda, 25 Nov 1954, Stauffer 957 (EA, K, P).
Yemen. Hadramaut: Al Mukalla, Arabia, West rd, 6 Sep 1949, Guichard KG/HAD 35 (BM); Kor Seiban, 14 Sep 2002, Killian et al. YP-3578 (B); Sa’dah: Sadah, 1 Jul 1984, Gordon 609 B (E); Sana’a: Sana’a, 12 Mar 1981, Miller 3004 (E); 14 Feb 1934, Rathjens s.n. (BM); Menacha, 7 Mar 1889, Schweinfurth 1476 (BM, GH, K, P); Ar Rowdah nr San’a, 23 Feb 1972, Wood 72-7 (BM); 10 Oct 1974, Wood Y/74-20 (BM); Ta’izz: Jabal Sabir, c. 15 km S of Taizz, 11 Jun 1982, Gordon 11 (E); Jabal Sabir, nr Taiz, 23 Sep 1977, Lavranos 15947 (E).
Solanum nigrum var. vulgare L., Sp. Pl. 186. 1753.
Type. “Solanum 3 α”; cultivated in George Clifford’s garden in Hartekamp, The Netherlands, Anon. s.n. (lectotype, designated here: BM [BM000558026]).
Solanum nigrum var. judaicum L., Sp. Pl. 186. 1753.
Type. Unknown (no specimens or illustrations cited).
Solanum vulgatum Baumg., Fl. Lips. 120. 1790, nom. illeg. superfl.
Type. Based on Solanum nigrum L. [S. nigrum L. from Syst. Veg. ed. 14: 224. 1784 cited in synonymy]
Solanum humile Salisb., Prodr. Stirp. Chap. Allerton 134. 1796, nom. illeg. superfl.
Type. Based on Solanum nigrum L. (cited in synonymy)
Solanum judaicum Besser, Prim. Fl. Galiciae Austriac. 1: 183. 1809.
Type. No localities of specimens cited; probably from what is now Ukraine (no specimens cited; no original material located).
Solanum morella Desv., Pl. d’Angers 113. 1818, nom. illegit. superfl.
Type. Based on Solanum nigrum L. (cited in synonymy)
Solanum parviflorum Moretti ex Badarò, Giorn. Fis. Chim. Storia Nat. Med. Arti Dec. 2, 7: 364. 1824, nom. illeg, not Solanum parviflorum Nocca (1793)
Type. Italy. [Liguria]“ in olivetis Liguriae occid[ose]”, 1824, G.B. Badarò s.n. (no specimens cited; lectotype, designated by
Solanum cestrifolium Jacq. ex Spreng., Syst. Veg., ed. 16 [Sprengel] 1: 680. 1825.
Type. “Patria?” [origins unknown, although probably from cultivation?] (no specimens cited; no original material found).
Solanum rhinozerothis Blume, Bijdr. Fl. Ned. Ind. 13: 695. 1826.
Type. Indonesia [no locality cited in protologue], C.L. Blume s.n. (no specimens cited; neotype, designated here: L [L2883159]).
Solanum vulgatum (L.) Spenn., Fl. Friburg. 2: 427. 1826.
Type. Based on Solanum nigrum var. vulgare L.
Solanum vulgatum var. nigrum (L.) Spenn., Fl. Friburg. 2: 427. 1826.
Type. Based on Solanum nigrum L.
Solanum vulgatum var. chlorocarpum Spenn., Fl. Friburg. 3: 1074. 1829.
Type. Switzerland. Fribourg: Sin. loc. (no specimens cited; no original material found).
Solanum moschatum C.Presl, Delic. Prag. 77. 1832.
Type. Italy. Sicily: Palermo (“in cultis ruderatis Panormi Siciliae”, Anon. s.n. (no specimens cited; original material at PR?, PRC?, not found).
Solanum nigrum var. perennans Bertol., Fl. Ital. [Bertoloni] 2: 634. 1836.
Type. Based on Solanum moschatum C.Presl.; Solanum parviflorum Moretti ex Badarò (no specimens cited; no original material found).
Solanum nigrum var. atriplicifolium G.Mey., Chloris Han. 265. 1836.
Type. France. Pays de la Loire: “St. Germain de Calberte, basse Lozère”/”Mans [Le Mans]”[annotation “Solanum atriplicifolium” in Desportes hand], 1806, [illegible] s.n. (lectotype, designated here: G-DC [G00144334]).
Solanum nigrum var. atriplicifolium Desp. ex G.Don, Gen. Hist. 4: 412. 1838, nom. illeg. (isonym), not Solanum nigrum var. atriplicifolium G.Mey. (1836)
Type. Based on same original material as Solanum nigrum var. atriplicifolium G.Mey.
Solanum vulgare Hegetschw., Fl. Schweiz 219. Dec 1838-Jan 1839, nom. illeg. superfl.
Type. Based on “Solanum nigrum Willd.” (= S. nigrum L.)
Solanum chenopodium Raf., Autik. Bot. 107. 1840.
Type. “Europa” (no specimens cited; original material probably lost).
Solanum exaratum Raf., Autik. Bot. 107. 1840.
Type. “Europa” (no specimens cited; original material probably lost).
Solanum bidentatum Raf., Autik. Bot. 108. 1840.
Type. “Italia, Sicilia” (no specimens cited; original material probably lost).
Solanum tauschii Opiz, Oekon.-techn. Fl. Böhm. [Berchtold & al.] 3: XX. 1843.
Type. Czech Republic. “Prag”, I.F. Tausch s.n. [Herb. Flor. Boehm. 1076] (no herbaria cited; no original material found, perhaps at PR?); “Luzic”, Sadel s.n. (no herbaria cited; no original material found, perhaps at PR?); “Folimanta bei Prag”, Tanbler s.n. (no herbaria cited; no original material found, perhaps at PR?); Sin loc., P.M. Opiz 10/8 40 (no herbaria cited; no original material found, perhaps at PR?).
Solanum reineggeri Opiz, Oekon.-techn. Fl. Böhm. [Berchtold & al.] 3(2): XIX. 1843.
Type. Austria. Niederösterreich; Sin. loc., Reinegger s.n. (no herbaria cited; no original material found, perhaps at PR?).
Solanum decipiens Opiz, Oekon.-techn. Fl. Böhm. [Berchtold & al.] 3(2): XXIV. 1843.
Type. Czech Republic. “Troja”, P.M. Opiz 10/838; “Ruchelbad”, P.M. Opiz 23/10 835; “Radlic”, P.M. Opiz 4/8 40; “Szaslau”, Janoti 804 (no herbaria cited; no original material found, perhaps at PR?).
Solanum schultesii Opiz, Oekon.-techn. Fl. Böhm. [Berchtold & al.] 3(2): XXIV. 1843.
Type. Czech Republic. “ Im Baumgarten” P.M. Opiz 10/10 35; “Prag”, P.M. Opiz s.n. (no herbaria cited; no original material found, perhaps at PR?).
Solanum nigrum forma stenopetalum A.Braun ex Döll, Rhein. Flor. 412. 1843.
Type. Germany. “Bei Ettlingen”, A. Braun s.n. (no herbaria cited; no original material found, not found at KR); Germany. “Bei Carlsruhe in der Nähe des Linkenheimer Thores”, A. Braun & J.C. Döll s.n. (no herbaria cited; no original material found, not found at KR).
Solanum nigrum forma chlorocarpum A.Braun ex Döll, Rhein. Flor. 413. 1843.
Type. Germany. “Bei Gerolsau und Lichtenthal unweit Baden”, A. Braun & J.C. Döll s.n. (no herbaria cited; no original material found, not found at KR); Germany. “Bei Carlsruhe, Knielingen”, A. Braun s.n. (no herbaria cited; no original material found, not found at KR); Germany. “Mannheim, Dossenheim, Oppenheim”, J.C. Döll s.n. (no herbaria cited; no original material found, not found at KR).
Solanum guineense var. nepalense Dunal, Prodr. [A. P. de Candolle] 13(1): 49. 1852.
Type. Cultivated in Avignon Botanic Garden, from eastern Nepal, Herb. Requien s.n. (lectotype, designated here: AV [Herb. E. Requien]).
Solanum pterocaulum var. deppei Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, as ‘pterocaulon’.
Type. Cultivated in France at Montpellier “Solanum deppei. In hortis bot. cultum” (no specimens cited, described from living plants “v.v. Hort. Monsp.”; neotype, designated here: MPU [MPU310704]).
Solanum paludosum Dunal, Prodr. [A. P. de Candolle] 13(1): 57. 1852.
Type. Myanmar “cat. itir. Burm.”, 1827, N. Wallich 402 (holotype: G-DC [G00144525]).
Solanum roxburghii Dunal, Prodr. [A. P. de Candolle] 13(1): 57. 1852.
Type. “In India orientali” (no specimens cited; lectotype, designated here: Wight, Icones plantarum Indiae Orientalis 2: t. 344. 1843) “Peninsula Ind. orientalis”, R. Wight s.n. [herb. Wight 2326] (epitype, designated here: E [E00718973]).
Solanum memphiticum var. repandum Dunal, Prodr. [A. P. de Candolle] 13(1): 47. 1852.
Type. Germany. Saxony: cultivated in Leipzig [fide JE where original set is stored] “colui”, 1824, W. Gerhard s.n. (holotype: G [G00144263]; isotypes: JE [JE00009838], W [1889-0232903, 1889-0283857]).
Solanum nigrum var. atriplicifolium Desp. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852, nom. illeg., Solanum nigrum var. atriplicifolium G.Mey. (1836).
Type. Based on same original material (specimen) as Solanum nigrum var. atriplicifolium G.Mey.
Solanum cechicum Opiz, Lotos 4: 94. 1854.
Type. Czech Republic. “Um Prag”, P.M. Opiz s.n. (no herbaria cited; no original material found, perhaps at PR?).
Solanum nigrum forma judaicum Miq., Fl. Ned. Ind. 2: 637. 1856.
Type. Indonesia. Java: [West Java] Buitenzorg [Bogor], C.L. Blume s.n. (lectotype, designated here: L [L2880952]).
Solanum nigrum forma paludosum (Dunal) Miq., Fl. Ned. Ind. 2: 637. 1856.
Type. Based on Solanum paludosum Dunal
Solanum
hirsutum
Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg., not Solanum hirsutum (Vahl)
Type. Slovakia. “ex itinere Arvensi”[Árva], P. Kitaibel s.n. (lectotype, designated here: BP [Herb. Kit. fasc. IX No. 103]).
Solanum acutifolium Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg., not Solanum acutifolium Ruiz & Pav. (1799).
Type. Slovakia. “ex itinere Arvensi”[Árva], P. Kitaibel s.n. (lectotype, designated here: BP [Herb. Kit. fasc. IX No. 103]).
Solanum nigrum var. macrocarpum Schur, Enum. Pl. Transsilv. 478. 1866.
Type. Romania. Sibiu: Sibiu “In ruderalis prope Cibinium Transilv.” [protologue “Schur sert. n. 1994, auf unbebauten Orten bei Hermannstadt, Aug.”], Oct, F. Schur s.n. (neotype, designated here: LW [LW00210121]; no duplicates found at BRNU).
Solanum nigrum var. chlorocarpum (Spenn.) Schur, Enum. Pl. Transsilv. 478. 1866.
Type. Based on Solanum vulgatum var. chlorocarpum Spenn.
Solanum nigrum var. glabrum Lowe, Man. Fl. Madeira 2: 73. 1872.
Type. Portugal. Madeira: Mr. Gordon’s kitchen garden, the Mt., 8 Dec 1932, R.T. Lowe 16 [a] (lectotype, designated by
Solanum nigrum var. hebecaulon Lowe, Man. Fl. Madeira 2: 73. 1872.
Type. Portugal. Madeira: [“Levada de Sta. Luzia, above Funchal, Feb”, protologue], R.T. Lowe [?] s.n. (no specimens corresponding to this locality and date located).
Solanum morella subsp. nigrum (L.) Rouy, Fl. France 10: 364. 1908.
Type. Based on Solanum nigrum L.
Solanum ganchouenense H.Lév., Repert. Spec. Nov. Regni Veg. 11: 295. 1912.
Type. China. Guizhou: Gan Chouen, Aug 1910, J. Cavalérie 3815 (holotype: E [E00284474]; isotypes: E [E00284475], K [K001080605], P [P00055234]).
Solanum chenopodiifolium H.Lév., Repert. Spec. Nov. Regni Veg. 12: 531. 1913.
Type. China. Yunnan: Tong-Tchouan plaine, Sep 1912, E.E. Maire s.n. (holotype: E [E00284477]).
Solanum nigrum subvar. atriplicifolium (Desp. ex Dunal) Schinz & Thell., Fl. Schweiz, ed. 3, 2: 295. 1914.
Type. Based on Solanum nigrum var. atriplicifolium Desp. ex Dunal (=Solanum nigrum var. atriplicifolium G.Mey.)
Solanum peregrinum E.P.Bicknell, Bull. Torrey Bot. Club 42: 332. 1915.
Type. United States of America. Massachusetts: Nantucket County, Nantucket street, E.P. Bricknell 7719 (holotype: NY [00138955]; isotype: NY [00073847]).
Solanum probstianum Polg., Mitteil. Naturfor. Gesellsch. Solothurn 12: 30. 1938.
Type. Cultivated in Hungary at Györ, from seeds sent by R. Probst from Switzerland (Solothurn: Derendingen [Kammgarnfabrik Derendingen] in 1932), 23 Aug 1933, S. Polgár s.n. [Herb. Polg. 4051] (no specimens cited; lectotype, designated here: BP [BP-272406]).
Solanum nigrum var. schultesii (Opiz) Rouy, Acta Horti Gothob. 10: 201. 1935.
Type. Based on Solanum schultesii Opiz
Solanum pseudoflavum Pojark., Bot. Mater. Gerb. Inst. Komarova Akad. Nauk S.S.S.R. 17: 338. 1955.
Type. Kazakhstan. Between Vernoy Alma-Ata, Vernenskiy area, between town of Vtrnym and the station Karasukskaya, the village of Dimitrivka, V.S. Titov 2220 (holotype: LE).
Solanum nigrum forma pallidum Wessely, Repert. Spec. Nov. Regni Veg. 63: 311. 1960, as “Solanum nigrum subsp. nigrum var. atriplicifolium forma pallidum”.
Type. Germany. Rhineland-Palatinate: Neuwied, Wirtgen s.n. (holotype: W [not seen]).
Solanum nigrum forma luridum Wessely, Repert. Spec. Nov. Regni Veg. 63: 311. 1960, as “Solanum nigrum subsp. schultesii forma luridum”.
Type. Germany. Saxony: Dresden, Trümmerstellen am Postplatz, 22 Sep 1957, I. Wessely 0.23 (holotype: GFW).
Solanum nigrum subsp. schultesii (Opiz) Wessely, Feddes Repert. Spec. Nov. Regni Veg. 63: 311. 1960.
Type. Based on Solanum schultesii Opiz
Solanum nigrum var. incisum Täckh. & Boulos, Publ. Cairo Univ. Cairo Herb. 5: 101. 1974 [“1972”].
Type. Egypt. Faiyum, Sinnuris, L. Boulos s.n. (holotype: CAI [CAI000175]).
“Habitat in Orbis totius cultis” [sheet marked with Θ, meaning central part of Asia = Middle East], Without collector (lectotype, designated by
Annual or short-lived erect to sprawling perennial herbs to 1.0 m tall, subwoody and branching at base. Stems spreading to decumbent, terete to sharply angled and ridged, green, the ridges often spinescent, older stems not appearing spinescent, not markedly hollow; new growth pubescent with simple, spreading, uniseriate, eglandular or glandular trichomes, these 1-6-celled, 0.5–0.6 mm long; older stems glabrescent, the trichome bases persisting as pseudospines. Sympodial units difoliate, the leaves not geminate. Leaves simple, 3.8–7.2(-14.5) cm long, 2.5–5.0(-9.5) cm wide, broadly ovate, membranous, green, concolorous, without smell or smell somewhat foetid; adaxial surface sparsely pubescent with spreading, simple, uniseriate trichomes like those on stem evenly scattered along veins and lamina; abaxial surface more densely pubescent along veins and sparsely along lamina with eglandular and/or glandular trichomes like those of the stems; major veins 5–7 pairs; base obtuse to truncate, somewhat attenuate; margins sinuate-dentate, especially in the lower 2/3, to occasionally entire or deeply toothed; apex acute; petioles 0.5–3.0 cm long, pubescent with simple uniseriate glandular and eglandular trichomes like those of the stems. Inflorescences 0.8–2.0 cm long, internodal, simple to occasionally furcate, the flowers spaced along the rhachis, with (3-)4–10 flowers, pubescent with spreading simple uniseriate trichomes like those on stem; peduncle 0.5–1.5 cm long, straight; pedicels 3–5 mm long, 0.2–0.3 mm in diameter at the base and 0.2–0.3 mm at the apex, spreading, articulated at the base; pedicel scars spaced 0.3–0.7 mm apart. Buds subglobose, the corolla approximately halfway exserted from the calyx before anthesis. Flowers 5-merous, all perfect. Calyx tube 0.8–1.0 mm long, conical, the lobes 0.5–0.8 mm long, 0.6–0.8 mm wide, triangular with acute or somewhat rounded apices, pubescent with spreading simple uniseriate eglandular and glandular trichomes like those of the pedicels. Corolla 10–12 mm in diameter, white with a yellow-green central portion near the base, stellate, lobed 1/2 to 2/3 of the way to the base, the lobes 4.0–5.0 mm long, 2.0–2.5 mm wide, strongly reflexed at anthesis, later spreading, densely papillate-pubescent abaxially with simple uniseriate eglandular trichomes. Stamens equal; filament tube very short to minute; free portion of the filaments 0.5–0.7 mm long, adaxially pubescent with spreading uniseriate simple trichomes; anthers 1.8–2.5 mm long, 0.8–1.0 mm wide, ellipsoid, very slightly wider at base, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.5–3.5 mm long, densely pubescent with tangled 2-3-celled simple uniseriate trichomes in the lower half where included in the anther cone, exserted 0–1 mm beyond anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 6–10 mm in diameter, purple-black or green to yellowish-green at maturity, the pericarp dull or slightly shiny; fruiting pedicels 10–12 mm long, 0.4–0.5 mm in diameter at the base and 1.0–1.1 mm at apex, generally spreading to occasionally recurved, spaced 1.0–2.0 mm apart, dropping with mature fruits, not persistent but occasionally remaining on the inflorescence; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes 1.0–2.0 mm long, spreading to reflexed in fruit. Seeds (15-)20–40 per berry, 1.8–2.0 mm long, 1.5–1.6 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells absent (North America and Europe) but usually 2(-8) per berry in other areas (Asia), ca. 0.5 mm in diameter, brown. Chromosome number: 2n=6x=72 (
Solanum nigrum L. A Habit B Inflorescence (dense indumentum) C Inflorescence (sparse indumentum) D Fully mature full-black berries, calyx lobes remaining appressed or slightly spreading (A Nijmegen acc. 824750016; B Nijmegen acc. A34750479; C Nijmegen acc. 824750029A, D Nijmegen acc. A44750150). Photos by S. Knapp.
Weed of cultivated land, open spaces in forests and roadsides; found in disturbed areas between 0–2,200 (3,500) m elevation.
Australia: blackberry, black berry nightshade (
Leaves used as vegetable in India, China, southeast Asia and in Europe (in local areas); thought to be poisonous by association with the deadly nightshade, Atropa belladonna (see Uses in introductory section). Berries sometimes used for jam (
(
Solanum nigrum, the type species of the genus Solanum, is a widespread weed with much morphological variation recognised at various infraspecific levels by many different authors (e.g.
We do not include material identified as S. nigrum in recent African regional floras (
Throughout its range in Europe and into Eurasia as far as western China, S. nigrum is sympatric with S. villosum. The simplest distinguishing character is mature berry colour; S. villosum has red, orange or yellow berries, while those of S. nigrum are black or green. Many old collections, however, do not state berry colour on the label, so identification can be difficult. Calyx lobes are useful for distinguishing these taxa; S. nigrum calyx lobes are usually deltate and acute, with sharp triangular sinuses, while those of S. villosum are longer, usually rounded at the tip and the sinuses are broad and quite transparent (see description of S. villosum), leaving a paler “window” just below the sinus in flower buds and early flowers. The shiny, translucent berries of S. villosum (mostly slightly ellipsoid) usually dry blackish-brown, but are distinct from the matte, more opaque berries of S. nigrum. Neither species has stone cells in Europe, but in Asia, S. nigrum usually has 2 (or occasionally more) stone cells in the berries. Both species retain pedicels after fruits drop, but S. nigrum is not as extreme in this regard and often plants are found with old inflorescences with no remaining pedicels.
Solanum nigrum has been considered native only to Europe, but our study of populations of this widespread weed across its range has shown that the largest morphological variation can be observed in Asia. Populations of S. nigrum from Asia have more stone cells in the fruit and the plants have a more delicate look overall with longer peduncles and often fewer flowers per inflorescence. Material from Asia has been described as different taxa (e.g. S. guanchounense, S. chenopodiifolium) but the variation is continuous across the range, with European populations being more invariant (except in leaf shape and indumentum). Populations in Europe represent a relatively monomorphic set of populations compared to material from China and eastern Asia. We therefore consider that the species is native to the entire Eurasian area, with limited introductions to North America. North American material we have seen appears most similar to European plants based on morphology, suggesting the introduction to North America came from European populations, but this has not been tested genetically.
Australian populations of S. nigrum could have originated either from Europe or from Asia. In general, S. nigrum in Australia does not seem to have stone cells in general and plants fall into two continuously variable groups: one is similar to classic European S. nigrum and the other is more similar to plants from eastern and south-eastern Asia.
Solanum nigrum is an autoalloploid species, now thought to have originated from a tetraploid S. villosum and a diploid S. americanum by spontaneous amphiploidy (
In parts of eastern England, S. × procurrens A.C.Leslie (
In describing S. judaicum,
The name S. cestrifolium has a complex history; we can find no place of publication for “Solanum cestrifolium Jacq.” as cited by
The varietal epithet “atriplicifolium” was used by many authors to refer to plants of S. nigrum with dentate leaf margins. It appears to have been used in reference to a specimen that was annotated “Solanum atriplicifolium” by Narcisse Desportes, probably seen in Paris or Geneva. Georg Meyer of Hannover (1818) was the first botanist to effectively and validly publish this epithet and did so at the varietal rank.
The various names coined by the amateur Czech botanist Philipp Maximilian Opiz and here recognised as synonyms of S. nigrum were all included by him in the “superspecies” S. nigrum and, from descriptions and key, represent leaf shape, pubescence and inflorescence size variations of that species. Opiz’s enormous herbarium is housed mostly in PR (
Solanum chenopodium, S. exarmatum and S. bidentatum were all coined by
A single collection of a plant from Nepal cultivated in Avignon was cited in the protologue of S. guineense var. nepalense (
Solanum roxburghii was coined by
The names coined by the Hungarian botanist Pál Kitaibel were published posthumously by
We have selected a specimen collected by Schur (LW00210121), but with a different date of collection and exact locality, as the neotype for S. nigrum var. macrocarpum; searches in the other relevant herbaria revealed no original material.
Solanum probstianum was described from material cultivated in Hungary from seeds sent by Rudolf Probst to Sandor Polgár in 1932. Although no specific specimens were cited in Polgár’s protologue (in
A total of 1,537 specimens were examined from 82 countries during the study across Africa, Asia, Australia, Eurasia and the Pacific. Adventive specimens from the New World were also studied in order to understand the full range of morphology within the species. All specimens examined can be seen in Appendix 2 (csv format) and Appendix 3 (traditional Specimens Examined list in pdf format).
Solanum styleanum Dunal, Prodr. [A. P. de Candolle] 13(1): 44. 1852.
Type. Chile. Sin. loc., J. Styles s.n. (holotype: G-DC [G00144016]).
Bosleria nevadensis A.Nelson, Proc. Biol. Soc. Washington 18(30): 175. 1905.
Type. United States of America. Nevada: Washoe County, Pyramid Lake, 9 Jun 1903, G.H. True s.n. (holotype: RM [RM0004387]).
Solanum nitidibaccatum forma integrifolium Blom, Acta Horti Gothob. 24: 118. 1961.
Type. Sweden. Göteborg, som ogräs i Göteborgs Botaniska Trädgård, 10 Sep 1948, C. Blom s.n. (lectotype, designated here: GB [GB-0146965]; isolectotype: E [E00593447]).
Solanum patagonicum C.V.Morton, Revis. Argentine Sp. Solanum 146. 1976.
Type. Chile. Región XII (Magallanes): Río Paine, 100m, 15 Jan 1931, A. Donat 415 (holotype: BM [BM000617673]; isotypes: BA, BAF, GH [GH00077732], K, SI [SI003331, SI003332], US [00027733, acc. # 2639758]).
Solanum physalifolium var. nitidibaccatum (Bitter) Edmonds, Bot. J. Linn. Soc. 92: 27. 1986.
Type. Based on Solanum nitidibaccatum Bitter
Chile. Sin. loc., 1829, E.F. Poeppig s.n. (lectotype, designated by
Annual prostrate or spreading herbs to 20 cm tall, branching at base. Stems decumbent or ascending, terete, green, not markedly hollow; new growth densely viscid-pubescent with simple, spreading, uniseriate, translucent, glandular trichomes, these 2-8(10)-celled, 1.5–2.0 mm long, with a glandular apical cell; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 2.0–5.5(-9.5) cm long, 1.5–5.0(-6.5) cm wide, ovate to broadly ovate, rarely elliptic, membranous, green, concolorous, without smell; adaxial surface sparsely pubescent with spreading 2–4-celled translucent, simple, uniseriate gland-tipped trichomes like those of the stem, these denser along the veins; abaxial surface more evenly densely pubescent on the lamina and veins; major veins 3–6 pairs, not clearly evident abaxially; base attenuate to cuneate, at times asymmetric, decurrent on the petiole; margins entire or sinuate-dentate; apex acute to obtuse; petioles 0.5–2.7(-4.5) cm long, sparsely pubescent with simple uniseriate glandular trichomes like those of the stems and leaves. Inflorescences 1.0–2.0 cm long, generally internodal but occasionally leaf-opposed, simple, the flowers spaced along the rhachis, with 4–8(-10) flowers, sparsely pubescent with spreading trichomes like those on stems and leaves; peduncle 0.6–1.3 cm long, straight; pedicels 4–12 mm long, 0.1–0.2 mm in diameter at the base and 0.2–0.4 mm at apex, straight and spreading, articulated at the base; pedicel scars spaced 0.3–1 mm apart. Buds subglobose, the corolla only slightly exserted from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 1–2 mm long, conical, the lobes 1.7–2.5 mm long, less than 1 mm wide, triangular with acute to obtuse apices, sparsely pubescent with 1–4-celled glandular trichomes like those of the pedicels. Corolla 4–6 mm in diameter, white with a yellow-green central eye with black “V” or “U” shaped margins in the lobe sinuses, rotate-stellate, lobed 1/3 of the way to the base, the lobes 2.3–3.2 mm long, 2.5–3.7 mm wide, spreading at anthesis, sparsely papillate-pubescent abaxially with 1–4-celled simple uniseriate trichomes, especially along tips and midvein. Stamens equal; filament tube minute; free portion of the filaments 1.5–2.0 mm long, adaxially sparsely pubescent with tangled uniseriate 4–6-celled simple trichomes; anthers 1.0–1.4 mm long, 0.5–0.8 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.5–3.0 mm long, densely pubescent with 2–3-celled simple uniseriate trichomes in the lower half where included in the anther cone, exserted 0.2–1.0 mm beyond the anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 4–13 mm in diameter, brownish-green and marbled with white (this not easily visible in herbarium specimens) at maturity, the surface of the pericarp usually shiny; fruiting pedicels 4–13 mm long, ca. 0.2 mm in diameter at the base, spaced 1–3 mm apart, reflexed and slightly curving, dropping with mature fruits, not persistent; fruiting calyx accrescent, becoming papery in mature fruit, the tube ca. 3 mm long, the lobes 2.5–3.5(-4) mm long, 3–4 mm wide, appressed against the berry, but the berry clearly visible. Seeds 13–24 per berry, 2.0–2.2 mm long, 1.2–1.4 mm wide, flattened and tear-drop shaped with a subapical hilum, brown, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells usually (1-)2–3 per berry, occasionally absent, ca. 0.5 mm in diameter. Chromosome number: 2n=2x=24 (
Grows along roadsides, in disturbed and cultivated areas in the shade of trees and shrubs, in rocky and sandy areas; between sea level and 2,000 (-2,700) m elevation in its native range, between sea level and 2,400 m in the Old World primarily as a weed in cultivations.
Australia: cherry nightshade (
None recorded; a weed of agriculture and actively controlled.
(
Solanum nitidibaccatum is morphologically similar to S. sarrachoides and has been treated under that taxon in many previous treatments (e.g.
Solanum nitidibaccatum is a diploid species native to the south-eastern parts of South America and within the Old World is morphologically most similar to S. sarrachoides, with which it has been confused. The two species are best distinguished by the complete inclusion of buds in calyx before anthesis, the larger stone cells in berries and the more erect habit of S. sarrachoides (see key in
Solanum nitidibaccatum has been introduced extensively to other parts of the world where it has become a prolific and successful weed of disturbed sites usually associated with agriculture of the wool trade. Trade with South America — particularly the importation of grain, seeds and the spreading of wool ‘shoddy’ — has been largely responsible for its introduction into Europe, with one of its common European names being the Argentinian nightshade. The taxon is now a widespread adventive in Europe where it is rapidly becoming naturalised and often forms extensive populations. It has been introduced into Australia on a number of occasions, where it persists as a weed of cultivation and is sparingly established in all States (
In parts of eastern England, S. × procurrens A.C.Leslie (
The amateur Swedish botanist Carl Blom rarely cited herbaria for his names; we have selected the sheet in GB, where he worked, as the lectotype of his var. integrifolium.
Australia. Australian Capital Territory: CSIRO Black Mountain site, Acton, Canberra, 9 Jan 1995, Lepschi 1729 (AD, CANB, HO,NSW); New South Wales: Canyonleigh via Marulan, May 1976, Cooper s.n. (NSW); White Rock, 8 km upstream Bathurst, 9 May 1982, Dellon s.n. (NSW); Queensland: The Summit c. 5 miles NNE of Stanthorpe, 8 Feb 1972, Henderson & Parham 1241 (AD, BRI, MEL, NSW); South Australia: Adelaide Hills, 20 Mar 1986, Jackson 5976 (AD, CANB); Tea Tree Gully, 12 Mar 1974, Spooner 3341 (AD); Mitcham, 18 Jan 1980, Winn s.n. (AD); Tasmania: Murphy’s Flat Reserve, 2010 (Bush Blitz), 25 Mar 2010, Baker 2203 (HO); 12 Jan 2001, Buchanan 15827 (HO); Victoria: Australian Alps, 19 Jan 1981, Beauglehole 68690 (MEL); Melbourne; Ripponlea Estate, Hotham Street, Elsternwick, 6 Mar 1984, Clarke 1682 (NSW); Milne’s property, 27 Jan 1966, Shepherd 215 (CANB); Riverina, Jan 1975, Without Collector s.n. (MEL); Western Australia: Badgerup Road opposite Jambaris Road, Wanneroo, 15 May 1997, Burt s.n. (PERTH); Pinjarra, 17 Nov 1987, Sheldow s.n. (PERTH).
Austria. Burgenland: Nordburgenland, WSW von Eisenstadt, S von Müllendorf, 2 Aug 2007, Barta s.n. (BM); Seewinkel, zw. Apetion u Frauernkirchen, knapp W v E-Teil d. Fuchslochlacke, W v Feldweg, 12 Oct 2008, Walter 7085 (W); Nieder-Österreich: Marchfeld, nahe Gänserndorf, Ackerrand neben der Strasse 1.5–1.7 km W der Kirche von Weikendorf, 28 Sep 2010, Barta s.n. (BM); Wiener Becken, Steinfeld, 7.5 km NNE Wiener Neustadt, 1.7 km NW Eggendorf, knapp WNW, 19 Sep 1998, Walter 4189 (K, W); Wien: 10 Bezirk, in Kurpark Oberlau, 22 Oct 2012, Barta 1219 (W); 22 Bezirk, knapp S der Ostbahngleitstrasse nahe der U2-Haltestelle Aspern-Nord, 6 Aug 2014, Barta 3270 (W).
Belgium. Wallonia: Vesdre, Sep 1958, Lousley s.n. (K).
France. Grand Est: Bas-Rhin, Strasbourg, Graffenstaden, Oct 1961, Patzak s.n. (W); Nouvelle Aquitaine: Gironde, Bordeaux, Bassens, 24 Jul 1927, Duffour 5538 (BM, MA); Gironde, Bassens, 9 Sep 1931, Jallu 1038 (H, P).
Germany. Hessen: Frankfurt am Main, Frankfurt-am-Main, Hessen-Nassau, Sep 1911, Peipers s.n. (BM); Niedersachsen: Leer, Ostfriesl, Blamgelände, 10 Oct 1955, Klimmek s.n. (W).
Ireland. Leinster: Kilkenny, Rosbercon Port, 5 Oct 1995, Reynolds s.n. (BM),
Netherlands. Gelderland: Nijmegen, prov. Gelderland, 23 Sep 1928, Kern & Reichgelt 14998 (BM, H).
New Zealand. North Island: Waikato, Pukekawa, 18 Dec 1986, Dawes s.n. (AK); Auckland, Arikikapakapa Golf Course, Rotorua Ecological District, Northern Volcanic Plateau Ecological Region, 15 Mar 2013, Hobbs 13297 (AK). South Island: Canterbury, Christchurch, waste land behind Fendalton Mall buildings, Fendalton, 2 Feb 1999, Healy 99/ 34 (AK, CHR).
Sweden. Prov. Halland, 26 Sep 1923, Jungner s.n. (BM); Haisenborg, vid fabrikenb Kärnan, 6 Oct 1938, Lange s.n. (BM); Skåne, Saxtorp, Flygeltofta, 28 Aug 1958, Nilsson s.n. (BM); Götaland: Halland, Halmstad, Gustavsfält, 22 Sep 1955, Blom s.n. (BM); Halland, pr. urben Halmiam, 26 Apr 1923, Jungner s.n. (BM); Halland, prope usbem Halmiam, 26 Sep 1923, Jungner s.n. (K); Halland, Halmstad, 26 Sep 1923, Jungner 1375 (K); Västra Götaland, Göteborg, Ringon, 29 Sep 1951, Blom s.n. (K).
United Kingdom. Channel Isles: Jersey, N of St. Ouen’s Bay, Sep 1996, Dupree s.n. (BM); England: Essex, vice county 18, Dagenham dumps, 1938, Airy Shaw s.n. (K); Bedfordshire, Flitton, 14 Oct 1950, Doug 1401 (K); Bedfordshire, nr Flitwick, 16 Sep 1988, Hanson 499 (BM); Hampshire, Blackmoor, 24 Oct 1959, Lousley s.n. (BM); Berkshire, Reading, 25 Nov 1999, Rutherford s.n. (BM); Gloucestershire, Wapping Wharf, Bristol Harbour, 14 Sep 1942, Sandwith s.n. (BM); Scotland: Highlands, Auldearn, Broomhill Farm, 27 Oct 1967, McCallum-Webster s.n. (BM); Midlothian, Railway Tip, Borthwick, 20 Jul 1963, McCallum-Webster s.n. (BM); Midlothian, Borthwicts, 20 Jul 1963, McCallum-Webster 8768 (K); Wales: Vale of Glamorgan, Barry Dock, 29 Sep 1923, Melville s.n. (BM).
Solanum forsteri Seem., J. Bot. 1: 207. 1863.
Type. Chile. Región V (Valparaíso): Easter Island, J.R. Forster & G. Forster s.n. (lectotype, designated here: BM [BM000900372]).
Solanum fauriei H.Lév., Repert. Spec. Nov. Regni Veg. 10: 152. 1911.
Type. United States of America. Hawaii: Oahu, Kaela, Nov 1909, U. Faurie 861 (lectotype, designated here: BM [BM000846671]; isolectotypes: P [P00315100, P03961997]).
Solanum apopsilomenum Bitter, Repert. Spec. Nov. Regni Veg. 12: 89. 1913.
Type. New Zealand. Mount [illegible, perhaps Zwart], 28 Sep 1838, E. Schwarz s.n. [Exped. Novara] (holotype: W [0022367]).
Solanum microtatanthum Bitter, Bot. Jahrb. Syst. 55: 63. 1919.
Type. Papua New Guinea. Kelil, 180 m, F.R.R. Schlechter 16407 (holotype: B, destroyed; lectotype, designated here: E [E00273859]; isolectotypes: G [G00343298], L [L0003621], GH [GH00077834], K [K001080534, K001080535]).
Solanum brachypetalum Bitter, Bot. Jahrb. Syst. 55: 64. 1919.
Type. Papua New Guinea. “Ssigaun, in Dorfern”, 600 m, Jun 1896, C. Lauterbach 2360 (holotype: B, destroyed; lectotype, designated here: WRSA).
Solanum insulae-paschalis Bitter, Nat. Hist. Juan Fernandez & Easter Island 2: 78. 1922.
Type. Chile. Región V (Valparaíso): Easter Island, Hanga ho ono (La Perouse), au linem actem Brunnen, C. Skottsberg & I. Skottsberg 663 (lectotype, designated here: GOET [GOET003528]; isolectotype: UPS [UPS104030]).
Solanum nigrum var. nihoense F.Br., Bernice P. Bishop Mus. Bull. 81: 36, pl. 16B. 1931.
Type. United States of America. Hawaii: Leeward Island, Nihoa, E.L. Caum 62 (holotype: BISH; isotypes: K [K000922201], NY [00172289]).
Solanum nigrum var. pitcairnense F.Br., Bernice P. Bishop Mus. Bull. 130: 255. 1935.
Type. Pitcairn Islands: Pitcairn island, 13 Mar 1922, E.H. Quayle 375a (holotype: BISH [BISH1005084]; isotype: BISH [BISH1005083]).
Solanum allanii Polgár, Trans.& Proc. Roy. Soc. N. Z. 69: 278. 1939.
Type. Cultivated in Györ Hungary from seeds of Miss L.B. Moore #8 (New Zealand, Auckland, Mt. Wellington), Jan 1938, S. Polgár s.n. (lectotype, designated here: BP [acc. # 146410]; isolectotype: BP [acc. # 261051]).
Cultivated in Germany at the Berlin Botanical Garden from seeds from Australia “New Holland (Listeman) hort. bot. Berlin”, Anon. s.n. (lectotype, designated here: HBG [HBG511471]).
Annual or short-lived sprawling to erect perennial herbs to 1 m tall, subwoody and branching at base. Stems spreading to decumbent, terete or sometimes slightly ridged, green to yellow-green, older stems greenish-grey, not or occasionally somewhat hollow; new growth densely to sparsely pubescent with simple, antrorse, uniseriate, translucent, eglandular or sometimes glandular trichomes, these 4–6-celled, 0.5–1 mm long; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.5–8.0(-17) cm long, 1–4 -(9) cm wide, elliptic to slightly ovate, very variable in size, membranous, green, concolorous, without smell; adaxial surface glabrous or sparsely and evenly pubescent with simple uniseriate ca. 4-celled trichomes to 0.5 mm long; abaxial surfaces glabrous or sparsely pubescent with simple uniseriate trichomes along the veins; major veins 3–8 pairs, not prominent; base cuneate, decurrent on the petiole; margins entire or shallowly toothed, if present the teeth acute; apex acute to acuminate; petioles 0.5–3 cm long, sparsely pubescent with antrorse simple uniseriate trichomes like those of the stems. Inflorescences 1–2 cm long, internodal, unbranched but very rarely furcate, umbelliform to sub-umbelliform, with 3–7 flowers clustered near the tip of the rhachis, sparsely pubescent with antrorse simple uniseriate 3–4-celled trichomes like those of the stems; peduncle 1–3 cm long, straight and stout; pedicels 0.4–0.8 cm long, < 0.3 mm in diameter at the base, ca. 0.3 mm in diameter at the apex, filiform, nodding, pubescent like the peduncle, articulated at the base; pedicel scars clustered near the tip of the rhachis, often the lowest flower ca. 0.5 mm spaced from the rest. Buds ellipsoid, the corolla strongly exserted from the calyx tube long before anthesis. Flowers 5-merous, all perfect. Calyx tube 1–2 mm long, conical, the lobes often unequal, the lateral two largest 1–1.2 mm long, 0.5–0.6 mm wide, the top and lowermost 0.4–1.0 mm long, 0.3–0.5 mm wide, long triangular often with a rounded tip, glabrous or sparsely pubescent with antrorse simple uniseriate trichomes ca. 0.5 mm long. Corolla 6–10 mm in diameter, white or white with a purplish tinge, stellate, lobed ca. 1/2 way to the base, the lobes 3.0–4.2 mm long, 1.0–1.2 mm wide, spreading to reflexed, densely papillate on tips and margins. Stamens equal; filament tube < 0.1 mm long; free portion of the filaments 0.5–1.0 (-1.5) mm long, glabrous or adaxially pubescent with tangled simple uniseriate trichomes; anthers 1.2–1.6 mm long, 0.7–1.0 mm wide, ellipsoid, yellow, somewhat sagittate at the base, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary rounded, glabrous; style 3–4 mm long, strongly curved in the distal 1/4, densely pubescent with simple uniseriate trichomes 0.2–0.5 mm long, these tangled in the basal 1/2 to 2/3 of the length, not exserted beyond anthers and only the stigma visible outside the anther cone; stigma capitate, the surfaces minutely papillate. Fruit a globose berry, 4–10 mm in diameter, green or bluish-black at maturity, the pericarp thin, matte; fruiting pedicels 0.7–1.5 cm long, ca. 0.5 mm in diameter at the base and the apex, erect or spreading, becoming yellow, falling with the mature fruits, not persistent; fruiting calyx lobes not accrescent, the tube less than 1 mm long, the lobes 1.2–1.7 mm long, appressed to the basal quarter of the berry, occasionally somewhat spreading, never strongly reflexed. Seeds 50–100 per berry, 1.2–2.2 mm long, 0.7–1.8 mm wide, flattened reniform, pale yellowish-tan, the surfaces minutely pitted, thin and the embryo clearly visible, the testal cells rectangular to pentagonal in outline. Stone cells (0-)2(-4) per berry, >0.5 mm in diameter, brown. Chromosome number: 2n=6x=72 (
Grows in disturbed areas, along roadsides and field edges; between sea level and 2,600 m elevation in a wide variety of habitats.
Australia: currant, green-berry nightshade (
Leaves eaten as greens throughout the species range.
(
Solanum opacum is extremely similar to S. americanum; both species have minute flowers with anthers usually less than 1.5 mm long. They co-occur across the Pacific, but can be distinguished easily, particularly in fruit. Solanum opacum has matte berries that are green or greyish-purple at maturity, with appressed or slightly spreading calyx lobes, while S. americanum has very shiny berries that are black or dark purple at maturity and calyx lobes that are very strongly reflexed. The pedicels of S. americanum remain on the plant as fruits fall, while those of S. opacum drop with the mature berries. In flower, the relative length of filament to anther is a useful distinguishing character; S. opacum has filaments that are as long as or slightly longer than the anthers, while the filaments of S. americanum are always shorter than the anthers.
Our circumscription of S. opacum encompasses a much wider distribution than that of
Solanum opacum is hexaploid and, based on network analysis of arbitrary amplified DNA markers,
Solanum forsteri was named in honour of the Forster’s (father Johann and son Georg, who accompanied Captain Cook on the HMS Resolution from 1772 to 1775) and we have selected the sheet collected by them on Easter Island (BM000900372) as the lectotype. This specimen is mounted together with a collection of S. americanum that is on different paper (white rather than the blue associated with Forster’s material), showing the difficulty that early botanists (and this holds true still today) had in distinguishing these two species with minute anthers that co-occur across the Pacific.
In describing S. microtatanthum,
Australia. New South Wales: Botany Bay, 28 Apr 1770, Banks & Solander s.n. (BM); Sans Souci, nr Georges River, 27 Sep 1961, Goode 346 (K, NSW); Kyogle, Toonumbar State Forest 26 km NW of Kyogle, 22 Feb 1972, Henderson 1261 (AD, AD, NSW); Upper Hunter Shire, Ben Halls Gap State Forest, 20 Feb 1991, Hosking 307 (NE); Wollongong, Bulli, 1875, Johnson s.n. (MEL); Wollongong, Mt Keira, 27 Jul 1983, Mills s.n. (WOLL); Norfolk Island: sin. loc, Oct 1805, Caley s.n. (BM); Kingston, 24 Aug 1964, Uhe 1126 (K); Northern Territory: Adelaide River, Northern Territory, grown in glasshouse at Indooroopilly, Brisbane, 18 Dec 1972, Henderson 1356 (CANB, K); Queensland: Nr border of Queensland/New South Wales, Main Range National Park, 10 Feb 2006, Bohs et al. 3561 (BM, UT); Birnam Range 7.5 km NE of Beaudesert, Tremayne rd, Jan 2001, Halford Q-3882 (BRI, K, MEL, NSW); c. 13 km WNW of Goondiwindi on rd to St. George, 30 Sep 1975, Henderson 2357 (AD, BRI, K); South Australia: Upper River Murray nr 375 mile peg 4 mi downstream from Little Hunchee Island, on North bank, 16 Sep 1979, Symon 11586 (AD, CANB, K); Kangaroo Island, Kangaroo Island, Waterhouse s.n. (MEL); Tasmania: Tasman, Port Arthur, 1893, Bufton 11 (MEL); sin. loc, 1842, Gunn 51[a] (BM); Latrobe, Harford on Solomans Hill, 6 Nov 1932, Hamilton 163 (CANB, HO); Van Diemen’s Land, Gunn, 1835, Without collector s.n. (K); King Island, Nr Sea Elephant River, 23 Mar 2009, Wapstra et al. 691 (AD); Victoria: Mt Buffalo access rd 12 km WNW of Bright, 1 Jan 2003, Lepschi 4916 (CANB, MEL); Latrobe, 18 Jul 1988, Thompson 157 (AD, AD, MEL); Cedar Creek, Bentleigh district, 12 Feb 1920, White s.n. (NSW); Western Australia: c. 200 mi N of Chesapeake rd 1.5 km E of Mt Chuladup, 7 Dec 2001, Barker 8355 (AD); Eyre, S of Cocklebeddy, 5 Dec 1962, MacDonald 29 (K); Bayswater, Lower Swan River, 20 Mar 1909, Morrison 19014 (K).
Chile. Región V (Valparaíso): Easter Island, von Chamisso s.n. (F, NY); Juan Fernández Islands, Masafuera, Quebrada de Veradero, 12 Mar 1917, Skottsberg & Skottsberg 568 (GOET).
Cook Islands. Mitiaro: just before entering swamp on rd to Atai Foodland, 22 Jul 1991, Luttrell 187 (FHO); rd from the village of Takaue to the marsh, 24 Apr 1985, Whistler 5576 (US).
Fiji. Yuen Yick’s farm, Nasinu, Naitasiri, 3 Dec 1957, Ledua s.n. (K); sin. loc., 1860, Seemann 344 (BM, K); Fulanga: Limestone Formation, 22 Feb 1934, Smith 1174 (K); Kandavu: Mount Mbuke Levu, 23 Oct 1933, Smith 206 (K); Viti Levu: Mba, summit of Mt. Nanggaranambuluta E of Nandarivatu, 19 Jun 1947, Smith 4849 (K, US); Hills E of Wainikoroiluva River, nr Namuamua, Namosi, 15 Oct 1953, Smith 9070 (K, US).
French Polynesia. Austral Islands: Raivavae, Vaiuru, 10 Aug 1934, Fosberg 11756 (US); Rurutu, du N de l’ile, 17 Apr 1981, Hallé 7032 (US); Rurutu, N Moerai, 25 Apr 1981, Hallé 7321 (US); Raivavae, Ahuoivi, 9 Aug 1934, St. John 16070 (US); Gambier Islands: Mangareva, 1833, Le Guillou s.n. (US); Aukena, Koiovao, 29 May 1934, St. John 14659 (US); Society Islands: Tahiti, 6 Apr 1840, Barclay 3309 (BM)Tahiti, plateau de Taravao, sentier du captage de l’Hamoa, 16 Sep 1982, Florence 3853 (US); Raiatea, first valley S of Uturoa, 14 Oct 1926, Moore 212 (US); Me’et’ia, Fatia-po to Fareura, 12 May 1934, St. John 14187 (US).
Indonesia. Bali: Kintamani, 14 Nov 1966, Schwabe s.n. (B); East Nusa Tenggara: Flores, nr Mataloko, Verheijen 24 (L); Flores, nr Mataloko, Verheijen 191 (L); Papua: Mimika Regency, PT-Freeport Indonesia Concession Area, 9 Aug 1998, Johns et al. 9565 (A, K); Mimika Regency, W and above Tembagapura, 23 Aug 1998, Sands 7296 (K); Sulawesi: Central Sulawesi, Sopu valley, 30 May 1975, Balgooy 3555 (A, K); West Papua: West-Irian, Eipomek-Tal, bei Malingdam, 24 Feb 1976, Hiepko & Schutze-Motel 1195 (B).
Japan. Nagasaki Pref., Katamatsuura-gun, Emukae-cho, Okugawachi-men, 17 Aug 1994, Yonekura 3230 (MO).
Malaysia. Sabah: Kampung Tiung, side of hill Tiung Valley, Tuaran Distr., 8 Aug 1985, Maikin Lantoh SAN-108976 (K)
Marquesas Islands. Hatutaa Island: vallon de la pointe SW, 10 Jul 1988, Florence & Teikiteetini 9399 (US)
Marshall Islands. Arno Atoll: Ine village, 12 May 1950, Anderson 3672 (L, US); Jaluit Atoll: Jabor, 28 Apr 1958, Fosberg 39475 (US); Kwajalein Atoll: Kwajalein Islet, 19 Jan 1950, Fosberg 31180 (US)
New Caledonia. sin. loc, Caldwell s.n. (K); NO de la Nouvelle Caledonie, 1867, Krieger s.n. (W); Nord: Ilôt Poudiou, observatory Island, Isle of Pines, Oct 1853, MacGillivray 805 (K); Sud: Nouméa, Îlot Freycinet, Ilot de Freycinet, Aug 1884, Grunow s.n. (W); Noumea, Baie Tina, 24 Jul 1973, MacKee 26964 (K).
New Zealand. Chatham Islands: Rekohu, (Chatham Island), Waitangi, Ellice Point, Chatham Islands Ecological Region and District, 1 Dec 2008, de Lange &Horne CH-2032 (AK). Kermadec Islands: Macauley Island, Southern Kermadec Islands Group, 29 Jun 2006, Barkla M- 26 (AK); Curtis Island, Nov 1900, Shakespear s.n. (AK). North Island: Northland, dunes near Te Arai Forest Sanctuary, Aupouri State Forest, N of Waimahru Stream, Mangonui County, 25 Sep 1985, Bellingham 0144 (AK); Taranaki, Sugarloaf Island, off Mangonui, Moturoa Group, off Karikari Peninsula, 14 May 1976, Wright 1252 (AK). Outlying Islands: Three Kings Islands, Great Island, above cliff S.E. Bay, 19 Apr 1946, Turbott & Bell s.n. (AK). South Island: Motuara, Banks & Solander s.n. (BM). Tokelau Territory: Fakaofu, Union Islands, Mar 1891, Lister s.n. (K).
Niue. Niue Island, “71 Polokai”, 1901, Smith s.n. (AK).
Norfolk Island. Rocky Point Reserve, 6 Oct 1989, Gardner 5866 (AK).
Papua New Guinea. Boridi, 6 Sep 1935, Carr 12985 (BM, K); Wiligimaan, Baliem, Div. Hollandia, 28 Jun 1961, Versteegh BW-12516 (A, L); Bougainville Island: Namatea, NW Bougainville, 7 Mar 1932, Waterhouse 691 B (K, L); Crown Prince Mountain, Oct 1960, Womersley NGF-13346 (A); Central: E side lake Myola No. 2, Distr. Central, Subdistr. Port Moresby, 13 Sep 1973, Croft NGF-34532 (E, K, L); Kokoda trail, eastern side lake Myola, 22 Jul 1974, Croft LAE-61910 (A, K, QRS); Madang: Saidor, Moro, Naho-Rawa Division, Finisterre mountains, 19 Nov 1964, Perumal 21473 (BM); Morobe: Mount Kaindi, Wau, 4 Oct 1977, Conn & Kairo 491 (CANB, K); Bulldog rd, nr Edie Creek, Wau, 13 Aug 1963, Millar & Holttum NGF-15825[b] (A, US); Southern Highlands: Hagen-Mendi rd, Mendi Subdistrict, Southern Highlands District, 21 Sep 1968, Vandenberg et al. NGF-40062 (K); Western Highlands: nr Ampyak Highlands, Ecological Site 1, 15 Dec 1964, Flenley ANU-2192 (K); nr Tomba village, S slope of Mount Hagen Range, 27 Aug 1956, Hoogland & Pullen 6014 (BM, US); Western Province: Western, 19 Jun 1979, Sohmer LAE-75542 (K); Kubor Range, Uinba, Nona-Minj Divide, 20 Aug 1963, Vink 16311 (K).
Philippines. Luzon: Cordillera (CAR), Mount Pulag, 26 Jan 1968, Jacobs 7173 (K).
Pitcairn Islands. Oeno Island: Close to hut, 23 Jun 1934, St. John & Fosberg 15195 (US); Pitcairn Island: Bounty Bay, baie de la Bounty, Cap Est, 22 Apr 1991, Florence 10774 (US); Bounty Bay, 15 Jun 1934, Fosberg & Clark 11337 (US).
Samoa. Apia: Apia, 6 May 1907, Vaupel Sol-3 (K); Savai’i: E of Olo, 8 Aug 1931, Christerpherson & Hume 2311 (K).
Solomon Islands. Guadalcanal: north central Guadalcanal, Tina River, 14 Sep 1967, Maurisi BSIP8109 (K).
Taiwan. Pingtung County: Wutain Hsiang, Tawu village, 13 Mar 1999, Shu-hui Wu 1161 (MO); Taoyuan County: Kuanyin Hsiang, Hsinpo, 28 Dec 1999, Ching-I Peng, 17879 (MO).
Tonga. Tofua: sin. loc, Jan 1967, Scarth-Johnson s.n. (K); Tongatapu: Kologa, Jun 1926, Setchell & Parks 15374 (US).
United States of America. Hawaii: Leeward Island, Nihoa, 17 Jun 1923, Caum 62 (US); Oahu, Kaela, Nov 1909, Faurie 861 (BM, P); Kauai, Kaholuamanoa, above Waimea, 1 Oct 1895, Heller 2867 (AK, BM, E, US); Pearl and Hermes Atoll, Southeast Island, midway between pond and tower, 18 Aug 1964, Young 112 (US).
Argentina. Tucumán: Dept. Tafí del Valle, Yerba Buena, 19 Jan 1919, S. Venturi 159 (holotype: US [00027724, acc. # 1548805]; isotypes: BA [BA2463], LIL [LIL001454], LP [LP010926], MA, SI [SI003329]).
Annual, decumbent or prostrate herbs, the young plants sometimes erect, up to 15 cm tall often rooting at the lower nodes, forming dense patches, the branches to ca. 1 m long. Stems decumbent or ascending, terete or somewhat angled with ridges, green, older stems yellowish-brown, not markedly hollow; new growth pubescent with simple, spreading, uniseriate, translucent, eglandular trichomes, these 0.5–1 mm long, to or nearly glabrous; older stems glabrous. Sympodial units difoliate, the leaves not geminate. Leaves simple, 2.5–9 cm long, 2.5–7.5 cm wide, broadly ovate, thinly membranous, green, concolorous, without smell; adaxial surfaces glabrous to sparsely pubescent with simple hairs to 0.5 mm on the major veins; abaxial surfaces glabrous; major veins 3–4 pairs; base long attenuate, decurrent on the petiole; margins 3-lobed nearly to the midrib, rarely the lateral lobes themselves lobed, the terminal lobe ovate, the lateral lobes asymmetrically ovate or lanceolate-ovate, acute at the tips, the sinuses sometimes sparsely ciliate; apex acute; petioles 0.5–2 cm, winged to the base, glabrous or sometimes sparsely ciliate near the base. Inflorescences 1.2–2.5 cm, internodal or often just below a node, unbranched or rarely with a few branches, the flowers spaced along the rhachis, with 4–9 flowers, glabrous to sparsely pubescent; peduncle 0.7–1.4 cm long, delicate; pedicels 3–5 mm long, 0.2–0.3 mm in diameter at the base and at the apex, filiform, spreading, articulated at the base; pedicel scars spaced 1–5 mm apart. Buds ellipsoid, the corolla completely covered by the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 1.5–2 mm long, cup-shaped, the lobes ca. 0.75–1.5 mm long, less than 1 mm wide, lanceolate-oblong, the tips acute, glabrous. Corolla ca. 7 mm in diameter, white or rarely light violet, rotate-stellate, lobed ca. 1/2 way to the base, the lobes 1.5–2.5 mm long, 1–2 mm wide at the base, reflexed or spreading at anthesis, abaxially minutely white-puberulent on the tips of the lobes, glabrous adaxially. Stamens equal; filament tube minute; free portion of the filaments 0.5–1 mm long, adaxially pubescent with tangled uniseriate trichomes; anthers 1.6–2 mm long, 0.7–0.8 mm wide, oblong or ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary glabrous; style 2.3–3.3 mm long, glabrous or sparsely pubescent in the lower part where included in the anther cone, exserted 0.5–0.8 mm beyond anthers; stigma capitate, the surface minutely papillate, green in live plants. Fruit a depressed-globose and bilobed (especially when young) berry, 6–8 mm in diameter, pale yellow, the pericarp thin and somewhat shiny; fruiting pedicels 4–7 mm long, 0.5–0.7 mm in diameter at the base and at the apex, spreading, recurved at the base to hold the fruit downwards, nearly in contact with the soil, dropping with the mature fruit, not persistent; fruiting calyx not markedly accrescent but the lobes somewhat elongating in fruit, the tube 2–3 mm long, the lobes 2–3(-4) mm long, covering the basal 1/3 of the berry, the tips somewhat recurved. Seeds 20???30 per berry, 1.5–1.6 mm long, 1.2–1.6 mm wide, flattened reniform, light yellow, the surfaces pitted, the testal cells sinuate in outline. Stone cells 2(4) per berry, 2 larger and apical (1–1.5 mm in diameter), the other 2 equatorial, smaller, 0.5–0.6 mm in diameter. Chromosome number: 2n=2x=24 (
Grows in disturbed sites, along roadsides and field margins, on rocky, sandy or clay soils; between (50-) 1,400 and 3,000 (-3,700) m in its native range, near sea level in Australia.
None recorded for the Old World.
None recorded for the Old World.
(
Solanum palitans is distinct amongst the species of morelloids occurring in the Old World in its combination of uniformly 3-lobed, thin membranous leaves and pale yellow mature fruits. The species has a creeping habit, with stems growing close to the ground extending up to 3 m and often rooting at nodes. Outside of its native range in South America, it is only found in Australia, but may appear in other areas of suitable habitat given time. Details of its variability and habitats in Argentina can be found in
Like other adventive members of this group, it is likely to have been introduced to Australia with wool waste from Argentina. The first collection cited in
Australia. New South Wales: Barallier via Moss Vale, May 1950, Carlon s.n. (NSW); Burragorang Lookout, 14 mi S of Yerranderie, 17 mi WNW of Bowral, 23 Mar 1966, Constable 6738 (CANB, K); Wingecarribee, 39 km WNW of Mittagong, 16 Mar 1975, Coveny et al. 6089 (AD, NSW); Tamworth Regional, Moorsville, The Forest, S of Moore Creek, 4 Sep 1996, Hosking 1282 (CANB, MEL, NS, NSW); Campbelltown, Showground, 26 Apr 1964, McBarron, 9021 (AD, NSW); Armidale Dumaresq, Hillgrove mine, S of Hillgrove, 15 Oct 2014, Without collector 16 (NE).
Solanum pachyarthrotrichum Bitter, Repert. Spec. Nov. Regni Veg. 10: 542. 1912.
Type. Cameroon. Sin loc., H. Deistel 631 (holotype: B, destroyed; no duplicates found).
Solanum hypopsilum Bitter, Repert. Spec. Nov. Regni Veg. 10: 543. 1912.
Type. Cameroon. Sud-Ouest: Buea, 1000 m, 8 Mar 1898, H. Lehmbach 175 (holotype: B, destroyed; no duplicates found).
Solanum molliusculum Bitter, Repert. Spec. Nov. Regni Veg. 10: 546. 1912.
Type. Cameroon. Sud-Ouest: Buea, canyon east of Mannsquelle, P. Preuss 740a (holotype: B, destroyed; no duplicates found).
Cameroon. Sud-Ouest: Mt Cameroon, Dec 1862, G. Mann 1938 (holotype: K [K000028649]; isotype: GH [00395064]).
Annual or short-lived prostrate or sprawling perennial herbs to 0.5–1.5 m tall, subwoody and branching at base. Stems spreading, usually strongly ridged with pseudospinose dentate processes, green, older stems brown or grey, usually hollow; new growth densely pubescent with simple, spreading, uniseriate, translucent, mixed glandular and eglandular trichomes, these 7–10-celled, 1.0–1.5 mm long; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 2.5–7 cm long, 1.3–4 cm wide, smaller at distal ends of branches, ovate, widest in the lower third, membranous, green, concolorous, smell not known; both surfaces evenly pubescent with simple uniseriate mostly eglandular trichomes like those on stem; major veins 4–5 pairs; base cuneate; margins entire or shallowly toothed in the basal half; apex acute to acuminate; petioles 0.5–2 cm long, pubescent with trichomes like the leaves. Inflorescences 1–2.5 cm long, opposite the leaves, unbranched, sub-umbelliform to racemose, with 2–5(-6) flowers clustered at the tip, pubescent with uniseriate trichomes like those of the stems; peduncle 0.3–1.5 cm long, straight; pedicels 0.7–0.8(-1) cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, slender, spreading, pubescent with simple uniseriate trichomes like the rest of the inflorescence, articulated at the base; pedicel scars clustered in the apical portion of the inflorescence rhachis, lowermost scars ca. 2 mm apart, the rest overlapping at the distal end. Buds ellipsoid, densely pubescent, the corolla at least 1/2 way exserted from the calyx tube until just before anthesis. Flowers 5-merous, all perfect. Calyx tube 1.5–2 mm long, conical, the lobes 1–1.5 mm long, 0.9–1.4 mm wide, elongate-deltate, rounded at the tips, densely pubescent with simple uniseriate trichomes like the rest of the inflorescence. Corolla 8–14 mm in diameter, white to pale violet (“blue”) or white with purple venation, stellate, lobed nearly to the base, the lobes 4–6 mm long, 2–2.5 mm wide, spreading at anthesis, abaxially moderately pubescent with simple uniseriate 2–3-celled trichomes, these denser near the tips and margins, also papillate along the margins and tips, adaxially glabrous. Stamens equal; filament tube minute; free portion of the filaments 0.5–1 mm long, densely pubescent adaxially with tangled simple trichomes; anthers 2–2.5 mm long, ca. 1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary conical, glabrous; style 3–5 mm long, densely pubescent with tangled simple trichomes in the basal portion inside the anther tube, exserted ca. 2 mm beyond the anther cone, often elongating early and protruding from unopened buds; stigma capitate, the surfaces minutely papillose, colour in live plants not known. Fruit a globose berry, 4–6 mm in diameter, purple or black at maturity, the pericarp thin and matte; fruiting pedicels 0.9–1.2 cm long, ca. 0.75 mm in diameter at the base and at the apex, spreading, not falling with the fruit, remaining on the plant and persistent on older inflorescences; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes 1.5–2.5 mm long, appressed to the berry, spreading and hyaline, the tips slightly reflexed. Seeds ca. 20 per berry, ca. 1.5 mm long, ca. 1 mm wide, not markedly flattened, tear-drop shaped with a subapical hilum, pale beige, the surface minutely pitted and the lateral testal cell walls elongate and the seed appearing hairy, the testal cells pentagonal in outline. Stone cells 4–6(-10), often found near the pedicel junction at the base of the berry. Chromosome number: 2n=4x=48 (
Solanum pseudospinosum A Flowering habit B Inflorescence C Bud D Flower at anthesis E Fruiting habit F Detail of adaxial leaf surface G Eglandular trichome H Glandular trichome I Infructescence J Seed (A, C, DCasas 10313; BCasas 10157; E, JBreteler et al. 39). Scale bar: 4 cm (A, E), 1 cm (B, I), 5 mm (C), 7 mm (D), 4 mm (F), 0.8 mm (G–H) and 2.5 mm (J). Drawing by L. Smith.
Grows at forest margins, in clearings and along roadsides; between 2,100 and 4,000 m elevation.
Cameroon: afom.
None recorded.
(
Solanum pseudospinosum is a high elevation species that, in our circumscription, only occurs along the so-called Cameroon line that is along the junction of the Congo and West African cratons (
The Cameroon line along which Solanum pseudospinosum occurs runs from the island of Bioko to the Tchabal Mbabo plateau and is of volcanic origin with activity beginning the Cretaceous period, but peaking in the Pliocene and continuing to the present (
Solanum pseudospinosum is a tetraploid species with unknown parentage (
All of the species described by
Specimens examined. Cameroon. N slope of Cameroun Mountain, 7 Apr 1985, Thomas 4642 (MO); Nord-Ouest: Piste Acha-Abaw au Lac Oku, 40 km NE Bamenda, 5 Dec 1974, Letouzey 13444 (K); Sud-Ouest: Mt. Cameroon, above Buea, nr Hut 2, 1 Apr 1952, Boughey GC6933 (K); Mt. Cameroon, Johann-Albrechtshöhe, 30 Jan 1962, Breteler et al. MC39 (K); Mt. Cameroon, 9 Oct 1992, Cheek & Sidwell 3664 (K, SCA, WAG, YA); Mt. Cameroon, 4 Nov 1993, Cheek et al. 5352 (K); Mt. Cameroon, 17 Feb 1927, Dalziel 8335 (E, K); Mt. Cameroon, nr Hut 2, Jan 1967, Guile, 1560 (MO); Bamenda, Southern Cameroons, Bamenda Division, Bafut-Ngemba forest reserve, 23 Feb 1958, Hepper 2146 (K); Mt. Cameroon, beside Hut 2, 6 Apr 1937, Hutchinson & Metcalfe 54 (K); Mt. Cameroon, To no 3 Hut, Jan 1931, Maitland 1301 (K); Mt. Cameroon, Feb 1931, Maitland 1333 (K); Mt. Cameroon, Jan 1862, Mann 1321 (K); Mt. Cameroon, NW de Buea, 31 Mar 1981, Meijer 15435 (MO); Mt. Cameroon, Kamerun-Berg, oberhalb Buea, 22 Dec 1928, Mildbraed 10888 a (K); Mt. Cameroon, Hut 2, 21 Dec 1958, Morton 758 (K); Mt. Cameroon 2 mi W of Mann’s Spring, 29 Dec 1958, Morton K874 (K); Mt. Cameroon, around Mann’s Spring, 3 Dec 1996, Nning et al. 52 (K, MO, SCA, YA); Mt. Cameroon, Bokwango, 5 Oct 1992, Thomas 9348 (K).
Equatorial Guinea. Bioko: cumbre del pico Basilé, 3 Jul 1986, Fernández-Casas 10156B (K, MO); Carretera del pico Basilé, 10 Jul 1986, Fernández-Casas 10313 (BM, K); Bioko Norte: Pico Basilé, Distr. Barney, cruce de la Virgen de Bisila, 12 Dec 2007, Cabezas et al. 907 (MA); Cumbre del pico Basilé, 8 Oct 1986, do Carvalho 2544 (BM, H, K, MA, MO); Cumbre del pico Basilé, 3 Jul 1986, Fernández-Casas 10157 (BM, K, MO); Pico Basilé, cumbre del pico, 5 Feb 1989, Fernández-Casas 11184 (K, MA).
Solanum pygmaeum var. hastatum Bonte ex Aellen, Ber. Schweiz. Bot. Ges. 50: 236. 1940.
Type. Argentina. Buenos Aires: Pergamino, J.A. de la Peña, 14 Jan 1925, L.R. Parodi 6107 (lectotype, designated here: BAA [BAA00004675]).
Solanum pygmaeum var. integrifolium Bonte ex Aellen, Ber. Schweiz. Bot. Ges. 50: 236. 1940.
Type. Switzerland. Basel-Stadt: Basel, Hof der Aktienmühle, Feb 1939, P. Aellen 3201 (lectotype, designated here: P [P03933622]; isolectotype: P [P03933624]).
Solanum pygmaeum var. sinuatodentatum Bonte ex Aellen, Ber. Schweiz. Bot. Ges. 50: 237. 1940, as “sinuato-dentatum”.
Type. Switzerland. Basel-Stadt: Basel, Hof der Aktienmühle, Feb 1939, P. Aellen 3202 (lectotype, designated here: P [P04477345]; isolectotypes: BM [BM001035970], P [P03933623]).
Solanum pygmaeum var. latifolium Bonte ex Aellen, Ber. Schweiz. Bot. Ges. 50: 237. 1940.
Type. Hungary. Györ: Oelfabrik, 1915 and 1918, S. Polgár s.n. (syntypes; no duplicates located, at BP?); Germany. Neuss: Oelfabrik, 1915, L. Bonte s.n. (syntype; no duplicates located); Germany. Emmerich: mit Oelfrucht, 1929, R. Scheuermann s.n. (syntype; no duplicates located).
Solanum pygmaeum var. suspensum C.V.Morton, Revis. Argentine Sp. Solanum 138. 1976.
Type. Argentina. Córdoba: Alta Córdoba, barrio de la ciudad de Córdoba, T. Stuckert 4713 (holotype: G [n.v.]; isotypes: CORD [CORD00004273, CORD00004274]).
Solanum deterrimum C.V.Morton, Revis. Argentine Sp. Solanum 138. 1976.
Type. Argentina. Buenos Aires: Sierra de la Ventana, 23 Feb 1944, H. Ruíz de Huidrobo 1332 (holotype: A [00077613]; isotypes: NY [00139129], S [acc. # 12-27773], SI [SI003308, SI003307]).
Argentina. Buenos Aires: “in Pampas de Buenos Ayres esquina de Ballesteros”, Sep, L. Née, s.n. (lectotype, designated by
Perennial small upright herbs up to 30 cm tall, subwoody at base, perennating via underground rhizomes. Stems decumbent or ascending, delicate, terete or somewhat angled with ridges, not markedly hollow; new growth pubescent with simple, appressed, uniseriate, translucent, eglandular trichomes, these 1–6-celled, 0.2–0.5 mm long or nearly glabrous; older stems glabrous or glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.0–5.0 cm long, 0.5–3 cm wide, ovate to narrowly elliptic, pale green, concolorous, without smell; adaxial surface glabrous or sparsely pubescent along leaf lamina and margins with simple, uniseriate trichomes like those on stem; abaxial surface sparsely pubescent with similar trichomes but the pubescence denser along the midrib; major veins 3–4 pairs; base attenuate, decurrent on the petiole; margins sinuate to entire; apex acute to obtuse; petioles 0.5–1.7 cm long, with scattered simple, appressed, uniseriate eglandular trichomes like those on stem. Inflorescences 1–3 cm long, generally internodal, simple or rarely furcate, umbelliform to sub-umbelliform, with (2-)4–6 flowers clustered at the tip, glabrous or with scattered simple, appressed, uniseriate eglandular trichomes like those on stem; peduncle (1.3-)1.5–2.6 cm long, delicate; pedicels 6–13 mm long, 0.5–1 mm in diameter at the base, ca. 1 mm in diameter at the apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0–2.5 mm apart. Buds globose to broadly ovoid, the corolla strongly exserted from the calyx tube but only halfway exserted beyond the elongate and reflexed calyx lobes before anthesis. Flowers 5-merous, all perfect. Calyx tube (0.5-)1.7–2.0(-2.2) mm long, conical, the lobes 1.5–1.8 mm long, 0.7–0.9 mm wide, narrowly elliptic with long-acuminate to acute apices, glabrous to sparsely pubescent with simple uniseriate eglandular trichomes like those on stem. Corolla 9–16 mm in diameter, white to pale lilac with a yellow-green central portion near the base, stellate, lobed 1/2 way to the base, the lobes 5.0–6.7 mm long, ca. 3.0–3.5 mm wide, strongly reflexed at anthesis, later spreading, glabrous to sparsely pubescent abaxially with simple uniseriate trichomes like those of the stem but shorter. Stamens equal; filament tube minute; free portion of the filaments 1.0–1.2 mm long, adaxially pubescent with tangled uniseriate 4–9-celled eglandular trichomes to 0.5 mm long; anthers (3.0-)3.5–3.8 mm long, 0.7–1.0 mm wide, oblong-ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style ca. 6.3 mm long, densely pubescent with (1-)2–3-celled simple uniseriate trichomes in the lower 4/5, exserted 1–2 mm beyond the anther cone; stigma capitate to clavate, bilobed, minutely papillate, green in live plants. Fruit a subglobose berry, 8–10 mm in diameter, greyish-green at maturity, the pericarp opaque and glaucous; fruiting pedicels 12–15 mm long, ca. 1 mm in diameter at the base and at the apex, deflexed and often somewhat curved, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes 1.5–2 mm long, appressed against the berry. Seeds >50 per berry, 1.8–2.0 mm long, 1.2–1.4 mm wide, flattened and tear-drop shaped with a subapical hilum, pale yellow, the surfaces minutely pitted, the testal cells irregularly quadrate in outline. Stone cells 6–8, the 2 apical ones 1.5–2 mm in diameter, usually very closely paired, the rest equatorial and 1–1.2 mm in diameter, pale whitish-brown. Chromosome number: 2n=2x=24 (one individual with n=18 and chromosomal anomalies with supernumerary bivalents or univalents not segregating;
Grows in sandy and clay soils, along rail road tracks and road sides; between 100 and 1,000 m elevation in its native range, but brought only sporadically to Europe and North America via wool shipments and trade to ports at sea level and has not naturalised widely.
None recorded in Old World.
None recorded.
(
Solanum pygmaeum is only occasionally collected in Europe, usually associated with wool waste. It is a plant that spreads by underground stems, so it is surprising that it has not become established; we include the species here so that old herbarium specimens can be identified and so that the species can be spotted if it does become established as climate changes.
The species is easy to distinguish from all other morelloids occurring in Europe by its large flowers (anthers > 3mm long), narrowly elliptic calyx lobes (1.5–1.8 mm long) and rhizomatous habit. Leaves are quite variable in size, but are usually narrowly elliptic, less often wider in the lower half.
We have selected the P duplicates that were part of the “Société cénomane d’exsicccata” as the lectotypes of var. integrifolium and var. sinuatodentatum because they are well-preserved and have both flowers and young fruits. The lectotype we have selected for var. hastatum is in an Argentinian herbarium (see best practice as outlined in
France. Provence-Alpes-Côte d’Azur: Hyères, Fleming 306 (BM).
United Kingdom. England: Cornwall, Phillack Towans, Hayle, 12 Sep 1929, Foggitt s.n. (BM); Greater London, Winchmore Hill, 9 Sep 1920, Herb. Hall s.n. (BM); Greater London, Edgware, 22 Aug 1950, Hill s.n. (BM); Greater London, Southall, 14 Aug 1945, Kent s.n. (BM); Greater London, Hanwell, 3 Sep 1947, Kent & Sandwith 3423 (K); Middlesex, 18 Aug 1945, Lousley 4710 (BM); Hayle Towans, West Cornwall, 21 Sep 1929, Melville & Smith 2922 (BM, K); Norfolk, Blickling, 3 Sep 1921, Robinson s.n. (BM).
Solanum retroflexum var. angustifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852.
Type. South Africa. Eastern Cape: Graaff Reinet (“Graafreynet”), 3000-4000 ft, 1838, J.F. Drège 7864b (holotype: G-DC [G00144331]; isotypes: K [K000414172], S [acc. # S-G-5707]).
Solanum retroflexum var. latifolium Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852.
Type. South Africa. Western Cape: Paarlberg, 1000-2000 ft, 1838, J.F. Drège 7864a (lectotype, designated here: G-DC [G00144331]; isolectotypes: BM [BM000943868], K [K000414175], P [P00341909]).
Solanum burbankii Bitter, Repert. Spec. Nov. Regni Veg. 12: 83. 1913, as ‘Burbanki’.
Type. Cultivated in Germany at Bremen, the seeds originally from Hamburg Botanical Garden thought to have been sent by Luther Burbank from United States of America. California, Santa Rosa (no specimens cited; no original material found); Cultivated in St. Louis, Missouri (USA) in the garden of Mr. Waldstein, seeds from Childs, 1 Sep 1909, Anon. [Waldstein] s.n. (neotype, designated here: MO [MO-3002957, acc. # 6651350]).
Solanum nigrum var. probstii Polg., Mitteil. Naturfor. Gesellsch. Solothurn 8: 71. 1928.
Type. Switzerland. Solothurn: Degend. K.K. [= Wollkompost der Kammgarnfabrik Derendingen], [in protologue “22, 24, 26, 27”], 24 Aug 1926, R. Probst s.n. (syntypes; no herbarium cited; lectotype, designated here: BP [acc. # 485285]).
Solanum burbankii var. glabrescens Polg., Mitteil. Naturfor. Gesellsch. Solothurn 8: 72. 1928.
Type. Switzerland. Solothurn: Degend. K.K. [= Wollkompost der Kammgarnfabrik Derendingen], [in protologue “22, 27”], 20 Oct 1922, R. Probst s.n. (syntypes; no herbarium cited; lectotype, designated here: BP [acc. # 485327]).
South Africa. Eastern Cape: Graaff Reinet (“Graafeynet”), 3000–4000 ft, 1838, J.F. Drège 7864b (lectotype designated here: G-DC [G00144331]; isolectotypes: K [K000414172], S [acc. # S-G-5707]).
Annual to perennial prostrate to erect herbs to 0.6 m tall, subwoody and branching at base. Stems sprawling, terete or ridged, 0.3–0.6 cm in diameter, if stems ridged the ridges sometimes spinescent, green to yellowish-brown, older stems straw coloured, not markedly hollow; new growth sparsely to densely pubescent with simple, spreading, uniseriate, translucent, glandular and/or eglandular trichomes, these 1–5(-8)-celled, 0.1–0.8 mm long; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, (0.5-) 1.5–7.5 cm long, 1.5–5.5 cm wide, rhomboidal to lanceolate, membranous, green, slightly discolorous, without smell; adaxial surface green, sparsely to densely pubescent with simple uniseriate trichomes like those on stem evenly spread along lamina and veins; abaxial surface slightly paler, more densely pubescent along veins and lamina; major veins 3–7 pairs; base truncate then abruptly attenuate along the petiole; margins shallowly toothed, the teeth rounded; apex acute, the tip sometimes rounded; petioles (0.5-) 1.5–3.5 cm long, sparsely to densely pubescent with simple uniseriate trichomes like those of the stems. Inflorescences 1.8–3.0 cm long, internodal, simple, sub-umbelliform, with 3–7 flowers clustered towards the tip of the rhachis, sparsely to densely pubescent with glandular and/or eglandular simple uniseriate trichomes like those on stems; peduncle 1.5–3.5 cm long, erect; pedicels 1.0–1.5 cm long, 0.3–0.6 mm in diameter at the base, 0.4–0.6 mm in diameter at the apex, recurving but not fully reflexed, pubescent like the peduncle, becoming woody, green or yellow-brown, articulated at the base; pedicel scars spaced 0–0.5 mm apart. Buds globose, the corolla 1/3 exserted from the calyx before anthesis. Flowers 5-merous, all perfect. Calyx tube 1.0–1.7 mm long, campanulate, the lobes equal, 1.0–1.5 mm long, less than 1 mm wide, oblong with rounded tips, green, sparsely pubescent with simple uniseriate trichomes like of the inflorescence. Corolla 11–16 mm in diameter, white, with a yellow basal star, stellate, lobed to 1/2–2/3 towards the base, the lobes 5.0–6.0 mm long, 2.5–2.7 mm wide, spreading to reflexed, densely papillate-pubescent abaxially with simple uniseriate trichomes, these denser on tips and margins. Stamens equal; filament tube minute; free portion of the filaments 1.2–1.5 mm long, glabrous or adaxially pubescent with tangled 6–8-celled simple uniseriate trichomes; anthers 1.3–1.8(-2.0) mm long, 1.0–1.5 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying, the connective becoming brownish in dry material. Ovary rounded, glabrous; style 1.9–2.2 mm long, slightly curved, pubescent with simple uniseriate trichomes 0.2–0.5 mm long in the basal 1/3 where included in the anther cone, exserted 0.5–1.5 mm beyond anther cone; stigma capitate, the surface minutely papillate. Fruit a globose berry, 6–10 mm in diameter, purple-black at maturity, the pericarp thin, matte with a glaucous cast; fruiting pedicels 1.0–1.5 cm long, 0.4–0.6 mm in diameter at the base, 1.0–1.2 mm at apex, becoming woody, recurving to deflexed, pale green to yellow-brown, spaced 0–0.5 mm apart, not falling with the fruit, remaining on the plant and persistent on older inflorescences; fruiting calyx not accrescent, the tube 1.0–1.5 mm long, the lobes 1.5–2.0 mm long, strongly reflexed. Seeds (5-)12–35 per berry, 1.6–1.8 mm long, 1.3–1.5 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow to brown, the surfaces minutely pitted, the testal cells rectangular to pentagonal in outline. Stone cells absent. Chromosome number: 2n=4x=48 (
Solanum retroflexum A Habit B Glandular indumentum present in some individuals C Flowers D Mature, slightly ovoid dull black-purple fruits with reflexed calyx lobes (A, C Nijmegen acc. A1450022; B Nijmegen acc. 944750163; C Nijmegen acc. A14750023; D Nijmegen acc. A14750025). Photos by S. Knapp and G. van der Weerden.
(Figure
Grows in disturbed soil at watering holes, along dry watercourses, in shady places, in long grass, in Euphorbia candelabrum-Buxus zone, in dry hillsides and in disturbed areas along roads; between sea level and 1,800 (-2,300) m elevation.
South Africa: mofhswe, nastegaal/nastergal, umsobo, gsoba, msoba (
Berries used raw and for jam (
(
Solanum retroflexum is a species that shows great variation in its indumentum, varying from nearly glabrous to densely pubescent with either eglandular or glandular trichomes. AFLP studies have shown that specimens of S. retroflexum with different indumentum types are genetically highly similar (
The species can be distinguished from other Old World morelloids based on the combination of characters of few-flowered inflorescences with 3–7 flowers, relatively long filaments (1.2–1.5 mm long) compared to anther length (1.3–1.8(-2.0) mm), strongly reflexed calyx lobes in fruit and matte purple-black berries that lack stone cells and drop without the pedicels. Leaf shape is generally rhomboidal, which is characteristic of the species and helpful in distinguishing it from closely related S. nigrum and S. villosum. Calyx lobes are generally longer than in S. villosum, S. scabrum or S. nigrum. It can be distinguished from the morphologically similar S. villosum based on its opaque purple-black fruit, its pedicels that remain behind after fruits drop, deeply stellate corolla with long corolla lobes and the long filaments as compared to anther length.
Solanum retroflexum is cultivated commercially in South Africa in KwaZulu-Natal and Free State for jam production (
Solanum retroflexum is a tetraploid species with unclear parentage. Previous studies have suggested that one of the contributing parents of the tetraploid S. retroflexum is the diploid S. americanum (
Solanum retroflexum or one of its close relatives has been suggested to have contributed to the origin of the hexaploid S. nigrum (
In describing S. retroflexum,
Solanum burbankii was described from living material sent to Bitter (“colui ipso in horto Bremeno complures per annos e seminibus ab horto Hamburgensi acceptis” [grew in the Bremen garden for several years from seeds originally accepted from Hamburg]) and thought to have been from California (
The Swiss botanist Rudolf Probst worked extensively on the adventive flora found around woollen mills in Solothurn (Switzerland) and sent much of his Solanum material to the Hungarian botanist Sandor Polgár, apparently both as specimens and as seeds. In his various publications (
Australia. New South Wales: Bouddi, 1970, Edmonds C74 (K); Queensland: Somerset, Kentville, 28 Oct 1965, Henderson 123 (BRI, K); South Australia: Lower Eyre Peninsula, Sec[tion] 10, Hundred of Flinders, 13 Nov 1966, Alcock 1268 (AD); Pilli Waterhole, sect. 10, Hundred of Flinders, 27 Apr 1968, Alcock 2099 (AD, CANB, K); Victoria: Mildura, Hattah/Kulkyne National Park, 11 Feb 1989, Browne s.n. (AD).
Botswana. Livingstone’s Cave, Melepelele, 35 mi NW of Gaberone, 21 Apr 1974, Mott 236 C (K).
Lesotho. Leribe: LHDA Phase 1A, 13 Jan 1996, Phillipson 4757 (MO); Basutoland, Dieterlen 157 (BM, K); Kolonyama plateau W border, 4 Oct 1969, Williamson 48 (K).
Malawi. Southern: Blantyre, Ndirande Mountain, 2 May 1970, Brummitt 10323 (K);
Mozambique. Zambezia: Mocuba, ao km 40, estrada de Milange, 18 Mar 1943, Torre 4957 (MO);
Namibia. Erongo Mountains, 28 Jun 1916, Pearson 9834 (K); Khomas: Windhuk Bergland, regio Finkenstein, 13 Dec 1963, Seydel 3783 (A, K, MO).
South Africa. Eastern Cape: turnoff to Carlisle Bridge from Grt Riebeek East Rd, Grahamstown grid, 31 Mar 1974, Bayliss BRI-B-761 (A, K); Annes Villa, Zuurberg, 2 Oct 1975, Bayliss 7182 (A, MO); Graaff Reniet, Bolus 50 (F, K); SA Nr camp site 1 km E of Steilkop, New Agatha Forest Reserve, 21 Apr 1971, Scheepers 54 (EA); Free State: Fauresmith, Apr 1937, Henrici 3080 (K); Mequatling’s Nek, Apr 1972, Jacobs 8533 (K); Gauteng: Pretoria, Carlisle Bridge, Albany, 2 Apr 1978, Bayliss 8679 (MO); Burttholm, Verreniging, 25 Apr 1918, Burtt Davy 17658 (K); Magaliesberge, Pretoria District, 8 Jun 1955, Schlieben 7009 (F, K, MO); KwaZulu-Natal: 2829 (Harrismith) Drakensberg, Royal Natal National Park, 19 Jan 1987, Goldblatt & Manning 8416 (MO); Drakensberg Range, Tugela Valley & Mont-aux-Source, 3 Apr 1934, Humbert 14849 (MA); Distr. Alexandra, Station Dumisa, Campbellson, 17 Dec 1912, Rudatis 1805 (W); Ngome, 15 Dec 1969, Strey 9409 (E, EA, K); Limpopo: Duiwelskop, grid no. 3219CA, Clamvillham District, 15 Oct 1983, Bean 1366 (MO); New Agatha Forest Reserve, nr campsite 1 km E of Steilkop, 21 Apr 1971, Múller & Schespere 54 (K); Mpumalanga: Ermelo, Nooitgedacht Research Station, 19 Jan 1976, Balsinhas 2901 (K, MO); Namaqualand, Klipfontein, 23 Dec 1949, MacDonald 117 (BM); Bei der Stadt Lydenburg, Mar 1884, Wilms 1022 a (BM, E); North West: head of Helskloof, Hottentosparadyskloof, 28 Aug 1977, Thompson & Le Roux 134 (K); Magaliesberg, Jackson’s kiln, 28 Feb 1957, Vueeden 97 (K); Northern Cape: Westeljike bellings van Rebunie, Calvinia, 27 Dec 1977, Hanekom 2499 (K); Namaqualand, Ai-Ansi, 1 Jan 1950, MacDonald 137 (BM); Western Cape: Pro Spei (Cape of Good Hope), 1771, Banks & Solander s.n. (BM); Nr drift of Cayman’s River, George, Barkly s.n. (BM); Table Mountain, Burchell 856 (K); Simon’s Bay, Cape of Good Hope, 1853, Wright s.n. (GH).
Swaziland. Ukulula, Mbabane Dist, 13 Mar 1955, Compton 25011 (K).
Zambia. Central: Copperbelt: Kitwe, 15 Jan 1960, Fanshawe, F-5353 (K); Northern: Kaloswe, Koloswe, 16 Jul 1930, Hutchinson & Gillett 3767 (K); Zimbabwe. Owelo Teachers College, 11 Nov 1966, Biegel 1413 (K); Vumba, Clouds Downs, 29 Feb 1960, Head 170 (BM); sin. loc, Hislop Z-155 (K); Bulawayo, Jan 1898, Rand 171 (BM); Bulawayo: Bulawayo, garden, 9 Oct 1946, Best 499 (K); Harare: Harare, Salisbury Expt. Station, 26 May 1943, Arnold 10206 (K); Harare, Salisbury Botanic Garden, 30 Jan 1969, Muller 743 (K); Manicaland: Umtali, Nuza plateau, Mutasa Distr., Oct 1934, Gilliland 974 (BM, K); Nyanga, Inyanga Distr., 11 Jan 1931, Norlindh & Weimarck 4199 (BM, MO); Mashonaland Central: Mazowe, southern Rhodesia, Mazoe, Umvukwes, Ruorka Ranche, 17 Dec 1952, Wild 3961 (MO); Mashonaland East: Pasture Research Station, Marandellas, 28 Mar 1934, Brain 10574 (MO); Masvingo: Victoria, Rhodesia, 1909, Monro 1024 (BM); Matabeleland North: Makoholi Experiment Station, Victoria District, 13 Mar 1978, Senderayi 226 (K); Matabeleland South: Matobo, Farm Beana, Kobila, Apr 1957, Miller 4341 (K); Gwanda, Tuli Experimental Station Reservior in pasture block, 16 Jan 1965, Norris-Rogers 599 (K); Midlands: Sable Park, QueQue District, 13 Mar 1978, Chipunga 168 (MO).
Solanum justischmidtii E.H.L.Krause, Deutschl. Fl. (Sturm), ed. 2, 10: 72. 1903.
Type. Germany. Hamburg: Hamburg, J. Schmidt s.n. (lectotype, designated by
Solanum sarachidium Bitter, Repert. Spec. Nov. Regni Veg. 11: 211. 1912.
Type. Paraguay. "Gran Chaco: Loma Clavel", Nov 1903, T. Rojas 2493 (lectotype, designated by
Solanum sarrachoides var. sarachidium (Bitter) C.V.Morton, Revis. Argentine Sp. Solanum 122. 1976.
Type. Based on Solanum sarachidium Bitter
Brazil. “Brasilia australis”, F. Sellow s.n. (lectotype, designated by
Annual erect to decumbent herbs 30–70 cm tall, rarely to 1 m, somewhat branching at base. Stems sprawling, terete, green, not markedly hollow; new growth densely viscid-pubescent with simple, spreading, uniseriate, translucent, glandular trichomes, the trichomes of two lengths, the shorter 1–4-celled, 0.2–0.5 mm long and the longer 5–14-celled, 1–2 mm long, both with glandular apical cells; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 3.0–7.5 cm long, 3.0–6.0 cm wide, broadly ovate, thin and membranous, concolorous, without smell; adaxial and abaxial surfaces sparsely to densely pubescent with spreading, simple, uniseriate glandular trichomes like those of the stem, evenly distributed on lamina and veins; major veins 3–4 pairs; base truncate to cordate, sometimes asymmetric; margins entire or regularly sinuate-dentate; apex acute; petioles 0.5–3.2 cm long, sparsely pubescent with trichomes like those of the stem and leaves. Inflorescences 0.7–1.7 cm long, usually leaf-opposed but occasionally internodal, simple, sub-umbelliform, with 2–5(-7) flowers clustered at the tip, sparsely pubescent with spreading trichomes like those of the stems; peduncle 0.7–1.0 cm long, straight; pedicels 5–7 mm long, 0.1–0.2 mm in diameter at the base, 0.3–0.4 mm in diameter at the apex, straight and spreading, articulated at the base; pedicel scars spaced ca. 0(-1) mm apart. Buds globose, the corolla included within the calyx lobes and only the tip of the bud showing. Flowers 5-merous, all perfect. Calyx tube 0.5–1.0 mm long, conical, the lobes 1.5–2.0 mm long, 0.5–0.7 mm wide, lanceolate to narrowly ovate with acute apices, sparsely pubescent with 1–4-celled spreading glandular trichomes like those on the pedicels but shorter. Corolla 5–8 mm in diameter, white with a yellow-green central eye, pentagonal-stellate, lobed 1/2–1/3 of the way to the base, the lobes 3.0–4.5 mm long, 5.0–7.0 mm wide, spreading at anthesis, sparsely papillate-pubescent abaxially with glandular 1–4-celled simple uniseriate trichomes and eglandular papillae, the denser along margins, tips and midvein. Stamens equal; filament tube minute; free portion of the filaments 1.0–1.5 mm long, adaxially sparsely pubescent with tangled uniseriate 4–6-celled simple trichomes; anthers 1.2–2.0 mm long, 0.4–0.8 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 3.0–3.5 mm long, densely pubescent with 2–3-celled simple uniseriate trichomes in the lower 1/2–2/3 where included in the anther cone, not usually exserted beyond the anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 6–9 mm in diameter, green-brownish grey at maturity, the pericarp thin and dull to somewhat shiny; fruiting pedicels 5–9 mm long, 0.2–0.3 mm in diameter at the base and at the apex, spaced 0–1 mm apart, reflexed, dropping with mature fruits, not persistent; fruiting calyx accrescent, becoming papery in mature fruit, the tube 3–4 mm long, the lobes 5.5–8.0 mm long and 3.5–4.0 mm wide, the tips slightly reflexed or spreading. Seeds (23-)59–69(-93) per berry, 1.3–1.7 mm long, 1.0–1.5 mm wide, flattened and tear-drop shaped with a subapical hilum, pale yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells 4–6 per berry, (0.5) 0.8–1 mm in diameter. Chromosome number: 2n=2x=24 (
Grows in urban areas, along riversides and other disturbed areas; between sea level and 2,300 m elevation in its native range, between sea level and 800 (1,400) m in the introduced range.
South Africa: umrobe wezinja; Sweden: Klibbnattskatta (
None recorded.
(
Solanum sarrachoides is morphologically similar to S. nitidibaccatum. The two taxa can be distinguished based on generally truncate leaf bases, leaf-opposed inflorescences that are umbellate to sub-umbellate with fewer flowers (2–5, rarely 6–7), shorter calyx lobes 1.5–2.0 mm long and a corolla with yellow-green central eye in S. sarrachoides, compared to S. nitidibaccatum which has attenuate to cuneate leaf bases, internodal inflorescences that are racemose with more flowers (4–8, occasionally up to 9–10), longer calyx lobes 1.7–2.5 mm long and a corolla with yellow-green central eye with black-purple “V”- or “U”-shaped margins. The buds of S. sarrachoides are included in the calyx until just before anthesis and the berries are usually matte instead of shiny as they are in S. nitidibaccatum.
Solanum sarrachoides is a diploid species native to north-eastern and central Argentina, Paraguay and southernmost Brazil. The introduction of the species to Europe and North America is largely due to trade with South America and the importation of seeds and grain together with the practice of spreading wool waste ('shoddy') as manure and, based on herbarium records, seems to have been introduced some time at the beginning of the 20th century. Despite its introduction in the early 1900s, the species remains relatively uncommon in both Europe and North America, with sporadic records from elsewhere, including South Africa (e.g. Eastern Cape, Phillipson & Hobson 5256). Solanum sarrachoides has not spread to Australasia; all literature records (e.g.
Within the Old World, S. sarrachoides has often been confused with S. nitidibaccatum and records of S. sarrachoides or S. nitidibaccatum in literature should be taken with caution due to common misidentification of voucher material. Many regional treatments do not separate between these morphologically very similar species (e.g.
Austria. Steiermark: Fussee, Mistablagerumgsplatz, 26 Aug 1950, Rechinger s.n. (MA).
France. Grand Est: Bas-Rhin, Strasbourg, Petrolhafen in Strassburg, 18 Oct 1953, Aellen s.n. (W); Nouvelle Aquitaine: Gironde, Gironde, Bassem, 19 Sep 1924, D’Alleizette s.n. (W); Gironde, Bordeaux, 6 Sep 1926, Herb. Guiol s.n. (BM).
Germany. Neuft, [Speten], 20 Sep 1917, Boutz s.n. (B); Hamburg: Hamburg, bei Wandobek, Sep 1900, Schmidt s.n. (W); Hessen: Herdingen, Rheinwerfl, 26 Sep 1920, Boutz s.n. (B).
South Africa. Eastern Cape: Amatole Mountains, Elandsberg, Coolin farm, 20 Mar 1986, Phillipson 1353 (K, MO); Rooiberge, nr Graaf Reinet Rooiskuur dam on Roodeberg Farm, 17 Dec 2000, Phillipson & Hobson 5256 (K, MO).
Spain. Andalucia: Granada, Cacín, 9 Nov 1970, Pérez Raya s.n. (MA); Castilla-La Mancha: Toledo, Montalbán, embalse de Castrejón, 18 Apr 1983, López s.n. (MA).
Sweden. Götaland: Skåne, Malmö, Sep 1906, Hylmö s.n. (BM); Västra Götaland, Mölndal, Svenska Oljeslageriet, 30 Aug 1936, Blom s.n. (BM); Västra Götaland, Angered, Agnesbergskvarn, 31 Aug 1938, Blom s.n. (W); Västra Götaland, Molndal, Sveska Oljeslageriet, 30 Aug 1936, Blom 1376 (CORD, K, W); Västergötland, Asfroued Au, Sjoliageu, 10 Sep 1966, Westfelt s.n. (BM).
United Kingdom. England: Bedfordshire, Cabbage field at Flitton, 8 Oct 1974, Hanson 107a (BM); Essex, Wasteground Dagenham, 2 Oct 1927, Melville s.n. (K); Essex, Dagenham, 6 Oct 1945, Sandwith & Milne-Redhead s.n. (BM, K); Essex, Barking Tip, 12 Sep 1953, Welch 5298 (BM); Gloucestershire, Avonmouth Docks, 29 Aug 1922, Polgár s.n. (BM); Hertfordshire, Wheathampstead, 14 Oct 1962, Dony 4165 (BM).
Solanum nigrum var. guineense L., Sp. Pl. 186. 1753.
Type. “Solanum guineense fructu magno instar cerasi nigerrimo umbellato” cultivated in England, at James Sherard’s garden in Eltham (Hortus Elthamensis) (lectotype, designated by
Solanum guineense (L.) Mill., Gard. Dict., ed. 8, no. 7. 1768, nom. illeg., not Solanum guineense L. (1753)
Type. Based on Solanum nigrum var. guineense L.
Solanum melanocerasum All., Auct. Syn. Meth. Stirp. Hort. Regii Taur. 664. 1774.
Type. “Solanum guineense, fructo magno, instar cerasi nigerrimo, umbellato” cultivated in England at John Sherard’s garden in Eltham (Hortus Elthamensis), Herb. Dillenius 336 (neotype, designated by
Solanum guineense (L.) Lam. Tabl. Encycl. 2: 18. 1794, nom. illeg., not Solanum guineense L. (1753), Solanum guineense (L.) Mill. (1768)
Type. Based on Solanum nigrum var. guineense L.
Solanum triangulare Lam., Tabl. Encycl. 2: 18. 1794.
Type. “Ex Ind. Orient”, Herb. Lamarck s.n. (lectotype, designated by
Solanum quadrangulare var. triangulare (Lam) Pers., Sys. 225. 1805.
Type. Based on Solanum triangulare Lam.
Solanum melanocerasum Willd., Enum. Pl. (Willdenow) 1: 237. 1809, nom. illeg., not Solanum melanocerasum All. (1773)
Type. Cultivated in Berlin Botanical Garden, from “Europa australi” [southern Europe] (no original material labelled as “melanocerasum” found in B-W; neotype, designated here: B-W [BW04368010]).
Solanum pterocaulum Dunal, Hist. Nat. Solanum 153. 1813, nom. illeg. superfl.
Type. Based on Solanum scabrum Mill. (cited in synonymy).
Solanum fistulosum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 749. 1814.
Type. “Originaire de l’Isle de France [Mauritius], est cultivée en Amerique [Brazil]”, Herb. Richard s.n. (lectotype, designated by
Solanum memphiticum Mart., Pl. Hort. Erlang. 63. 1814, nom. illeg., not Solanum memphiticum J.F.Gmel. (1791)
Type. Cultivated in Erlangen, Bavaria, Germany, origins not known (no specimens cited; no original material located, possibly at ER?).
Solanum nigrum var. melanocerasum (Willd.) Dunal, Solan. Syn. 12. 1816.
Type. Based on Solanum melanocerasum Willd.
Solanum nitens Opiz, Oekon.-techn. Fl. Böhm. [Berchtold & al.] 3(2): XXI. 1843.
Type. Czech Republic. Prague, “in Semubackern por dem Reuthore Prag”, P.M. Opiz 10/840 (no herbaria cited; no original material found, perhaps at PR?).
Solanum oleraceum var. macrocarpum Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852.
Type. Brazil. Bahia: Ilheus, 1841, C.F.P. Martius 1255 (lectotype, designated by
Solanum tinctorium Welw., Apont. 590. 1859 [1858].
Type. Angola. Golungo Alto, 1856, F.M.J. Welwitsch 6103 (lectotype, designated here: BM [BM000942995]; isolectotypes: BM [BM000942996], K [K001029777]).
Solanum nigrum var. pterocaulum (Dunal) Schur, Enum. Pl. Transsilv. 478. 1866, as ‘pterocaulon’.
Type. Based on S. pterocaulum Dunal [“excl. German floras”]
Solanum boerhaavii Thell., Rep. Bot. Soc.& Exchange Club Brit. Isles 8: 187. 1927, as “Boerhaavii”.
Type. Based on Solanum nigrum var. guineense L. [as replacement name for Solanum guineense (L.) Mill.]
Solanum nigrum var. pterocaulum (Dunal) Domin, Biblioth. Bot. 89: 1127. 1928.
Type. Based on S. pterocaulum Dunal
Solanum intrusum Soria, Baileya 7: 33. 1959.
Type. Based on (replacement name for) Solanum nigrum var. guineense L., and Solanum guineense (L.) Lam.
Solanum scabrum subsp. laevis Olet, Novon 16(4): 510. 2006.
Type. Uganda. Buganda: Kampala district, Kawempe div., Kawempe North, Kalerwe, Tula road, 1220 m, 14 Feb 2001, E.A. Olet 88 (holotype: MHU; isotypes: H, K, MO [ex descr.]).
Cultivated in Chelsea Physic Garden, said in protologue to “grow naturally in North America”, Herb. Miller s.n. (lectotype, designated by
Annual or short-lived erect or ascending perennial herbs to 1.5 m tall, often woody at the base. Stems spreading, terete, ridged or winged, green to purple, if ridged or winged the stems later spinescent, older stems with or without prominent pseudospines, usually somewhat hollow; new growth puberulent with simple, spreading, uniseriate, translucent, eglandular trichomes, these 2–8-celled, 0.3–0.8 mm long; older stems glabrescent. Sympodial units difoliate, the leaves usually not geminate, but if leaves paired, then one is usually smaller. Leaves simple, 4–15(-20) cm long, 3–10(-16) cm wide, broadly ovate to elliptic, very variable in size depending on cultivars and growth conditions, membranous to somewhat fleshy, green to dark green to somewhat purple coloured above, slightly paler below, without smell; adaxial and abaxial surfaces glabrous or sparsely pubescent with simple uniseriate trichomes like those on the stem mainly along veins and scattered along lamina; major veins 3–6(-8) pairs, paler green or often purple tinged; base abruptly acute or truncate, narrowly winged on to the petiole; margins entire or rarely shallowly sinuate; apex rounded to acute; petioles 1–5(-8) cm long, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the stem. Inflorescences 1–2(-4) cm long, internodal, simple, furcate or many times branched (in cultivars), sub-umbelliform, with 4–10(-30+) flowers clustered towards the tip(s) of the rhachis, glabrous or sparsely pubescent with simple uniseriate trichomes like those on the stem; peduncle 1–5(-8) cm long, erect and thick, much thickened at apex, subwoody; pedicels 0.4–1 cm long, 0.3–0.5 mm in diameter at the base, 0.75–0.9 mm in diameter at the apex, abruptly expanding to the calyx tube, stout, erect and/or spreading, articulated at the base; pedicel scars tightly clustered near the tip of the rhachis, spaced 0–2 mm apart, sometimes with short stumps ca. 0.5–1.0 mm long. Buds globose to subglobose, the corolla exserted 1/2–1/3 from the calyx tube before anthesis. Flowers 5-merous or occasionally fasciated and 6–7-merous in cultivars, all perfect. Calyx tube 0.9–1.1 mm long, abruptly cup-shaped with a broad base, the lobes slightly unequal, 0.9–1.5 mm long, 0.8–1.4 mm wide, broadly deltate with a rounded tip, green or purple-tinged, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the pedicels, the margins often drying scarious and white. Corolla 7–12 mm in diameter, white, purple-tinged or occasionally lilac to dark purple, with a yellow basal star, stellate, lobed ca. 1/2 way to the base, the lobes 2.5–4 mm long, 1.5–3 mm wide, spreading or reflexed, densely papillate on tips and margins. Stamens equal; filament tube very short, to 0.1 mm long; free portion of the filaments 0.5–0.8 mm long, glabrous or pubescent with tangled uniseriate simple trichomes; anthers 2–3 mm long, ellipsoid or slightly tapering towards the tips, yellow, orange or brown, poricidal at the tips, the pores lengthening to slits with age and drying, the connective often becoming brownish-black in dry specimens. Ovary rounded, glabrous; style 2.5–5 mm long, densely pubescent with simple uniseriate trichomes 0.2–0.5 mm long in the basal 1/2 where included in the anther cone, exserted beyond anthers 0–1.5 mm; stigma capitate, the surface minutely papillate. Fruit a globose to depressed-globose berry, 10–20 mm in diameter, purplish-black at maturity, the pericarp thick, shiny or somewhat matte, not transparent; fruiting pedicels 0.7–1.5(-2.0) cm long, 0.5–1 mm in diameter at the base, 1.1–1.5 mm in diameter at the apex, stout, erect and spreading, purple or brown, usually not falling with the fruit, remaining on the plant and often persistent on older inflorescences; fruiting calyx not accrescent, the tube 1.5–2 mm long, usually tearing unevenly, the lobes 2–3 mm long, usually with thicker white margins in dry material, appressed or spreading to slightly reflexed. Seeds (20-)100–150 per berry, 2–2.8 mm long, 1.5–1.8 mm wide, flattened and tear-drop shaped with a subapical hilum, yellow-brown or purple, the surfaces minutely pitted, thin and the embryo clearly visible, the testal cells rectangular to pentagonal in outline. Stone cells absent. Chromosome number: 2n=6x=72 (
Solanum scabrum A Habit of wild form B Flower of wild form C Infructescence of wild form D Habit of cultivated form E Inflorescence of cultivated form F Fruit of cultivated form G Seed (A–CPilz 2108; D–GNee 16088). Scale bar: 4 cm (A, D), 3.3 mm (B), 1.5 cm (C, F), 7 mm (E) and 2 mm (G). Drawing by L. Smith.
Solanum scabrum A Common habit B Habit in taller varieties C Flowers of the larger berried variety at full anthesis D Fruits of a larger berried variety E Flowers of the smaller berried variety at full anthesis F Fruits of a smaller berried variety (A Nijmegen acc. BG13; B Nijmegen acc. A34750072; C Nijmegen acc. GB22; D Nijmegen acc. H065; E Nijmegen acc. A34750067; F Nijmegen acc. 2010/3). Photos by S. Knapp.
Grows in open areas, in a wide variety of habitats, in wet forests or drier areas, along roads and at field edges; often cultivated; between sea level and 2,300 m elevation.
Benin: odu, ogomo, feibii (
Leaves used as greens (spinach) and cooked, sold in markets; berries used as ink; in Benin, the powdered leaves are used to cure dysentery (
(
Solanum scabrum is the most commonly cultivated and widespread of the African black nightshades. It is the most important indigenous leafy vegetable in the black nightshade group and is commonly known as the African nightshade (
The species has also been introduced to Europe, North America, Australia and New Zealand. In the United States of America, S. scabrum is known as “Garden Huckleberry” and its identity, origin and suitability to human consumption were the subject of great interest in the 1960s (as S. guineense, e.g.
Different cultivars are recognised, the late flowering plants cultivated for their larger abundance of leaves with smaller number of fruits and earlier flowering plants that have larger inflorescences which are cultivated for their fruits (
The presumed wild forms were described at the infraspecific rank by
Solanum scabrum can be distinguished from the somewhat similar S. americanum by its larger anthers (2–3 mm long versus 0.8–1.5 mm long). In both these species, as well as S. retroflexum, the berries usually lack stone cells (some populations of S. americanum can have up to 4 stone cells) and drop without the pedicels at maturity, leaving the pedicels behind on old inflorescences. In both S. scabrum and S. americanum, berries are purple-black and shiny, while S. retroflexum has dull purple-black berries with a distinct grey bloom. The pedicels of S. scabrum are usually erect or spreading.
Plants described as S. fistulosum and S. oleraceum var. macrocarpon by
Solanum scabrum is closely related to S. nigrum and S. villosum based on molecular data (
The nomenclature and typification of the various synonyms of S. scabrum and the earliest name applied to the taxon, have been extensively discussed by others (e.g.
Solanum pterocaulum is illegitimate because
No original material has been traced for S. memphiticum Mart., but the emphasis placed on the dark purplish colour of the stems and leaves by
Of the several species in this group described by P.M. Opiz (see discussion under S. nigrum), we are convinced from the description that S. nitens is a synonym of S. scabrum, rather than of S. nigrum, although
The lectotype (G00144295) chosen for S. oleraceum var. macrocarpum is the better preserved of the two duplicates of Martius 1225 cited by
The lectotype chosen for S. tinctorium (Welwitsch 6103) is the more clearly duplicated of the collections cited in the protologue; we have selected the best preserved sheet (BM000942995) as the lectotype. The name apparently comes from the staining berries that were used as ink.
Angola. Bengo: Ambriz, Abriz, Dec 1872, Monteiro s.n. (K); Benguela: Hochland zwischen Ganda und Caconda, Dec 1933, Hundt 797 (BM); Cuanza Norte: Varzea do Isidoro ad rivum Cuango, Dto. Golungo Alto, Jul 1855, Welwitsch 6100 (BM); Malanje: Distr. Pungo Andongo, prope Lusillo, Jan 1857, Welwitsch 6108 (BM); Namibe: Mofsamedes, Herb. Moira, Habit Cavalheiros, Jul 1859, Welwitsch 6033 (BM).
Benin. Atlantique: Wida, Dahomey, 26 Aug 1903, Estève 111 (BM).
Botswana. Ghanzi: Ghanzi camp, 4 May 1969, Brown 6019 (K); Ghanzi camp, 4 May 1969, Cole 6019 (K); North West: Mutsoi NE of Nokaneng, 21 Mar 1967, Lambrecht 90 (K).
Burkina Faso. Seno: Dori Dam, 16 Oct 2007, Sanou & Leonard BUR-596 (K).
Cameroon. Adamaoua: Mayo-Banyo, Mambilla Plateau Cameroon, around Somie village, 21 Jun 2009, Komaromi 48 (K); Centre: Mimboman area, Yaounde, 16 Sep 1986, Manning 235 (MO); Extreme-Nord: Monts Mandara, Hossere Oupay, 15 km NNO de Mokolo, 15 Sep 1964, Letouzey 6880 (K, P); Littoral: Donala, 16 Aug 1986, Johns 86-481 (K); Nord-Ouest: Mezam, Above Bamenda, 20 Jan 1928, Migeod 358 (BM, K); Sud-Ouest: Mt Kupe, Kupe Village, 24 May 1996, Ryan 289 (K, MO, P).
Central African Republic. Mboukou Griko, Territoire du Haut-Obangi, 24 Sep 1902, Chevalier 5518 (K).
Comoros. Anjouan: Anjouan (Comoro-Insel Johanna), Jun 1875, Hildebrandt 1626[b] (BM, W); Moheli Island: Moheli, NE center of island, 14 Aug 1987, D’Arcy 17617 (MO, P); 1 Nov 1990, D’Arcy 17765 (MO); Njazídja: Grande Comore, 9 Aug 1981, Doutrelepont 1203 (MO, P).
Côte d’Ivoire. Abidjan: Adjame, 31 May 1973, De Koning 1747 (MO); Dimbokro: c. 5 km W of Dimbokro, 13 Oct 1975, van der Burg 1177 (MO); M’Bahiakro: Koffi-Akakro (s. Prikro), 25 Jul 1973, Smittenberg-Visser 61 (MO); Montagnes: Man, 6 km N nr Yebegouin, 27 Jan 1984, Hepper & Maley 7849 (K).
Democratic Republic of the Congo. Katanga: Lukonzolwa, Moero, 1933, Quarré 3297 (K); Kongolo, 5 Feb 1920, Schantz 646 (K); Kinshasa: Maluku, Route Menkao-Kingankati, 5 Nov 1971, Breyne 2227 (MO); Nord-Kivu: Lubarika, 1953, Gilon 315[a] (MO); Orientale: Ituri, w.v. Albert-See, Oct 1934, Gusinde s.n. (W); Yangambi, 26 Jul 1938, Louis 10507 (K, P); Sud Kivu: Kizozi, Jul 1933, Lejeune 55 (K).
Egypt. Giza: Faculty of Agriculture, Giza, 13 Jun 1971, Sisi s.n. (MO).
Equatorial Guinea. Annobon: Pico de Fogo, SW side, 25 Jul 1959, Melville 188 (BM, K, MA, P); Bioko: entre Moca y Riaba por el camino viejo, 20 Feb 1989, Fernández-Casas 11820 (K, MA, MO, P); Bioko Norte: Pico Basilé, Carretera del pico Basile, 28 Mar 1990, Do Carvalho 4305 (K, MA, MO, P); Bioko Sur: Moca, camino de Ureca, 18 Feb 1989, Fernández-Casas 11725 (K, MA, MO).
Eritrea. Maekel: Asmara, 1 May 1892, Terracciano & Pappi 187 [2206] (FT); Semienawi Keyih Bahri: Asmara, Beless, Hamasen, 4 May 1892, Terracciano & Pappi 2536 (FT).
Gabon. Estuaire: Jardin Cenarest, Libreville, Poubelle, 16 Jan 1986, Louis 1990 (MO).
Ghana. Ashanti: nr Mampong, Ashanti, 7 Aug 1963, Darko 5115 (K); Central: Aswansi, W.P, 12 Oct 1954, Darko 1032 (K); Eastern: Akosombo, 25 May 1970, Enti 1723 (MO); Akwapim, Mampong Scarp, 14 Jun 1953, Morton s.n. (K); Greater Accra: Accra, May 1961, Irvine 5095 (K); Volta: Adzido, Keta, 16 Sep 1960, Akpabla 2117 (K).
Guinea. Nzérékoré: Lola, Mt. Nimba, 1 Nov 2012, Diabate & Mas 1419 (MO).
Indonesia. Java: Central Java, Mt. Slamet, 15 Mar 2004, Hoover et al. 89 (A); Seram: Manusela National Park, 12 Sep 1987, Argent C87-179 (A, E); Sumatra: E of Berastagi, Karo Highlands, 4 Jun 1928, Si Toroes 407 (A, GH).
Lesotho. Sehlabathebe, 4 Jan 1973, Bayliss 5476 (MO).
Liberia. Central: 3 mi NE of Suacoco, Gbarnga, 7 Feb 1951, Daniel 117 (B, BM, MO); Grand Gedeh: Tchien, Mim Timber Co (Fijnhout), 16 May 1970, de Koning 519 (MO); Lofa: mi along rd to Wologesi, 15 Jul 1970, Jansen 2015 (K, MO, P); Nimba: Nimba Mountains, 27 Jul 1962, Leeuwenberg & Voorhoeve 4668 (B, K, MO, P).
Madagascar. Antananarivo: Grande Terre-Ouangani, Apendzo-Jivany. Barakani, 7 May 2002, Barthelat et al. 885 (K, MO, P); Antsiranana: Mt. d’Ambre, partie centrale, Prov. Diego-Suarez, 11 Nov 2007, Gautier 5198 (K, MO); Fianarantsoa: Ankafana, 1880, Cowan s.n. (BM); 10 km W of Ivato on Route 35, 25 Jan 1975, Croat 29606 (MO); Toamasina: au km 26 de la route de Tamatave, 18 Oct 1951, Benoist 147 (P); Toliara: Beroroha, Vallée du Mangoky et de l’Isahaina aux environs de Beroroha, Oct 1933, Humbert 11309 (K, MO, P); Andohahela RNI, Mt. Trafonaomby, Taolagnaro, 7 Apr 1994, Randriamampionona 692 (MO, P).
Malawi. Central: Salima Distr., Liganga village A. Mpemba, 14 Jun 1985, Kwatha et al. 210 (MO); Salima Distr., Luwadzi stream, 14 Jun 1985, Salubeni et al. 4242 (MO); Northern: Nkhata Bay, Musalowa village, Chizumulu Island, 25 Mar 1989, Balaka & Patel 2025 (K, MO); Southern: Malawe Hill, W of Port Herald, 23 Mar 1960, Phipps 2653 (K, MO).
Mauritania. Trarza: Rosso Administrative area, Village Rosso, 9 Oct 1962, Adam 18728 (MO).
Mozambique. Baroma: N’kanya, N of Zambesi River, 25 Jul 1950, Chase 2857 (BM); Cabo Delgado: Pemba, Jun 1909, Rogers 8256 (K); Manica: Lower slopes of Chimanimani Mountains, Apr 1967, Westwater 192954 (K).
Namibia. Onjossariviers, im Ufergestrupp, 21 Jun 1957, Seydel 1162 (A, K).
New Zealand. North Island: Bay of Plenty, Ohope, Whakatane Ecological Region, Taneatua Ecological District, 13 Mar 1979, Grant s.n. (AK).
Nigeria. Delta: [Ogwashi-Ukwu], 25 Nov 1912, Thomas 2042 (K); Edo: Nikrowa Forest Reserve, Mid-west State, Iyekovia Distr., 8 Oct 1973, Daramola FHI-72483 (K, MO); Enugu: 10 mi E of Erugu, Mar 1948, Irvine 3607 (K); Gongola: Gemu Distr., Mambilla plateau, 18 Aug 1977, Fagbemi 438 (MO); Jos: Naraguta, Lely 31 (K); Kaduna: Zaria, 1975, Magaji MG-725 (K); Kano: Kano, Wudil Distr, 50 km SE of the city of Kano, 16 Mar 1988, Etkin 63[b] (MO); Kogi: Odu, SW Nigeria, Van Eyenhuisen 7 (K); Lagos: Lagos, Dalziel 1188a (K); Lagos, Dalziel 1188b (K); Niger: Nupe, Barter 1054 (K, P); Ondo: between Ikare and Oke-Agbe, Ikare District, 2 May 1979, Daramola & Osanyinlusi 154 (K); Oyo: Ibadan, 7 km W of Polytechnic, 11 Jun 1977, Pilz 2108 (K, MO); Taraba: State of Gongola, Distr. Mambilla Plateau, Nguroje township, 23 May 1982, Odewo 130 (MO).
São Tome e Principe. Bom Sucesso, 27 Jan 1949, Espírito Santo 218 (BM); Vanhulst (Macambrará), 28 Oct 1932, Exell 89 (BM).
Senegal. In paludilb[us] N’Boro nec non ins[ula] Bonavista, 1838, Brunner 108 (BM, G, W); Sin. loc., 1828, Perrottet 555 (BM, W); Perrottet 556 (BM);
Seychelles. Morne Blanc, 2 Oct 1970, Schlieben 11670 (B, K).
Sierra Leone. Eastern: Kailahun, Musaia, 19 Dec 1946, Deighton 4571 (K); Northern: Port Loko, Rokupr, Magbema, 18 Apr 1959, Jordan, 1059 (K); Yonibana, 12 Nov 1914, Thomas 4909 (W); Western: Freetown, 28 Apr 1965, Morton s.n. (K).
South Africa. Eastern Cape: Grahamstown, 7 Sep 1970, Bayliss 4594 (A); KwaZulu-Natal: 2632 CD Bella Vista grid, between Ndumu Store and the Game Reserve, 30 Oct 1969, Moll 4138 (A, K); Limpopo: Mopani, Shiluvane, Aug 1899, Junod 575 (K); North West: Okawango Delta, Okavango Delta, Delta camp, 16 Jun 1994, Cole 923 (K); Northern Cape: Augrabies Falls National Park, Orange river bank, 14 Mar 1978, Balsinhas & Harding 3290 (K, MO); Western Cape: Dowweklip, Voelklip, Hermanus, 6 Jun 1980, Williams 287 (K, MO); Skeleton Ravine, 3 Oct 1897, Wolley-Dod 3180 (K).
South Sudan. Bahr El Ghazal: Anglo-Egyptian Sudan, Bahr El Ghazal Prov., Ibba, 10 Mar 1934, Dandy 624 (BM, EA); Equatoria: R. Napere, 25 Nov 1937, Wyld 347 (BM).
Tanzania. Dodoma: Mpwapwa, Mbuga Village, Kibakwe Division, Mbuga Ward, 31 May 2005, Kindeketa et al. 2549 (EA, MO); Iringa: Ruaha National Park, FTEA region T7, top of Mpululu mountain, 21 May 1968, Renvoize & Abdallah 2312 (EA, K); Kagera: Bukoba, 30 Oct 1992, Breteler 11599 (MO); Mbeya: Rungwe, Ngumbulu Village, NE part of Rungwe Forest Reserve, 14 Mar 2008, Abeid et al. 2848 (MO); Morogoro: Tanganyika, Ulugurus, Jan 1935, Bruce 524 (BM, K); Shinyanga: Shinyanga, Nov 1938, Koritschoner 2191 (EA, K).
Togo. Lomé, 22 Sep 1976, Ern et al. 883 (B); slopes of Bauman Peak, 14 Aug 1962, Morton A4277 (MO).
Uganda. Central: Kyadondo Mengo (U4); Nr Kanyanya, 16 Jun 1990, Rwaburindore 2992 (MO); Mengo: Distr. W. Mengo, 4 mi Gayaza rd, 5 Jul 1980, Rwaburindore 701 (MO); Northern: Yumbe, 26 Nov 1941, Thomas 4070 (EA, K); Western: Kigezi DFI, 28 Aug 1972, Goode G3-72 (K); Kigezi, Bugangari, Rhuzumbura, Kigezi, Feb 1949, Purseglove 2712 (EA, K).
United Kingdom. England: Hertfordshire, Rye Meads Sewage Works, nr Rye House, 1 Oct 1996, Hanson s.n. (K).
Zimbabwe. Mashonaland Central: Imayanga, Nyamaropa TTk, 16 Jan 1967, Biegel 1767 (MO); Matabeleland North: Binga Distr., Sanyam R. and Zambezi R. confluence, Sep 1955, Davies 1514 (MO); Matabeleland North: Binga, Chizarira Game Reserve, Busi River, 10 Nov 1971, Thomson 481 (K); Wankie: Wankie Distr., Victoria Falls, Elephant Hills Hotel, 19 Dec 1978, Mshasha 144 (MO).
Solanum dasytrichum Bitter, Bot. Jahrb. Syst. 49: 568. 1913.
Type. Tanzania. Tanga: Usambara, Kwai, E. Eick 227 (holotype: B, destroyed; no duplicates found).
Solanum florulentum Bitter, Repert. Spec. Nov. Regni Veg. 10: 544. 1912.
Type. Tanzania. Tanga: Lushoto, Distr. Kwai, 1600 m, E. Albers 189 (holotype: B, destroyed; no isotype at EA).
Solanum kifinikense Bitter, Repert. Spec. Nov. Regni Veg. 10: 545. 1912.
Type. Tanzania. Kilimanjaro: Kifinika volcano, Mar 1894, G. Volkens 1909 (lectotype, designated by
Solanum pentagonocalyx Bitter, Repert. Spec. Nov. Regni Veg. 10: 544. 1912.
Type. Tanzania. Tanga: Usambara, Kwa Mstuza, Handei Kewegolot, Aug 1893, C. Holst 9021 (lectotype, designated by
Solanum tetrachondrum Bitter, Bot. Jahrb. Syst. 49: 565. 1913.
Type. Tanzania. Kilimanjaro: near Marangu, G. Volkens 623 (holotype: B, destroyed; no duplicates found).
Solanum tetrachondrum var. subintegrum Bitter, Bot. Jahrb. Syst. 49: 566. 1913.
Type. Tanzania. Kilimanjaro: Marangu, G. Volkens 622 (holotype: B, destroyed; no duplicates found).
Solanum viridimaculatum Gilli, Ann. Naturhist. Mus. Wien 77: 43. 1973.
Type. Tanzania. Njombe: Madunda, Livingstone-Gebirge am Nordostufer des Nyassasees, bei Madunda, Nebelwald, 2200 m, 29 Jul 1958, A. Gilli 499 (holotype: W [1973-0001020]).
Tanzania. Kilimanjaro: Marangu, 1600 m, 10 Sep 1910, H.J.P. Winkler 3856 (holotype: WRSL).
Annual to short lived erect to weakly scrambling perennials to 1.5 m tall, subwoody and somewhat branching at base. Stems spreading to decumbent, usually somewhat winged and with spinescent processes, fleshy, green to somewhat purple tinged, older stems drying pale yellowish-brown or whitish-grey, markedly hollow and even older stems collapsing in herbarium specimens; new growth glabrous or sparsely to moderately pubescent with simple, spreading, uniseriate, translucent, usually eglandular (sometimes tipped with a single-celled gland) trichomes, these 3–5-celled, 0.2–1.0 mm long; older stems glabrous. Sympodial units difoliate, the leaves usually not geminate. Leaves simple, 3–13 cm long, 1.5–2.5 cm wide, ovate to elliptic, very variable in size even on an individual plant, membranous, concolorous, without smell; adaxial surface glabrescent to sparsely pubescent with simple uniseriate trichomes like those on stem; abaxial surface usually less pubescent that the adaxial ones, glabrous to sparsely pubescent with simple uniseriate trichomes; major veins 4–8 pairs; base abruptly attenuate; margins entire or very rarely shallowly sinuate to lobed, if so the tips of the lobes acute or rounded; apex acute to attenuate; petioles (0-)4–8 cm long, pubescent with simple uniseriate trichomes like the leaves. Inflorescences 1–4(-6) cm long, internodal, unbranched or with up to 5 branches, but if branched usually only furcate, the flowers spaced along the rhachis, with 10–40 flowers, glabrous or sparsely pubescent like the stems; peduncle 1–3(14) cm long; pedicels 0.7–0.8 cm long, ca. 0.3 mm in diameter at the base, ca. 0.5 mm in diameter at the apex, slender, nodding at anthesis, sharply bent just above the insertion point so the base of the pedicel is an acute angle, this especially noticeable in fruit, articulated at the base or ca. 1 mm from the rhachis leaving a small stump, falling with the fruit at maturity; pedicel scars 1–2 mm apart in the distal part of the inflorescence, further apart towards the base. Buds ellipsoid, the corolla exserted ca. 1/2 way from the calyx until just before anthesis. Flowers 5-merous, perfect (although style length differs in flowers along the rhachis). Calyx tube ca. 1 mm long, conical with hyaline sinuses, the lobes 0.5–1 mm long, 0.4–0.9 mm wide, ovate to broadly triangular, glabrous to sparsely pubescent like the rest of the inflorescence. Corolla (6-)8–10 mm in diameter, white, stellate, lobed ca. 1/2 way to the base, the lobes 3–4 mm long, 1.5–2 mm wide, spreading or reflexed (apparently reflexed in older flowers), minutely papillate on the tips and margins. Stamens equal; filament tube ca. 0.25 mm long; free portion of the filaments 0.5–0.75 mmm long, adaxially pubescent with tangled simple uniseriate trichomes; anthers 1.5–2.5 mm long, 0.75–1 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary rounded, glabrous; style 3.5–5 mm long, bent or straight, densely pubescent in the basal 1/2 to 1/3, exserted to ca. 1 mm or occasionally included in the anther cone and the stigma flush with the pores; stigma capitate, the surface minutely papillate. Fruit a globose berry, 4–6 mm in diameter, pale green or translucent blackish-grey when ripe, the pericarp thin, matte; fruiting pedicels 0.8–1.1 cm long, ca. 1 mm in diameter at the base, sharply bent at the base and strongly pendent, dropping with mature fruits, not persistent; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes 0.5–1 mm long, appressed to the berry, the calyx from above pentagonal or stellate. Seeds 20–40 per berry, 1.5–2 mm long, 1–1.5 mm wide, flattened and tear-drop shaped with a subapical hilum, tan or pale brown, the surfaces minutely pitted, the testal cells elongate-rectangular. Stone cells (0-)1–5(-10) per berry. Chromosome number: 2n=4x=48 (
Solanum tarderemotum A Habit of glabrescent form B Habit of pubescent form C Detail of adaxial leaf surface D Eglandular trichome E Glandular trichome F Inflorescence G Bud H Flower at anthesis I Furcate inflorescence J Infructescence (A, JFriis et al. 2047; B–EBurger 2006; F, GGilbert & Thulin 651; H Nijmegen acc. A34750039; IGoode G/72). Scale bar: 4 cm (A–B), 4 mm (C), 0.75 mm (D–E), 1 cm (F, I), 5 mm (G–H) and 1.5 cm (J). Drawing by L. Smith.
Solanum tarderemotum A Habit B Inflorescence C Buds D Flowers at full anthesis E Infructescence F Mature fruits (A Nijmegen acc. A54750214; B Nijmegen acc. A14750164; C Nijmegen acc. A34750035; D Nijmegen acc. A34750039; E Nijmegen acc. A54750214; F Nijmegen acc. HNSAR22). Photos by S. Knapp.
Grows in a wide variety of disturbed habitats, along forest margins and roadsides and in clearings, often cultivated; between (0-) 500 and 3,300 m elevation.
Burundi: isogo [Kirundi language], urusongo; Cameroon: kifon, kre’ fom; Democratic Republic of the Congo: mulunda; Ethiopia: chau, murenchiwa/marenchiwa, nech tunaye; Kenya: abune, isoiik, isoyiot, isusa, litusta/liususa, lokitoemenyan, managu, mnavu, ol’ momoye, osuga, sujet, usuga; Rwanda: isogo, ururandayri [Kinyarwanda language]; Tanzania: enyafui [Kimaasai language], mhaki, mnavu, msogo, muhaka, ormomoi aves, soko, suga; Uganda: ensugga, enswiga/eshwiga/eswiga [Kigezi language], nouiga, nsuggaenzirugavu; Zimbabwe: m’sungu sungu-m’adzizambito.
Leaves eaten as a vegetable throughout the range; also used as a medicinal plant (in Karamoja Distr., Uganda).
(
Solanum tarderemotum is a morphologically variable species.
Solanum tarderemotum is distinguished from other sympatric species by its usually many-flowered inflorescences, long-exserted style that often is bent rather than perfectly straight (easier to see in live plants), spathulate calyx lobes that are rounded at apex, very strongly reflexed fruiting pedicels and berries that drop with the pedicel attached. Stems of S. tarderemotum are hollow and collapse upon drying, unlike the stems of the similar S. villosum that are more solid. Solanum scabrum also has hollow stems, but fewer-flowered inflorescences and larger fruits that drop without the pedicels. Solanum nigrum (North African populations), S. scabrum and S. villosum all lack stone cells, while the fruits of S. tarderemotum consistently have 2–6 stone cells in each berry. Most material of S. tarderemotum is either glabrous or has eglandular pubescence, but plants from higher elevations in the eastern African mountains are glandular pubescent; these were identified as S. pseudospinosum by
Previous molecular studies identified S. tarderemotum form C (Clade IV) as a separate entity based on AFLP data that was analysed using Maximum Parsimony phylogenetic method (
Solanum tarderemotum is a tetraploid species with unknown parentage (
Both names S. tarderemotum and S. florulentum were published in the same publication. No publication has yet synonymised the two. Most of the types for all names associated with our circumscription of this species were destroyed in Berlin. We have chosen to use the name S. tarderemotum because the holotype (WRSL) is still extant, while no known duplicates of the type of S. florulentum have been located.
No duplicate material has been found for the type collection of S. dasytrichum that was described from a specimen in B that is no longer extant;
Angola. Huila: Ha, juxta ripas rivi de Sopollo, Dec 1859, Welwitsch 6034 (BM, K); Uíge: Santa Cruz Mission, 12 Aug 1962, Codd 7521 (K).
Botswana. North-West: Ngami, Central Management Unit, Selinda Reserve, 8 Apr 2005, Heath & Heath 1023 (K); Burundi. Rwegura, Territoire Kayenza, 28 May 1969, Lewalle 3620 (K); Bugarama: Teza, 29 Dec 1978, Reekmans 7400 (EA, K, MO); Bujumbura: Katumba, 12 Dec 1979, Reekmans 8444 (MO); Muramvya: Nyabigondo, 2 Feb 1967, Lewalle 1536 (MO).
Cameroon. Nord-Ouest: Bui, Oku-Elak, 27 Oct 1996, Cheek et al. 8455 (K, MO); Boyo, Ijim Mountain Forest, 21 Nov 1996, Kamundi, et al. 673 (K, MO); Sud-Ouest: Mt Kupe, Nyasoso, 23 Oct 1995, Cheek et al. 7470 (K, MO); Mt. Cameroon R.C., 24 Mar 1961, Swarbrick SCA271 (E).
Cape Verde. Santiago: valley of Santo Domingo, 3 Nov 1839, Hooker 117 bis (K).
Chad. Lac: Koulfe, Chari Central, 28 Jun 1903, Chevalier 8788 (K, P); N’Djamena: c. 65 km S of Fort Lamy, 1 Jan 1965, de Wilde et al. 5118 (K).
Comoros. Iles Comores, Boivin s.n. (W); Mohilla Island, 5 Apr 1861, Meller s.n. (K); Anjouan: sin. loc, Jun 1875, Hildebrandt 1626[a] (BM); Mwali: Insul. Mohely, 1854, Boivin s.n. (BM).
Democratic Republic of the Congo. Katanga: a 3 km de Lukuni (Katanga), source de la Kasapa, 12 May 1961, Poelman 7 (K); Katuba, ferme Droogmums, Kaletele, Jan 1927, Quarré 18 (GH); Kivu Nord: Ngungu, 18 km SW Sake, 11 Aug 1954, Stauffer 48 (K, P); Kibati, Nord-Kivu: Butembo, 21 Feb 1974, Baudet 487 (K); NW slope of Mt. Vislke, 24 Feb 1975, D’Arcy 8089 (MO); Orientale: Ituri, Kibali, Aru, Aug 1931, Lebrun 3575 (GH, K); Sud Kivu: Lac Tsimuka, plaine de la Ruzizi, Jan 1950, Germain 5593 (K); Kalongé, Kalonge, long riviere Nyamwamba endroit fran et ombrage, 12 Feb 1953, de Witte 10477 (MO).
Eritrea. Semienawi Keyih Bahri: c. 10 km S of Nefasit, 2 Feb 1969, de Wilde 4502 (K).
Ethiopia. Amhara: South Gondar, 4 km N of Debre Tabor, 13 Sep 2004, Friis et al. 11552 (K); Semien Mountains, 1 Oct 2003, Wieringa 4971 (K); Dire Dawa: Dire Dawa, 16 Oct 1969, Parker 580 (K); Harari: between Harrar and Abbaba, Sep 1901, Wellby s.n. (K); Oromia: Gara Mullata, c. 50 km due W of Harar, 2 Aug 1962, Burger 2006 (FT, K); Hana, E part of the Omo, 23 Mar 1976, Fukui 15 (EA, K); c. 5 km N of Addis Ababa, 3 May 1965, de Wilde & de Wilde-Duyfjes 6507 (K, MO); Somali: Harerge, on the rd from Alemaya to Asbe Tafari, 6 km from Kobbo, 16 Aug 1967, Westphal & Westphal-Stevels 1227 (K); Southern Nations (SNNP): Lower Omo River, 7 Oct 1970, Carr 875 (EA, K); rd from Jimma to Serbo, 14 km from Jimma, 1 Aug 1968, Westphal & Westphal-Stevels 5499 (K, MO).
Ghana. Ohamu, Agric. Res. Station, Jul 1961, Irvine 4957 (K).
Guinea. Nzérékoré: Nzo, Mt Nimba, 5 Nov 1969, Adam 24671 (MO).
Kenya. Central: vicinity of Lake Naivasha, 17 Jul 1909, Mearns 842 (GH); Meru North, Mt. Kenya area, 18 Apr 2010, Vorontsova et al. 198 (BM, BR, EA, K, MO, NY); Coast: Mwanda, Mgange Nyika, 3 Oct 1971, Klungness 74 (K); Eastern: Makueni, Chyulu Hills, 15 May 1938, Bally 7787 (EA, K); Nairobi: Nairobi, 1924, McDonald 808 (K); nr Nairobi, Whyte s.n. (K); Rift Valley: Mt. Elgon, Kitale, 27 Dec 1960, Löffler E-107 (W); Kericho, Southwestern Mau Forest Reserve, camp 7, river Dimbilil, 7 Aug 1949, Maas Geesternanus 5603 (K, MO); Katilia forest, 12 mi NNE of Kangetet, Kerio River, 25 May 1970, Mathew 6379 (EA, K); Western: Kakamega, Kaptiki Secondary School, 11 Nov 1984, Hohl 157 (EA, W); pr. Forest Station ad mar, Mt Aberdare Expedition, 14 Jan 1922, Rob & Fries 921 (MO).
Lesotho. Sehlabathebe, 4 Jan 1973, Guillarmod et al. 138 (K).
Madagascar. Antananarivo: Ambohidratrimo, Ambohimanga, près de Tananarive, 19 Apr 1928, Decary 6179 (P); Antananarivo-Nord, Tananarive, Waterlot s.n. (P); Toliara: valley half a mi W of Ampoza, 5 Sep 1929, White s.n. (BM).
Malawi. Central: Dedza Distr., Dedza mountain forest, 19 Jan 1987, Balaka & Patel 1861 (MO); Northern: Rumphi Distr., Nykia Plateau, 2 mi E of Chelinda, 4 Mar 1977, Pawek 12428 (K, MO); Southern: Mt. Mulanje, foot NE slopes of Namasile opposite Sombani hut, 3 Jan 1971, Hilliard & Burtt 6134 (E).
Mali. Ackerrand kurz vor Koulikoro, 29 Sep 1992, Ehrich 336 (B).
Mozambique. Maputo: Lourengo Marques, Namaacha, 1 Aug 1967, Marques 2142 (MO); Zambezia: Namuli Mountain, Muretha Plateau, 26 May 2007, Harris 186 (K).
Namibia. Kavango West: Tondoro Mission, Tondoro Camp 1 km E of Mission, 15 Dec 1955, de Winter 3955 (K); Zambezi: E Caprivi, Zipfel, Lizazuli, 2 Jan 1959, Killick & Leistner 3253 (K).
Nigeria. Bauchi: Toro, Panshanu Pass, 15 Aug 1962, Lawlor & Hall 410 (K); Cross River: probably collected nr Obudu Cattle Range, 25 May 1971, Meer van, 1861 (MO); Enugu: Nsukka, University of Nigeria campus, 23 Jan 1962, Okigbo, 57 (K); Osun: Shasha Forest Reserve, 23 Apr 1968, Gledhill, 998 (K).
Rwanda. Northern: Mt. Visoke, 6 Apr 1970, Fossey 16 (EA, K).
Senegal. Nr. Bono, nec non Ins. Bonavista, Brunner s.n. (K); Dec 1823, Roger 17 (K).
Sierra Leone. Northern: Mt. Bintumani, Kabala (admin), Mt Loma, Mira, 30 Nov 1965, Adam 22258 (GH, MO).
Somalia. Banaadir: 3 km from Muqdisho airport along rd to Jasiira, 4 May 1990, Thulin & Hedrén 7172 (K); Togdheer: Wagga Mt., 1905, Bury s.n. (BM); Woqooyi Galbeed: Murak, Sep 1933, Godding 162 (K); mountains above Qoton, 11 Feb 2002, Thulin 10906 (K).
South Africa. Eastern Cape: Somerset, 1860, Cooper 528 (K); Fort Beaufort, 1860, Cooper 554 (K, W); Uitenhage, Enon, Drège s.n. (K); Gauteng: Pretoria, Region SWA, Kaokoveld, Otjomborombonga on Kunene River, 13 Jul 1976, Leistner et al. 104 (K, MO); KwaZulu-Natal: Weza, Ingeli, 5 Mar 1972, Strey 10899 (K, MO); Mpumalanga: Matebe Valley, May 1883, Holub s.n. (K); North West: nr Vryburg, 60 mi NW of Vryburg, Kalahari Desert, 5 Feb 1948, Rodin 3500 (K, MO).
South Sudan. Bahr El Ghazal: Isablei [?], Lande der Bongo, 26 Nov 1869, Schweinfurth 2649 (K); Equatoria: Imatong Mountains, Mt. Angargi, 14 Jun 1939, Andrews 1947 (K); Distr. Torit, Lowiliwili, Imatongs, 14 Nov 1949, Jackson 902 (BM); Greater Upper Nile: Upper Nile, Boing, 26 Oct 1951, Sherif A2891 (K).
Sudan. Blue Nile: Sennar, Würtemmburg s.n. (W); Darfur: Jebel Marra, Nyertete, 21 Jan 1964, Wickens 1044 (K); Jebel Marra, Zalingei, Wickens 1776 (K); Kassala: Erkowit, Red Sea Hills, Mar 1929, Lady Maffey 39 (K); Nr Kamobsana, Red Sea Prov., 25 Jan 1912, MacDougal & Sykes 137 (BM); Kamobsana, Red Sea Prov., 25 Jan 1912, MacDougal & Sykes 139a (BM). Kurdufan: ad Cordofanum Milbes, 4 Dec 1839, Kotschy 291 (BM, E, GH, K, P, W).
Tanzania. Arusha: Mbulmbul, Block DL, 49, 24 Jun 1944, Greenway 6951 (EA, K); Ngorogoro Crater, 24 Jun 1938, Pole Evans & Erens 936 A (E, K, P); Buha: Kakombe Valley, E shore of Tanganyika Lake from Gombe stream to Missonge, 25 Dec 1963, Pirozynski P86 (EA, K); Eastern: Morogoro, Lukwangulu Plateau, Uluguru Mts, 19 Sep 1970, Thulin & Mhoro 1048 (K); Iringa: Kidatu, Iringa District, E dam site, 28 Mar 1971, Mhoro 864 (EA, K); Kagera: Karagwe, 1 Mar 1862, Speke & Grant 453 (K); Kigoma: Mpanda, Sisaga, Mahali Mountains, 27 Aug 1958, Jefford et al. 1818 (K); Kakombe, 10 mi W of Kigoma, 7 Jul 1959, Newbould & Harley 4285 (K, MO); Kilimanjaro:, Chome Forest Reserve, Namboja, 30 Mar 2001, Mlangwa et al. 1506 (MO); Mbeya: Mbeya, 3 Mar 1932, Davies 461 (EA, K); Rungwe, Nyassa Hochland-Station Kyimbila, 1911, Stolz 384 (B, BM, K, MO, W); Mbeya/Njombe: Poroto Mountains, Mbeya Distr., 16 May 1957, Richards 9735 (EA, K); Mbulu: Lake Manyara National Park, 2 Dec 1963, Greenway & Kirrika 11113 (EA, K); Hanang Mt., 3 May 1962, Polhill & Paulo 2302 (EA, K); Morogoro: Kiberege, Mar 1936, Culwick 2 (K); Kanga Mountains, 11 May 2007, Luke et al. 12026 (EA, K, MO); Mwanza: Mwanza, Davis 201 (K); Njombe: Stromgebeit des obern Ruhudje, Landschaft Lupembe, nordlich des Flusses, Mar 1931, Schlieben 413 (K); Northern: Moshi, Weru-Weru gorge, 22 Feb 1955, Huxley 116 (EA, K); Rukwa: Nsanga Forest, Mpanda, Ufipa, 8 Aug 1960, Richards 13004 (K); Ruvuma: Songea, Matagoro Hills, just S of Songea, 22 Feb 1956, Milne-Redhead & Taylor 8869 (EA, K); Shinyanga: Shinyanga, Nov 1938, Koritschoner 1903 (EA, K); Tabora: Unyamwezi, Mininga, 1860, Speke & Grant 79 (K); Tanga: Usambaras, between Ngua and Magunga Estates, 17 Jul 1953, Drummond & Hemsley 3347 (EA, K); Ufipa: Sumbawanga, 3 km SE of Moravian Mission at Nkutwe nr Tatanda, 31 Oct 1992, Harder & Kayombo 1351 (EA, MO); Ulanga: Ifakara, 16 Jul 1959, Haerdi 286o (EA, K).
Uganda. Central: Mukono, Kipayo, Dec 1913, Dümmer 563 (BM, K, P); Mengo, 12 mi to Kampala, Entebbe rd, May 1932, Eggeling 697 (EA, K); Eastern: Mt. Elgon, 25 Jan 1993, Katende & Sheil 1099 (K); Northern: Karamoja, Moroto Township, Sep 1958, Wilson 609 (EA, K); Western: Kigezi D.F.I, 28 Aug 1972, Goode G-2-72 (K); Mihunga, Ruwenzori, 12 Jan 1939, Loveridge 350 (A, K, MO); Kigezi, Muhavura Hill, 11 Jan 1933, Rogers 337 (BM, EA, K).
Zambia. Lusaka: Mt. Makulu, Kafue Basin, 10 Apr 1963, van Rensberg 1886 (K); North-Western: Mwinilunga, 4 Oct 1937, Milne-Redhead 2565 (K); Northern: below Kwimbi Mission, 10 Feb 1955, Richards 4426 (K); Southern: Livingstone Distr., Victoria Falls, 21 Feb 1997, Luwiika et al. 460 (MO).
Zimbabwe. Harare: Salisbury, 26 Feb 1927, Eyles 4712 (K); Manicaland: Nr Chirinda, May 1906, Swynnerton 481 (BM, K); Mashonaland Central: Mazowe, Umvukwes, 17 Dec 1952, Wild 40771 (K); Mashonaland East: Distr. Salisbury, Mandara, 25 Sep 1974, Bisgel 4632 (MO); Marandella, 8 Apr 1948, Corby 1-20917 (K); Matabeleland South: Mazowe, Matobo National Park, Matopus NP, 24 Feb 1981, Philcox & Leppard 8823 (K).
Solanum triflorum var. majus Hook., Fl. Bor.-Amer. 2: 90. 1837, as “major”.
Type. Canada. Saskatchewan: “Carleton House Fort, Saskatchewan River”, J. Richardson s.n. (lectotype, designated here: BM [BM000934745]; isolectotype: K [K001159656, large plants]).
Solanum triflorum var. minus Hook., Fl. Bor.-Amer. 2: 90. 1837, as “minor”.
Type. Canada. Saskatchewan: “In the Garden (a weed) of Carleton House Fort, entrance of Badger’s Hole, and Saskatchewan River to Edmonton House [protologue]”, T. Drummond s.n. (lectotype, designated here: E [E00526685]; isolectotypes: BM [BM000934744], K [K001159656]).
Solanum mendocinum Phil., Anales Univ. Chile 21(2): 403. 1862.
Type. Argentina. Mendoza: Mendoza, 1860–1861, W. Díaz s.n. (lectotype, designated by
Solanum calophyllum Phil., Anales Univ. Chile 21(2): 403. 1862.
Type. Argentina. Mendoza: Mendoza, 1860–1861, R. Philippi s.n. (lectotype, designated here [cited as holotype in
Solanum pyrethrifolium Griseb., Abh. Königl. Ges. Wiss. Göttingen 24: 250. 1879.
Type. Argentina. Tucumán: Lules, Dec 1873, P. G. Lorentz & G. Hieronymus 1132 (lectotype, designated by
Solanum gaudichaudii var. pyrethrifolium (Griseb.) Kuntze, Revis. Gen. Pl. 3(3): 226. 1898.
Type. Based on Solanum pyrethrifolium Griseb.
Solanum triflorum var. calophyllum (Phil.) Bitter, Abh. Naturwiss. Vereine Bremen 23: 144. 1914.
Type. Based on Solanum calophyllum Phil.
Solanum triflorum var. pyrethrifolium (Griseb.) Bitter ex Probst, Mitteil. Naturfor. Gesellsch. Solothurn 9: 41. 1932.
Type. Based on Solanum pyrethrifolium Griseb.
Solanum ponticum Prodan, Bul. Fac. Agron. Cluj 7: 49. 1938.
Type. Romania. Satul noi, in arenosis, I.E. Nyárádu s.n. (no original material located; BUCA?, CLA?).
Solanum triflorum var. ponticum (Prodan) Borza, Bul. Grad. Bot. Univ. Cluj 22: 20. 1942.
Type. Based on Solanum ponticum Prodan
Solanum triflorum var. dentatum Ooststr., Gorteria 3: 90. 1966.
Type. Netherlands. Gelderland: Nijmegen, bij molen aan de Graafsweg, Jul 1938, B. Reichgelt & T.J. Reichgelt s.n. [Herb. Kern & Reichgelt 4610] (holotype: L [L3384989 [old barcode L0860668], acc. # 951.219 101]).
Solanum triflorum forma malvinum De Langhe & D’hose, Belg. J. Bot. 119(2): 160. 1987 [“1986”].
Type. Belgium. Antwerpen, 14 Oct 1984, J.-E. De Langhe & R. D’hose s.n. [219/84] (holotype: “in herbario De Langhe sub numero 219/84”, not seen, BR?).
United States of America. North Dakota: Nr Fort Mandan, Anon. [Lewis & Clark] s.n. (lectotype, designated by
Annual, decumbent and prostrate herbs to 40 cm tall, much branched at the base, to 70 cm in diameter. Stems decumbent to ascending, terete, green, forming adventitious roots at the nodes, not markedly hollow; new growth glabrous to sparsely pubescent with simple, spreading, uniseriate, translucent, eglandular trichomes, these (3-)4–10-celled, 0.5–2.0 mm long, occasionally with a few glandular trichomes with a 1-many-celled apical gland; older stems glabrescent. Sympodial units difoliate or trifoliate, the leaves not geminate. Leaves simple and shallowly lobed to deeply pinnatifid, (0.5-)2.0–4.0(-5.0) cm long, 0.2–2.9 cm wide, narrowly elliptic to oblong- to ovate-elliptic, fleshy in texture, concolorous, without smell; adaxial surface glabrous to sparsely pubescent with simple, uniseriate trichomes like those on stem, scattered along lamina and more densely along the veins; abaxial surface more densely pubescent on veins and lamina; major veins 3–6 pairs, not clearly evident abaxially; base cuneate, decurrent on the petiole; sinuate-lobate to deeply pinnatifid to near-pinnate, with 3–6 linear to triangular pairs of lobes; apex acute; petioles (0.5-)1.0–2.0(-2.4) cm long, pubescent with simple uniseriate trichomes like those of the stems. Inflorescences 1.0–2.0 cm long, internodal, simple, umbelliform to sub-umbelliform, with 1–5(-6) flowers clustered near the tip, glabrous to sparsely pubescent with spreading trichomes like those of the stems; peduncle 0.8–3.5 cm long, straight, often with apical leafy “bracteoles”; pedicels 3–12 mm long, 0.4–0.5 mm in diameter at the base and 0.4–0.5 mm at apex, straight and spreading, articulated at the base; pedicel scars spaced 0(-0.5) mm apart. Buds narrowly ellipsoid or occasionally narrowly ovoid, the corolla exserted 1/5–2/5 from the calyx tube before anthesis. Flowers 5-merous, all perfect. Calyx tube 1.0–1.5 mm long, conical, the lobes 2.5–3.5(-7) mm long, 0.8–1.0(-4) mm wide, triangular-oblong with acute apices, densely pubescent with simple, uniseriate eglandular trichomes like those of the stem. Corolla 10–14 mm in diameter, white to lilac with a yellow-green central eye with black-purple colouration at the base, stellate, lobed 1/2–1/3 to the base, the lobes 4.0–5.0 mm long, 1.8–2.2 mm wide, reflexed at anthesis, densely pubescent abaxially with short simple uniseriate eglandular trichomes like those on stems and leaves. Stamens equal; filament tube minute; free portion of the filaments 0.6–1.0 mm long, adaxially sparsely pubescent with tangled simple, uniseriate trichomes; anthers 2.8–3.1 mm long, 0.4–0.5 mm wide, narrowly ellipsoid, pale yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.5–3.5 mm long, densely pubescent with 2–3-celled simple uniseriate trichomes in the lower half, not exserted beyond the anther cone; stigma capitate, minutely papillate, green in live plants. Fruit a globose berry, 8–10(-20) mm in diameter, fleshy, dark green at maturity, the pericarp usually shiny; fruiting pedicels 12–17 mm long, 0.5–1.0 mm in diameter at the base, 1.0–1.5 mm at apex, spaced 0–0.5(-1.0) mm apart, reflexed and becoming woody, dropping with mature fruits, not persistent; fruiting calyx not markedly accrescent but the lobes slightly elongating in fruit, the tube less than 1 mm long, the lobes (4.0-)4.5–5.5(-8.0) mm long, strongly reflexed to spreading. Seeds 40–60 per berry, 2.0–2.5 mm long, 1.7–2.0 mm wide, subglobose, yellow, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells 13–30, 1.0–1.5 mm in diameter. Chromosome number: 2n=2x=24 (
(Figure
Grows across a wide range of habitats, along road sides, sandy soils, in cultivations and in salt plains; between sea level and 2,300 (-2,900) m elevation in its native range, between sea level and 1,800 m in the introduced range.
Australia: cut-leaf nightshade (
None recorded; a weed of agriculture.
(
Solanum triflorum is a weed of agricultural areas and appears not to spread aggressively based on collection numbers (see
In its native range, S. triflorum has a very large range of berry sizes; most introduced material has larger berries, but it is possible we have not seen material that has the smaller berry size from the Old World. There is also large variation in the indumentum and leaf shape in S. triflorum in its native range (
The varieties of S. triflorum described by
Australia. New South Wales: Queanbeyan, Queanbeyan Stockyards, 5 Feb 1931, Calvert KEW-84 (CANB, PERTH); Murrumbidgee River, Dew’s Corner 4 km S of Fairvale Homestead, 8 May 1987, Canning & Lloyd 6421 (AD, CANB, NSW); Gundaroo Creek, 4.7 km N of Gundaroo on rd to Gunning, 23 Mar 1991, Lepschi 542 (AD, CANB, K); Fyshwick, Australian Capital Territory, 7 Apr 1991, Lepschi 555 (CANB, HO); Mid-Western Regional, Bylong district, 11 Oct 1953, McKee 654 (MEL); Armidale Dumaresq, Armidale, 10 Jan 1973, Williams s.n. (NE); Queensland: Southern Downs, Stanthorpe, 24 Feb 1954, Alison s.n. (BRI); Southern Downs, Ballandean, 23 Jan 1974, Booth s.n. (BRI); Southern Downs, 8 km of Ballandean, 20 Feb 1974, Swann s.n. (BRI); Southern Downs, Wallangarra, 22 Dec 1958, Taylor s.n. (BRI); South Australia: Region 9, Murray, 28 May 1972, Alcock 3939 (AD, CORD); Culburra, c. 80 km NW of Bordertown, 12 Feb 1963, Fry s.n. (E); Keith, 5 Apr 1965, Kain s.n. (K); NSW Finley, 30 Jan 1975, Symon 9807 (AD, MO, NSW); Murray Bridge, northern fenceline of Monarto Conservation Park, 6 Feb 1998, Taylor 291a (AD, MEL); Tasmania: Pitt Water, Seven Mile Beach Protected Area, 31 Mar 2006, Baker 1707 (CANB, HO, MEL); Clarence, Seven Mile Beach, 3 Apr 2000, Buchanan 15695 (AD, HO); Victoria: East Gippsland, 6 km NW of Mt Deddick, 17 Apr 1981, Forbes & Rees 909 (AD, MEL); Banyule, nr Montpelier Billabong, 7 Mar 2011, Lynch s.n. (CANB, HO, MEL); Western Australia: Wickepin, 17 Jan 1949, Eccles s.n. (PERTH)Beverley, 25 km E of Brookton, Hambly s.n. (PERTH); Steerdale, John Forrest rd, Hopetoun, 3 Dec 1997, Hill s.n. (PERTH); Swan, Midland, 12 mi E of Perth, Jan 1968, Moir s.n. (AD); Brookton, Williams & Williams s.n. (PERTH).
Austria. Wien: Wien, Müllplatz Laaerberg, 1 Sep 1969, Forstner & Melzer s.n. (W).
Belgium. Flanders: De Panne, 15 Aug 1976, Goetghebeur 2605 (H, MO); Gent, 7 Aug 1983, Goetghebeur & Pauw De 5252 (H, MO, UT); Westhoek, 15 Aug 1976, Goetghebeur 8626 (W); Antwerp, Linkeroever, Middenvijver, 22 Sep 2009, Wieringa 6785 (BM, W).
France. Hauts-de-France: Pas-de-Calais, Camiers, au NW d’Etaples, chemin conduissant a la plage Saint Gabriel, 30 Oct 1973, Lambinon et al. 1461 (BM, H).
Germany. Nordrhein-Westfalen: Niederrhein, Emmerich, 20 Oct 1929, Krüger 700 (W).
Morocco. Oriental: Oued Charef, au nord de Ain-Benimathar, 21 Jun 2008, Calvo et al. 2501 (MA).
Netherlands. Zuid Holland: Isle of Rosenburgh, De Beer, 26 Aug 1948, van Hattum 4073 (MO); Isle of Rozenburg, De Beer, 26 Aug 1948, van Hattum 4073 (K); Goeree, Havenhoofd, 13 Aug 1977, Podlech 8624 (BM, H, MO).
Romania. Moldova, Distr. Covurlui, stationem viae ferreae Fulgeresti, 6 Sep 1922, Petruscu & Borza 2465 (W).
South Africa. Eastern Cape: Morgendal, 10 Feb 1960, Acocks 21049 (EA).
Sweden. Götaland: Skåne, Malmö, Sep 1906, Hylmö s.n. (BM).
Tunisia. Gabes: Gabes, Djebel Dirra, Apr 1909, Pitard s.n. (MA).
United Kingdom. England: nr Hartley, Wintney Hants, 17 Aug 1949, Hodge 4288 (K); Essex, Barking, 17 Oct 1953, Lousley s.n. (BM); Northumberland, Holy Island, 5 Sep 1936, Hutton s.n. (K); Northumberland, Holy Island, 13 Sep 1955, Lousley s.n. (BM); Suffolk, Woodbridge, 14 Sep 1932, Milne-Redhead & Airy Shaw 1834 (K); Scotland: Possil, Glasgow, 11 Sep 1918, Grierson 955-18 (K); Wales: Gwynedd, Caernarfonshire, Dinas Dinlle, Herrington s.n. (BM); Cardiff, Cardiff, 22 Aug 1925, Smith s.n. (BM).
Tanzania. Mbeya: Mbeya District, Umalila Forest Reserve, ca. 7 km W of Ruanda II on road to Izumbwe, 2 km SSE of Mbogo Mtn. main peak, 9°11’S, 33°18’E, 2180 m, 14 Nov 1992. R.E. Gereau, D.K. Harder, C.J. Kayombo & M.J. Kayombo 5084 (holotype: DSM; isotypes: EA, K [K001081869], MO [MO-2293667, acc. # 4281580], NHT).
Annual to short-lived erect perennial herbs to 0.5 m tall, erect, predominantly branching from the base. Stems spreading, ridged or winged, dark purple to green, erect, not markedly hollow; new growth puberulent with simple, spreading, uniseriate, translucent, eglandular trichomes, these 2–8-celled, 0.3–0.8 mm long; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.8–2.8 cm long, 0.1–1.7 (-2.1) cm wide, ovate to elliptic, membranous, green to dark green above and below, without smell; adaxial and abaxial surfaces glabrous or sparsely pubescent with simple uniseriate trichomes like those on the stem mainly along veins; major veins 3–4 pairs, paler green or often purple tinged; base truncate, narrowly winged on to the petiole; margins entire; apex acuminate to acute; petioles 0.6–1.3 cm long, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the stem. Inflorescences 2–6 cm long, internodal, simple or usually branched and often leafy, the flowers spaced along the rhachis, with 2–9(-20) flowers, glabrous or sparsely pubescent with simple uniseriate trichomes like those on the stem; peduncle (0.5-)1.1–4 cm long, erect, ca. 0.5 mm thick at apex, stout; pedicels 5–7(-9) mm long, ca. 0.2 mm in diameter at the base, ca. 0.5 mm in diameter at the apex, stout, pendent, articulated at the base; pedicel scars widely spaced (1-)3–8 mm apart. Buds subglobose, the corolla exserted 1/2 from the calyx tube before anthesis. Flowers 5-merous, perfect. Calyx tube 0.8–1.0 mm long, cup-shaped, the lobes equal, 0.1–0.5 mm long, 0.5–0.7 mm wide, broadly deltate with an acute tip, green or purple-tinged, glabrous or sparsely pubescent with simple uniseriate trichomes like those of the pedicels. Corolla 7–11 mm in diameter, white, margins occasionally tinged with purple, with a yellow basal star, stellate, lobed ca. 1/2 way to the base, the lobes 2.5–3.5 mm long, 2–3 mm wide, spreading or reflexed, densely papillate abaxially. Stamens equal; filament tube to 0.1 mm long; free portion of the filaments 0.5–1.0 mm long, pubescent adaxially with tangled uniseriate simple trichomes; anthers 1.8–2 mm long, 0.5–0.6 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary rounded, glabrous; style 3.1–3.5(-4.2) mm long, densely pubescent with simple uniseriate trichomes 0.2–0.5 mm long in the basal 1/2–2/3, exserted 1.0–1.5 mm beyond anther cone; stigma clavate, the surface minutely papillate. Fruit a globose berry, 3–4(-5) mm in diameter, pale yellow-brown at maturity, aromatic, the pericarp soft, matte, not transparent; fruiting pedicels 0.8–1.3 cm long, 0.3–0.5 mm in diameter at the base, 0.5–1.0 mm in diameter at the apex, stout and becoming woody, erect or spreading, pale brown, falling with the mature fruit, not persistent; fruiting calyx not accrescent, the tube ca. 1 mm long, the lobes ca. 1 mm long, appressed to the berry. Seeds (5-)6–12 per berry, 1.6–2.1 mm long, 1.3–1.8 mm wide, flattened and tear-drop shaped with a subapical hilum, brownish, the surfaces minutely pitted, thin, the testal cells rectangular to pentagonal in outline. Stone cells 9–22 per berry, ca. 0.5 mm in diameter. Chromosome number: 2n=4x=48 (
Grows on volcanic soils, frequent on the ash layer in charcoal-burning areas, also commonly cultivated; between 1,700 and 2,200 m elevation.
Tanzania: insungwe [Malila people] (
Leaves used as spinach; berries eaten raw.
(
Solanum umalilaense can be distinguished from other African species of morelloids by its simple to branched and often leafy inflorescences, flowers with very short rounded calyx lobes and persistent, small, light yellowish-brown fruits. Solanum umalilaense produces a large number of inflorescences such that, at full anthesis, the plant appears to be covered with white flowers, strikingly different from other species in the section (
More field work is needed in southern Tanzania and adjacent Zambia to determine the true range and morphological variation of S. umalilaense; collections analysed to date come from what might be a single population.
Specimens examined. Tanzania. Iringa: Mufindi, Igowole, 10 Mar 1989, Kayombo & Kayombo 215 (MO); Mbeya: Maganjo village, Lwindi ward, 8 Jul 2010, Manoko 2010-1 (DSM, NIJ, WAG); Isangati village, Iyunga ward, 8 Jul 2010, Manoko 2010-2 (DSM, NIJ, WAG); Igala village, Holondo ward, 8 Jul 2010, Manoko 2010-8 (DSM, NIJ, WAG); Isangati village, Iyunga Mapinduzi ward, 9 Jul 2010, Manoko 2010-11 (DSM, NIJ, WAG); Isangati village, Iyunga Mapinduzi ward, 9 Jul 2010, Manoko 2010-12 (DSM, NIJ, WAG); nr Umalila Forest reserve, 9 Jul 2010, Manoko 2010-14 (DSM, NIJ, WAG); Uyole, Mbeya, 22 May 1968, Mwambunga 6 (DSM).
Solanum nigrum var. villosum L., Sp. Pl. 186. 1753.
Type. “Solanum annuum hirsutus, baccis luteis”, cultivated in England, at James Sherard’s garden in Eltham (Hortus Elthamensis) (lectotype, designated by
Solanum luteum Mill., Gard. Dict. ed. 8, no. 3. 1768.
Type. “Solanumofficinarum lacinis luteis 1735, G.B. 166”, cultivated in England at the Chelsea Physic Garden, seeds said to come from America, Herb. Miller s.n. [691 on sheet] (lectotype, designated by
Solanum rubrum Mill., Gard. Dict. ed. 8, no. 4. 1768.
Type. Cultivated in England at the Chelsea Physic Garden, origin said to be the West Indies, Herb. Miller s.n. (lectotype, designated by
Solanum nigrum subsp. villosum (L.) Ehrh., Hannover. Mag. 1780, 14: 218. 1780.
Type. Based on Solanum nigrum var. villosum L.
Solanum villosum (L.) Willd., Prodr. Fl. Berol.: 87. 1787, nom. illeg., not Solanum villosum Mill. (1768)
Type. Based on Solanum nigrum var. villosum L.
Solanum nigrum var. rubrum (Mill.) Aiton, Hort. Kew. [W.Aiton] 1: 234. 1789.
Type. Based on Solanum rubrum Mill.
Solanum aegyptiacum J.F.Gmel., Syst. Nat., ed. 13 [bis] 2: 385. 1791.
Type. Egypt. Cairo: “circa Cairo”, 1762, Herb. P. Forsskål 420 (lectotype, designated here: C [C10013600]).
Solanum villosum Lam., Tabl. Encycl. 2: 18. 1794, nom. illeg., not Solanum villosum Mill. (1768)
Type. “Ex Gallia australi, &c.”, Anon. s.n. (lectotype, designated by
Solanum villosum Moench, Methodus (Moench) 474. 1794, nom. illeg., not Solanum villosum Mill. (1768).
Type. Germany. Hessen [?], C. Moench s.n. (no original material located).
Solanum alatum Moench, Methodus (Moench) 474. 1794, nom. cons. prop
Type. Pakistan. Balochistan: Quetta, 30 km E Gumbaz, 1050 m, 30°02’N, 69°00’E, 17 May 1965, K.H. Rechinger 29684 (type cons. prop. by
Solanum villosum (L.) Willd., Enum. Pl. (Willdenow) 236. 1809, nom. illeg., not Solanum villosum Mill. (1768).
Type. Based on Solanum nigrum var. villosum L.
Solanum humile Bernh. ex Willd., Enum. Pl. (Willdenow) 236. 1809.
Type. Cultivated in Berlin Botanic Garden, from “Europa australi” [southern Europe], Anon. s.n. (neotype, designated by
Solanum suffruticosum Schousb. ex Willd., Enum. Pl. (Willdenow) 236. 1809.
Type. Cultivated in Berlin Botanic Garden, said to be originally from “Barbaria” [northern Africa west of Egypt], Herb. Willdenow s.n. (lectotype, designated here: B [B-W04363010]).
Solanum incertum Dunal, Hist. Nat. Solanum 155. 1813.
Type. “Nelen-Tsjunda” from Malabar [western India] (lectotype, designated here: Rheede von Draakestein, Hortus Malabaricus 10: t. 73. 1690).
Solanum erythraeum Dunal, Hist. Nat. Solanum 238. 1813, nom. illeg. superfl.
Type. Based on Solanum rubrum L. (as S. rubrum “Murr. Syst. Veg 183, Gmel. Syst. Nat 384”)
Solanum kitaibelii Schult., Öster. Fl., ed. 2, 1: 395. 1814.
Type. Hungary. Sin. loc. “Pannonia”, Without collector [P. Kitaibel] s.n. [349] (lectotype, designated by
Solanum flavum Kit. ex Schult., Öster. Fl., ed. 2, 1: 394. 1814.
Type. Hungary. Sin. loc. [“Pannonia”], Without collector [P. Kitaibel] s.n. (lectotype, designated by
Solanum ochroleucum Bastard, J. Bot. (Desvaux) 3: 20. 1814.
Type. France. Pays de la Loire: Maine-et-Loire, Anjou [Angers], 1813, Without collector [prob. T. Bastard] s.n. (lectotype, designated here: G-DC [G00144474]; isolectotype: ANG [acc. # 2006.213]).
Solanum nigrum var. miniatum (Bernh. ex Willd.) Fr., Nov. Flor. Suec.: 111. 1823.
Type. Based on Solanum miniatum Bernh. ex Willd.
Solanum villosum var. miniatum (Bernh. ex Willd.) Kostel., Clav. Anal. Fl. Bohem. 34. 1824.
Type. Based on Solanum miniatum Bernh. ex Willd.
Solanum luteovirescens C.C.Gmel., Fl. Bad. 4: 177. 1826, nom. illegit. superfl.
Type. Based on S. humile Bernh. ex Willd. (cited in synonymy).
Solanum puniceum C.C.Gmel., Fl. Bad. 4: 176. 1826, nom. illegit. superfl.
Type. Based on S. miniatum Bernh. ex Willd. (cited in synonymy).
Solanum vulgatum var. luteum Spenn., Fl. Friburg. 2: 427. 1829, nom. illeg. superfl.
Type. Based on Solanum nigrum var villosum L. (cited in synonymy)
Solanum nigrum subvar. ochroleucum (Bastard) Coss. & Germ., Intr. Fl. Anal. et Descr. Paris, 91. 1842.
Type. Based on Solanum ochroleucum Bastard
Solanum villosum subvar. miniatum (Bernh. ex Willd.) Germ. & Wedd., Introd. Fl. Anales Paris 91. 1842.
Type. Based on Solanum miniatum Bernh. ex Willd.
Solanum villosum forma miniatum (Bernh. ex Willd.) Döll, Rhein. Flor. 413. 1843.
Type. Based on Solanum miniatum Bernh. ex Willd.
Solanum nigrum forma humile (Bernh. ex Willd.) Döll, Rhein. Flor. 413. 1843.
Type. Based on Solanum humile Bernh. ex Willd.
Solanum nigrum forma villosum (Lam.) Döll, Rhein. Flor. 413. 1843.
Type. Based on Solanum villosum Lam.
Solanum nigrum forma angulosum Döll, Rhein. Flor. 413. 1843.
Type. Germany. “Bei Carlsruhe beim Linkenheimer Thore, A. Braun” (no specimens cited; no original material located, not found at KR); Germany. “Bei Philippsburg, Stud. Bauer” (no specimens cited; no original material located, not found at KR): Germany. “Bei Mannheim, Oppenheim, J.C. Döll” (no specimens cited; no original material located, not found at KR).
Solanum nigrum subsp. miniatum (Bernh. ex Willd.) Hartm., Sv. Norsk Exc.-Fl.: 35. 1846.
Type. Based on Solanum miniatum Bernh. ex Willd.
Solanum nigrum subsp. villosum (Lam.) Hartm., Sv. Norsk Esc.-Fl. 34. 1846.
Type. Based on Solanum villosum Lam.
Solanum nigrum subsp. humile (Bernh. ex Willd.) Hartm., Sv. Norsk Exc.-Fl. 34. 1846.
Type. Based on Solanum humile Bernh. ex Willd.
Solanum sinaicum Boiss., Diagn. Pl. Orient. ser. 1, 11: 135. 1849.
Type. Egypt. [Sinai] “Arabia petraea, Wadi Sheikh” [protologue “in fissuris rupium jugi Sianaitici in valle Wadi Scheick”], Mar 1846, E. Boissier s.n. (lectotype, designated here: G [G00343377] isolectotypes: DD, K [K000759386]).
Solanum plebeium A.Rich., Tent. Fl. Abyss. 2: 100. 1850.
Type. Ethiopia? [“Abyssinia”], Prov. Chiré [Shire?], R.Q. Dillon & A. Petit s.n. (lectotype, designated by
Solanum nigrum subsp. luteovirescens Kirschl., Fl. Alsace 1: 532. 1852.
Type. Based on Solanum luteovirescens C.C.Gmel. (nom. illeg.)
Solanum nigrum subsp. luteum (Mill.) Kirschl., Fl. Alsace 1: 532. 1852.
Type. Based on Solanum luteum Mill.
Solanum nigrum subsp. puniceum Kirschl., Fl. Alsace 1: 532. 1852.
Type. Based on Solanum puniceum C.C.Gmel. (nom. illeg.)
Solanum ochroleucum var. flavum Dunal, Prodr. [A. P. de Candolle] 13(1): 56. 1852.
Type. Greece. Peloponnese: “in vineis agri Argolidis” [Argolis], J. Sartori 86 (lectotype, designated by
Solanum miniatum var. stenopetalum Dunal, Prodr. [A. P. de Candolle] 13(1): 56. 1852.
Type. France. Auvergne-Rhône-Alpes: Isère, “circa Montaud prope Salon”, J.L. Castagne s.n. (lectotype, designated here: G-DC [G00144520]; isolectotype: MPU not seen).
Solanum miniatum var. villosissimum Dunal, Prodr. [A. P. de Candolle] 13(1): 56. 1852.
Type. France. Provence-Alpes-Côte d’Azur: Var, Miraval, a l’entre de la grotte, 8 May 1807, A.P. de Candolle s.n. (lectotype, designated here: G-DC [G00144520]).
Solanum miniatum var. subserratum Dunal, Prodr. [A. P. de Candolle] 13(1): 57. 1852.
Type. “Arabie”, Without collector[No. 26] (lectotype, designated here: MPU [MPU091575]).
Solanum minutiflorum Dunal, Prodr. [A. P. de Candolle] 13(1): 58. 1852.
Type. France. Hyères, A. Richard s.n. (lectotype, designated here: G-DC [G00144587]; isolectotype: P [P00582961]).
Solanum nilagiricum Schltdl., Linnaea 26: 484. 1855.
Type. Cultivated in Germany at Halle [from Nilgiri Hills, India?] (no specimens cited; no specimens located [HAL ? not seen]).
Solanum suffruticosum Schousb. ex Lange, Linnaea 28: 361. 1856, nom. illeg., not Solanum suffruticosum Schousb. ex Willd. (1809)
Type. Cultivated in Denmark at Copenhagen, seeds from “Pontevedra Galleciae Aug 1852 [Galicia, Spain], 1855, Anon. s.n. (lectotype, designated here: C [C10019275]).
Solanum nigrum var. croceum Neilr., Fl. Nied.-Oesterr.: 535. 1858, nom. illeg. superfl.
Type. Based on Solanum nigrum var. villosum L.
Solanum nigrum var. miniatum (Bernh. ex Willd.) Neilr., Fl. Nied.-Oesterr. 535. 1858, nom. illeg., not Solanum nigrum var. miniatum (Bernh. ex Willd.) Fr. (1823)
Type. Based on Solanum miniatum Bernh. ex Willd.
S. nigrum var. luteum Neilr., Fl. Nied.-Oesterr. 535. 1858.
Type. Based on Solanum flavum Kit. ex Schult.
Solanum nigrum var. suffruticosum (Schousb. ex Willd.) Moris, Fl. Sardoa 3: 148. 1858–1859.
Type. Based on Solanum suffruticosum Schousb. ex Willd. (cites Solanum suffruticosum Schousb. concept of Dunal)
Solanum nigrum var. luteum Döll, Fl. Baden [Döll] 767. 1859, nom. illeg., not Solanum nigrum var. luteum Neilr. (1858).
Type. Germany. “in der Carlsruher Gegend bei Dachslanden und Mühlburg, bei Schwetzingen, Mannheim und Dossenheim” (no specimens cited; no original material located at KR).
Solanum
canescens
Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg., not Solanum canescens
Type. Hungary. “E Cottu Thurócz. v Barodiensi” [protologue “in Cottu Thurociensi et Borsodiensi”, P. Kitaibel s.n. (lectotype, designated here: BP [herb. Kit. fasc. IX: 105, numbered sheet]; isolectotypes: BP [unlabelled sheets in fasc. IX: 105]).
Solanum flavum Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg, not Solanum flavum Kit. ex Schult.(1814)
Type. Hungary. “E Cottu Thurócz. v Barodiensi” [protologue “in Cottu Thurociensi et Borsodiensi”, P. Kitaibel s.n. (lectotype, designated here: BP [BP [herb. Kit. fasc. IX: 105, numbered sheet]; isolectotypes: BP [unlabelled sheets in fasc. IX: 105]).
Solanum incanum Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg. superfl., also not S. incanum L. (1753)
Type. Based on Solanum kitaibelii Kit. ex Schult. (cited in synonymy).
Solanum olivaceum Kit. ex Kanitz, Linnaea 32: 440. 1863, nom. illeg. superfl.
Type. Based on Solanum humile Bernh. ex Willd. (cited in synonymy).
Solanum pseudovillosum Schur, Enum. Pl. Transsilv. 477. 1866.
Type. Transylvania [Romania], Hermannstadt [Sibiu] and Kronstadt [Brașov], Aug-Sep, P.J.F. Schur s.n. (no herbaria cited; no original material located; LW?, BRNU?).
Solanum nigrum subsp. alatum (Moench) Čelak., Prodr. Fl. Böhmen [2:?] 309. 1871.
Type. Based on Solanum alatum Moench
Solanum nigrum subsp. villosum (Lam.) Čelak., Prodr. Fl. Böhmen 309. 1871.
Type. Based on Solanum villosum Lam.
Solanum patens Lowe, Man. Fl. Madeira 2: 74. 1872.
Type. Portugal. Madeira: Ribeiria do Sta. Luzia, 1 Jul 1833, R.T. Lowe 547 [a] (lectotype, designated by
Solanum villosum var. velutinum Lowe, Man. Fl. Madeira 2: 76. 1872, as “velutina”
Type. Portugal. Madeira: Valle [garden], 24 Aug 1831, R.T. Lowe [722] (lectotype, designated here: BM [BM000056199]).
Solanum villosum var. laevigatum Lowe, Man. Fl. Madeira 2: 78. 1872, as “laevigata”
Type. Portugal. “Lisbon, court of the Museum [Jan 1865]”, R.T. Lowe s.n. (lectotype, designated by
Solanum hildebrandtii A.Braun & C.D.Bouché, Ind. Sem. Hort. Berol. 18. 1874.
Type. Somalia. Saanag: Yafir, Ahlgebirge [Karin Yafir, Ahl Mountains, N Somalia], J.M. Hildebrandt 865 (neotype, designated by
Solanum nigrum var. humile (Bernh. ex Willd.) Boiss., Fl. Orient. 4: 284. 1879.
Type. Based on Solanum humile Bernh. ex Willd.
Solanum nigrum var. induratum Boiss., Fl. Orient. 4: 284. 1879, nom. illeg. superfl.
Type. Based on Solanum nigrum var. suffruticosum (Schousb. ex Willd.) Moris (cited in synonymy)
Solanum villosum subsp. ochroleucum (Bastard) Nyman, Consp. Fl. Eur. 3: 526. 1881.
Type. Based on Solanum ochroleucum Bastard
Solanum nigrum subsp. miniatum (Bernh. ex Willd.) Arcang., Comp. Fl. Ital. (Arcangeli): 497. 1882.
Type. Based on Solanum miniatum Bernh. ex Willd.
Solanum morella subsp. luteum (Mill.) Rouy, Fl. France 10: 366. 1908.
Type. Based on Solanum luteum Mill.
Solanum plebeium var. brachysepalum Bitter, Bot. Jahrb. Syst. 49: 564. 1913.
Type. Ethiopia. Oromia: “Gallahochland, Ego”, 2000 m, H. Ellenbeck 358 (holotype: B, destroyed; no duplicates located).
Solanum plebeium var. subtile Bitter, Bot. Jahrb. Syst. 49: 564. 1913.
Type. Ethiopia. Harari: “Harar, Hararmaja-See”, H. Ellenbeck 462 (holotype: B, destroyed; no duplicates located).
Solanum nigrum forma glabrum Chiov., Res. Sci. Somalia Ital. 128. 1916.
Type. Somalia. Baay: Berdale [Berdaale], 12 Oct 1913, G. Paoli 935 (holotype: FT [FT003070]).
Solanum nigrum forma humile (Bernh. ex Willd.) Lindm., Svensk Fan. Fl. 480. 1918.
Type. Based on Solanum humile Bernh. ex Willd.
Solanum nigrum var. sancti-marini Pamp., Boll. Mus. Republ. San Marino 4(3–4): 116. 1920.
Type. San Marino. Sin. loc., Without collector s.n. (no specimens cited; possibly at FI?; no original material located).
Solanum nigrum var. alatum (Moench) Fiori, Nuov. Fl. Italia 2: 311. 1926.
Type. Based on Solanum alatum Moench
Solanum nigrum var. aurantium Maire, Bull. Soc. Sci. Nat. Maroc 8(4–6): 139. 1928.
Type. Morocco. Souss-Massa: Anti-Atlas Mountains “Djebel Inter à Talaïnt” [Djebel Inter à Talguit (=Tafraoute?)], 600 m, 4 Apr 1922. R. Maire s.n. (lectotype, designated here: RAB [RAB-44657]; isolectotypes: MPU [MPU001957], P [P00489756]).
Solanum villosum var. miniatibaccatum Blom, Act. Hort. Gothoburg. 10: 207. 1935.
Type. Sweden. Göteborgstrakten: Göteborg, som ogräs i Botaniska Trädgården, 1935, C. Blom s.n. (lectotype, designated here: GB; isolectotypes: LD, S, UPS).
Solanum luteum subsp. alatum (Moench) Dostál, Květena Č.S.R. 1270. 1949.
Type. Based on Solanum alatum Moench
Solanum transcaucasicum Pojark., Bot. Mater. Gerb. Inst. Komorova Akad. Nauk S.S.S.R. 17: 332. 1955.
Type. Azerbaijan. Lankaran: Talysh, near Tatuni “Talysh bei Tatuni”, R.F. Hohenacker s.n. (holotype: LE).
Solanum olgae Pojark., Bot. Mater. Gerb. Inst. Komorova Akad. Nauk S.S.S.R. 17: 333. 1955.
Type. Uzbekistan. Samarkandaya Oblast’, [Tadzkikistan, Hissar, near Kschtut in protologue] Kschtut, on the road to Sherbut, Fedotov & Gol’bek s.n. (holotype: LE; isotype: LE).
Solanum zelenetzkii Pojark., Bot. Mater. Gerb. Inst. Komorova Akad. Nauk S.S.S.R. 17: 336. 1955.
Type. Ukraine: Crimea, “Tauria Australis”, Jalta [Yalta in protologue], Oreanda, 21 Aug 1906, K. Golde s.n. (holotype: LE).
Solanum woronowii Pojark., Bot. Mater. Gerb. Inst. Komorova Akad. Nauk S.S.S.R. 17: 337. 1955.
Type. Georgia. Abkhazia AR: “ Abchazia [Abkhazia], Gagry”, 30 Oct 1954, A. Pojarkova 6 (holotype: LE).
Solanum luteum var. calvum Wessely, Feddes Repert. Spec. Nov. Regni. Veg. 63: 315. 1960.
Type. Cultivated in Germany at Greifswald, seeds from Hort. Bot. Marseille, I. Wessely 214 (holotype: GFW).
Solanum luteum var. procerum Wessely, Feddes Repert. Spec. Nov. Regni. Veg. 63: 315. 1960.
Type. Cultivated in Germany at Greifswald, seeds from München Botanical Garden, 1957, I. Wessely 320 (holotype: GFW).
Solanum luteum forma miniatibaccatum (Blom) Wessely, Feddes Repert. Spec. Nov. Regni. Veg. 63: 315. 1960.
Type. Based on Solanum villosum var. miniatibaccatum Blom
Solanum purpureilineatum Sabnis & Bhatt, Bull. Bot. Surv. India 12(1–4) 258. 1972 [“1970”].
Type. India. Gujarat: Baroda District, Baroda, L.V. Palace compound, 2 Oct 1960, S.D. Sabnis 2762, 2763 (holotype: BARO [two sheet holotype, single gathering).
Solanum carmanicum Schönb.-Tem., Fl. Iranica 100: 8. 1972.
Type. Iran. Kerman: Bam, ca. 900 m, 7 May 1948, K.H. Rechinger & F. Rechinger 3630 (holotype: W [1958-0003176]; isotype: E [E00251353], G [G00343074], US [00992058, acc. # 2316288]).
Solanum villosum subsp. alatum (Moench) Edmonds, Bot. J. Linn. Soc. 75: 171. 1977.
Type. Based on Solanum alatum Moench
Solanum nigrum var. humile (Bernh. ex Willd.) C.Y. Wu & S.C.Huang, Acta Phytotax. Sin. 16(2): 72. 1978, nom. illeg., not Solanum nigrum var. humile (Bernh. ex Willd.) Boiss. (1879).
Type. Based on Solanum humile Bernh. ex Willd.
Solanum villosum subsp. puniceum (Kirschl.) Edmonds, Bot. J. Linn. Soc. 78: 215. 1979.
Type. Based on Solanum nigrum subsp. puniceum Kirschl.
Solanum villosum subsp. miniatum (Bernh. ex Willd.) Edmonds, Bot. J. Linn. Soc. 89: 166. 1984.
Type. Based on Solanum miniatum Bernh. ex Willd.
Solanum
villosum
subsp.
alatum
(Moench) Dostál, Folia Mus. Rerum Nat. Bohemiae Occid., Bot. 21: 11. 1984, nom. illeg., not Solanum villosum subsp. alatum (Moench)
Type. Based on Solanum alatum Moench
Solanum luteum var. miniatibaccatum (Blom) Hyl., Uppsala Univ. Årsskr. 7: 279. 1945.
Type. Based on Solanum villosum var. miniatibaccatum Blom
Cultivated in Chelsea Physic Garden from “Barbadoes where it is supposed to grow naturally”, Herb. Miller s.n. (lectotype, designated by
Annual to short lived erect or weakly scrambling perennial herbs up to 0.5 m tall, subwoody at base and much branched. Stems spreading to decumbent, terete to ridged, green to purple, not hollow; new growth densely pubescent with simple, spreading, uniseriate, translucent, eglandular and/or glandular trichomes, these 3–10-celled, 0.2–2.0 mm long; older stems glabrescent. Sympodial units difoliate, the leaves not geminate. Leaves simple, 1.5–5.0(-10.0) cm long, 0.7–2.5(-6.5) cm wide, broadly to narrowly ovate to elliptic, membranous, green on both sides, without smell or somewhat pleasant smelling; adaxial surfaces sparsely to densely pubescent with spreading, simple, uniseriate eglandular and/or glandular trichomes like those on stem evenly along veins and lamina; abaxial surfaces more densely pubescent on veins and lamina; major veins 4–6 pairs; base acute to truncate, short-attenuate, often asymmetric; margins sinuate-dentate to rarely entire; apex acute; petioles 0.5–3.0(-4.5) cm long, pubescent with simple uniseriate glandular and/or eglandular trichomes like those on stems. Inflorescences 0.4–2.0 cm long, internodal, simple, the flowers spaced along the rhachis, with (2-)3–5(-8) flowers clustered at the tip or more commonly spaced along the rhachis, pubescent with spreading simple glandular and/or eglandular uniseriate trichomes like those of the stems; peduncle 0.4–1.5 cm long, straight; pedicels 4–7 mm long, 0.2–0.3 mm in diameter at the base and 0.4–0.5 mm at apex, spreading, articulated at the base; pedicel scars spaced 0–1.0 mm apart. Buds globose, the corolla exserted ca. 1/5 from the calyx before anthesis. Flowers 5-merous, all perfect. Calyx tube 1.2–1.5 mm long, conical, the lobes 0.8–1.5 mm long, 0.5–0.8 mm wide, elliptic to triangular with obtuse thickened apices and paler (almost scarious) sinuses, pubescent with spreading simple uniseriate eglandular and/or glandular trichomes like those on stem. Corolla 8–15(-20) mm in diameter, white with a yellow-green central portion near the base and occasionally with purple stripes along lobe midveins abaxially, stellate, lobed 1/2 way to the base, the lobes 2.5–4.5 mm long, 2.0–3.5 mm wide, strongly reflexed at anthesis, later spreading, densely papillate-pubescent abaxially with simple uniseriate eglandular trichomes. Stamens equal; filament tube minute, pubescent with spreading uniseriate simple eglandular trichomes adaxially; free portion of the filaments 1.0–1.3 mm long, pubescent like the tube; anthers 1.8–2.2(-2.4) mm long, 0.5–0.7 mm wide, ellipsoid, yellow, poricidal at the tips, the pores lengthening to slits with age and drying. Ovary globose, glabrous; style 2.8–3.5(-4.0) mm long, densely pubescent with 2–3-celled simple uniseriate trichomes in the lower half, exserted 0–1 mm beyond anther cone; stigma capitate, the surface minutely papillate, green in live plants. Fruit an ellipsoid berry, usually somewhat longer than broad, 8.5–10 mm long, 8.0–9.5 mm wide, (red-)orange to yellow at maturity, the pericarp thin, shiny and translucent; fruiting pedicels 8–14 mm long, 0.4–0.5 mm in diameter at the base, 0.7–1.5 mm at apex, strongly reflexed, becoming woody, spaced 1.0–2.0 mm apart not falling with the fruit, remaining on the plant and always persistent on older inflorescences; fruiting calyx not accrescent, the tube 1.0–1.5 mm long, the lobes 2.0–3.0 mm long, strongly reflexed in fruit. Seeds 20–40 per berry, 1.8–2.2 mm long, 1.5–1.7 mm wide, flattened and tear-drop shaped with a subapical hilum, brown, the surfaces minutely pitted, the testal cells pentagonal in outline. Stone cells absent, but occasionally 1–2 found in North African and Arabian material, ca. 0.5 mm in diameter. Chromosome number: 2n=4x=48 (
Solanum villosum A Habit B Inflorescence in hairy individual C Flowers and buds D Immature infructescence E Fully mature fruits F Fully mature fruits showing reflexed calyx lobes (A Nijmegen acc. 884750135; B Nijmegen acc. 914750047; C Nijmegen acc. A34750061, D Nijmegen acc. A34750043; E Nijmegen acc. A14750048; F Nijmegen acc. A34750038). Photos by T. Särkinen and G. van der Weerden.
(Figure
Grows in open and disturbed areas in a wide variety of habitat types, including in cities; between sea level and 3,000 (-3,600) m elevation.
Austria: Rotbeeren-Nachtschatten, Gelbbeeren-Nachtschatten, Gelbrotbeeren-Nachtschatten (
Leaves used as spinach (usually boiled, often in milk) and as a pot-herb in Europe, Africa and the Middle East; berries eaten raw (especially by children, see Uses above) and cooked (
(
This common European species was long known as S. luteum (e.g.
Variation in pubescence and stem characteristics within S. villosum has been recognised at many ranks from subspecies, varieties and different species (
Solanum villosum can be easily recognised from similar species by its orange-red berries (ranging from yellow to bright red) that are slightly elongate and ellipsoid. The only other species present in the Old World with pale yellow fruits is S. palitans, but this taxon differs in being a prostrate plant with thinly membranous 3-lobed leaves and is easy to tell apart. The rest of the morelloids in the Old World have rounded or subglobose black or green berries. Solanum villosum is much more common in southern Europe than is S. nigrum, while S. nigrum becomes more common towards northern and central Europe. These latter two species are difficult to tell apart based on herbarium material alone, especially when mature fruit colour is not available. Calyx lobes of S. villosum are generally longer than those of S. nigrum (relative to width), strongly reflexed at fruit maturity and the sinus is rounded (as opposed to acute in S. nigrum) and, in flower, the calyx tube has pale patches that look like small windows or white/translucent areas below the sinus itself. This is particularly easy to see in herbarium specimens. The leaves of S. villosum are similar to those of S. retroflexum in being usually more or less rhomboid with dentate margins, but leaves are more narrowly rhomboid than those of S. retroflexum (i.e. narrower relative to length). Specimens from the southern part of the range of S. villosum and the northern range of S. retroflexum (e.g. Malawi and Mozambique) can be difficult to distinguish in the absence of information on fruit colour (S. retroflexum has dull purple-black berries, also with strongly reflexed calyx lobes in fruit).
Solanum villosum has generally distinctly longer filaments relative to anther length than other species (except S. opacum and S. retroflexum, see descriptions of these species), although large variation can be observed within and between individuals especially in herbarium specimens. We have observed in the field that filaments elongate during flower maturation, where younger flowers in the same inflorescence can be seen with shorter filaments and older flowers with fully extended and distinctly longer filaments (SK photo 101532). Solanum villosum has a large capitate stigma that does not generally extend much beyond the anthers, in contrast to S. tarderemotum with which it often grows in sympatry in Africa, where the style is long-exserted and often bent and the stigma is smaller.
A set of herbarium and seed bank accessions of S. villosum from Kenya have narrower leaves with entire margins and larger, branched inflorescences with fruits in which calyx lobes do not recurve as strongly as in more typical populations, but that lack stone cells in the berries. These accessions are somewhat morphologically intermediate between S. villosum and S. tarderemotum and could present man-made or natural introgression between the two species (see discussion of S. tarderemotum).
The origin of the tetraploid S. villosum has been the focus of previous crossing and cytological studies (e.g.
Although
The name S. alatum Moench has been commonly used for densely pubescent eglandular populations of S. villosum (
Solanum suffruticosum apparently was a “herbarium” name of Peter Schousboe, a Danish botanist who worked in Tangiers in the early part of the 19th century. The epithet was used and attributed to Schousboe by
Solanum kitaibelii and S. flavum are names coined by
We have designated a specimen with the exact locality as the lectotype of S. ochroleucum; the G-DC duplicate is better preserved (G00144474).
Döll’s various infraspecific taxa described in his Rheinische Flora (1843) as “Formen” (here taken as the rank of forma) are mostly new combinations of previously published epithets, but his forma angulosum is based on several cited localities in Germany. We have not been able to locate specimens for these collections and defer typification until more in-depth searches are done. It may be that these names are based on field observations and not herbarium material.
Solanum sinaicum was described by
The spelling of
The names coined by the Hungarian botanist Pál Kitaibel were published posthumously by
In his Manual flora of Madeira, (
Two un-numbered collections were cited in the protologue of S. nigrum var. aurantium (
The names coined by
Solanum purpureilineatum (
A total of 1,287 specimens were examined from 86 countries during the study across Africa, Asia, Australia, Eurasia and the Pacific. Specimens from the entire range (including adventives) were studied in order to understand the full range of morphology within the species. All specimens examined from the Old World can be seen in Appendix 2 (csv format) and Appendix 3 (traditional Specimens Examined list in pdf format).
Solanum cestrifolium Jacq., mentioned by
Solanum foetidum Rottb., Acta Lit. Univ. Hafn. 1: 287. 1778. Type. “Surinam”, C.F. Rottbøll s.n. (no original material located); this name has sometimes been referred to the morelloids, but is likely not to even be a Solanum, it might refer to a plant of Datura.
Solanum foliosum Link, Phys. Beschr. Canar. Ins. [Buch] 144. 1825. Type. Spain. Islas Canarias: Tenerife, Puerto Orotava, L.C. Buch s.n. (no specimens cited; original material seen by Link in B, destroyed). Link’s description suggests this plant is like, not the same as, S. nigrum, but then goes on to describe a plant with pinnatifid leaves and flowers larger than those of S. nigrum. Possibly S. nava Webb & Berthel. (1845) or perhaps the cultivated S. laxum Spreng. (1824)
Solanum nigrum var. aspergilliflorum Sendtn., Fl. Bras. (Martius) 10: 16. 1846. Type. Sin. loc. [Brazil?] (no specimens or collectors cited). Might be S. chenopodioides Lam. based on Sendtner’s notes “S. chenopodioides Lam.?” beneath the description in the original publication, but the description mentions stellate hairs on the leaf undersides, a character that does not occur in the Morelloid clade.
Solanum nigrum var. humile Macloskie, Rep. Princeton Univ. Exped. Patagonia 8: 707. 1905. The “description” consists solely of a reference to “S. humile Salisb.” an illegitimate superfluous name citing S. nigrum L. in synonymy. Solanum nigrum does not occur in Patagonia, so this name cannot be considered homotypic; identity uncertain, but certainly a species native to the New World.
Solanum nigrum var. stylosum Witasek ex Reiche, Anales Univ. Chile 124: 456. 1909. Type. Chile. “Provincias Centrales” (no specimens cited; no original material found). The description is very short and no specimens annotated as this from Chile have been found at W or WU; it could be S. alphonsei Dunal of the Dulcamaroid clade; the description of the peduncle as twice as long as the branches is indicative of that species, rather than a morelloid.
Solanum rubrum L., Syst. Nat. ed. 12: 173. 1767, nom utique rej. prop. No original type material traced,
Solanum uliginosum Blume, Bijdr. Fl. Ned. Ind. 13: 695. 1826. No specimens or localities cited, not treated by
Solanum nigrum forma uliginosum (Blume) Miq., Fl. Ned. Ind. 2: 637. 1856.
Type. Based on Solanum uliginosum Blume
Designations are listed in alphabetic order; Code Articles are numbered as in
Solanum series Luteum Pojark., Flora SSSR 22: 32. 1955, as “Lutea”, not validly published; in Russian, no diagnosis or description in Latin (Art. 39.1). [only included species S. luteum Mill. = S. villosum Mill.]
Solanum aegyptiacum Forssk., Fl. Aegypt.-Arab. 46. 1775, nomen nudum (see
Solanum aegyptiacum Forssk., var. b) Fructu nigro; foliis integris villossissimus, Fl. Aegypt.-Arab. 46. 1775, polynomial, not validly published (see
Solanum amaranthifolium Gillies ex Rusby, Bull. Torrey Bot. Club 26: 152. 1899, nomen nudum; based on a Gillies manuscript name at Kew; two specimens by Gillies (K001166701, K001166704) are annotated “S. amaranthifolium Gill.” in Gillies’ hand = S. chenopodioides Lam.
Solanum arenarium Schur, Enum. Pl. Transsilv. 477. 1866, pro syn. Solanum humile Bernh. ex Willd. = S. villosum Mill.
Solanum asperum Hornem. ex Walp., Repert. Bot. Syst. (Walpers) 3: 49. 1844, pro syn. Solanum rumphii Dunal = S. americanum Mill.
Solanum atriplicifolium Desp. ex Dunal, Solan. Syn. 12. 1816, herbarium name pro syn. Solanum nigrum L.= S. nigrum L.
Solanum atriplicifolium Desp. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852, pro. syn. Solanum nigrum var atriplicifolium Dunal = S. nigrum L.
Solanum besserianum Weinm., Elench. Hort. Pawlowsk.: 100. 1824, nomen nudum, no description or diagnosis; Cult. “hort. Pawl.”, said to be “S. cestrifolium Jacq.?” [no such published name found; validated by
Solanum cestrifolium Gussone, Ind. Sem. Hort. Boccadifalco 11. 1825, nomen nudum = S. nigrum L.
Solanum chlorocarpum Schur, Enum. Pl. Transsilv. 478. 1866, pro. syn. Solanum nigrum var. chlorocarpum Koch. = S. nigrum L.
Solanum chousboe var. merrillianum (T.N.Liou) C.Y.Wu & S.C.Huang. This citation from Flora of China (
Solanum cuneifolium Hort. Avens ex Dunal, Prodr. [A. P. de Candolle] 13(1): 53. 1852, pro syn. Solanum nigrum L. = S. nigrum L.
Solanum decurrens Wall. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852, pro syn. Solanum rhinozerothis Blume = S. nigrum Mill.
Solanum deppei Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, pro syn. Solanum pterocaulon var. deppei Dunal = Solanum scabrum Mill.
Solanum gracile Otto ex Baxter, Hort. Brit. [Loudon] 2, Suppl.: 639 1850, nomen nudum; also later homonym of S. gracile Sendtn. (1846) = identity uncertain
Solanum heterogonum Dunal, Prodr. [A. P. de Candolle] 13(1): 52. 1852, pro syn. Solanum pterocaulon var. heterogonum Dunal = S. scabrum Mill.
Solanum judaicum Hack. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 53. 1852, pro syn. Solanum tauschii Opiz. = S. nigrum L.
Solanum kraussianum Buchinger ex Krauss, Flora 27: 831. 1844, pro syn. of polynomial Solanum nigrum L. var. fol. dentato-lobatis puberulis and not intended as a new name, refers to J.F. Drège 7864a = S. retroflexum Dunal
Solanum muricatum Bertero ex Dunal, Prodr. [A. P. de Candolle] 13(1): 150. 1852, pro syn. Solanum rancaguense Dunal = S. furcatum Dunal
Solanum nigrum var. aegyptiacum Lam. ex C.C.Gmel., Hort. Carlsruh. 253 1811, nomen nudum, no description or diagnosis; the epithet “aegyptiacum” was not used by Lamarck in the genus Solanum and must be an error. = identity uncertain
Solanum nigrum L. var. angustifolium Filov, Kult. Fl. SSSR (Zhukovskii) 10: 382. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. villosum Mill.
Solanum nigrum var. atriplicifolium Desp. ex Filov, Kult. Fl. SSSR (Zhukovskii) 10: 370–386. 1958, not validly published, no direct citation of basionym (Art. 39.1); also nom. illeg., later homonym of S. nigrum var. atriplicifolium G.Mey. = S. nigrum L.
Solanum nigrum subsp. australiense Filov, Kult. Fl. SSSR (Zhukovskii) 10: 380. 1958, not validly published; no diagnosis or description in Latin (Art 39.1) = S. nigrum L.
Solanum nigrum var. burbankii (Bitter) Filov, Kult. Fl. SSSR (Zhukovskii) 10: 383. 1958, as “berbankii”, not validly published, no direct citation of basionym (Art. 38.1) = S. nigrum L.
Solanum nigrum subsp. chinense Filov, Kult. Fl. SSSR (Zhukovskii) 10: 382. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. americanum Mill.
Solanum nigrum var. chlorocarpum Koch, Syn. Fl. Germ. Helv., ed. 2: 600. 1846, not a new name, clear citation of Solanum nigrum var. chlorocarpum Spenn. (“S. nigrum chlorocarpum Spenn. fl. fri.”) = S. nigrum L.
Solanum nigrum subsp. cultum Filov, Kult. Fl. SSSR (Zhukovskii) 10: 383. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. scabrum Mill.
Solanum nigrum var. frutescens Macloskie, Rep. Princeton Univ. Exped. Patagonia 8: 707. 1905, nomen nudum = S. nitidibaccatum, S. triflorum, S. chenopodioides, or S. pygmaeum that all occur in Northern Patagonia
Solanum nigrum forma genuinum Döll, Rhein. Flor. 412. 1843, not validly published (Art. 24.3) = S. nigrum L.
Solanum nigrum var. genuinum Godr., Fl. Lorraine 2: 133. 1843, not validly published (Art. 24.3) = S. nigrum L.
Solanum nigrum var. genuinum Hassl., Trab. Mus. Farm. Fac. Cienc. Med. Buenos Aires 21: 104. 1909, not validly published (Art. 24.3) = S. nigrum L.
Solanum nigrum var. griseocarpum Filov, Kult. Fl. SSSR (Zhukovskii) 10: 378. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. nigrum L.
Solanum nigrum var. humile F.M.Bailey, Queensland Philos. Soc. 3: 1–4. 1881, not validly published; no diagnosis or description (Art. 38.1) = S. nigrum L.
Solanum nigrum var. inerme K.Koch, Linnaea 17: 283. 1843, not intended as a new name (also nomen sub nudum); protologue “Solanum nigrum L. β inerme. Rami non murieulati. In Transcaucasia abundat” is much less detailed than where in the same work Koch describes new taxa, neither does he attribute the name to himself as he does for other new taxa in this paper.
Solanum nigrum subsp. irano-algiricum Filov, Kult. Fl. SSSR (Zhukovskii) 10: 378. 1958, not validly published; no diagnosis or description in Latin (Art. 39.1) = S. villosum Mill.
Solanum nigrum var. legitimum Neilr., Fl. Nied.-Oesterr. 535. 1859, not validly published (Art. 24.3) = S. nigrum L.
Solanum nigrum subsp. memphiticum (Mart.) Marzell, Ill. Fl. Mitt.-Eur. [Hegi] 5(4): 2593. 1927, not intended as a new name; Marzell only says “Zur gesamtart S. nigrum gehört anscheinend auch S. memphiticum Mart.” [to the total S. nigrum also belongs S. memphiticum Mart.] = S. nigrum L.
Solanum nigrum var. merrillianum (Liou) Filov, Kult. Fl. SSSR (Zhukovskii) 10: 383. 1958, not validly published; no direct citation of basionym (Art. 38.1) = S. americanum Mill.
Solanum nigrum var. villosum (Mill.) Filov, Kult. Fl. SSSR (Zhukovskii) 10: 380. 1958, not validly published; no direct citation of basionym (Art. 38.1) = S. villosum Mill.
Solanum nigrum var. violaceum Chen ex Wessely, Feddes Repert. Spec. Nov. Regni Veg. 63(3): 293. 1960, nomen nudum; not intended as a new name, listed as one of the taxa accepted by
Solanum nigrum var. vulgatum L., Sp. Pl. ed. 2: 266. 1762, evidently an orthographic variant of Solanum nigrum L. var. vulgare L. (1753). = S. nigrum L. (see
Solanum nigrum var. vulgatum Dunal, Prodr. [A. P. de Candolle] 13(1): 50. 1852, not a new name, referring to Solanum nigrum var. vulgatum L. (and of others “auct.”) = S. nigrum L.
Solanum nodiflorum Desv. ex Dunal, Prodr. [A. P. de Candolle] 13(1): 46. 1852, pro syn. Solanum desvauxii Ham. = S. americanum Mill.
Solanum nodiflorum var. acuminatum Chodat, Bull. Herb. Boissier, sér. 2, 2: 811. 1902, not intended as a new name, as “acuminatum (?)”, with no specimen cited. In the rest of this work the new taxa are clearly indicated with “nob.” and a specimen (or several) cited = S. americanum Mill.
Solanum nodiflorum var. fauriei (H.Lév.) O.Deg. & O.Deg., Plants Hawaii Natl. Parks 419. 1930, no evidence in any editions of Degener’s book (1930, 1945, 1974) that this name, occurring in several indices and based on Solanum fauriei H.Lév. was ever published; probably = S. opacum A.Braun & Bouché
Solanum nodiflorum var. sativum A.Chev. Expl. Bot. Afrique Occ. Franç. 1: 464. 1920. nomen nudum; based on “Guinea. Kankan, dans les jardins cultives, 16 Mar 1899, A.Chevalier 594” = S. scabrum Mill.
Solanum pigmaeum Larrañaga, Escritos Damaso Antonio Larrañaga 2: 88. 1923, almost certainly not intended as a new name, an orthographic variant of Solanum pygmaeum Cav. (1806). In other parts of Larrañaga’s work (e.g. S. trilobum in vol. 1: 12), he indicates new taxa with “sp. n.” or gen. n.”; none of the names in vol. 2 is so designated, leading us to believe he was not coining new names, but instead using those already published. This work was compiled from his notes and diaries and these were heavily annotated (see intro in vol. 1 http://bdh-rd.bne.es/viewer.vm?id=0000016651&page=1) up to his death in 1848. = S. pygmaeum Cav.
Solanum photeinocarpum var. violaceum C.Y.Wu & S.C.Huang, Acta Phytotax. Sin. 16(2): 72. 1978, nomen nudum; incorrectly cited in this publication as violaceum Chen ex Wessely but
Solanum pterocaulon Rchb. ex Nyman, Consp. Fl. Eur. 3: 526. 1881. (1881), pro syn. Solanum melanocerasum Willd. = Solanum scabrum Mill.
Solanum repens Noronha, Verh. Batav. Genootsch. Kunsten 5: 84. 1791, nomen nudum = S. nigrum L.
Solanum rubrum Murray, Syst. Veg., ed. 14 (J. A. Murray) 224. 1784, not intended as a new name; Murray repeats
Solanum villosum Opiz ex Dunal, Prodr. [A. P. de Candolle] 13(1): 51. 1852, pro syn. Solanum decipiens Opiz. = S. nigrum L.
Solanum villosum var. alatum (Moench) Marzell, in Hegi, Ill. Fl. Mitt.-Eur. 5(4): 2594. 1927, not intended as a new name, citation of Solanum alatum Moench in text = S. villosum Mill.
Solanum villosum var. glabrescens Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 592. 1819. This name was cited in
Solanum viridescens Hort. ex Rchb., Fl. Germ. Excurs. 391. 1831, pro syn. Solanum humile Bernh. ex Willd. = S. villosum Mill.
Solanum virgatum Endl. ex Sendtn., Fl. Bras. (Martius) 10: 13. 1846, pro syn. Solanum gracile Sendtn. = S. chenopodioides Lam.
We thank the curators of herbaria cited in the text for permission to examine or borrow the specimens in their care, especially Thomas Rouillard (ANG), Pierre Moulet (AV), Lajos Somlyay (BP), Mbaluka Kimeu (EA), Nicolas Fumeaux (G), Jim Solomon (MO), Caroline Loup (MPU), Michael Nee (ex NY, MO), Laura Guglielmone (TO) Mats Hjertson (UPS) and Krysztof Swierkosz (WRSL) who kindly searched for duplicates and original material to facilitate lectotypification; Sarah Ficinski for preparing all maps and for assistance in data recording; Graham Hardy (E) for assistance in finding protologues and access to relevant library resources; Ranee Prakash (BM) for help navigating the complex world of Ayuvedic medicine and Indina common names; Brooke Micke for investigating common names and uses across Africa; Rosemary Wise, Lucy Smith, Maria Louise Szent-Ivany, Maria Teresa Cabrera and Laura Ribulgo for line drawings of the taxa treated here; Jean-François Butaud (UPF), Paul Gonzales (USM), Franco Chiarini (CORD) and Alicia Sérsic (CORD) for allowing us to use their photographs in illustrating the taxa treated here; the Board of the Instituto Darwinion (San Isidro, Buenos Aires, Argentina) and the Flora of Argentina editorial committee for allowing us to use illustrations that had been previously published; David Lorence (PTBG) for helping us to understand diversity in Hawaii; Maria Vorontsova (K) for field assistance in Africa; Gao Tiangang and Wang Jinxiu (both PE) for field assistance in China; and the governments of China, Kenya and Tanzania for permission to collect plants in their respective countries. Careful editing and reviewing by Eric Tepe and Lynn Bohs greatly improved the treatment. This work was in part funded by the National Science Foundation’s Planetary Biodiversity Inventory programme (DEB-0316614 ‘PBI Solanum – a worldwide treatment’ http://www.solanaceaesource.org) to SK; the SYNTHESYS Project http://www.synthesys.info/ which was financed by European Community Research Infrastructure Actions under the FP6 and FP7 “Structuring the European Research Area” Programme; Natural History Museum Special Funds awards; and the Royal Society through travel grants.
Index to numbered collections
Specimens examined (CSV format)
Specimens examined (PDF format)