PhytoKeys 22: 1–432, doi: 10.3897/phytokeys.22.4041
A revision of the Dulcamaroid Clade of  Solanum L. (Solanaceae)
Sandra Knapp 1
1 Department of Life Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom

Corresponding author: Sandra Knapp (s.knapp@nhm.ac.uk)

Academic editor: L. Penev

received 25 September 2012 | accepted 20 February 2012 | Published 10 May 2013


(C) 2013 Sandra Knapp. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


For reference, use of the paginated PDF or printed version of this article is recommended.

Abstract

The Dulcamaroid clade of Solanum contains 45 species of mostly vining or weakly scandent species, including the common circumboreal weed Solanum dulcamara L. The group comprises members of the previously recognised infrageneric groupings sect. Andropedas Rusby, sect. Californisolanum A. Child, sect. Dulcamara (Moench) Dumort., sect. Holophylla (G.Don) Walp., sect. Jasminosolanum (Bitter) Seithe, sect.Lysiphellos (Bitter) Seithe, subsect. Nitidum A.Child and sect. Subdulcamara Dunal. These infrageneric groups are not monophyletic as traditionally recognised, and the complex history of the classification of the dulcamaroid solanums is reviewed. Many of the species in the clade are quite variable morphologically; plants are shrubs, herbaceous vines or woody canopy lianas, and habits can vary between these states in a single locality. Variation in leaf shape and pubescence density and type is also extreme and has lead to the description of many minor morphological variants as distinct species. The flowers of members of the group are generally very showy, and several species (e.g., Solanum crispum Ruiz & Pav., Solanum laxum Spreng., Solanum seaforthianum Andrews) are popular ornamental plants that have occasionally escaped from cultivation and become naturalised. The clade is here divided into five morphologically and geographically delimited species groups to facilitate further study. One new species from southern Ecuador, Solanum agnoston S.Knapp sp. nov., is described here. Full descriptions and synonymies (including designations of lectotypes or neotypes), preliminary conservation assessments, illustrations, distribution maps, and an extensive list of localities are provided for all species.

Keywords

Asia, classification, monograph, new species, Solanum, South America, taxonomy, vines, USA

Introduction

Solanum L. is one of the ten most species-rich genera of flowering plants (Frodin 2004). With approximately 1400 species (J. Bennett and S. Knapp, unpubl.) occurring on all temperate and tropical continents, the genus occupies an incredibly wide range of habitats and habits, but the highest diversity of both groups and species occurs in circum-Amazonian tropical South America. Solanum is recognised by its usually pentamerous flowers with fused sepals and petals, stellate to pentagonal corollas, and stamens with short filaments and anthers opening by terminal pores. The genus was one of Linneaus’s (1753) larger, with 23 species mostly described from European or African material. The French botanist Michel-Félix Dunal included 235 species in his thesis (Dunal 1813), mostly the result of extensive European exploration of the Americas. By 1816, when Dunal revised this work (Dunal 1816), this number had risen to 321; many of these additional taxa were based on specimens collected by Alexander von Humboldt and Aimé Bonpland in tropical South America (see Knapp 2007b). The last time the genus was monographed in its entirety was in Candolle’s Prodromus (Dunal 1852) which included 900 Solanum species. Subsequent work on the taxonomy of Solanum has largely been limited to rearrangements of infrageneric taxa, or to the species-level revisions of smaller groups within the genus (see references in Table 1) and floristic treatments. The combination of large numbers of species with relatively poorly circumscribed groups within the genus has meant that Solanum taxonomy has proceeded in a piecemeal fashion until relatively recently. A project funded by the United States National Science Foundation’s Planetary Biodiversity Inventory program begun in 2004 sought to redress this situation by attempting to accelerate species-level taxonomic work across the genus as a whole and at the same time providing a robust phylogenetic framework for this taxonomy. Work by participants of the ‘PBI Solanum’ project (see http://www.solanaceaesource.org ) will result in a modern monographic treatment of the entire genus available on-line. This treatment is part of this collaborative effort.

Table 1.

The major clades of Solanum (after Bohs 2005). For revisions of individual species published on-line see Solanaceae Source (http://www.solanaceaesource.org ).

Clade name (Bohs 2005) Approx. # of species Recent taxonomic monographs
Thelopodium clade 3 Knapp 2001
Regmandra clade 11 Bennett 2008
Archaesolanum clade 9 Symon 1994
Normania clade 3 Bohs and Olmstead 2001
African Non-Spiny clade ca. 15 in progress, Knapp and Vorontosova
Potato clade ca. 200 Knapp and Helgason 1997; Spooner et al. 2004; Peralta et al. 2008; Tepe and Bohs 2011
Morelloid clade ca. 75 in progress, Barboza and Särkinen
Dulcamaroid clade 43 This treatment
Wendlandii/Allophyllum clade ca. 12 Bohs 1990; Clark et al. in prep.
Cyphomandra clade 50 Bohs 1994; Bohs 2001
Geminata clade 145 Knapp 2002a; Knapp 2008a
Brevantherum clade ca. 80 Roe 1967, 1972; Carvalho 1996; Stern et al. 2012
Leptostemonum clade ca. 450 Whalen 1979; Whalen et al. 1981, Vorontosova and Knapp 2012, in review; many other groups with monographs in progress
History of Solanum Classification

Dunal (1813, 1816) divided the genus into two major groups, Inermia (unarmed solanums) and Aculeata (armed solanums), based on presence or absence of prickles. He renamed these at the sectional level (but illegitimately, as he cited groups he had previously named as sections, e.g., Pteroidea Dunal [Dunal 1816] in synonymy) “Pachystemonum”, for species with stout anthers and no prickles, and “Leptostemonum”, for species with tapering anthers, usually with stellate hairs and usually possessing prickles (Dunal 1852); these divisions were essentially the same as his Inermia and Aculeata. In Candolle’s Prodromus, Dunal (1852) erected a classification of subsections and ambiguous grades to reflect morphology, mostly of leaf division and inflorescence position. Dunal (1852) maintained as separate the genera Lycopersicon Mill. and Cyphomandra Sendtn., both now subsumed in a monophyletic Solanum (see Bohs 2005; Peralta et al. 2008). The German botanist Georg Bitter, who worked in Berlin, Bremen and Göttingen in the years between the two World Wars, published extensively in Solanum (see complete bibliography in Weber 1928) but never constructed a classification of the entire genus. In the mid-20th century, Seithe (1962) compiled Bitter’s voluminous work on Solanum classification in a comprehensive system based largely on hair types; she recognised two major divisions, “chorus subgenerum Solanum (L.) Seithe”, the simple and branched hair solanums, and “chorus subgenerum Stellatipilum Seithe”, the stellate haired solanums. Danert (1967, 1970) used his own research on growth form and branching patterns in Solanaceae to propose a new subgeneric classification system for Solanum; he largely kept the same groupings as Seithe (1962) but recognised them at different ranks. D’Arcy (1972) assembled all the subgeneric names in Solanum and provided lectotypes for those that remained untypified. He divided the genus (in what he called a “provisional conspectus”) into seven subgenera (Solanum L., Archaesolanum Marzell, Bassovia (Aubl.) Bitter, Brevantherum (Seithe) D’Arcy (=Minon Raf.), Lyciosolanum Bitter, Potatoe (G.Don) D’Arcy and Leptostemonum Bitter [cited by D’Arcy as Leptostemonum (Dunal) Bitter]) and 52 sections based on combinations of characters he felt were more realistic than those used in the past. His subgenus Leptostemonum is equivalent to that of previous workers, but his divisions of the “non-spiny solanums” were quite different. Like Dunal (1852), but not Seithe (1962), D’Arcy excluded Lycopersicon and Cyphomandra from Solanum. Nee (1999) used only the New World species of Solanum to construct a simplified system in which he recognised 3 subgenera – Bassovia (comprised of the species previously recognised as Cyphomandra and their relatives, in which he included section Pteroidea, see Knapp and Helgason 1997), Solanum (the rest of the “non-spiny” solanums) and Leptostemonum (the “spiny” solanums). He recognised 21 sections with numerous subsections, and listed component taxa in each. Since the largest species diversity of Solanum occurs in the New World, Nee’s (1999) system has been very useful for focusing in-depth taxonomic studies on smaller, putatively monophyletic groups. Child and Lester (2001) provided a synopsis of infrageneric classification of Solanum, based largely on the work of Bitter and Seithe (1962). They did not indicate component species of any of their recognised sections, which comprise fewer species than those of Nee (1999).

DNA sequence data has revolutionised hypotheses of angiosperm relationships (APG III 2009) and Solanum is no exception. Based on cladistic analyses of DNA sequence data Solanum can be divided into 13 well-supported monophyletic clades (Bohs 2005; Weese and Bohs 2007; see Table 1), the largest of which is the group commonly known as the spiny solanums (the Leptostemonum clade), which is largely coincident with Dunal’s (1852) “Leptostemonum”. The largest non-spiny solanum clades (see Table 1) are the Geminata clade, the Potato clade, the Brevantherum clade and the Morelloid/Dulcamaroid clade; this last comprises all the herbaceous species previously placed in section Solanum (see Weese and Bohs 2007) and allied groups, plus the species previously placed in sections Jasminosolanum Bitter, Dulcamara (Moench) Dumort., Lysiphellos Bitter, Nitidum A.Child and Andropedas Rusby plus a variety of other taxa (see Bohs 2005 for discussion). Most current taxonomic work in Solanum is being undertaken in this cladistic framework underpinned by molecular data and concomitantly, more taxa are being added to molecular analyses to test its stability and robustness (e.g., Stern et al. 2011). As more taxa have been added the thirteen clades have generally been supported, but many species have not yet been included in molecular analyeses and their relationships remain to be tested.

History of taxonomy of species of the Dulcamaroid Clade

The Dulcamaroid clade as recognised by Bohs (2005) and treated here is comprised of elements from several previously recognised subgenera and sections of Solanum. Species of the group are morphologically quite variable (see Morphology below); this in part accounts for these species’ chequered classification history and extensive synonymy. The common European woody nightshade or bittersweet, Solanum dulcamara L., has a long history of use in local medicine (Dunal 1813) in Eurasia (see species treatment of Solanum dulcamara). It was the only member of this group described by Linnaeus (1753), who recognised its extreme variability but did not explicitly name any of these variants, although he did explicitly name variants of other highly variable taxa. He did differentiate an African polynomial taken from Dillenius (1732) as “ß”. This polynomial refers to the South African species Solanum africanum Mill. (long known as Solanum quadrangulare Thunb.), provisionally a member of the African Non-Spiny clade (sensu Bohs 2005), further underlining the morphological similarities between the Dulcamaroid clade as here recognised and the unrelated “African Non-Spiny” (ANS) clade (see below).

Dunal (1813) reviewed the medicinal history of Solanum dulcamara in his thesis, and included it in his Inermia in a group characterised as “Foliis lobatis….” together with Solanum lyratum Thunb. and Solanum triquetrum Cav. In another group – “Foliis integerrimus…” he included an additional eight species now included in the Dulcamaroid clade as defined here; Solanum dichotomum Lour. (=Solanum lyratum) was put into a subgroup with lateral inflorescences, while the rest were said to have terminal inflorescences. Dunal had only seen herbarium specimens or live plants of three of these taxa (Solanum dulcamara, Solanum triquetrum and Solanum pubigerum Dunal) and relied on published descriptions for the rest. The greatly increased number of species (19 names recognised as members of the Dulcamaroid clade as treated here) included in the updated synopsis (Dunal 1816) were scattered over three groups based on leaf division. Solanum seaforthianum Andrews was part of Dunal’s (1816) group containing members of section Pteroidea (sensu Knapp and Helagson 1997, part of the Potato clade, Bohs 2005), Solanum cyrrhosum Dunal (=Solanum seaforthianum) was included in a group with Solanum dulcamara and Solanum lyratum; and Solanum nitidum Ruiz & Pav. (together with other members of the Solanum nitidum species group sensu Knapp 1989) with various members of the Geminata clade (sensu Knapp 2002a, 2008a) such as Solanum sessile Ruiz & Pav. and Solanum oblongifolium Dunal. Dunal’s division of Solanum into groups was quite detailed, but definition of the groups themselves was not completely parallel (see Knapp 1983 for a discussion of sectional nomenclature in Solanum). It is clear that the species now recognised here as members of the Dulcamaroid clade were in different groups due to their highly variable leaf morphology that can be pinnatifid to simple on a single stem (see below). Dunal’s (1852) Prodromus treatment included many more species of the Dulcamaroid clade as defined here; he mostly treated these taxa as members of his subsections Dulcamara (in the groups Dulcamara and Subdulcamara) and Micranthes (in the group Oppositifolia * Indubitaria). He included Solanum viscosissimum Sendtn. in his section Tuberarium (group Potatoe with the potatoes and tomatoes), based on its deeply pinnatifid leaves. Difficulty in grouping these taxa which today we recognise as closely related is perhaps understandable as label data from 19th century herbarium specimens is sparse and usually does not include any mention of habit. Dependence upon leaf morphology and inflorescence position to determine relationships in Dunal’s systems would easily be foiled by the extremes of variability in leaf shape and inflorescence size found in the Dulcamaroid clade.

Seithe (1962) examined 36 species (representing 24 of the species recognised here and 12 species relegated to synonymy here) of the Dulcamaroid clade in her analyses of hair types in the genus. Of these, 11 were classified only as members of subgenus Solanum without assignment to a section; Seithe was unsure as to the affinities of these taxa. The rest of the species were included in her sections Dulcamara (e.g., Solanum dulcamara, Solanum lyratum), Lysiphellos (e.g., Solanum decorticans Sendtn. [=Solanum inodorum Vell.]), Jasminosolanum (e.g., Solanum seaforthianum, Solanum jasminoides Paxt. [=Solanum laxum Spreng.]), Anthoresis (e.g., Solanum aureum Dunal, Solanum storkii C.V.Morton & Standl.) and Pseudocapsicum (e.g., Solanum triquetrum). Those taxa included in Anthoresis are all the members of the Solanum nitidum species group (sensu Knapp 1989) and were grouped together based on the possession of highly branched trichomes. Danert (1970) recognised the same set of sections (renaming section Anthoresis as “section Holophyllum Walp.”), but did not list any of the component species in his system, so understanding his concepts of specific relationships is difficult. He did suggest a close relationship between sections Dulcamara, Jasminosolanum and Aculeigerum Seithe (equivalent in part to the Wendlandii/Allophyllum clade of Bohs 2005) and the vining taxa from Africa (recognised as sections Afrosolanum Bitter, Lemurisolanum Bitter, Benderianum Bitter, Macronesiotes Bitter, and Quadrangulare Bitter). He pointed out the extreme morphological similarity between members of his section Holophyllum and section Brevantherum Seithe; species in both these groups have dichasial branching, but they differ in their trichome morphology. Danert (1970), despite his emphasis on architecture in classification, put more weight on the trichome characters and agreed with Seithe (1962) that his sections “Holophyllum” (the same as Seithe’s Anthoresis) and Brevantherum, while similar morphologically, were not close evolutionarily.

D’Arcy (1972) included most of the vining dulcamaroid species in his subgenus Potatoe (G.Don) D’Arcy, along with the potatoes (section Petota Dumort.) and section Regmandra (Dunal) Ugent, in sections Dulcamara and Jasminosolanum (typified with Solanum dulcamara and Solanum jasminoides [=Solanum laxum] respectively). This placement was based on their possession of a scandent habit, pinnate leaves and articulation of pedicels above the base (D’Arcy 1972). Species such as Solanum pulverulentum Pers. (=Solanum cutervanum Zahlbr.) were included in his subgenus Brevantherum (=subgenus Minon) in section Holophylla (G.Don) Walp. due to their possession of complex branched trichomes (D’Arcy 1972). Solanum decorticans (=Solanum inodorum) was segregated as section Lysiphellos of subgenus Solanum. He included all of the African vining species (which all have simple leaves) in subgenus Solanum as Bitter’s original sections (sensu Seithe 1962). D’Arcy’s separation of the members of the Dulcamaroid clade in two different subgenera was the first explicit hypothesis as to the affinities of these taxa on a higher level.

Knapp (1989) studied the members of section Holophylla and recognised that the section was morphologically heterogeneous and almost certainly not monophyletic. She segregated a group of 11 species, the Solanum nitidum species group, and identified the unusual pedicel insertion morphology (see below) as a morphological synapomorphy for the group. Her cladistic treatment of the Solanum nitidum species group based on morphology was one of the first explicitly phylogenetic analyses of an entire clade in Solanum. Solanum pubigerum Dunal and Solanum aligerum Schltdl. were suggested as sister taxa to the Solanum nitidum group, but Knapp (1989) suggested the entire group was more closely related to other woody tree solanums such as the members of section Geminata (G.Don) Walp. (the Geminata clade, see Knapp 2002a, 2008a) rather than to species associated with Solanum dulcamara and its relatives. Child (1998) established subsection Nitidum A.Child based on this monograph (Knapp 1989), but did not suggest further relationships or examine material in detail. Child (1998) also segregated the Californian species Solanum xanti A.Gray (=Solanum umbelliferum Eschsch.) and its relatives as section Californisolanum A.Child and followed D’Arcy (1972) in suggesting these species were members of subgenus Potatoe and related to “dulcamaroid taxa”.

In his treatment Solanum in the New World Nee (1999) included the species of the Dulcamaroid clade in sections Dulcamara (the majority of species included here) and Holophylla (members of the Solanum nitidum species group sensu Knapp 1989). His circumscription of section Holophylla also includes all of the members of the Geminata clade as treated by Knapp (2008). All the vining species (plus other species previously recognised as section Parasolanum Bitter, e.g. Solanum corymbosum Jacq.) were combined into a single section Dulcamara by Nee (1999); included in his circumscription are the African Solanum terminale Forssk. and its relatives, which in molecular analyses form the distinct African Non-Spiny clade (Bohs 2005). Although the two groups are morphologically quite similar, in that both contain species that climb with the aid of petioles (see below), sequence differences thus far clearly differentiate them into two clades. Solanum corymbosum and other members of section Parasolanum group are resolved unambiguously as part of the Morelloid clade in all molecular analyses (Bohs 2005; Weese and Bohs 2007).

Two years after the publication of Nee’s (1999) system for New World taxa, Child and Lester (2001) provided a synopsis of infrageneric taxa in Solanum worldwide. This system is an attempt to bring together information from the scattered publications on Solanum taxonomy at the time, rather than a comprehensive reanalysis of infrageneric classification based on new data. Child and Lester (2001) were inspired by Bitter’s many works (see above) and their system is in large part based on his divisions of the genus. No component species in any groups are listed, and although “type” species are given, these are not validly published. They treat members of the Dulcamaroid clade (as far as one can tell from the designations of “types”) in their subgenus Solanum, section Holophyllum (as subsection Nitidum and the “species group Solanum valdiviense sensu A. Child”) and subgenus Potatoe (as “Dulcamaroid taxa”, sections Dulcamara, Jasminosolanum and Californisolanum). Following D’Arcy (1972), they treated all of the African vining species as members of subgenus Solanum.

The unexpected relationship of the Solanum nitidum species group (sensu Knapp 1989) with the vining dulcamaroid solanums was revealed in analyses of DNA sequence data (Bohs and Olmstead 1999; Bohs 2005; Weese and Bohs 2007). The monophyly of the Dulcamaroid clade as recognised here had previously never been suggested. Earlier ideas about the relationships of these groups had always emphasised their habit differences and segregated them clearly, but the presence of a “pedicel sleeve” (sensu Knapp 1989) is a clear synapomorphy uniting these seemingly very different taxa. Knowledge of plants in the field also has helped to find characters uniting these taxa; many of the shrubby taxa are scandent or scrambling, a character only apparent with good field notes or observations. The distant relationship of these taxa and the vining species from Africa and Madagascar (the African Non-Spiny clade sensu Bohs 2005) is somewhat surprising, given that they share the unusual characters of twining petioles and pedicel insertion into a small “sleeve”. Only three species (albeit from three of Bitter’s sections!) from the African Non-Spiny clade have been included in molecular analyses to date and with more in-depth sampling relationships are likely to become clearer. What is clear from analyses using molecular sequence data is that the members of the Dulcamaroid clade are not closely related to the potatoes as had suggested D’Arcy (1972) and Child and Lester (2001). In the analysis using the plastid gene ndhF the sister group to the Dulcamaroid clade is the Morelloid clade, comprising members of section Solanum and relatives (Bohs 2005). A subsequent analysis (Weese and Bohs 2007) using two plastid regions (ndhF and trnT-F) and a nuclear region (waxy, GBSSI or Granule Bound Starch Synthetase) resolved the Dulcamaroid clade (with 93% bootstrap support) as part of a polytomy involving the Morelloid, Normania, Archaesolanum and African Non-Spiny (ANS) clades (see Figure 1 in Weese and Bohs 2007).

Although the structure of relationships within the Dulcamaroid clade is very poorly resolved and the taxa sampled relatively few, a few sister relationships are apparent within the clade. In the ndhF analysis (Bohs 2005) five groups form a polytomy: (Solanum ipomoeoides Chodat & Hassl. [=Solanum uncinellum Lindl.]) + (Solanum calileguae Cabrera+ Solanum jasminoides [=Solanum laxum]) + (Solanum crispum Ruiz & Pav. + Solanum amygdalifolium Steud.) + (Solanum aligerum + Solanum nitidum) + (Solanum dulcamara + (Solanum seaforthianum + Solanum wallacei (A.Gray) Parish)). Weese and Bohs (2007) used fewer species in their three-gene analysis, and recovered Solanum pubigerum + (Solanum nitidum + Solanum aligerum) as sister to a polytomy of (Solanum calileguae + Solanum ipomoeoides [=Solanum uncinellum]) + (Solanum crispum + Solanum amygdalifolium) + (Solanum dulcamara + Solanum seaforthianum). These relationships are consistent with those recovered using only ndhF sequences and support the distinction of a group corresponding to section Holophylla in the narrow sense including Solanum nitidum, Solanum pubigerum and Solanum aligerum, but not Solanum crispum. Relationships amongst the vining dulcamaroid species are less clear, but additional sampling will certainly help with the delimitation of smaller groups within the clade. Most of these relationships are very poorly supported in both analyses, so any sister taxon relationships should be regarded as preliminary.

Morphology

Habit. Members of the Dulcamaroid clade are all woody plants and vary from shrubs or lax shrubs to vines (see Figure 1). Some species (e.g., Solanum dulcamaroides Poir., Solanum uncinellum) are large canopy lianas, while other vining species are woody only at the base (e.g., Solanum dulcamara, Solanum lyratum), especially in temperate climates. Solanum umbelliferum in California has sprawling herbaceous stems arising from a thick woody rootstock or base. Some species spread by underground stems; Solanum dulcamara is often considered a pest of gardens in Europe for this reason. Many of the species of the group are small shrubs that only become vining as they grow (i.e., Solanum viscosissimum) while others spread over adjacent vegetation, but never truly climb (e.g., Solanum angustifidum Bitter) and others are erect and never twine (e.g., Solanum aligerum, Solanum endoadenium Bitter, Solanum storkii).

Figure 1.

Habit of members of the Dulcamaroid clade. A Twining petioles that aid in climbing (Solanum laxum, cultivated in London UK) B Solanum angustifidum (Barboza et al. 3489) as a lax shrub on a fence in Argentina C Erect shrub of Solanum nitidum (Nee et al. 51755, Cochabamba, Bolivia).

Climbing members of the group have petioles that coil around supports (Solanum laxum [as Solanum jasminoides] was characterised as a “leaf-climber” by Darwin 1865) anchoring the ascending stems (Figure 1). This climbing mechanism is not found elsewhere in Solanum except in the African Non-Spiny (ANS) clade, which in molecular phylogenetic analyses is not closely related to the Dulcamaroid clade (Weese and Bohs 2007; T. Särkinen and S. Knapp, unpublished data). Darwin (1865) measured the time taken for petioles to clasp supports and found it very slow compared to other twining plants; once a petiole has clasped a support it accumulates woody tissue and thickens considerably.

Stems. The main axis of the plant is a typical Solanum sympodium formed by a succession of lateral axes with alternate leaves arranged in a 1/3 phyllotaxic spiral, and inflorescences are terminal at the end of each sympodial unit (Danert 1958). Danert (1958, 1967, 1970) and Child and Lester (1991) documented sympodial units and anthoclades (patterns of foliar lateral branches and associated inflorescences) in the Solanaceae. The number of leaves per sympodium is very regular and of major taxonomic significance in the entire genus Solanum (see Knapp 2001 for a further discussion).

In the Dulcamaroid clade sympodial units are almost always plurifoliate with many (or an irregular number) of leaves between each inflorescence in contrast to other groups such as the tomatoes (Peralta et al. 2008) or the Geminata clade (Knapp 2008) in which sympodial units are composed of 3 or fewer leaves. Plurifoliate sympodial units have the leaves arranged in a spiral fashion along the stems, and the leaves in members of the dulcamaroids are never geminate (paired, as seen in the Geminata clade, Knapp 2008). Most members of the group have monochasial branching, with a single axillary branch arising from below the inflorescence, giving the stems a zig-zag appearance (Danert 1958). Members of the Solanum nitidum group are either monochasial or some species have dichasial branching (see Knapp 1989); in species with dichasial branching two axillary shoots develop and the branching is dichotomous (e.g., Solanum stenophyllum Dunal). Most of these species have a mixture of the two branching types but are predominantly one or the other.

Leaves. The leaves of members of the Dulcamaroid clade are extremely variable, both across the group and within individual plants (see Figure 2 and individual species illustrations). Leaves on reproductive stems are either simple or variously pinnatifid/pinnate. There is always a wing of leaf tissue along the midrib between the lobes connecting all the dissections that is more pronounced than that found in the tomatoes and potatoes (see Peralta et al. 2008 for discussion of tomato leaf morphology), but some species such as Solanum angustifidum and Solanum viscosissimum or some forms of Solanum lyratum have such deeply divided leaves that they are best characterised as pinnate as is commonly done in other such taxa (see Peralta et al. 2008).

Figure 2.

Leaf variation in members of the Dulcamaroid clade. A Leaf variation along a single short stem, Solanum salicifolium (Barboza et al. 3467, Catamarca, Argentina) B Thick leathery leaves of Solanum macbridei (Nee & Solomon 30182, La Paz, Bolivia) C Uneven leaf lobing in Solanum dulcamara (Knapp IM-10102, Tuscany, Italy) D Variable membranous leaves of Solanum lyratum (Knapp et al. 10139, Jiangxi, China).

Figure 3.

Trichome types found in members of the Dulcamaroid clade. A Loose dendritic B Simple glandular C Echinoid D Furcate.

Leaf polymorphism is common in the group and in many species individual stems bear both simple and pinnatifid leaves (e.g., Solanum dulcamara, Solanum lyratum, Solanum salicifolium Phil.). The control of this has not been investigated, but may have to do with hormonal balance due to light or nutrients. In other species, most notably in some plants of Solanum viscosissimum, lateral inflorescence-bearing shoots have simple leaves while main axis leaves are pinnatifid or pinnate. This variability has lead to the many synonyms in some species; different leaf morphologies collected from different parts of the plant have been described as separate taxa. In some taxa there seems to be some genetic control of leaf morphology; variants with deeply dissected leaves of Solanum lyratum all come from a few localities and may represent local populations with fixed genetic differences. A set of specimens of the otherwise consistently simple-leaved Solanum uncinellum from near Iquitos have deeply dissected leaves but are identical in flower and fruit characteristics. Leaf morphology in pinnately leaved species of Solanum is complex, and governed by a set of interacting genes (Holtan and Hake 2003); this variation can make identification difficult in some species.

Leaf divisions are narrowly elliptic, elliptic to broadly elliptic, or obovate; the divisions rarely are petiolulate. The base is usually asymmetric, and varies from truncate or rounded, to cordate or lyrate. The apex (of leaves and leaflets) is rounded, acute or acuminate. The margins are entire, never serrate or crenate and the margins are straight or revolute.

The juvenile leaves of most of the species for which they have been observed are usually pinnate (pinnatifid), but because botanists usually only collect reproductive stems, this is not known for many species. Where juvenile leaves are known I have included this information in species descriptions. A sterile specimen with pinnate leaves from Iquitos (Williams 8481, F) was identified by MacBride (1964) as Solanum dulcamara. In my view this is a specimen of the pinnate-leaved seedling of Solanum uncinellum, not a distinct species, nor is it a southern range extension of Solanum dulcamara.

Leaf size and texture can also vary enormously depending on the environment or position on the stem. In Solanum triquetrum, which grows in arid environments in the southwestern USA, leaves formed in wet periods are larger, sometimes by several times, than those from dry periods (see Figure 97). Leaf texture varies across the group from very delicate and membranous in some temperate species (e.g., Solanum lyratum) to thick and coriaceous in taxa from high elevations (e.g., Solanum macbridei Hunz. & Lallana). Turreson (1922) grew thick-leaved plants of Solanum dulcamara from seaside habitats in Sweden in common inland gardens with thin-leaved plants from more inland forest areas and found that the coastal forms developed thinner (and less pubescent) leaves away from their natural habitat. Clough et al. (1979), also working with Solanum dulcamara, found that light levels were a primary factor influencing leaf morphology and physiological performance; a major effect was on leaf specific weight, or thickness. Leaf texture and morphology are both important species specific characters in the group, although within species variation in some taxa is very large.

Pubescence. Trichome types and density are very useful for species recognition in Solanum. Previous classifications of the genus (Seithe 1962) have been constructed using primarily trichome types, with groups defined by possession of stellate, branched or simple trichomes. Later research (e.g., Knapp 2002a; Bohs 2005) has shown that trichome types do not define monophyletic groups in Solanum, but that they can be useful at the species level. As with many of the vegetative features of members of the Dulcamaroid clade, trichomes are very variable (see Figure 3 for trichome types). Trichomes of members of the Dulcamaroid clade are either uniseriate (consisting of a single file of cells) and either simple or branched, or multiseriate and echinoid and densely dendritic (Roe 1971; Knapp 1989; Mentz and Stehmann 1992). Species of the Solanum nitidum group (sensu Knapp 1989) have branched or densely branched trichomes; some of these approach the echinoid and tree-like trichomes characteristic of the Brevantherum clade (see Roe 1971) with numerous very short branches clustered at the tip (e.g., Solanum stenophyllum and Solanum storkii).

Simple trichomes are either eglandular or glandular, with the glands usually consisting of a single cell. Solanum kulliwaita S.Knapp has unusual bead-like terminal cells on the trichomes of the inflorescence that all appear to be glandular (Figure 52); this type of trichome has also been observed in Nicotiana L. in Australia (Marks et al. 2011). Glandular trichomes are very consistently present in some species (e.g., Solanum lyratum, Solanum viscosissimum), while in others possession of glandular or eglandular trichomes appears to be polymorphic within species (e.g., Solanum endoadenium, Solanum umbelliferum). Possession of glands has often been used as an infraspecific marker in other groups of Solanum (for example some species of section Solanum, Edmonds 1982) but molecular work with these taxa has shown that the trichome types do not define monophyletic groups within these species but instead are polyphyletic and represent polymorphism (Oyet 2004; Manoko 2007).

Terminal glands also occur on branched trichomes, but these trichomes are more typically eglandular. Several different branched trichome types occur in the Dulcamaroid clade: 1) uniseriate trichomes with few branches (e.g., Solanum sanchez-vegae S.Knapp, Figure 84), 2) uniseriate trichomes with many congested branches (e.g., Solanum aureum, Figure 20), 3) multiseriate or uniseriate trichomes with densely congested very short branches (tree-like or “Tannenbaumartig” sensu Seithe 1962, e.g., Solanum stenophyllum, Figure 94) and 4) multiseriate echinoid trichomes (e.g., Solanum storkii, Figure 95). These latter two types have very short, often unicellular branches and are found only in the Solanum nitidum group.

Inflorescences. The inflorescences of members of the Dulcamaroid clade, as in most Solanaceae, are terminal with a subtending lateral bud. The inflorescence usually occupies a position at the end of branches, but if growth continues from the axillary bud it can appear lateral. In some species the inflorescences are borne on what appear to be short shoots with very reduced leaves (e.g., Solanum inodorum, Solanum valdiviense Dunal); the flowering and non-flowering shoots of these plants are very divergent morphologically and the short shoots often have smaller leaves (see Figures 49, 104). Solanum inodorum is extreme in this regard with the flowering shoots seeming to be axillary in larger leaves. In fact, the inflorescence is terminal on a short shoot with much reduced, almost bract-like, leaves.

The basic inflorescence, as in all other species of Solanum, is a scorpoid cyme with a variety of branching patterns. In the Dulcamaroid clade inflorescences are usually very large and highly branched. Turrill (1935) made crosses between a form of Solanum dulcamara with 1-flowered inflorescences he called “uniflora” and the more typical many-branched form, and suggested that the branches of the complex inflorescence were dichotomous and that subtending bracts were suppressed. Lippman et al. (2008) found that the highly branched inflorescence of Solanum crispum was developmentally similar to compound inflorescence (s) mutants of tomato; each shoot inflorescence meristem produces multiple flowers in a proliferating manner. They suggest that delays in floral termination (perhaps mediated by S, or the genetic pathway that S defines) coupled with sequential expression of AN (anantha, another tomato inflorescence mutant) provide a developmental framework for the modulation of sympodial branching in the entire family. In the Dulcamaroid clade flower number varies from 2-3 (Solanum salicifolium, Figure 82) to more than 100 (Solanum sanchez-vegae, Figure 84); within a plant inflorescence size can vary greatly with inflorescence age as each inflorescence continues to branch and produce flowers throughout its “life”.

In any given inflorescence only a few flowers are open at a time (up to about 40-50 in species with large inflorescences like Solanum sanchez-vegae). Flowers are generally more congested near the inflorescence branch tips; in Solanum aligerum and Solanum pubigerum the flowers are grouped onto tiny “platforms”, or highly congested groups of 3-5 flowers (see Knapp 1989: Figure 2b; Figures 10, 74). The pedicel scars are irregularly and relatively widely spaced, and the space between scars is generally greater more basally in the inflorescence. Differentiation between the peduncle and the end of the shoot is difficult to determine in most species, but in some species (e.g., Solanum kulliwaita), the pubescence of the inflorescence is distinct from that of the shoot.

Pedicels. The pedicels of species in the Dulcamaroid clade are distinctive in being inserted into a tiny sleeve of tissue that appears to arise from the inflorescence rhachis (see Figure 2c in Knapp 1989); this is one of the few morphological synapomorphies for the clade. This is most pronounced in the species of the Solanum nitidum group, in which the sleeve is sometimes up to 2 mm long and forms a small cup in which the base of the pedicel sits (Figure 2C). In most species the pedicel is articulated slightly above the base, leaving a small peg or cup on the inflorescence axis; this accentuates the zig-zag appearance of the inflorescence architecture that results from sympodial growth. The pedicel articulation in Solanum boldoense Dunal & A.DC. of Cuba is unusual in being in the distal third of the pedicel, often directly beneath the calyx tube (see Figure 22). The colour of the pedicel sleeve is sometimes (e.g., Solanum dulcamara) different from the pedicel (see Figure 4D), further supporting the idea that the sleeve is anatomically part of the inflorescence rhachis.

Figure 4.

Inflorescences of members of the Dulcamaroid clade. A Open cyme of Solanum dulcamaroides (cultivated in Nijmegen, The Netherlands A44750197) B Grouped flowers on platforms of Solanum aligerum (Knapp et al. 10436, Cusco, Peru) C Pedicel sleeve of Solanum nitidum (Knapp et al. 10222, Huancavelica, Peru) D Contrasting pedicel colour in Solanum valdiviense (cultivated in Royal Botanic Garden, Edinburgh, UK, Knapp IM-10105).

Pedicel orientation in the species of the group is generally deflexed or slightly nodding at anthesis. In fruit, those species with pendent inflorescences generally have deflexed pedicels, but shrubby species such as those of the Solanum nitidum group have erect pedicels in fruit (e.g., Solanum coalitum S.Knapp).

Calyces. The calyx is typically sympetalous and 5-merous with the tube and lobes more or less equal in size. The shape of the calyx lobes varies considerably and is a useful character for species identification. Lobes vary from mere undulations of the calyx rim (e.g., Solanum boldoense, Solanum uncinellum) to deltate (many species), quadrate (e.g., Solanum aligerum, Solanum pyrifolium Lam.) or lanceolate (e.g. Solanum imbaburense S.Knapp); apices are usually acute to acuminate, but are occasionally rounded (e.g., Solanum angustifidum) or elongate (e.g., Solanum viscosissimum). Pubescence of the calyx tube and lobes usually parallels that of the pedicels and inflorescence rhachis, but is generally sparser.

Corollas. The corolla of members of the Dulcamaroid clade is sympetalous, 5-merous and regular, in common with the vast majority of Solanum species. Color is either white or varying shades of purple; many species have populations of both color forms, and in Solanum dulcamara these have been described many times at the infraspecific level. Solanum laxum growing in full sun often has pale violet rather than the more commonly encountered white flowers, but the degree to which flower color is environmentally or genetically determined is not clear. At the base of the corolla tube is often a ring or irregular area of differently colored tissue refered to as the eye. In the group of species found in the temperate zone (e.g., Solanum dulcamara, Solanum lyratum, Solanum umbelliferum) at the base of each corolla lobe is a small pair of shiny green dots often ringed with darker green or white tissue. These have been variously interpreted as “pseudo-nectaries” functioning in pollinator attraction (Buchmann et al. 1977), but specific field studies have not been undertaken to test this. This character can be difficult to see in herbarium specimens. The center part of Solanum flowers is usually UV absorbent and seen by bees as a dark guide to the prominent anther cone (Buchmann et al. 1977). Solanum endoadenium has a prominent green eye, but the tissue is not shiny, perhaps indicating it is not homologous with the similar structure in Solanum dulcamara (see Figure 5).

Figure 5.

Flowers of members of the Dulcamaroid clade. A Solanum lyratum (Knapp et al. 10142, Hubei, China) B Solanum nitidum (Knapp et al. 10222, Huancavelica, Peru) C Solanum endoadenium (Barboza et al. 3476, Catamarca, Argentina) D Solanum dulcamara (Kent, UK) E Solanum laxum (cultivated in London, UK) F Solanum uncinellum (cultivated in Nijmegen, The Netherlands 994750113).

Corolla shape varies from deeply stellate (Solanum uncinellum, Figures 5, 101) to pentagonal (Solanum wallacei, Figure 108). The lobes are triangular and usually either spreading (Solanum dulcamaroides, Figures 5, 38) or strongly reflexed (Solanum lyratum, Figures 5, 61) at anthesis. Solanum macbridei is unusual in having campanulate corollas (Figure 63). The tips are usually somewhat cucullate (e.g., Figure 14D) and are papillose to densely pubescent. In those species with dense overall pubescence, the abaxial surface of the corolla lobes is also usually densely pubescent. Corolla diameter varies from approximately 1 cm (Solanum endoadenium) to almost 5 cm (Solanum wallacei); most species have corollas that range from 1.5–3 cm in diameter.

Anthers. Anthers of members of the Dulcamaroid clade conform to the typical poricidal morphology of all other species of “non-spiny” Solanum (see Knapp 2001, 2002a). In most taxa the pore usually “unzips” during anther dehiscence to form a tear-drop shaped slit (Figure 5 and species illustrations), from which pollen is shed during vibratile pollination (Buchmann 1986). Some species of the Dulcamaroid clade have pores that do not elongate at all (e.g., Solanum dulcamaroides, Solanum seaforthianum), or only elongate slightly with age (e.g., Solanum valdiviense). Anthers in the group are generally ellipsoid, but those of Solanum dulcamara and Solanum uncinellum are elongate and tapering and those of Solanum dulcamaroides and Solanum boldoense are almost spheroidal. Anthers of members of most species are loosely connivent, while those of Solanum dulcamara are tightly connivent, and are held together physically with a sticky “glue” (Glover et al. 2004) such that the pores act as a single opening to a visiting bee. The anthers of Solanum aureum occasionally appear pubescent due to the prescence of elongate papillae on the dorsal surface (see Figure 20).

All five anthers in most of the species of the group are morphologically identical; some taxa, however, have filaments of different length, making the flowers zygomorphic. This filament length difference ranges from quite subtle (Solanum seaforthianum, Figure 86) to very conspicuous (Solanum uncinellum, Figure 5F, 101). Flowers of Solanum flaccidum Vell. that have just opened have filaments of equal length; the long filament apparently elongates over the course of anthesis. This has been observed in other species of Solanum (Solanum turneroides Bitter and Solanum evolvuloides Giacomin & Stehmann, members of section Gonatotrichum Bitter of the Brevantherum clade, see Stern et al. in press) and in the genus Lycianthes Hassl. (Dean 2001).

Pollen of members of the group is similar in size and shape to that of other species of Solanum (Buchmann 1986); it is tricolporate with a relatively smooth exine and held in yellow “pollen shamming” anthers. Some plants of Solanum lyratum have dark purple anthers; this appears not to be related to light or exposure but may be genetic. I have not seen dark anthers in any other member of the group, but anther color is variable in the Cyphomandra clade (section Pachyphylla Dunal, Bohs 2004). Nitrogen and protein content of pollen grains is the same as for other species of Solanum (Buchmann 1986), and is typical for a buzz-pollinated plant.

Gynoecium. The gynoecium is typically bicarpellate; the carpels are fused in a superior ovary with axillary placentation. The ovary is usually conical to globose or slightly ellipsoid and usually glabrous; if pubescent; the trichomes are only sparsely present at the apex (e.g., Solanum cutervanum). The flowers lack nectaries, as do all other species of Solanum. The style is simple, straight or slightly curved, glabrous or variously pubescent, and is exserted beyond the anthers in all but Solanum luculentum S.Knapp (see below). The stigma is capitate (e.g., Solanum valdiviense, Solanum umbelliferum) to clavate (e.g., Solanum amygdalifolium) and is sometimes distinctly bilobed (e.g., Solanum uncinellum). The ovules are anatropous and non-arillate.

Fruits. As with all species of Solanum, the fruit is a bicarpellate berry. Fruits of members of this group are usually brightly colored and juicy (see Figure 6 for respresentative photographs). Many species change fruit color through maturation; for example, Solanum nitidum has green immature fruits that pass through a stage of being bright red before turning dark purple at full maturity. This makes it difficult to use fruit color as an identification aid, and in Solanum uncinellum fruit color has been variously recorded on labels as blue, dark purple, red and white. Without in-depth field studies it is unclear whether these represent populational or regional differences or maturational changes. Species here regarded as synonyms of Solanum umbelliferum in California have sometimes been distinguished using “mature” fruit color (either green or black; Nee 1993b), but such color polymorphisms are found in other Solanum species (most notably Solanum nigrum L. in Europe, see Manoko 2007). Fruit surface morphology is usually shiny, but in some species the pericarp is distinctly dull and non-shiny (e.g., Solanum sanchez-vegae). Solanum endoadenium has bright orange fruits that appear to dry on the plant and release the seeds (see Figure 6D); this has not been studied in detail. Fruit diameter varies from less than 1 cm (Solanum angustifidum, Solanum viscosissimum, Solanum pittosporifolium Hemsl.) to approximately 4 cm (Solanum wallacei).

Figure 6.

Fruits of members of the Dulcamaroid clade. A Solanum pittosporifolium (Knapp et al. 10136, Jiangxi, China) B Solanum salicifolium (Barboza et al. 3467, Catamarca, Argentina) C Solanum storkii (Knapp & Monro 10053, Bocas del Toro, Panama) D Solanum angustifidum (Barboza et al. 3489, Argentina) F Solanum nitidum (Knapp et al. 10136, Cusco, Peru).

Seeds. Seed morphology has proved very useful in Solanum taxonomy. Enzymatic digestion of the outer testal walls reveals great variety in the morphology and structure of the lateral testal cell walls, varying both between and within groups (Whalen 1979; Lester and Durrands 1984; Knapp and Helgason 1997). Souèges (1907) provided an early study of Solanum seeds, including Solanum dulcamara. He described the development of the integument and recognised outer and inner layers of the seeds. Seeds of members of the Dulcamaroid clade are oval, obovate or kidney-shaped in outline and flattened laterally. The cells of the outer epidermal layer in some species (e.g., Solanum dulcamaroides, Solanum lyratum, Solanum uncinellum, Solanum umbelliferum) develop radial wall thickenings that form as “hair-like outgrowths” or “pseudohairs” in mature seeds (Lester and Durrands 1984; Lester 1991). These hair-like outgrowths often greatly enlarge the outer layer of the integument and the seed coat appears pubescent; seed measurements here include these projections. In seeds from mature fruits the pseudohairs are translucent, connate or fused laterally to each other and tightly adpressed to the epidermis giving a silky appearance to the seed surface, or if long and distinct produce a hairy and shaggy seed surface. In some species (e.g., Solanum dulcamaroides) the pseudohairs form a prominent wing around the margin of the seed. The testal cells form a reticulate or honeycomb pattern with cell outlines at the basal portions deeply sinuous and irregular (e.g., Solanum boldoense), pentagonal/rectangular (e.g., Solanum seaforthianum, Solanum pubigerum) or somewhat intermediate (e.g., Solanum triquetrum).

Seed number per berry varies from few (fewer than 10, Solanum leiophyllum Benth., Solanum triquetrum) to many (more than 100, Solanum wallacei), but most species fall in the range of 20-50 seeds per berry. Seed size varies from 1.5 mm length and 1 mm width to 8 mm length and 6 mm width. Color varies from yellow or pale brown to dark brown.

Biology and natural history

Habitats and distribution. Members of the Dulcamaroid clade are both tropical and temperate, and occur in a wide range of habitats from deserts to lowland rainforest. Taxa occurring in the temperate areas of Asia and Europe occupy a wide range of habitats and elevations; Solanum dulcamara, for example, is found on sea coasts and extends into high elevations in the mountains of Italy and the Alps. Genetic work done in the early 20th century (Turesson 1922) revealed some hereditary component to the immense morphological variation in this species, but habitat is also important (Turrill 1935). Solanum lyratum has been collected from sea level to more than 2000 m and is found in many forest types and even in weedy gardens in cities. Most of the species of the group are not of conservation concern (see Table 3 and species treatments) but some have narrow distributions and are assessed as endangered or vulnerable.

Most species in the group are South American, and in the Americas the species of the group range from southern Oregon (USA) to central Argentina and onto the islands of the Caribbean. In South America highest species diversity occurs in two areas, the forests of Atlantic coastal Brazil and the eastern slope of the Andes from Colombia to Argentina (see Table 2). This circum-Amazonian pattern of species richness is common in other groups of Solanum (Knapp 2002b); endemism follows the same pattern. Argentina has the highest species diversity in the Dulcamaroid clade (with Peru and Ecuador very close behind), due in part to the mixing of species from these two centres of species richness, in addition to several endemics or near endemics along the Andean flanks.

Table 2.

Geographical distribution of New World species of the Dulcamaroid clade (species in square brackets are introduced or cultivated).

Country Species
United States of America [Solanum dulcamara], [Solanum laxum], Solanum triquetrum, Solanum umbelliferum, Solanum wallacei
Mexico Solanum aligerum, Solanum dulcamaroides, Solanum pubigerum, Solanum sousae, Solanum triquetrum, Solanum umbelliferum
Guatemala Solanum dulcamaroides, Solanum muenscheri, Solanum pubigerum, Solanum aligerum
Honduras Solanum pubigerum, Solanum aligerum
El Salvador Solanum dulcamaroides, Solanum pubigerum, Solanum aligerum
Nicaragua Solanum dulcamaroides, Solanum pubigerum, Solanum aligerum
Costa Rica Solanum aligerum, Solanum pubigerum, Solanum seaforthianum, Solanum storkii, Solanum uncinellum
Panama Solanum aligerum, Solanum uncinellum
Colombia Solanum aspersum, Solanum aureum, Solanum dichroandrum, [Solanum laxum], Solanum luculentum, Solanum seaforthianum, Solanum stenophyllum
Venezuela Solanum dichroandrum, Solanum luculentum, Solanum seaforthianum, Solanum uncinellum
Guyana Solanum uncinellum
Surinam Solanum uncinellum
French Guiana Solanum uncinellum
Ecuador Solanum agnoston, Solanum aspersum, Solanum aureum, Solanum coalitum, Solanum imbaburense, Solanum leiophyllum, Solanum nitidum, Solanum stenophyllum, Solanum uncinellum
Peru Solanum aligerum, Solanum aureum, Solanum cutervanum, Solanum kulliwaita, Solanum macbridei, Solanum nitidum, Solanum ruizii, Solanum sanchez-vegae, Solanum uncinellum
Bolivia Solanum aligerum, [Solanum angustifidum], Solanum endoadenium, Solanum nitidum, Solanum uncinellum
Brazil Solanum amygdalifolium, Solanum flaccidum, Solanum inodorum, Solanum odoriferum, Solanum laxum, Solanum uncinellum, Solanum viscosissimum
Paraguay Solanum amygdalifolium, Solanum flaccidum, Solanum laxum, Solanum uncinellum
Argentina Solanum aligerum, Solanum amygdalifolium, Solanum angustifidum, Solanum calileguae, Solanum crispum, Solanum endoadenium, Solanum laxum, [Solanum odoriferum], Solanum salicifolium, Solanum uncinellum, Solanum valdiviense
Uruguay Solanum amygdalifolium, Solanum laxum
Chile Solanum alphonsei, Solanum crispum, Solanum valdiviense
Cuba Solanum boldoense, Solanum seaforthianum
Haiti/Dominican Republic Solanum pyrifolium, Solanum seaforthianum
Lesser and Greater Antilles Solanum seaforthianum
Table 3.

Conservation assessments for members of the Dulcamaroid clade (calculated using methods in Moat 2007). For details see individual species treatments.

Species Preliminary Conservation Assessment
Solanum agnoston S. Knapp DD (Data Deficient)
Solanum aligerum Schltdl. LC (Least Concern)
Solanum alphonsei Dunal EN (Endangered)
Solanum amygdalifolium Steud. LC (Least Concern)
Solanum angustifidum Bitter LC (Least Concern)
Solanum aspersum S. Knapp VU (Vulnerable)
Solanum aureum Dunal LC (Least Concern)
Solanum boldoense Dunal & A.DC. NT (Near Threatened)
Solanum calileguae Cabrera EN (Endangered)
Solanum coalitum S. Knapp CR (Critically Endangered)
Solanum crispum Ruiz & Pav. LC (Least Concern)
Solanum cutervanum Zahlbr. LC (Least Concern)
Solanum dichroandrum Dunal LC (Least Concern)
Solanum dulcamara L. LC (Least Concern)
Solanum dulcamaroides Poir. LC (Least Concern)
Solanum endoadenium Bitter LC (Least Concern)
Solanum flaccidum Vell. LC (Least Concern)
Solanum imbaburense S. Knapp DD (Data Deficient)
Solanum inodorum Vell. LC (Least Concern)
Solanum kulliwaita S.Knapp NT (Near Threatened)
Solanum laxum Spreng. LC (Least Concern)
Solanum leiophyllum Benth. NT (Near Threatened)
Solanum luculentum S. Knapp LC (Least Concern)
Solanum lyratum Thunb. LC (Least Concern)
Solanum macbridei Hunz. & Lallana NT (Near Threatened)
Solanum muenscheri Standl. & Steyerm. EN (Endangered)
Solanum nitidum Ruiz & Pav. LC (Least Concern)
Solanum odoriferum Vell. LC (Least Concern)
Solanum pittosporifolium Hemsl. LC (Least Concern)
Solanum pubigerum Dunal LC (Least Concern)
Solanum pyrifolium Lam. NT (Near Threatened)
Solanum ruizii S. Knapp VU (Vulnerable)
Solanum salicifolium Phil. LC (Least Concern)
Solanum sanchez-vegae S. Knapp NT (Near Threatened)
Solanum seaforthianum Andrews LC (Least Concern)
Solanum septemlobum Bunge LC (Least Concern)
Solanum sousae S. Knapp NT (Near Threatened)
Solanum stenophyllum Dunal LC (Least Concern)
Solanum storkii C.V.Morton & Standl. VU (Vulnerable)
Solanum triquetrum Cav. LC (Least Concern)
Solanum umbelliferum Eschsch. LC (Least Concern)
Solanum uncinellum Lindl. LC (Least Concern)
Solanum valdiviense Dunal LC (Least Concern)
Solanum viscosissimum Sendtn. LC (Least Concern)
Solanum wallacei (A. Gray) Parish CR (Critically Endangered)

Floral biology and pollination. Most Solanum species have hermaphroditic flowers, but in some groups derived sexual systems are common (Whalen and Costich 1986; Anderson and Symon 1989; Knapp et al. 1998; Martine et al. 2009). Andromonoecy is very common in the Leptostemonum clade (Whalen and Costich 1986; Levin et al. 2006), as is dioecy (Anderson and Symon 1989; Martine et al. 2009). Dioecy is phylogenetically widespread in Solanum, with dioecious species occurring primarily in the spiny solanums (subgenus Leptostemonum Bitter, Anderson and Symon 1989), but also in the non-spiny solanums in the Potato (Anderson and Levine 1982), the Geminata (Knapp et al. 1998) and Brevantherum (S. Knapp, pers. obs., Solanum punctulatum Dunal from Jamaica) clades. Solanum luculentum of the Dulcamaroid clade has all the morphological correlates of a dioecious species, but has not yet been examined in detail to determine whether or not it is indeed truly dioecious. If so, Solanum luculentum represents the independent evolution of dioecy in an otherwise completely hermaphroditic group, rather than within a predominantly andromonoecious clade, as is common in the spiny solanums (Martine et al. 2009). The evolution of dioecy in a group with strictly hermaphroditic flowers has also occurred in the Potato clade (e.g., Solanum appendiculatum Dunal, see Anderson and Levine 1982).

Self-compatibility (SC) is widespread in Solanum (Whalen and Anderson 1981) and is essentially irreversible once acquired from the ancestral self-incompatible (SI) state ( 2003, 2006). Only three species of the Dulcamaroid clade have been assessed for their compatibility status: Solanum dulcamara, Solanum laxum and Solanum wallacei (Goldberg et al. 2010). Of these, only Solanum laxum has been shown to be SI (Whalen and Anderson 1981), but Solanum dulcamara flowers failed to set seed when isolated from visiting insects (Turrill 1935) and Cruden and Lyon (1985) scored Solanum dulcamara from Iowa as SI (“xenogamous”). Other authors, however, have stated that Solanum dulcamara is SC (Eijlander and Stiekema 1995; Vallejo-Marín and O’Brien 2006). In greenhouses and field plots in Nijmegen (Netherlands) self-crosses of Solanum dulcamara proved difficult and set few seed (G. van der Weerden, pers. comm., October 2010). There may be a geographic component to self-compatibility in Solanum dulcamara; those reporting SC have used American collections (naturalised) and those reporting SI have used European accessions. Buchmann et al. (1977) report that bagged flowers of Solanum umbelliferum (as Solanum xanti) failed to set fruit and concluded it was SI; this species, however, has not been included in recent large scale analyses of SI in the family (Goldberg et al. 2010). Solanum crispum grown in European gardens rarely sets fruit, and attempts to self flowers resulted in 0% fruit set (pers. obs.), but these experiments were not done in a controlled manner. That Solanum wallacei, an island endemic with a very narrow range, is SC is not at all surprising; it will be interesting to assess again the compatibility status of its close relative Solanum umbelliferum, which has been thought to be SI (Buchmann et al. 1977). Goldberg et al. (2010) have shown that lineages with self-incompatibility in the Solanaceae are more species-rich, due in part to higher extinction rates in SC lineages. It appears from the limited data available in the Dulcamaroid clade that widespread, clonal species (e.g., Solanum laxum, Solanum dulcamara, Solanum crispum) are SI, supporting the results of Vallejo-Marín and O’Brien (2006).

Flowers of all members of the Dulcamaroid clade are typical for Solanum and are buzz-pollinated by a wide variety of bees. In temperate regions species are visited by various species of bumblebees (Bombus); Solanum dulcamara and cultivated Solanum laxum and Solanum crispum in Europe are both visited by several species of bumblebees (pers. obs.). I have also observed bumblebees visiting several of the montane tropical species of the group (e.g., Solanum nitidum in Peru and Bolivia, Solanum storkii in Costa Rica), and Bombus species were common vibratile pollinators of Solanum umbelliferum (as Solanum xanti) in southern California (Buchmann et al. 1977). Pollen grains left on the anthers or near the pores by vibratile bees were collected by the gleaning bees from Solanum umbelliferum (Andrena spp., Andrenidae; Stenodynerus cochisensis (Viereck), Eumenidae; Buchmann et al. 1977). These bees are not as efficient at effecting pollination, however, as they are not necessarily in contact with the stigma on every visit.

Figure 7.

Habitats of members of the Dulcamaroid clade. A Gravel shingle at Rye Harbour Nature Reserve, coastal Sussex, UK (with Solanum dulcamara) B Dry forest grassland chaco habitat in Formosa, Argentina (with Solanum amygdalifolium) C Ceja de selva (near timberline) in Andean Peru (Cusco above Ollantaytambu) D High elevation dunes in Catamarca, Argentina (with Solanum endoadenium) E Eastern Andean slopes of Peru, premontane cloud forest F Rocky river course in the Lushan Mountains of Jiangxi, China.

Species concepts

My goal for this revision has been to provide species circumscriptions for the members of this large and morphologically variable clade, while clearly highlighting those taxa where further in-depth research would be useful. Delimitation of species here basically follows what is known as the “morphological cluster” species concept (Mallet 1995): i.e., “assemblages of individuals with morphological features in common and separate from other such assemblages by correlated morphological discontinuities in a number of features” (Davis and Heywood 1963). Biological (Mayr 1982), phylogenetic (Cracraft 1989) and the host of other finely defined species concepts (see Mallet 1995) are almost impossible to apply in practice when dealing with large, complex groups such as the clades of Solanum, and are therefore of little utility in a practical sense. It is important, however, to clearly state the criteria for the delimitation of species, rather than dogmatically follow particular ideological lines (see Luckow 1995; Davis 1997). My decisions relied on clear morphological discontinuities to define the easily distinguished species. Specific characters used for recognition are detailed with each species description and in the keys. Potential reasons for variability and intergradation are recent divergence and hybridization. In this revision I have tried to emphasise similarities between populations instead of differences, which so often reflect incomplete collecting or local variation. I have not recognised subspecies or varieties, as I do not feel these are useful categories, either in a taxonomic or evolutionary sense. The variation is better described and documented, rather than formalised with a name which then encumbers the literature. I have been conservative in my approach, recognising as distinct entities those population systems (sets of specimens) that differ in several morphological characteristics. Minor differences in morphology, distribution, habitat, and ecology are important in some groups, where the common groundplan for the species is very similar. On the other hand, I have recognised several extremely widespread, polymorphic species, e.g., Solanum dulcamara, Solanum lyratum, Solanum umbelliferum, Solanum uncinellum and Solanum viscosissimum. In these species variation exists in certain characters, but the pattern of variation is such that no reliable units can be consistently extracted. In these cases the variability within and between populations seems more important than the variations of the extremes other taxonomists have recognised as distinct. In these cases I have described the variation, realizing that others may wish to interpret it differently.

Materials and methods

Approximately 9000 collections were examined for this monograph from herbaria worldwide; specimens were seen at the following herbaria (all abbreviations from Index Herbariorum, http://sciweb.nybg.org/science2/IndexHerbariorum.asp ): A, AAU, B, B-W, BH, BM, BOLV, BRI, C, CAS, CICY, COL, CORD, DS, E, EA, ECON, EIU, F, FCQ, G, G-DC, GH, GOET, HA, HIB, HITBC, HBR, HUA, JBSD, JEPS, K, L, LAGU, LE, LIL, LOJA, LPB, MA, MBM, MEXU, MO, MOL, NY, OXF, P, PE, PMA, POM, Q, QCA, QCNE, RSA, S, SGO, SI, TI, UC, US, USM, UT, W, WIS, WU. Many of the species of the Dulcamaroid clade are common and widespread (e.g., Solanum dulcamara, Solanum laxum, Solanum uncinellum), so only selected specimens are cited; these represent the entire species range. All numbered collections are cited in the Index to Numbered Collections. Any collections without specific numbers are cited as s.n. (sine numero) in the Specimens examined sections of each species. Collection details for all specimens used for this monograph can be found on the Solanaceae Source website (http://www.solanaceaesource.org ) and in the data file available with this paper (see Supplementary Material). The specimens of Solanum at the New York Botanical Garden (NY) were databased as part of the PBI project; those specimens are best accessed through the NY Virtual Herbarium (http://sciweb.nybg.org/science2/VirtualHerbarium.asp ).

The JStor Plant Science website (http://plants.jstor.org ), established to display the images of type specimens digitised as part of the GPI (Global Plants Initiative) project funded by the Andrew W. Mellon Foundation, was an invaluable resource for images of type specimens, and many of the isotypes identified here were discovered via the website. Many type specimens in European herbaria were photographed in the early 20th century by J.F. Macbride of the Field Museum (http://fieldmuseum.org/explore/our-collections/berlin-negatives ), including those at B, since destroyed. These photographs are cited here with a F negative number (F neg. #), but herbaria in which I have seen the photographs (usually mounted on sheets) are not cited. Conrad V. Morton of the Smithsonian also took photographs of many type and other nomenclaturally important specimens, these are cited as Morton negative numbers.

With all specimens of types I have cited the barcode or accession number of each sheet if possible. Barcodes are cited with no space between the herbarium acronym and the number (e.g., BM000845345), while for accession numbers I have inserted a hyphen between the acronym and the number (e.g., MO-00678987). Some herbaria such as MA, have barcodes and accession numbers that are the same, these are cited as on the barcode label (e.g., MA-634567); other herbaria such as MO have barcodes and accession numbers that differ, here I have cited the accession number as it is the only element visible without a barcode reader.

In the specimen citations I have selected specimens that represent the species range, and present these citations in alphabetical order by country rather than a geographical order. For those taxa that are introduced and often naturalised (Solanum laxum and Solanum seaforthianum) I have separated the specimen citations into those from countries in the native range (or presumed native range) and those from countries where the species is introduced or naturalised.

Citation of literature follows BPH-2 (Bridson 2004) with alterations implemented in IPNI (International Plant Names Index, http://www.ipni.org ) and Harvard University Index of Botanical Publications (http://kiki.huh.harvard.edu/databases/publication_index.html ). In particular, I have used the square bracket convention for publications in which a species is described by one author in a publication edited or compiled by another – the traditional “in” attributions such as Dunal in DC. for those taxa described by Dunal in Candolle’s Prodromus Systematis Naturalis Regni Vegetabilis; this work is cited here as Prodr. [A.P de Candolle] and the names thus attributed only to Dunal. For “ex” attributions I have cited only the publishing author, as suggested in the Code (McNeill et al. 2012). Author names are abbreviated according to IPNI (International Plant Names Index, http://www.ipni.org ).

Each species treated here has been assessed for a preliminary conservation status using the GIS protocols of Moat (2007) that involve calculation of AOO (Area of Occupancy) and EOO (Extent of Occurrence) from georeferenced specimen data. A grid cell size of 10% of the EOO was used (see Moat 2007); additional analyses using the 2 km2 grid size as recommended by IUCN (2001) were done, but specimen collection density made the results difficult to interpret. When threat status as determined by EOO and AOO differ, I have been conservative and selected the higher (more threatened) assessment as the overall assessment of the species (for an example, see Solanum alphonsei), and have used my knowledge of the biology of these species to suggest amendments to these calculated values. These assessments are not complete in the sense of IUCN (2001), but are indicative of the threat status for the taxa treated here. Any species known from fewer than 5 collections has been assessed as Data Deficient (DD); this does not mean these are not of conservation concern, but that the available data are not sufficient for their status to be modelled using the AOO/EOO method.

Taxonomic treatment
Generic and clade descriptions
Solanum L., Sp. pl. 184. 1753.

http://species-id.net/wiki/Solanum

Lectotype species

designated by Henderson 1974: Solanum nigrum L. [I accept here the generic synonymy of D’Arcy (1972, 1974) with the addition of Amatula Moench, Cyphomandra Sendtn., Lycopersicon Mill., Normania Lowe, and Triguera Cav.].

Description.

Herbs, shrubs, trees, or vines, with or without prickles, glabrous or pubescent with unbranched or branched (including stellate), often glandular hairs. Leaves alternate or paired and frequently unequal in size, simple to pinnately lobed or compound, petiolate or sessile, without stipules, but sometimes with “pseudostipules” (Potato clade). Inflorescences cymose, branched or unbranched. Flowers usually perfect, (4-) 5-merous, actinomorphic or zygomorphic; calyx campanulate, sometimes accrescent in fruit, corolla rotate, campanulate, stellate, or urceolate, white, green, yellow, pink, or purple; stamens equal or unequal, the filaments generally short and inserted at the corolla base, the anthers basifixed, equal or unequal, blunt or tapered toward apex, opening by terminal pores, these sometimes expanding into longitudinal slits, or introrsely longitudinally dehiscent with age in sect. Lycopersicon; ovary 2-carpellate; ovules many; style articulated at base or above the base, usually slender; stigma capitate to elongate-clavate. Fruit a berry, usually fleshy but occasionally dry, usually many-seeded, the seeds often flattened; embryo curved, embedded in abundant endosperm. Chromosome number: n = 12, 23, 24, 48.

Discussion.

The generic description applies to Solanum including all those genera traditionally segregated from it: Cyphomandra (Bohs 1995), Lycopersicon (Spooner et al. 1993; Peralta et al. 2008), Normania, and Triguera (Bohs and Olmstead 2001). Data from chloroplast DNA sequences strongly support the inclusion of these segregates in a monophyletic Solanum (Bohs 2005). Some workers (e.g., Hunziker 2001) maintain these taxa as distinct genera.

The “Dulcamaroid clade” sensu Bohs (2005)

Dulcamara Moench, Meth. 514. 1794.

Lectotype species, designated by D’Arcy 1972: Dulcamara flexuosa Moench (=Solanum dulcamara L.)


Solanum section Dulcamara (Moench) Dumort., Fl. Belg. 39. 1827.

Lectotype species, (designated by D’Arcy 1972: Dulcamara flexuosa Moench (=Solanum dulcamara L.)


Solanum subsection Holophylla G.Don, Gen. Hist. 2: 414.1832.

Lectotype species, designated by Seithe 1962: Solanum cervantesii Lag. (=Solanum pubigerum Dunal); superfluous lectotype designated by D’Arcy 1972: Solanum pulverulentum Pers. (=Solanum cutervanum Zahlbr.)


Solanum section Holophylla (G.Don) Walp., Repert. Bot. Syst. 3: 51. 1844.

Lectotype species, designated by Seithe 1962: Solanum cervantesii Lag. (=Solanum pubigerum Dunal); superfluous lectotype designated by D’Arcy 1972: Solanum pulverulentum Pers. (=Solanum cutervanum Zahlbr.)


Solanum subsection Dulcamara (Moench) Dunal, Prodr. [A.P. de Candolle] 13 (1): 28. 1852.

Lectotype species, designated by D’Arcy 1972: Dulcamara flexuosa Moench (=Solanum dulcamara L.)


Solanum grad. ambig. Dulcamara (Moench) Dunal, Prodr. [A.P. de Candolle] 13 (1): 28. 1852.

Lectotype species, designated by D’Arcy 1972: Dulcamara flexuosa Moench (=Solanum dulcamara L.)


Solanum grad. ambig. Anthoresis Dunal, Prodr. [A.P. de Candolle] 13(1): 28, 84. 1852.

Lectotype species, designated by Seithe 1962: Solanum cervantesii Lag. (=Solanum pubigerum Dunal); superfluous lectotype designated by D’Arcy 1972: Solanum pulverulentum Pers. (=Solanum cutervanum Zahlbr.)


Solanum grad. ambig. Subdulcamara Dunal, Prodr. [A.P. de Candolle] 13(1): 28, 84. 1852.

Lectotype species, designated by D’Arcy 1972: Solanum ipomoea Sendtn. (=Solanum uncinellum Lindl.)


Solanum section Andropedas Rusby, Mem. Torrey Bot. Club 4: 231. 1895.

Type species: Solanum styracioides Rusby (=Solanum uncinellum Lindl.)


Solanum section Dulcamara (Moench) Bitter, Bot. Jahrb. 54: 428. 1917, non section Dulcamara (Moench) Dumort. 1827.

Lectotype species, designated by D’Arcy 1972: Dulcamara flexuosa Moench (=Solanum dulcamara L.)


Solanum section Anthoresis (Dunal) Bitter, Bot. Jahrb. Syst. 54: 489. 1917, pro parte.

Lectotype species, designated by Seithe 1962: Solanum cervantesii Lag. (=Solanum pubigerum Dunal); superfluous lectotype designated by D’Arcy 1972: Solanum pulverulentum Pers. (=Solanum cutervanum Zahlbr.)


Solanum subsection Lysiphellos Bitter, Repert. Spec. Nov. Regni Veg. 16: 90. 1919.

Type species: Solanum decorticans Sendtn. (=Solanum inodorum Vell.)


Solanum section Lysiphellos (Bitter) Seithe, Bot. Jahbr. Syst. 81: 288. 1962.

Type species: Solanum decorticans Sendtn. (=Solanum inodorum Vell.)


Solanum section Jasminosolanum Seithe, Bot. Jahbr. 81: 291. 1962.

Type species: Solanum jasminoides Paxton (=Solanum laxum Spreng.)


Solanum subsection Nitidum A.Child, Feddes Repert. 109: 409. 1998.

Type species: Solanum nitidum Ruiz & Pav.


Solanum section Californisolanum A.Child, Feddes Repert. 109: 410. 1998.

Type species: Solanum umbelliferum Eschch.


Description. Vines or lax shrubs, woody at the base; stems weak and clambering or erect, sometimes winged, pubescent or glabrous, the trichomes unbranched or branched, never strictly stellate. Sympodial units plurifoliate. Leaves simple or pinnatifid, sometimes deeply so such that the leaves appear pinnate, green, glabrous or variously pubescent with branched or unbranched trichomes, the branched trichomes sometimes very compact and echinoid (multangulate); petioles well developed, in vines often twining to aid in climbing. Inflorescences terminal, sometimes appearing lateral due to stem growth, simple or more usually many times branched, with up to 100 flowers, not bracteate; peduncle present, often poorly distinguished from the leafy stem; pedicels inserted in a short sleeve of inflorescence tissue, articulated at the base inside the sleeve or just beneath the calyx (Solanum boldoense). Flowers 5-merous, actinomorphic or slightly zygomorphic, usually perfect, one species probably dioecious (Solanum luculentum); calyx 5-parted, glabrous or pubescent; corolla 5-parted, rotate to stellate, white to purple, sometimes with spots at the base of the lobes; stamens 5, the filaments equal or unequal, glabrous or pubescent; anthers yellow or occasionally purple, ellipsoid or tapering, separate to tightly connivent, dehiscing by terminal pores, these usually elongating to slits with age; ovary globose to conical, usually glabrous, occasionally pubescent; style straight or slightly curved; stigma capitate to clavate. Fruit a globose berry, yellowish green to red, orange or black when ripe, glabrous, the pericarp thin, shiny or matte; calyx lobes in fruit not accrescent. Seeds flattened, often appearing hairy from outgrowths of the lateral cell walls.

Discussion. As with many of the major clades of Solanum morphological synapomorphies in the Dulcamaroids are few (Bohs 2005), but members of the clade all have a distinctive pedicel “sleeve” (Knapp 1989) that appears to be an extention of the inflorescence axis in which the pedicel base sits. This “cushion of tissue” was described by Turrill (1935) in Solanum dulcamara as being of the color of the rhachis rather than the pedicel (see Figure 4D); this is the case in most species of the group, lending support to the idea that the “sleeve” is part of the inflorescence axis. Most members of the clade possess a combination of other characters that, although occurring elsewhere in Solanum, together distinguish the group; these are, 1) vining habit, 2) twining petioles, 3) large, terminal, branched inflorescences, 4) brightly colored fruits, 5) pinnatifid leaves at least in juvenile plants.

In the absence of a well-sampled phylogenetic study (see above), I have divided the Dulcamaroid clade into a series of morphologically and geographically delimited groups (see Table 4). These should not be interpreted as monophyletic groups or taxa, and future phylogenetic analyses including more species will certainly reveal structure within this large and variable clade. Preliminary phylogenetic analyses with a number of plastid regions (T. Särkinen, pers. comm.) have revealed that the division between the Morelloid and Dulcamaroid clades is not completely clear; for example, Solanum salicifolium and Solanum valdiviense are of ambiguous position. I have included them here on gross morphological grounds whilst realising their evolutionary relationships may be with other taxa. Species are presented here in alphabetical order without regard to putative evolutionary relationships.

Table 4.

Informal species groups in the Dulcamaroid clade. For country distribution see Table 2 and individual species descriptions.

Species group Component taxa Distribution
“dulcamara group” Solanum dulcamara, Solanum lyratum, Solanum pittosporifolium, Solanum septemlobum, Solanum triquetrum, Solanum umbelliferum, Solanum wallacei Northern Hemisphere
“South American oddball group” Solanum amygdalifolium, Solanum aspersum, Solanum endoadenium, Solanum inodorum, Solanum luculentum, Solanum odoriferum Andes, SE Brazil
“aureum group” Solanum agnoston, Solanum aureum, Solanum calileguae, Solanum dichroandrum, Solanum kulliwaita, Solanum sanchez-vegae Andes
“laxum group” Solanum alphonsei, Solanum laxum, Solanum pyrifolium, Solanum sousae, Solanum viscosissimum SE Brazil, West Indies, Mexico
“dulcamaroides group” Solanum boldoense, Solanum dulcamaroides, Solanum seaforthianum West Indies, Mexico and Central America
“Andropedas” Solanum angustifidum, Solanum flaccidum, Solanum uncinellum South and Central America
“nitidum group” Solanum coalitum, Solanum crispum, Solanum cutervanum, Solanum imbaburense, Solanum leiophyllum, Solanum macbridei, Solanum muenscheri, Solanum nitidum, Solanum ruizii, Solanum stenophyllum, Solanum storkii Montane South and Central America
“aligerum group” Solanum aligerum, Solanum pubigerum Mexico to southern South America, large disjunction
Incertae sedis (possibly Morelloid clade) Solanum salicifolium, Solanum valdiviense Southern South America

In the species treatments here I have cited all type specimens I have seen with barcodes or accession numbers if possible (see Materials and Methods for citation style). Those specimens I have not seen but whose existence is known from the literature or from others are indicated as n.v. (non visi, not seen). For many names I have been unable to find specimens and in many cases specimens were not cited explicitly, but if an author worked primarily in a particular herbarium, I have indicated types with ‘?’ to help in future tracing of type specimens; many of these are for the synonyms of the widespread Solanum dulcamara, synonymy decisions in these cases were based on descriptions. Some of the infraspecific taxa, particularly in the widespread European species Solanum dulcamara were described from floristic accounts and did not have specimens cited in their original descriptions; I have not neotypified these.


Artificial key to species of the Dulcamaroid Clade
1 Stems and/or leaves pubescent with variously branched trichomes 2
Stems and leaves glabrous or the majority of trichomes simple and uniseriate (not branched) 28
2 Trichomes congested (tightly branched or echinoid) on stems and leaf undersides 3
Trichomes loosely branched on stems and leaf undersides 9
3 Trichomes of stems and leaves dendritic with short branches, not echinoid 4
Trichomes of stems and leaves echinoid (many branches from a single point or along an apparent “stem”) 5
4 Vines with golden trichomes; inflorescences many times branched; Andes Solanum aureum
Shrubs with white or greyish white trichomes; inflorescences simple or at most furcate; Pacific coast of North America Solanum umbelliferum
5 Stem not winged from decurrent leaf bases; style pubescent 6
Stem winged from decurrent leaf bases; style glabrous (if style pubescent, then the leaves densely golden pubescent abaxially) 7
6 Flowers 4–4.5 cm in diameter; bark of older stems pale grey or white; central Peru Solanum ruizii
Flowers 1.5–2 cm in diameter; bark of older stems reddish brown; northern Peru and Ecuador Solanum cutervanum
7 Leaves strongly discolorous when dry, the abaxial lamina surfaces completely obscured by the golden pubescence Solanum stenophyllum
Leaves not strongly discolorous when dry, the abaxial lamina surface clearly visible 8
8 Calyx lobes deltate; trichomes on abaxial leaf surfaces in shallow pits; Central America Solanum storkii
Calyx lobes long-triangular; trichomes scattered on stems and leaf surfaces, not in pits abaxially; Andes (southern Ecuador) Solanum stenophyllum
9 Leaves coriaceous and shiny adaxially; venation often impressed or not easily visible above 10
Leaves membranous or somewhat fleshy, not markedly coriaceous and shiny; venation not impressed above 15
10 Trichomes sparse, confined to the leaf margins or in the axils near the midveins abaxially 11
Trichomes sparse to dense on the leaf lamina, at least abaxially 14
11 Leaves less than 2 cm long; corolla campanulate at anthesis; high Andes, Peru and Bolivia Solanum macbridei
Leaves longer than 2 cm long; corolla spreading at anthesis; Andes of Ecuador 12
12 Vines; leaves with dendritic pubescence in the primary vein axils beneath Solanum agnoston
Shrubs; leaves glabrous or with trichomes only at the margins 13
13 Calyx lobes long-triangular; northern Ecuador Solanum imbaburense
Calyx lobes deltate; southern Ecuador Solanum coalitum
14 Stem winged; corolla with dendritic trichomes abaxially; shrubs or possibly lax shrubs Solanum leiophyllum
Stem not winged; corolla papillate abaxially; woody vines Solanum sanchez-vegae
15 Trichomes of leaf undersides confined to the axils of the primary veins Solanum agnoston
Trichomes variously distributed on leaf undersides, not in tufts in primary vein axils 16
16 Buds turbinate and pointed; anthers elongate 17
Buds ellipsoid or globose, not pointed; anthers ellipsoid or globose, not tapering at the tips 18
17 Anthers borne on unequal filaments, free; corolla densely papillate externally; Neotropics Solanum uncinellum
Anthers borne on equal filaments, tightly fused into a cone; corolla variously papillate on margins; Eurasia, North America Solanum dulcamara
18 Stems strongly angled or winged 19
Stems round, not angled or winged 20
19 Inflorescences terminal, many times branched; berries black; Mexico to Argentina Solanum aligerum
Inflorescences borne on short shoots, simple or at most furcate; berries green or red; Chile Solanum valdiviense
20 Glandular trichomes present (on any part of the plant) 21
Glandular trichomes absent 22
21 Glandular trichomes on inflorescence only (these simple); Andes Solanum kulliwaita
Glandular trichomes on stems and leaves (these dendritic or simple); Pacific coast of North America Solanum umbelliferum
22 Pubescence of lax transparent trichomes, not greyish white when dry 23
Pubescence greyish white when dry (probably pale beige on live plants), the trichome walls not transparent 24
23 At least some leaves lobed to pinnatifid (some appearing completely pinnate); flowers white; stem smooth; Argentina Solanum calileguae
Leaves simple; flowers violet or white; stem warty from persistent leaf bases; Venezuela and Colombia Solanum dichroandrum
24 Flowers with shiny green and white spots at the base of the corolla lobes; Pacific coast of North America Solanum umbelliferum
Flowers uniformly purple or white (occasionally with a white “eye”, but not with shiny green spots); tropical Mexico to southern South America 25
25 Calyx lobes very small, broadly deltate; corolla fleshy, the tips of the lobes cucullate; anthers globose; style glabrous; woody vines; Mexico to Central America Solanum dulcamaroides
Calyx lobes of various sizes, not minute; corolla membranous; anthers ellipsoid; style pubescent; shrubs or lax shrubs 26
26 Mature berries black; leaves densely pubescent on both surfaces; Guatemala Solanum muenscheri
Mature berries red; leaves sparsely pubescent on both surfaces, or the abaxial surfaces more densely pubescent; South America 27
27 Leaf bases acute; leaf blades with 11–25 pairs of veins, these strongly parallel; Andes from Ecuador to Bolivia Solanum nitidum
Leaf bases truncate to cordate; leaf blades with 6–12 pairs of veins, not strongly parallel; coastal Chile to Andean Argentina, widely cultivated Solanum crispum
28 Leaves glabrous on the lower surface (usually both surfaces glabrous) 29
Leaves with at least some simple uniseriate trichomes on the surfaces (usually denser beneath or of equal density on both surfaces, if trichomes branched go to 1a) 47
29 Leaves of reproductive stems lobed or divided (if some leaves simple, then rarer than lobed leaves) 30
Leaves of reproductive stems simple (if some leaves lobed or divided, then rarer than simple leaves, both leaf types occasionally present in equal numbers in Solanum viscosissimum) 37
30 Anthers borne on filaments of differing lengths 31
Anthers borne on equal filaments 33
31 Flower buds turbinate and pointed; anthers elongate; abaxial surface of corolla lobes densely papillate (glabrous plants with lobed leaves extremely rare) Solanum uncinellum
Flower buds not pointed; anthers ellipsoid; abaxial surface of corolla lobes glabrous or sparsely papillate at the tips 32
32 Vines; leaf lobe apices acute; berries red; anthers 2–3 mm long; Caribbean, widely cultivated Solanum seaforthianum
Shrubs; leaf lobe apices rounded; berries black; anthers 5–6 mm long; Argentina and Bolivia Solanum angustifidum
33 Inflorescences simple or at most once-forked, with < 10 flowers; shrubs 34
Inflorescences many times branched, usually open, with > 10 flowers; vines or lax shrubs 35
34 Leaf base attenuate; leaves lanceolate to narrowly elliptic; Andes of Argentina Solanum salicifolium
Leaf base acute; leaves elliptic to ovate; Pacific coast of North America Solanum umbelliferum
35 Leaves shallowly lobed, rhombic in outline; Chile Solanum alphonsei
Leaves deeply lobed (pinnatifid), ovate or elliptic in outline 36
36 Plants completely glabrous; leaf lobe apices rounded; Argentina and Bolivia Solanum angustifidum
Plants with some hyaline simple trichomes (if lower surface glabrous then upper leaf surface with simple uniseriate trichomes); leaf lobe apices acute; Brazil Solanum viscosissimum
37 Leaves coriaceous and shiny; venation usually not easily visible on upper surface 38
Leaves membranous; venation visible on both surfaces 41
38 Shrubs or small trees; stems warty from raised leaf bases; flowers purple, fleshy; Ecuador Solanum coalitum
Vines; stems not warty; flowers white, membranous 39
39 Bark green and shiny, not exfoliating; calyx truncate or with minute lobes; southeastern Brazil Solanum odoriferum
Bark exfoliating; calyx variously lobed, never truncate 40
40 Inflorescence apparently axillary; flowers all perfect; southeastern Brazil Solanum inodorum
Inflorescence terminal; flowers with either short or long styles, these apparently on different plants and the species dioecious; northern Colombia and Venezuela Solanum luculentum
41 Inflorescences simple or at most once-forked, with < 10 flowers 42
Inflorescences many times branched, with > 10 flowers 43
42 Leaf base acute; Pacific of coast North America Solanum umbelliferum
Leaf base attenuate, winged onto the stem; Andes of Argentina Solanum salicifolium
43 Leaves linear to lanceolate (narrowly elliptic), more than twice as long as wide; leaf base acute or attenuate 44
Leaves ovate to elliptic, not more than twice as long as wide; leaf base truncate or cordate (occasionally acute) 46
44 Stems strongly angled or winged; flowers rotate Solanum amygdalifolium
Stems rounded; flowers stellate 45
45 Inflorescences open, the flowers widely spaced; delicate often herbaceous vines; southeast Asia Solanum pittosporifolium
Inflorescences congested, with the flowers clustered; robust vines or lax shrubs; Chile, widely cultivated Solanum crispum
46 Pedicel articulation in the distal half, often near the base of the calyx tube; calyx truncate, with minute undulations rather than lobes; Cuba Solanum boldoense
Pedicel articulation at the base in a small cup; calyx lobes deltate to narrowly deltate, irregular in size and shape; Brazil Solanum viscosissimum
47 Trichomes glandular (at least in part) on leaves and/or inflorescences 48
Trichomes not glandular on any part of plant 53
48 Glandular trichomes on inflorescence only, absent from leaves; Andes Solanum kulliwaita
Glandular trichomes present on stems and leaves 49
49 Shrubs; inflorescences simple or once branched; flowers purple with green spots at the base of the corolla lobes 50
Vines or lax shrubs; inflorescences many times branched; flowers usually lacking green spots, if purple with green spots, the lobes strongly reflexed 52
50 Flowers stellate; berries orange; leaves narrowly elliptic; northern Argentina and Bolivia Solanum endoadenium
Flowers rotate to rotate stellate; berries green or purplish black; leaves elliptic to ovate in outline; Pacific coast of North America 51
51 Flowers 1–2.6 cm in diameter; berry 1–2 cm in diameter, with 50–60 seeds; mainland Solanum umbelliferum
Flowers 3–4.5 cm in diameter; berry 3–4 cm in diameter, with >100 seeds; Santa Catalina Island, California Solanum wallacei
52 Corolla 0.8–1.3 cm in diameter, stellate; berry bright red; southeast Asia Solanum lyratum
Corolla 1.5–1.8 cm in diameter, rotate-stellate; berry black or purple-black; southeastern Brazil Solanum viscosissimum
53 Flower buds turbinate; anthers elongate and tapering 54
Flower buds ellipsoid or globose, not pointed and turbinate; anthers ellipsoid 55
54 Anthers tightly fused in a cone; filaments equal; corolla lobes with green spots at the base; Eurasia and North America Solanum dulcamara
Anthers not tightly fused; filaments unequal; corolla lobes uniform in colour; widespread in Neotropics Solanum uncinellum
55 Leaves linear to lanceolate (narrowly elliptic), more than twice as long as wide 56
Leaves elliptic to ovate, less than twice as long as wide 60
56 Delicate vines or small lax shrubs; seeds appearing “pubescent”; southeast Asia Solanum pittosporifolium
Shrubs, usually erect; seeds with the testa various, not apparently “pubescent”; Neotropics 57
57 Stems warty from prominent leaf bases; berries orange Solanum endoadenium
Stems rounded, angled or winged, not warty; berries red 58
58 Stems rounded; inflorescences many times branched; fruiting pedicels erect; Mexico to Costa Rica Solanum pubigerum
Stems strongly angled or winged; inflorescences simple or at most once-forked; fruiting pedicels nodding, not erect 59
59 Inflorescences lateral or leaf opposed; calyx lobes long-triangular; Argentina Solanum salicifolium
Inflorescences borne on short shoots; calyx lobes deltate or quadrate; Chile Solanum valdiviense
60 Leaves lobed on reproductive stems (if leaves simple then the stems winged) 61
Leaves simple on reproductive stems (if lobed leaves occur then only rarely and most leaves simple) 63
61 Stems rounded, not angled Solanum alphonsei
Stems angled or slightly winged 62
62 Leaves hastate or basally lobed; inflorescences simple; corolla stellate; North America Solanum triquetrum
Leaves deeply pinnatifid; inflorescence branched; corolla rotate-stellate; northern Asia Solanum septemlobum
63 Leaf surfaces more or less equally and densely pubescent abaxially and adaxially 64
Leaf surface pubescence density on adaxial and abaxial surfaces differing, the abaxial surface usually more densely pubescent 65
64 Upper leaf surfaces bullate; flower buds narrowly ellipsoid; Andes Solanum aspersum
Upper leaf surfaces not bullate; flower buds inflated, globose; Mexico and Central America Solanum dulcamaroides
65 Anthers borne on unequal filaments 66
Anthers borne on equal filaments 67
66 Corolla 2–2.5 cm in diameter; Brazil Solanum flaccidum
Corolla 1.5–2 cm in diameter; Mexico Solanum sousae
67 Flower buds strongly inflated; corollas fleshy; anthers globose 68
Flower buds ellipsoid or slightly inflated; corollas membranous; anthers ellipsoid 69
68 Pedicel articulation in distal half, often at base of calyx tube; Cuba Solanum boldoense
Pedicel articulation at base; Mexico and Central America Solanum dulcamaroides
69 Pubescence of leaf undersides in distinct tufts in the vein axils; calyx lobes long-triangular; southern South America, widely cultivated Solanum laxum
Pubescence of leaf undersides along veins, not concentrated in tufts in the vein axils; calyx lobes quadrate (square) or lacking (the calyx rim merely undulate) 70
70 Shrubs or small trees; flowers 1–1.4 cm in diameter; berries red; fruiting pedicels erect Solanum pubigerum
Woody vines; flowers 1.5–2.5 cm in diameter; berries black or purplish black; fruiting pedicels pendent 71
71 Calyx lobes quadrate (square); style glabrous; Hispaniola Solanum pyrifolium
Calyx lobes minute to lacking; style pubescent; central Mexico Solanum sousae
Synoptic character list for species of the Dulcamaroid Clade

This synoptic set of observations is intended to simplify the task of identifying a member of this large and highly variable group. Sterile plants of members of the Dulcamaroid clade are often difficult to identify, and even plants with either flowers or fruit can be difficult. I have included many leaf and whole plant characters so that sterile plants can be in many cases identified to a choice of a few taxa. Once this step has been done the user can read the descriptions and match distributions considering other characters not used in this list to the plant in hand. In the following list, character states are followed by a list of species epithets in alphabetical order. Epithets in parentheses indicate that the state is relatively uncommon in that species. Species that vary in a particular character, for example in leaf trichome distribution, will be listed for each state present in that species. Not all relevant characters are considered; for example, I have listed tapering and globose anthers, but not ellipsoid anthers – common in most species in the group. In this way diagnostic states can be easily seen. A question mark (?) following a species epithet indicates that the character is likely to occur, but has not been absolutely verified. Many species of the group are rather narrow endemics, occurring in only one country or region. Table 2 lists species recorded for each country of the Neotropics; this list must be used with care, as new collecting is constantly revealing range extensions.


Shrubs or small trees: aligerum, angustifidum, coalitum, crispum, cutervanum, endoadenium, imbaburense, leiophyllum, macbridei, muenscheri, nitidum, pubigerum, ruizii, salicifolium, stenophyllum, storkii, umbelliferum, valdiviense, viscosissimum, wallacei


Canopy lianas: dulcamaroides, uncinellum


Stems winged or strongly angled: aligerum, amygdalifolium, valdiviense, salicifolium


Stems warty or knobbly from persistent leaf bases: coalitum, dichroandrum, endoadenium


Bark shiny and exfoliating: inodorum, luculentum


Stems and leaves with echinoid and/or tree-like trichomes: cutervanum, ruizii, stenophyllum, storkii


Stems and leaves with golden trichomes: aureum, ruizii, cutervanum, stenophyllum, storkii


Glandular trichomes present anywhere on the plant (incl. only on the inflorescences): coalitum, endoadenium, kulliwaita, lyratum, umbelliferum, wallacei, viscosissimum


Dendritic (deer-antler-like) trichomes present: agnoston, (aligerum), aureum, calileguae, coalitum, (crispum), dichroandrum, imbaburense, (kulliwaita), leiophyllum, macbridei, muenscheri, (nitidum), sanchez-vegae, (stenophyllum), (umbelliferum), (uncinellum), (valdiviense)


Most leaves pinnatifid or lobed on mature reproductive stems: alphonsei, angustifidum, calileguae, lyratum, seaforthianum, septemlobum, salicifolium, triquetrum, uncinellum, umbelliferum, viscosissimum


Leaves coriaceous and shiny: agnoston, coalitum, imbaburense, inodorum, leiophyllum, luculentum, macbridei, (odoriferum), sanchez-vegae


Venation not visible on upper surface: agnoston, inodorum, luculentum


Lower surface of leaves with tufts of trichomes in vein axils: agnoston, aligerum, flaccidum, laxum


Both leaf surfaces densely pubescent with simple (never branched) trichomes: aspersum, endoadenium, umbelliferum, wallacei


Inflorescences simple: umbelliferum, salicifolium, valdiviense


Flower buds inflated: boldoense, dulcamaroides, seaforthianum, laxum


Flower buds narrowly ellipsoid: aspersum, uncinellum


Flowers with shiny green spots at the base of the corolla lobes: dulcamara, endoadenium, lyratum, pittosporifolium, septemlobum, umbelliferum, wallacei


Calyx lobes long-triangular or with long tips: aligerum, alphonsei, crispum, imbaburense, kulliwaita, leiophyllum, muenscheri, nitidum, ruizii, salicifolium, stenophyllum, triquetrum


Calyx lobes square (quadrate): aligerum, pyrifolium, valdiviense


Calyx lobes lacking (mere enations of the calyx rim): boldoense, dulcamaroides, sousae


Filaments unequal: angustifidum, flaccidum, seaforthianum, sousae, uncinellum


Anthers globose: boldoense, dulcamaroides, seaforthianum


Anthers tapering (buds turbinate): dulcamara, uncinellum


Mature berries > 2 cm in diameter: dichroandrum, dulcamaroides, luculentum, uncinellum, wallacei


Mature berries red: boldoense, crispum, dulcamara, dulcamaroides, inodorum, lyratum, nitidum, pittosporifolium, pubigerum, salicifolium, seaforthianum, septemlobum, triquetrum, (uncinellum), valdiviense


Mature berries orange or yellowish green: aspersum?, (crispum), endoadenium, luculentum?, (umbelliferum), valdiviense, (wallacei)


Mature berries black or purple: aligerum, amygdalifolium, angustifidum, aureum, coalitum, cutervanum, dichroandrum, flaccidum, imbaburense, kulliwaita, laxum, leiophyllum, macbridei, muenscheri, (nitidum immature), odoriferum, pyrifolium, ruizii, sanchez-vegae, sousae?, stenophyllum, storkii, (umbelliferum), (uncinellum), viscosissimum, (wallacei)


North America and Mexico (native or naturalised, not in cultivation): aligerum, dulcamara, dulcamaroides, pubigerum, sousae, triquetrum, umbelliferum, wallacei


Caribbean islands: boldoense, pyrifolium, seaforthianum


Central America: aligerum, dulcamaroides, muenscheri, pubigerum, seaforthianum, storkii


SE Brazil: flaccidum, inodorum, laxum, odoriferum, viscosissimum


Above treeline in Andes: aureum, coalitum, imbaburense, leiophyllum, macbridei, nitidum, stenophyllum


Eurasia: dulcamara, lyratum, pittosporifolium, septemlobum


Cultivated for ornament in both New and Old World (often escaped): laxum, seaforthianum

Species descriptions
Diagnosis.

Differs from Solanum sanchez-vegae S. Knapp in bearing tufts of dendritic trichomes in the vein axils of leaf undersides and elongate, densely pubescent buds; differs from Solanum laxum Spreng. in having shiny upper leaf surfaces, dendritic trichomes rather than simple trichomes in the vein axils of leaf undersides, and densely pubescent buds.

Type.

Ecuador. Loja: km 86 de Saraguro, localidad entre Susudel y el Progreso, 3°20'S, 79°15'W, 5 Aug 1986, J. Jaramillo et al. 8832 (holotype: QCA [QCA183731]; isotypes: MO [MO-5203618], NY [NY00854692]).

Description.

Woody vine or lax shrub, 1–3 m long/tall; stems glabrous, thin and flexuous; new growth glabrous, with a few dendritic trichomes abaxially. Bark of older stems pale greyish brown, glabrous. Sympodial units plurifoliate. Leaves simple, 2–6.5 cm long, 0.8–3.5 cm wide, elliptic to narrowly elliptic, coriaceous or somewhat fleshy, the upper surfaces glabrous and shiny, occasionally with sparse simple uniseriate trichomes to 0.5 mm long along the midvein and the petiole groove, the lower surfaces sparsely pubescent with dendritic trichomes, the trichomes to 0.5 mm long, confined to tufts in the primary vein axils near the midrib; primary veins 3–4 pairs, in the type completely obscured on the upper surface, the midrib yellowish green; base obtuse to truncate; margins entire, sometimes slightly revolute; apex acute; petiole 0.5–1.5 cm long, sparsely pubescent with simple uniseriate trichomes 0.5–1.5 mm long in the groove adaxially, possibly twining. Inflorescences terminal or lateral on short shoots, 2–5 cm long, branched 2–3+ times, with 7–10 flowers, glabrous of with a few dendritic trichomes at the branching points; peduncles ca. 1 cm long or less; pedicels 1–1.2 cm long, < 0.5 mm in diameter at the base, ca. 1.5 mm in diameter at the apex, filiform, erect to nodding, glabrous, articulated at the base and inserted into a short sleeve; pedicel scars spaced 0.5–1 cm apart in clusters of 2 or 3. Buds narrowly ellipsoid, the corolla strongly exserted from from calyx before anthesis. Flowers all perfect, 5-merous. Calyx tube 1–1.5 mmm long, conical, the lobes 0.5–1.5 mm long, triangular and keeled, scarious and transparent, glabrous, but with a tuft of simple uniseriate trichomes < 0.5 mm long at the apex. Corolla known only from buds, ca. 2 cm in diameter, violet, stellate, lobed ca. 3/4 of the way to the base, the lobes not known from mature flowers, densely pubescent abaxially with weak, simple uniseriate trichomes on exposed bud surfaces, glabrous adaxially. Filaments with the tube ca. 0.5 mm long, the free portions minute, glabrous; anthers ca. 5 mm long, 1–1.5 mm wide, ellipsoid, sagittate at the base, minutely pubescent within between the sutures. Ovary glabrous; style and stigma not seen. Fruit a globose berry, ca. 1 cm in diameter, purple or violet when ripe, the pericarp thin and somewhat shiny; fruiting pedicels 1.8–2 cm long, ca. 1 mm in diameter the base, slightly expanded at the apex, pendent. Seeds ca. 5.5 mm long, 5 mm wide, flattened reniform, pale brown, the surfaces minutely pitted, the testal cells pentagonal in outline. Chromosome number not known.

Figure 8.

Solanum agnoston S.Knapp. (A, B drawn from Holm-Nielsen et al. 5115 C drawn from Jaramillo et al. 8832). Illustration by Bobbi Angell.

Etymology.

The specific epithet comes from the Greek “agnostos”, meaning unknown, referring to the paucity of specimens and information on this species.

Distribution

(Figure 9). Southern Ecuador in the provinces of Loja and Azuay, between Loja and Cuenca.

Figure 9.

Distribution of Solanum agnoston S.Knapp

Ecology.

Growing in dry scrub and high elevation forests in interAndean valleys at near 3000 m elevation.

Conservation status.

Data deficient (DD); known from only two specimens, assessment not possible.

Discussion.

Solanum agnoston is known from only two localities, both along the main highway from Loja to Cuenca in southern Ecuador. It superficially resembles Solanum laxum, a species of southern Brazil that is widely cultivated in tropical and subtropical environments, but differs from it in the dendritic trichomes of the lower leaf surfaces and the elliptic, pubescent buds. The leaves of the paratype specimen (Holm-Nielson et al. 5115) are thick and very shiny above, such that the venation of the upper surface is not visible; the leaves of Jaramillo et al. 8832 are not as shiny, but match in all other aspects. Label data on both specimens indicate the flowers are violet, but both duplicates of Jaramillo et al. 8832 I have been able to find are in fruit.

Jaramillo et al. 8832 is the only collection of this species with duplicates in Ecuadorian herbaria, and have therefore chosen this specimen as the type so the holotype is in an Ecuadorian institution.

Specimens examined.

Ecuador. Azuay: km 91 of Pan-American Highway N of Loja, 2900 m, 3°25'S, 79°10'W, 5 May 1973, L. Holm-Nielson et al. 5115 (S [S-11-33845]).

2. Solanum aligerum Schltdl., Linnaea 19: 301. 1847.

http://species-id.net/wiki/Solanum_aligerum

Figure 10
Solanum pterocladum van Heurck & Müll.-Arg., Observ. Bot. 44. 1870.
Type: Bolivia. La Paz: Prov. Larecaja, Sorata, 3000 m, May 1858, G. Mandon 415 (lectotype, designated here: G [G00016904]; isolectotypes: AWH [n.v.], F [F-760412, F-876198, F-1588536], G-DC [Morton neg. 8550], GH [GH00077740], BM [BM000815930], K [K000590235], NY [NY00172146], P [P00445070, P00445071, P00445072], S [S04-2969]).
Solanum manicatum Bitter, Bot. Jahrb. Syst. 50, Beibl. 111: 63. 1913.
Type: Peru. Ayacucho: Prov. Huanta, road to Tambo above Osno, Río Apurímac, 2600-2700 m, A. Weberbauer 5643 (holotype: B [F neg. 2620], destroyed; lectotype, designated here: MOL; isolectotypes: F [F-647977, frag.], G [G00016963], GH [GH00077711]).
Solanum dotanum C.V. Morton & Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1079. 1938.
Type: Costa Rica. San José: Laguna de La Chonta, NE of Santa María de Dota, 2000-2100 m, 18 Dec 1925, P. Standley 42265 (holotype: US [US-1252694]).
Solanum grossum C.V. Morton, Revis. Argentine Sp. Solanum 178. 1976.
Type: Argentina. Tucumán: Dpto. Chicligasta, Estancia Las Pavas, 2000 m, 22 Nov 1926, Solanum Venturi 4632 (holotype: US [US-1548932, barcode US0027006]; isotypes: F, GH [GH00077675, GH00077676], NY [NY00172009, NY0017210], SI, US [US-1343321, barcode US01014178]).
Type

. Mexico. Michoacán: Angangueo, Oct, C. Schiede s.n. (holotype: B, destroyed, no duplicates found). Mexico. Michoácan: Wooded slopes 8–10 miles NW and WNW of Ciudad Hidalgo, among mountains west of Cerro San Andrés and 6-7 miles N of village of San Pedro Aguaro, in steep ravine along brook, 2850-3000 m, 19 48'N, 100 40'W, 18 Mar 1949, R. McVaugh & R.L. Wilbur 9917 (neotype, designated here: MEXU [MEXU-92021]; isoneotypes: BM [BM000578989], NY [NY00961953], US [US-2452307]).

Description.

Small tree or shrub, 1.5–5 m tall, the branches arching. Stems glabrous to densely pubescent with simple or dendritic uniseriate trichomes 0.5–1 mm long, glabrescent, often strongly winged from the decurrent leaf bases, the wings to 0.5 cm wide; new growth pubescent with simple or dendritic uniseriate trichomes 0.5–1 mm. Bark of older stems dark reddish brown, shiny. Sympodial units plurifoliate. Leaves simple, (3.5-)6–15 cm long, 1.2–5 cm wide, narrowly elliptic to lanceolate, membranous to slightly fleshy, the upper surfaces glabrous or sparsely (to occasionally densely) pubescent with simple or less often dendritic trichomes on the veins and lamina, the lower surfaces with dense tufts of dendritic trichomes 0.5–1 mm long in the vein axils, these trichomes occasionally extending along the midrib or to the lamina and the leaves more densely pubescent; primary veins 10–15(-20) pairs, the midrib keeled above, the veins often drying yellowish brown; base attenuate, often winged onto the stem; margins entire, sometimes revolute; apex acuminate; petioles 0.7–2 cm long, glabrous or pubescent like the stems, never twining. Inflorescences terminal or lateral, 4–15 cm long, many times branched, with 10–60 flowers, glabrous to pubescent, the trichomes simple or dendritic, ca. 0.5 mm long, denser at branch tips and at flower insertion points; peduncle 1.5–3.5 cm long; pedicels 1–1.2 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, slender, erect to nodding, glabrous, articulated at the base and inserted into a short sleeve; pedicel scars mostly clustered at the tips of inflorescence branches on small platforms or irregularly spaced 2–5 mm apart more basally. Buds pointed ovoid or turbinate with an apical nipple, the corolla strongly exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 1.5–2.5 mm long, cup-shaped, narrowing abruptly to the pedicel, the lobes 1.5–2 mm long, quadrate with an apiculate tip to 0.5 mm long, the tips densely pubescent with simple uniseriate trichomes, these extending adaxially, the adaxial surface densely papillate. Corolla 1.8–2 cm in diameter, white, often tinged with purple, stellate, lobed 2/3 to 3/4 of the way to the base, the lobes ca. 7 mm long, 5 mm wide, spreading, minutely pubescent abaxially, the tips cucullate, densely pubescent, the sinuses broad and thin, glabrous adaxially. Filament tube minute, the free portion of the filaments 0.5–1 mm long, glabrous; anthers 4.5–5.5 mm long, 1–1.5 mm wide, ellipsoid, loosely connivent, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 6.5–7 mm long, glabrous; stigma minutely capitate to slightly clavate, the surface minutely papillose. Fruit a globose berry, 1–1.2 cm in diameter, black or greenish black when ripe, the pericarp thin and shiny, glabrous; fruiting pedicels 1.5–1.7 cm long, ca. 1.5 mm in diameter at the base, woody, more or less nodding with the weight of fruits. Seeds 20–40 per berry, 2.5–3 mm long, 2–2.5 mm wide, flattened reniform, reddish brown or pale yellow, the surfaces minutely pitted, the testal cells square, the cell walls disintegrating and the seeds appearing hairy. Chromosome number: not known.

Figure 10.

Solanum aligerum Schltdl. (A drawn from Martinez S. 4741 B–D drawn from Torres B. 2022 E drawn from Nee 26870). Illustration by Bobbi Angell.

Distribution

(Figure 11). Common from Mexico to Argentina, from 1500–3300 m. A gap in the distribution of Solanum aligerum occurs from Panama to Peru (see discussion).

Figure 11.

Distribution of Solanum aligerum Schltdl.

Ecology.

Solanum aligerum has been collected most commonly in pine-oak forests in Mexico and adjacent Central America, and in cloud forests in the Andes.

Common names.

Guatemala: seconillo (Steyermark 34672).

Conservation status.

Least Concern (LC); EOO >50, 000 km2 (LC) and AOO >5, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum aligerum has the widest latitudinal range of all the New World members of the Dulcamaroid clade, occurring all down the cordilleras of both continents, but with a distinct gap between western Panama and Peru (see below). In spite of this huge range, the species is remarkably uniform in general aspect, but varies considerably in pubescence. In Bolivia, several populations have been found with plants that are uniformly and densely pubescent with dendritic trichomes over all plant parts; at one time I thought these were a new species and annotated some material as such, but in studying the Dulcamaroid clade in detail it has become apparent that these plants are merely pubescence extremes in this widespread species. They agree with other Bolivian material of Solanum aligerum in all other respects. It would be interesting to investigate if this pubescence variation is due to some habitat variation not apparent from herbarium labels.

Solanum aligerum is extremely similar to Solanum pubigerum, with which it broadly overlaps in central Mexico and Central America. The two species can be very difficult to distinguish, but Solanum aligerum has dendritic trichomes concentrated in the vein axils or over the entire lamina, while Solanum pubigerum has simple trichomes found all along the midrib on the leaf undersides. Flowers are larger in Solanum aligerum with the calyx lobes quadrate rather than deltate (this can be difficult to see), and the berries of Solanum aligerum are also larger and usually black or blackish green, rather than red, when ripe. The stems of S. aligerum are often (but not always) prominently winged, while those of Solanum pubigerum are never so. In general the two species appear to not occupy the same forest types where their ranges overlap.

The absence of Solanum aligerum in Colombia and Ecuador could indicate the northern and southern populations are in the process of differentation. This disjunction is unusual, but has also been found in the potato species Solanum morelliforme Bitter that is disjunct between Guatemala and Bolivia (Simon et al. 2011). Solanum morelliforme populations from south of the disjunction were morphologically (as here in Solanum aligerum) and molecularly identical. Without the locality being revealed, it is impossible to distinguish a plant of Solanum aligerum as coming from any particular area, so I am confident that these all represent a single taxon. Molecular studies at the population and geographical level may reveal some differentiating characters.

The holotype of Solanum aligerum was destroyed in Berlin and no duplicates have been traced. I have used a widely distributed collection (McVaugh & Wilbur 9917) from the same region of Mexico as a neotype; this typification also honors Rogers McVaugh, whose many years of collecting and study have so improved our knowledge of the flora of Mexico. Many syntypes exist for Solanum pterocladum, as Mandon’s collections were widely distributed and no herbarium was cited in the original publication. I have selected that in G as a lectotype, as that is where Mueller worked.

Specimens examined.

Argentina. Catamarca: Andalgalá, sobre la ruta no. 65, muy cerca del limite ca. de Tucumán, 4 Nov 1967, Hunziker 19509 (CORD); Andalgalá, Esquina Grande, 29 Apr 1915, Jorgensen 88791 (SI, US); Andalgalá, Valle del Río Chacra, 1650 m, 12 Feb 1950, Sleumer 18 (B). Jujuy: Ledesma, Abra de las Cañas, Parque Nacional Calilegua, 1750 m, 27 Nov 2004, Barboza et al. 1070 (CORD); Ledesma, Abra Colorado, 23 km de la entrada del Parque Nacional Calilegua, 16 Feb 2012, Barboza et al. 3549 (BM, CORD); Valle Grande, camino a Valle Grande, Río Jordan, 1400 m, 18 Oct 1964, Cabrera & Fabris 16142 (SI); Capital, Lagunas de Yala, 11 Nov 1978, Cabrera et al. 29778 (MO, SI); Capital, Sierra de Zapia, Mina 9 de octubre, camino a la antena, 16 Nov 1980, Cabrera et al. 32061 (SI); Santa Bárbara, S de El Fuerte, Finca Confines, 17 Nov 1992, Kiesling et al. 8332 (MO, SI); Palpalá, above Quebrada “Los Tomates”, on road to Cerro Zapla, 1850 m, 13 Apr 2000, Nee et al. 50757 (NY); Santa Bárbara, Centinela, 11 Dec 1983, Rotman et al. 930 (SI). Salta: Parque Nacional El Rey, Cerro Chañar, 20 Nov 1981, Brown et al. 1638 (NY); Cafayate, 1400 m, 24 Nov 1949, Araque M. & Barkley 19Ar 353 (F); Santa Victoria, Los Toldos, Rio Toldos, frente a Quebrada del Astillero, 3-5 km al E del pueblo, 30 Oct 1987, Novara 7100 (CORD, G); Santa Victoria, Los Toldos, cerros al E del pueblo, margen S del Río Toldos, 1700 m, 2 Nov 1989, Novara et al. 9107 (G); Santa Victoria, Camino al Lipeo, a 15 km de Los Toldos, 1800 m, 8 Nov 1975, Schiavone et al. 11901 C. (F, US); Capital, Quebrada de San Lorenzo, Oct 1926, Venturi 5080 (NY, US); Guachipas, Alemania, 1500 m, 16 Dec 1929, Venturi 9934 (US). Tucumán: Tafí, Apeadero Muñoz, 13 Feb 2012, Barboza et al. 3498 (BM, CORD); Trancas, Quebrada de Rearte, 1800 m, 15 Jan 1946, Bellonio 253 (LE, W); Cuesta del Clavillo, algo más abajo de La Banderita, 1750 m, 10 Nov 1952, Hunziker 10005 (CORD); Monteros, camino a Tafí del Valle, km 41-42, 12 Feb 1986, Hunziker 24873 (CORD); Chichigasta, Las Pavas, Oct 1911, Jorgensen 36423 (SI); Chichigasta, Cuesta Rio Cochuna, 1200 m, 28 Jan 1937, O’Donell 18158 (F); Chichigasta, Las Lenguas, cuesta del Rio Cochuma, 1400 m, 3 Nov 1930, Schreiter 6470 (B, SI, US); Tafí, Garabatal, Siambón, 1600 m, 25 Nov 1930, Schreiter 6552 (SI, US); Tafí, La Hoyada, Casa Arce, 1460 m, Nov 1932, Schreiter 8812 (CORD, GH); Tafí, Cumbre de Garabatal (Porteruelo), 1750 m, 23 Oct 1939, Schreiter 9809 (F, GH); Burruyacú, Cerro del Campo, 1800 m, 14 Dec 1928, Venturi 7755 (US); Tafí, cerca de Taficillo, 1200 m, 5 Oct 1929, Venturi 9609 (LE, NY, US).

Bolivia. Chuquisaca: Punilla, Guerrabamba, 3200 m, Feb 1949, Cárdenas 4136 (US); Azurday, La Angostura, ca. 15 km de Azurduy hacia el cañon de la Angostura, 2500 m, 11 Jan 2004, Huaylla & Guachalla 676 (K); Azurday, Tarvita, 2 km SW on Tarabuco-Azurday road, 2750 m, 25 Sep 1991, Kessler 3235 (GOET); Oropeza, entre el km 11 y Ravelo, 3300 m, 19 Nov 1993, Kiesling & Metzing 8425 (NY); Oropeza, Maragua, cercanias de Sucre, 3062 m, 4 Aug 2007, Velayos et al. 11136 (MA); ca. 8 km from Punilla towards Ravelo, 3300 m, 2 Oct 1994, Wood 8708 (K, NY); Oropeza, 2 km de la cruce a Potolo y Punilla, 2994 m, 29 Mar 2005, Wood et al. 22011 (K). Cochabamba: Carrasco, Montepunco, entrando por los Yungas, 2700 m, 18 Oct 1984, Beck 8931 (F, NY); Carrasco, ca. 75.5 km E of Epizana on Carretera Fundamental 4, 2439 m, 21 Nov 1976, Davidson 5095 (MO, US); Carrasco, Serrania Siberia, 20-35 km W of Comarapa (Prov. Santa Cruz) on the old Cochabamba-Santa Cruz Road (Hwy 4), 2000 m, 14 Jan 1990, Dorr & Barnett 7009 (MO, NY, US); Carrasco, Aguarrica, a 43 km de Epizana por la carretera que conduce a Santa Cruz, cerca de Aguarrica, 3030 m, 26 Dec 1982, Fernández Casas 7793 (MA, NY); Ayopaya, 10 km Cocapata-Cotacajes, 2750 m, 8 May 1997, Kessler et al. 9340 (NY); Carrasco, 5 km (by air and road) SE of bridge at Lopez Mendoza, 19 km by road NW of Epizana, on road from Comaropa to Cochabamba, 2900 m, 11 Feb 1987, Nee & Solomon 34093 (CORD, MO, NY, US); Carrasco, 5 km (by air and road) SE of bridge at Lopez Mendoza, 19 km by road NW of Epizana, on road from Comaropa to Cochabamba, 2900 m, 11 Feb 1987, Nee & Solomon 34093 (NY); Carrasco, narrow canyon of Rio Monte Puncu, 5 km NE of Monte Puncu, 10 km (by air) NW of Epizana, 2700 m, 10 Mar 1988, Nee & Solomon 36607 (CORD, NY, US); Chapare, trail leading NW from the road to tablas Monte along Río Corani Mayu, 2280 m, 2 May 1994, Ritter 919 (GH, MO); Chapare, 23.8 km N of Colomi (junction of the road to Candelaria) on road to Chapare, then 2.2 km NW (left) on side road. Upper Rio Cayani, 2700 m, 19 Oct 1985, Solomon 14386 (G, MO, NY); Chapare, km 104 del camino a Chapare, 3100 m, 2 Dec 1966, Steinbach 563 (F, MO, NY, S, US); Carrasco, Totora-Duraznillo, 2700 m, 30 Mar 1920, Steinbach 3826 (NY); Chapare, Sacaba, Cerro de Incachaca, 2500 m, 4 Sep 1921, Steinbach 5752 (F, G, MO, NY); Pojos, 2600 m, 5 Nov 1928, Steinbach 8621 (BM, E, MO, NY, S); Totora-Pocona, 2700 m, 7 Nov 1928, Steinbach 8685 (BM); Chapare, Incachaca-La Aduana (Sanctutonis), 2700 m, 8 Mar 1929, Steinbach 9550 (BM, E, F, G, MO, NY, S); Chapare, Durazno, Quebrada de Coraní, 2800 m, 18 Jun 1928, Steinbach 9854 (BM, E, F, G, MO, NY, S); Ayopaya, Independencia, c. 1 km before La Mina, 38 km north of Independencia along road to Sailapata, 2856 m, 15 May 2002, Wood et al. 8479 (BOLV, K); Chapare, descent from the dam towards Villa Tunari on road from Cochabamba to the Chapare, 3000 m, 25 Jun 1994, Wood 8545 (K, LPB). La Paz: Larecaja, Laripata, 2995 m, 13 Aug 2007, Aedo et al. 14644 (MA); Larecaja, Sorata, 2439 m, Feb 1886, Rusby 781 (NY); Murillo, Zongo, 2760 m, 15 Aug 2007, Aedo et al. 14734 (MA); Inquisivi, Rio Calachaca Jahuira-down river 1 km from Aguas Calientes de Calachaca, 10 km NE of Choquetanga, 3200 m, 17 Jul 1991, Lewis 39251 (G, MO); Larecaja, along road to Consata, ca. 4-15 km above (N of) Sorata, 2896 m, 24 May 1990, Luteyn & Dorr 13794 (NY, NY, US); Sud Yungas, La Paz Calacoto 69 km hacia el Este, pasando el Nevado Illimani, Estacion general Ikiko, 3100 m, 31 Dec 1980, Beck 3904 (F); Inquisivi, 3 km (by air) and 8 km (by road) SW of Inquisivi, above road from Quime to Inquisivi, 2950 m, 12 Mar 1988, Nee & Solomon 36672 (CORD, F, G, MEXU, MO, NY, US); Inquisivi, 6.5 km SE of Inquisivi, 1.2 km SW of Machacamarca, 3000 m, 18 Mar 1988, Nee 36718 (CORD, MO, NY, US); Larecaja, above Rio Lakhathiya, just above junction with Rio Tusca Jahuira, 19 May 2001, Nee et al. 51833 (NY); Inquisivi, comunidada Coquetanga-Aguas Calientes-Calachaca, cuenca del rio Calachaca-Jahura, 9 km de Choquetanga, 3400 m, 20 Jun 1994, Salinas 3275 (K, NY); Sud Yungas, Mina Chojilla camino de acceso de vehiculos a Kacapi, 2700 m, 4 Jul 2000, Siñani RS 230 (NY); Pedro Domingo Murillo, Rio Zongo Valley, 22.5 km below dam at Lago Zongo, 3000 m, 9 Oct 1982, Solomon 8431 (K, MO, NY); Inquisivi, Camino de Quime a La Paz a unos 2–3 km arriba de Quime, 3221 m, 16 Mar 2003, Wood & Ortuño 19411 (K, LPB). Santa Cruz: Florida, Comarapa Road, 28 km from Comarapa, 2650 m, 15 Aug 1991, Acevedo et al. 4623 (NY, US); Caballero, Siberia, 2–2.5 km debajo de la escuela de Siberia, camino al valle de Saipina, 2850 m, 30 Nov 2003, Jordan & Vargas 253 (MO); Vallegrande, on road from Guadalupe to Pucará, 2725 m, 4 May 2001, Nee et al. 51736 (BM, NY); Caballero, loc 2-3 km S de Siberia sobre el camino a Larkapampa, 3000 m, 25 Jul 1996, Saldías & Fernández 4607 (NY); Caballero, Parque Nacional Amboró, Siberia, 25 km al NW sobre la carretera Comarapa-Cochabamba, 3000 m, 5 May 1993, Vargas C. et al. 2302 (NY); Caballero, Parque Nacional Amboró, proximadades de Cerro Bravo a 10 km al N de Comarapa, alrededores de la Parcela permanente, 2400 m, 18 Oct 1993, Vargas C. & Jardim 3002 (NY); Caballero, Siberia, ca. 1–2 km arriba de la comunidad de Siberia, sobre un camino vecinal, entrando hacia el Parque Nacional Amboró, 3001 m, 12 Apr 2003, Wood et al. 19700 (BOLV, K);. Tarija: route Villa Montes to Tarija, 15 km apres Entre Rios, 1850 m, 4 Nov 1993, Billiet & Jadin 6067 (K, MO); Aniceto Arce, 39.9 km S of jct of road to Entre Rios, on road to Padcaya, 2100 m, 29 Apr 1983, Solomon 10249 (MO, NY).

Costa Rica. Cartago: Volcán Irazu and Volcán Turrialba, along road from Pacayas to Hacienda Central, 31 Mar 1982, Barringer et al. 2233 (F); SE slope of Cerro de la Muerte, Cordillera del Talamanca, along Interamerican Hwy 63 miles from downtown San Jose, 2700 m, 23 May 1976, Croat 35427 (MO); Cantón Paraíso, Reserva Forestal Los Santos, Cueva del Savegre, carretera Interamericana de la Georgina hasta la entrada a San Gerardo de Dota, 3180 m, 18 Mar 1997, González & Hammel 1840 (G, MO); Cantón de El Guarco, Cordillera de Talamanca. Carretera Interamericana, entre E1 Empalme y La Chonta, 2400 m, 10 May 1995, Hammel 19815 (BM, MO); Irazú, 2134 m, 24 Jun 1974, Kuntze 2300 (NY); Macho Gap Camp, 39 kilometers S. of Cartago and 3 miles N of Copey, 2500 m, 19 Feb 1943, Little 6022 (F, US);. Heredia: Cantón de Barva, P.N. Braulio Carrillo, Cordillera Central, potreros La Georgina, 2600 m, 29 Oct 1993, Fernández & García 1503 (MO); Puntarenas: Monteverde, at edge of Continental Divide and of Pacific side of slope, 1300 m, 18 Aug 1976, Solomon 5365 (MO); N. San Isidro del General, 12 Aug 1971, Spellman et al. 653 (MO); San José: northern Cordillera Talamanca, region of Cerro de la Muerte, on carretera Nacional 34.5 km N. of San Isidro and 2.6 km S of La Georgina Inn, 2652 m, 4 Apr 1978, Davidson 7235 (F, MO, NY); Cantón de Pérez Zeledón, Cordillera de Talamanca. Estación Cuericí, 2600 m, 29 Aug 1995, Gamboa 281 (BM, MO); Villa Mills, Cerro de la Muerte, Carretera Panamericana Sur, 2800 m, 26 Feb 1965, Jiménez M. 2989 (F, NY); Z.P. Cerros de Escazú, Cedral, falda norte del cerro Rabo de Mico, cuenca del Río Poás, 1600 m, 9 Oct 1991, Morales 168 (MO); Santa María de Dota, Laguna de la Chonta, northeast of Santa María, 2000 m, 18 Dec 1925, Standley 42265 (F, US); near Finca La Cima, above Los Lotes, North of El Copey, 2100 m, 21 Dec 1925, Standley 42764 (US); San Gerardo de Dota, 8 to 10 km down from the Interamerican Highway, 18 Sep 1975, Utley & Utley 3083 (F, MEXU, MO, NY).

El Salvador. Santa Ana: Near summit of Cerro Monte Cristo, 2134 m, 18 Jan 1959, Allen 7172 (F, NY, US); Montecristo, P.N. Montecristo, plan de los helechos, 2051 m, 13 Mar 2002, Carballo et al. 281 (LAGU); Cordillera Miramundo, mountain of Montecristo, 2000 m, 27 Jan 1966, Molina R. et al. 16706 (F, NY, US); P.N. Montecristo, al final del atajo para el Trifinio, 2000 m, 24 Jan 2002, Monterrosa et al. 215 (B, BM, LAGU).

Guatemala. Chimaltenango: Chichavae, 2400 m, 5 Dec 1933, Skutch 738 (US). Huehuetenango: San Juan Ixcoy, Sierra de los Cuchumatanes, along road to Huehuetenango, 5 miles S of San Juan Ixcoy, 4 Feb 1965, Breedlove 8542 (F); San Mateo Ixtatán, Sierra de los Cuchumatanes, 4 miles east of San Mateo Ixtatan on road to Barillas, 7 Feb 1965, Breedlove 8744 (F); Cerro Cananá, between Nucapuxlac and Cananá, Sierra de los Cuchumatanes, 2500 m, 18 Jul 1942, Steyermark 49046 (F); wet cloud forest at Cruz de Limon, between San Mateo Ixtatan and Nuca’, Sierra de los Cuchamatanes, 2600 m, 31 Jul 1942, Steyermark 49870 (F, NY). Quetzaltenango: highway km 172 junction Quetzaltenango, Huehuetenango and Totonicapan, 10 Jan 1974, Molina R. et al. 30200 (F, MO); Volcán Zunil, 2500 m, 22 Jan 1940, Steyermark 34672 (F). Quiché: sin. loc., 1942, Aguilar s.n. (F). Totonicapán: Sierra Madre Mountains south of Totonicapán, near Mirador (km 170), 2800 m, 20 Dec 1972, Williams et al. 41447 (BM). Zacapa: Jones, 2020 m, 5 Apr 2000, CDC-CECON 1535 (BM).

Honduras. Francisco Morazán: along road to Parque Nacional La Tigra, 22-25 km NE of Tegucigalpa, 1850 m, 1 Feb 1987, Croat & Hannon 64047A (MO, NY); Tegucigalpa, 11 km NE of Tegucigalpa. La Tigra National Park Summit, 2000 m, 26 May 1992, D’Arcy 18006 (MO); La Tigra, 25 km NE de Tegucigalpa, 2105 m, 24 May 1986, Manzano 177 (MA); Montaña La Tigra S.O. de San Juancito, 2000 m, 23 Apr 1964, Molina R. 13744 (F, NY, US); Cerro Uyuca, 8 km O. del Zamorano, 1100 m, 27 Apr 1984, Soihet M. 142 (MO).

Mexico. Chiapas: Chamula, Yalal Chin, 1829 m, 3 Apr 1965, Breedlove 9540 (MEXU); San Cristóbal de las Casas, SW slope of Zontehuiz, 2774 m, 21 Jun 1965, Breedlove 10435 (F, MEXU, US); Chenalho, in paraje Los Angeles Chiste, 1700 m, 22 Oct 1976, Breedlove 40863 (MEXU, MO); La Independencia, 6-10 km north-northeast of La Soledad along logging road from Las Margaritas to Campo Alegre, 1600 m, 26 Nov 1980, Breedlove & Almeda 47819 (MEXU, MO, NY); Zinacantán, on NW side of Muk’ta vits (Cerro Huitepec), 2743 m, 18 Feb 1966, Laughlin 125 (MEXU, US). San Cristóbal de las Casas, Santa Cruz en San Felipe, 15 Nov 1986, Mendez Ton 9743 (MEXU, NY); Pueblo Nuevo Solistahuacán, on ridge north of Clinica Hierba Buena near Pueblo Nuevo Solistahuacán, 1981 m, 25 Jan 1965, Raven & Breedlove 19978 (F); Tenejapa, at Colonia Ach’lum, 2652 m, 27 Dec 1965, Shilom Ton 454 (F, MEXU, NY, US); San Andres, NW of San Cristobal, 579 m, 26 Feb 1931, Souviron & Erlanson 81 (BH, US); Tenejapa, a 20 km al SW de Tenejapa camino a San Cristóbal de las Casas, 25 Sep 1983, Téllez V. & Pankhurst 7262 (MEXU, MO); Mazapa de Madero, Granada de Tlalcanaque, 10 km al Noreste, 2500 m, 22 Feb 1987, Ventura & López 4352 (MO, NY). Colima: Nevado de Colima, Nevado de Zapotlan, a few miles south of Ciudad Guzman (Zapotlan), 3000 m, 2 Jul 1956, Gregory & Eiten 292 (P); Guerrero: Taxco de Alarcón, Taxco. 19.9 km al NO, 2360 m, 28 May 1998, Cruz Durán 2260 (MEXU). Chilpancingo de los Bravo, a 6 km al W de Omiltemi, camino a la Soledad La Joyas, 27 Mar 1982, Martínez S. & Téllez V. 272 (MO); Chilpancingo, laderas superiores del Cerro Alquitran, cerca de Mazatlan, 2600 m, 11 Feb 1970, Rzedowski 27036 (F, MO x2); Chichihualco, a 8 km al SO de Filo de Caballo, 2435 m, 21 Apr 1985, Soto N. & Aureoles C. 8345 (MEXU). Hidalgo: Lolotla, carr. Pachuca-Huejutla, 3.5 km al N de Ixtlahuaco, 1670 m, 5 Apr 1986, Baker et al. 054 (NY); 17 mi N of Jacala, ca 56 mi from Zimapan, 1 Aug 1972, Dunn 19059 (MO); Zacualtipán, 1 km al SSE de Zacualtipan, 2400 m, 19 May 1976, Flores F. 234 (MO, US); Zacualtipán, Zacualtipán, 1 km al SSE, 2400 m, 19 May 1976, Flores M. 234 (CORD, MEXU); Tenango de Doria, Cirio, 8 km al este de Tenango de Doria, 1700 m, 27 Mar 1980, Hernández M. & Hernández V. 4189 (MEXU); Tianguistengo, 10 km al oeste de Tianguistengo (Tepeoco), 2100 m, Apr 1981, Hernández M. et al. 5806 (MEXU, MO); Metztitlán, noreste de Hidalgo, Rincón de los Ahuajes, 3.5 km de la desviación a Zoquizoquipan, 6 Jul 1992, López García 113 (MEXU); Zacualtipán, Paraje Cumbre de Tlahuelompa, 2 km al SW del ejido de Tlahuelompa, 18 Dec 1992, López García 392 (MEXU); El Alcalaque, 2 km al NW del pobaldo Eloxochitlán, 29 Dec 1992, López G. 433 (MEXU); Tlanchinol, a 4 km al E de Tlanchinol, camino a Apantlasol, 3 Sep 1997, Martínez S. 28444 (MEXU, NY); Palo Hueco, km 301 on highway between Jacala and Santa Ana, 1524 m, 28 Apr 1947, Moore 2672 (BH); Barranca below Trinidad Iron Works, 1524 m, 10 May 1904, Pringle 8835 (BH, BM, F, GOET, K, LE, MEXU, MO, US); Molango, Ismolinta, 2 km S de Ismolinta rumbo a Eloxochitlán, 1580 m, 19 Jun 1995, Sousa Peña 577 (MEXU); Zacualtipán, vereda entre El Reparo y Zahuastipan, 3 km de la Majonera, 29 Jun 1988, Vázquez et al. 4626 (F, NY); Zoquizoquipan, 1940 m, 2 Sep 1962, Vela G. 977 (MEXU). Jalisco: Yolox, Sierra de Juárez, Carretera Ciudad de Oaxaca. a Tuxtepec, cerca de Cerro Pelon, a 300 m. antes de San Pedro Yolox, 2850 m, 18 Jan 1989, Chazaro et al. 5815 (MEXU); Cuatitlán, 19-20 km al NE de Cuatitlan, 500 m al W de Llanos de San Miguel, 2200 m, 29 Apr 1988, Cuevas & Guzmán 2871 (MEXU); along lumber road, 0.3 km E of fork Cerro La Cumbre/Rincón de Manantlán, at crossing of first small stream, N of Sierra de Manantlán Central, 17.7 km S of El Chante, 2250 m, 6 Jan 1980, Iltis et al. 2336 (F x2, MEXU, US); northwestern slopes of Nevado de Colima, above Jazmín, 2-3 km above settlement of El Isote, 2600 m, 26 Mar 1949, McVaugh & Wilbur 10066 (BM); Sierra de Manantlán (15-20 miles southest of Autlán), about 2 miles from Aserradero San Miguel Uno, west and south of divide towards Manzanillo, 2250 m, 4 Nov 1952, McVaugh & Sooby 13923 (BM); Venustiano Carranza, Puerto El Floripondio, camino a la Est. de Microondas Las Viboras, 2450 m, 22 Aug 1987, Rodríguez C. & Suárez J. 939 (MEXU). Michoacán: Zinapécuaro, Los Azufres, alrededores de la laguna larga, 31 May 1988, Díaz Barriga 4680 (MEXU); Zinapécuaro, Cañada La Hierbabuena, al SW de la Presa Laguna Larga, 2750 m, 17 Nov 1988, Jasso 532 (MEXU, MO); Melchor Ocampo, Cerro Camacho, Los Remedios, a 4 km al W de Ocampo, 2320 m, 26 Apr 1982, Martínez S. et al. 386 (BH, MEXU); Tzitzio, En Mil Cumbres a 31 km al SW de Cd Hidalgo, carretera a Morelia, 2500 m, 11 Oct 1983, Martínez S. et al. 4741 (MEXU, NY). Morelos: Tepoztlán, Santo Domingo Ocotitlán, en el cerro al N de la población, 2200 m, 26 Apr 1984, Gutiérrez 203 (MEXU). México: Temasceltepec, Rincón, 3 Oct 1932, Hinton 365 (BM, K, US); Temascaltepec, Comunidad, 2480 m, 5 Oct 1933, Hinton 3852 (BM, F, K, MO, US); Oaxaca: Comaltepec, immediately to the right of Hwy. 175, just beyond first major switchback on decent from Mirador below Cerro Humo Chico, in quebrada of Río Cerro Pelón, 2740 m, 1 Nov 1993, Boyle & Massart 2512 (BM, MEXU); Sierra Juárez, 2.5 miles by road. SW of summit at Cerro Pilon, on hwy 175 between Ixtlan de Juarez and Valle Nacional, 2550 m, 19 Jul 1972, Breckon & Breckon 1364 (F); San Martín Peras, Dto. Juxtlahuaca, a 2.7 km adelante de la desviacion de San Martin Peras, carretera para Coycoyan de las Flores, 2490 m, 20 Jun 1993, Calzada 18460 (MEXU, NY); Comaltepec, 46.5 km al NE de Istlán, carretera Oaxaca-Tuxtepec, 2650 m, 17 Dec 1987, Campos V. & Torres 886 (MEXU); Mixistlán, 30 km al S de Totontepec, Distr. Mixe, 2600 m, 10 Nov 1983, García Mendoza et al. 1287 (MO, NY); Patio de Arena, vicinity of Cerro Zempoaltepetl, ca. 5 km E of summit, 2800 m, 9 Aug 1950, Hallberg 879 (MEXU); Ixtlán, Sierra de Juárez, parte alta y cima de Cerro Pelon, 2900 m, 5 Aug 1985, Lorence et al. 4754 (MEXU); Yolox, distrito de Ixtlan, 8 km E of Yolox on road between Yolox and highway 175, 2900 m, 13 Apr 1981, Martín 531 (MO); Ixtepeji, La Botuda, Dto. Ixtlán, 10 May 2001, Santiago 49 (MEXU); Totontepec, Villa de Morelos, distr.: Mixe Tsa, 19 Aug 1991, Rivera Reyes 2814 (MEXU, NY); Santa María Teopoxco, Sierra Mazateca, near Plan de Guadalupe, 39 km by road (16 km by air) west o Huatla on road to Teotitlan, 2200 m, 17 Jan 1984, Solheim & Reisfield 1400 (NY); Puerto de la Soledad, 3 km al S de Huautla, sobre el camino a Teotitlán del Camino-Huautla, 10 Sep 1981, Téllez V. & Elisens 4698 (MEXU); Totontepec, 13 km al O de Totontepec, carretera a Villa Alta, Dto. Mixe, 28 Mar 1986, Torres C. & Ramírez 8450 (MEXU, MO, US); San Juan Tepeuxila, San Juan Tepeuxila, Dto. Cuicatlan, arroyo la primera toma, hacia Llano Chiflido, por Arroyo Paloma, 2369 m, 18 May 2002, Torres Colin et al. 16202 (MEXU). Puebla: Tlatlauqui vertiente norte, 1800 m, 31 Apr 1971, Boege 1738 (MEXU); Equimita, km 35 de la carretera que va hacia Cuetzalan, 22 Jun 1976, Inzunza 114 (MO); Huauchinango, 1951 m, 7 Oct 1944, Sharp 441218 (MEXU, US); Coxcatlán, L y Griega, desviación a Coyomeapan, de La Brecha a Zoquitlán, 2540 m, 28 Sep 1984, Tenorio L. & Romero de T. 7502 (MEXU); Tezuitlán, Coaxisco 12 km al NW de Tezuitlan, 2030 m, 12 Apr 1985, Tenorio L. et al. 8630 (MEXU); Pahuatlán, Acahuales, 1900 m, 22 May 1986, Tenorio L. & Romero de T. 11383 (MEXU); Tezuitlán, Río Frío, 12 km al N de Teziutlan, carr. a Nautla, 1500 m, 7 Jul 1987, Tenorio L. et al. 14020 (MEXU, NY). Querétaro: Pinal de Amoles, El Ranchito, 1 km de El Rnachito camino a San Pedro Escanela, 1980 m, 10 Nov 1988, Carranza 1251 (MEXU). San Luis Potosí: 22 km W of Santa Catarina on highway 86 at km 49, 2200 m, 29 Sep 1965, Roe & Roe 2191 (BM). Veracruz: Xico, Tonalasco, 25 Jun 1986, Arriaga C. 389 (MEXU); Huayacocotla, Vereda Tzimentey, cerca del limite SE de la Reserca propuesta, 1750 m, 25 Apr 1981, Bastelleros & Ballesteros 421 (MEXU); Acultzingo, La Laguna, carretera Puerto del Aire, poblado, 2320 m, 16 Dec 1977, Calzada & Delgado 4181 (MEXU); Huayacocotla, Agua de la Calabaza, carretera Huayacocotla-Chicontepec, 1800 m, 20 Jul 1979, Calzada 5464 (F); Coatepec, Cerro Huilotepec entre Mesa de los Laureles y Tierra Grande, 2500 m, 27 Jul 1987, Cházaro B. et al. 5275 (MEXU, NY); Las Minas, al SE de Rinconada por el Cerro La Tolva, 2300 m, 11 Aug 1988, Duran E. & Burgos 570 (MEXU); Huayacocotla, vicinity of a large shrine and “Bienvenidos a Huayacocotla” arch over the highway, SW entrance to Huayacocotla on road from Palo Bendito, 2150 m, 21 Jul 1982, Diggs & Nee 2958 (BH, CORD, F); Huayacocotla, Agua de la Calabaza, 16 km NE de Huayacocotla, carretera a Zilacatipan, 1750 m, 24 Apr 1981, Juárez G. & Vasquez B. 68 (F); Huayacocotla, Xalapa, INIREB, along Huayacocotla-Zontecomatlan road, between Barro Colorado and Tepozanes, 2 km by road NE of Agua de la Calabaza and 5 km by road SW of Zilacatipan, 1800 m, 27 Apr 1983, Nee 26894 (F, MO, NY); Puerto del Aire, just W and high above Acultzingo, top of very steep pass on Highway 150 (Tehuacán to Orizaba) near boundary between states of Veracruz and Puebla. Cumbres de Acultzingo, 2409 m, 27 Sep 1962, Ugent & Flores C. 2466 (BM); entre Los Ocotes y Helechales, 1 Jun 1980, Vargas et al. 306 (MEXU); Jalacingo, Ocotepec, cerca de cerro, 1750 m, 4 Mar 1970, Ventura A. 1008 (F, MO, NY); Acajete, Acajete, 1850 m, 1 Apr 1974, Ventura A. 9810 (MEXU, MO); Atzalán, La Florida, 1650 m, 16 Jun 1976, Ventura A. 12871 (F, MEXU); Altotonga, San Miguel Tlalpoalan, 1950 m, 2 Jun 1981, Ventura A. 18538 (MEXU); Acajete, La Joya, 2100 m, 1 Apr 1982, Ventura A. 19577 (MEXU, MO); Las Vigas, Rancho San Miguel, barranca El Cable, 2400 m, 11 Sep 1989, Zamora C. 1096 (MEXU);

Peru. Cusco: Urubamba, Ollantaytambo, Dist. Ollantaytambo, Abra de Malaga, 3600 m, 2 Jun 2002, Galiano et al. 4225 (MO); Hacienda Huyucalla, valle del Paucartambo, 3400 m, Jul 1930, Herrera 2986 (US); Paucartambo, 10 km from Puesto de Vigilancia (PN Manu) towards Paucartambo, 3313 m, 15 Mar 2012, Knapp et al. 10436 (BM, USM); Quispicanchis, near Marcapata, 24 Jul 1942, Metcalf 30729 (G, MO, US); Quispicanchis, entre Abra Walla Walla y Marcapata a 210 km de Cusco, 2800 m, 21 Apr 1988, Núñez V. et al. 8994 (F, MO x2); Pautarcambo to Tres Cruces, Cerro de Cusilluyoc, 3200 m, 2 May 1925, Pennell 14148 (F, NY); Calca, Dist. Lares, Mantto, 2100 m, 23 Jan 2003, Valenzuela et al. 1236 (MO); La Convención, Espiritupampa, Dist. Vilcabamba, 3300 m, 14 Oct 2003, Suclli et al. 1277 (BM, NY); La Convención, San Luis, valley of Lucumayu River, on road from Ollantaytambu to Quillabamba, environs of San Luis (former resturan) above Alfamayo, along roadside with 2 km of San Luis back up towards Abra de Malaga, 2800 m, 3 Apr 1997, Tupayachi & Emshwiller 3424 (BH); Paucartambo, Challabamba, Pesto Grande, 2900 m, 20 Apr 1986, Zimmerer 80 (NY). Junín: Tarma, Carpapata, al este de Carpapata, 8 Dec 2005, Marcelo Peña et al. 1920 (MOL). Puno: Carabaya, Ollachea to San Goban road, 2.5 km north of Ollachea, 15 Aug 1980, Boeke 3049 (F, MO, NY, US); Ollachea to San Gabon, 17 Jul 1978, Dillon et al. 1107 (BM, MO, NY).

3. Solanum alphonsei Dunal, Prodr. [A.P. de Candolle] 13(1): 70. 1852, as “ alphonsi

http://species-id.net/wiki/Solanum_alphonsei

Figure 12
Solanum alphonsei Dunal var. taguatagua Dunal, Prodr. [A.P. de Candolle] 13(1): 70. 1852.
Type: Chile. Sin. loc., C. Bertero 635 (holotype: P [P00319395, Morton neg. 8142, G neg. 39141]).
Solanum germainii Phil., Linnaea 29: 23. 1858, as “germaini
Type: Chile. Región Metropolitana: Prov. Santiago, mountains near Aculeo [“Aculco”], R.A. Philippi s.n. (holotype: SGO [SGO-55447, Dept. Invest. Agrícolas neg. s.n.]; isotypes: B [destroyed, F neg. 2732], LE, MA [MA-533038]).
Solanum tenuicaule Phil., Anales Univ. Chile 91: 13. 1895.
Type: Chile. sin. loc., Anon. [R.A. Philippi?] s.n. (lectotype, designated here: SGO [SGO-55451, barcode SGO000004604]).
Type.

Switzerland. Cultivated in Geneva, 1834, originally from Chile (lectotype, designated here: G-DC [G00144607].

Description.

Woody vine or lax shrub to 1 m tall. Stems erect or spreading, glabrous or with simple uniseriate glandular trichomes to 0.5 mm, these denser at the nodes, the glands 1-celled and soon deciduous; new growth glabrous to minutely glandular puberulent. Bark of older stems green to reddish brown, glabrous. Sympodial units plurifoliate. Leaves simple or more often shallowly and irregularly pinnatifid, (0.5-)1.5–5 cm long, (0.5-)1–3(-5) cm wide, narrowly elliptic to elliptic or rhomboid, membranous, occasionally slightly fleshy, both surfaces glabrous or with scattered simple uniseriate trichomes on the lamina, these denser along the margins; primary veins 4–5(-7) pairs, often drying reddish brown; base acute or truncate; margins entire or shallowly lobed in the basal half, the lobes divided less than halfway to the midrib; apex acute to more often rounded; petioles 0.5–1 cm long, glabrous to pubescent with simple uniseriate glandular trichomes, apparently twining. Inflorescences terminal or later lateral, 2–6.5 cm long, many times branched, openly divaricate, with up to 30 flowers, glabrous or pubescent with glandular simple uniseriate trichomes to 1 mm long; peduncle 0.5–3 cm long; pedicels 0.6–1 cm long, ca. 0.5 mm in diameter, slender, nodding at anthesis, glabrous, articulated at the base from a small sleeve, leaving a tiny peg on the inflorescence axis; pedicel scars irregularly spaced 2–12 mm apart. Buds globose when young, later ellipsoid, the corolla strongly exserted from the calyx before anthesis. Flowers all perfect, 5-merous. Calyx tube ca. 1.5 mm long, shallowly cup-shaped, the lobes 1–1.5 mm long, deltate with elongate tips, the sinuses splitting irregularly, glabrous or minutely glandular puberulent. Corolla 1–1.5 cm in diameter, white, stellate, lobed nearly to the base, the lobes 5–7 mm long, 3–4 mm wide, planar or slightly cupped at anthesis, densely papillate-puberulent on the tips and margins, the trichomes sometimes extending over entire abaxial surface. Filament tube minute, the free portion of the filaments ca. 0.5 mm long, glabrous; anthers ca. 3 mm long, 1.5 mm wide, ellipsoid, loosely connivent, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 6–7 mm long, glabrous; stigma capitate, to ca. 0.5 mm in diameter, the surface minutely papillose. Fruit a globose berry, ca. 0.8 cm in diameter, orange or reddish orange when ripe, glabrous, the surface shiny and thin; fruiting pedicel ca. 1 cm, deflexed to spreading. Seeds 15–20 per berry, ca. 1.5 mm long, 2 mm wide, flattened reniform, pale tan, the surfaces minutely pitted, the testal cells broadly sinuate in outline. Chromosome number: not known.

Figure 12.

Solanum alphonsei Dunal. (A–H drawn from Anon. s.n., Mar 1858, BM). Illustration by Bobbi Angell.

Distribution

(Figure 13). Solanum alphonsei occurs in southern Chile and possibly also adjacent Argentina, from sea level to the summit of the Andes at ca. 3000 m.

Figure 13.

Distribution of Solanum alphonsei Dunal.

Ecology.

In Nothofagus Blume (Nothofagaceae) forest and forest margins.

Conservation status.

Vulnerable (VU); EOO <20, 000 km2 (EN) and AOO <2, 000 km2 (VU). See Moat (2007) for explanation of measurements.

Discussion.

Solanum alphonsei is one of three species of the Dulcamaroid group found in the Nothofagus forests of southern Chile (and probably also from adjacent Argentina); the others are Solanum valdiviense and Solanum crispum. Solanum alphonsei differs from Solanum valdiviense in being a vine with non-angled stems and twining petioles, in having open and divaricately branched inflorescences and in its leaves that are rhombic or deltate in outline. The corollas of Solanum alphonsei are in general smaller than those of Solanum valdiviense, and are less deeply lobed. Solanum crispum is much more common than either of the other two species, and can be distinguished from Solanum alphonsei by its larger flowers, denser pubescence and larger inflorescences. Solanum alphonsei is much less commonly collected than is Solanum valdiviense. An extreme form of Solanum alphonsei with tiny leaves was given the herbarium name of “myrtilloides” by Witasek on a specimen in Vienna.

Dunal named Solanum alphonsei for Alphonse de Candolle, the editor of the Prodromus and son of his mentor from Montpellier, Agustin Pyramus de Candolle. Philippi named Solanum germainii after Philibert Germain, the Chilean botanist and collector. Both original spellings are correctable (McNeill et al. 2012).

No specimens were cited in the original description of Solanum alphonsei, but the material was said to have been cultivated in Geneva in 1834. The specimen in G-DC [G00144607] is dated 1834, and is chosen here as the lectotype. Other sheets in P, G, G-DC and BM from plants cultivated in Geneva are dated differently and are thus not type material. They could, however, have been collected from the same individual plant, and could be considered topotypes.

Specimens examined.

Chile. Región VI (O’Higgins): O’Higgins, Rancagua, La Leonera, 760 m, 9 Dec 2001, Aedo 7072 (MA); Rancagua, Mar 1828, Bertero 639 (P); San Fernando, Philippi s.n. (G, K). Región VII (Maule): Cauquenes, Hacienda de Cauquenes, , 20 Aug 1896, Dusén 57 (S); prov. Curico, Cordillera de la Costa, Sep 1897, Witasek s.n. (W).

4. Solanum amygdalifolium Steud., Nomencl. Bot. ed. 2, 2: 600. 1841

http://species-id.net/wiki/Solanum_amygdalifolium

Figure 14
Solanum persicifolium Mart., Flora 21, Beibl. 2: 78. 1838, as “persicaeifolium”, non Solanum persicifolium Dunal, 1813.
Type: Brazil. Rio de Janeiro: São Christovão near Sebastianopolis, C. Martius 255 (lectotype, designated here: M [M0171803]; isolectotypes: BR, LE, K [K000196335]).
Solanum angustifolium Lam., Tabl. Encycl. 2: 18. 1794, non Solanum angustifolium Mill., 1768.
Type: Argentina. Buenos Aires: Buenos Aires, P. Commerson s.n. (holotype: P-LA [P00357623, Morton neg. 8363]; isotypes: G [G00070233, G00070234], P [Morton neg. 8143], P).
Solanum angustifolium Lam. var. macrophyllum Dunal, Prodr. [A.P. de Candolle] 13(1): 90. 1852.
Type: Brazil. Bahia: sin. loc. [possibly Rio de Janeiro], J. Guillot s.n. (holotype: P [P00319635, Morton neg. 8144]).
Solanum brittonianum Morong, Ann. New York Acad. Sci. 7: 174. 1893.
Type: Paraguay. Pilcomayo River, 10 Jan 1888-1890, T. Morong 1531 (holotype: NY [NY00139076]; isotypes: MO [MO-1787131], NDG, US [US-1324465], WIS).
Solanum handelianum Morong, Ann. New York Acad. Sci. 7: 175. 1893.
Type: Based on Solanum angustifolium Lam., non Solanum angustifolium Mill., 1768.
Type.

Based on Solanum persicifolium Mart., non Solanum persicifolium Dunal, 1813.

Description.

Woody vine to 5+ m long, scrambling in low vegetation, semi-aquatic and along water courses. Stems strongly ridged with 4 whitish green wings along the entire length, completely glabrous; new growth minutely papillose, occasionally pubescent with tangled simple uniseriate trichomes, these soon deciduous. Bark of older stems green to pale yellowish green, the bark not markedly exfoliating. Sympodial units plurifoliate, not geminate. Leaves simple, 2–6 cm long, 0.5–2 cm wide, lanceolate to linear (very occasionally with a few shallow lobes, but these lobed leaves always accompanied by simple ones on the same stem), somewhat fleshy to chartaceous, glabrous on both surfaces; primary veins 4–6 pairs, not prominent on either surface; base attenuate; margins entire, not markedly revolute; apex tapering to acute, the ultimate tip rounded; petiole 0.1–0.5 cm long, glabrous or occasionally with a few scattered simple trichomes adaxially, twining to aid climbing. Inflorescences terminal, becoming lateral and sometimes leaf-opposed, 4–13 cm long, usually 4–5 times branched, with 8–15 flowers, glabrous except for a few weak simple uniseriate trichomes at the tips of the branches; peduncle 0.5–2.5 cm long, occasionally absent and the branching beginning at the base of the inflorescence; pedicels 1–1.5 cm long, ca. 1 mm in diameter at the base and apex, slender, spreading at anthesis, glabrous, articulated at the base from a small sleeve and leaving a peg to 1.5 mm high on the inflorescence axis; pedicel scars widely spaced 3–10 mm apart. Buds ellipsoid, the corolla ca. 3/4 exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 1.5–3 mm long, conical, the lobes 1–1.5 mm long, deltate to broadly semi-circular, glabrous, the tips papillate. Corolla 2.5–4 cm in diameter, violet, rotate-stellate, lobed ca. 1/2 of the way to the base, the lobes 8–10 mm long, 8–9 mm wide, planar to spreading at anthesis, abaxially densely pubescent with minute simple uniseriate trichomes ca. 0.2 mm long, adaxially glabrous with a few simple trichomes along the midvein. Filament tube minute, the free portion of the filaments ca. 1 mm long, densely pubescent with tangled weak simple uniseriate trichomes to ca. 0.5 mm adaxially so the ovary obscured; anthers 5–6 mm long, 1–1.5 mm wide, ellipsoid, loosely connivent, yellow, poricidal at the tips, the pores usually lengthening to slits with age. Ovary glabrous; style 11–15 mm long, sparsely pubescent in the lower 1/3 to 1/2; stigma clavate, the surface minutely papillate. Fruit a globose to ellipsoid berry, 1–1.2 cm in diameter, to 1.5 cm long, black and dull when mature (yellowish fide Cabrera 1983), glabrous, the pericarp thin; fruiting pedicels 1.2–1.5 cm long, ca. 1.5 mm in diameter, more or less woody, pendent from the weight of the berry. Seeds > 40 per berry, 1.5–2 mm long, 1–1.5 mm wide, rounded to flattened-reniform, pale yellow, the surfaces minutely pitted, the testal cells circular. Chromosome number: not known.

Figure 14.

Solanum amygdalifolium Steud. (A, C–E drawn from Nee 52046 B drawn from Mendoza 102). Illustration by Bobbi Angell.

Distribution

(Figure 15). In the Río de la Plata drainage from Buenos Aires, Argentina and adjacent Uruguay to the upper Río Pilcomayo in Paraguay, and in coastal Brazil from Bahia south to Rio Grande do Sul, from 0-700 m elevation. Solanum amygdalifolium is also cultivated outside of its native range for its showy flowers (Bolivia, Andean Argentina).

Figure 15.

Distribution of Solanum amygdalifolium Steud.

Ecology.

Occurs in chaco vegetation along streams and rivers, in thickets and in open vegetation.

Common names:

Argentina. duraznillo (Morton 1976); Jujuy: jazmín (Cabrera 1983); Tucumán: amor porteño (Morton 1976).

Conservation status.

Least Concern (LC); EOO >50, 000 km2 (LC) and AOO >5, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum amygdalifolium occurs over a very broad geographical range in association with fresh (non-brackish) water, and has been characterised as semi-aquatic by some authors (Mentz and Oliveira 2004). In chaco habitats in Paraguay it always grows in riverside thickets, and forms loose scrambling tangles. With its very large, showy flowers, strongly angled stems and narrow, simple leaves, it is not easily confused with any other species of the Dulcamaroid clade; it is somewhat similar to other dulcamaroids from southern South America, particularly narrow-leaved specimens of Solanum flaccidum, but the smaller flowers, more pubescent leaves and unequal filaments of the latter species are distinguishing features. Solanum flaccidum grows in completely different types of habitats than does Solanum amygdalifolium, so confusion in the field is unlikely.

In general, Solanum amygdalifolium is quite monomorphic vegetatively over its broad range, varying only in degree of pubescence between individuals and somewhat in inflorescence size depending on plant age, but the flowers of plants from the Chaco regions of Argentina and Paraguay are much larger than those of plants from more coastal populations near the mouth of the Río de la Plata in Buenos Aires and adjacent Uruguay. Flower size may be related to water availability, as plants collected from near streams and wet places in the wet season all appear to have larger flowers than those from drier areas. The specimens collected in the foothills of the Andes in the provinces of Jujuy and Salta, Argentina appear to have all been from cultivated plants (Cabrera 1983).

Two collections were cited in Martius’s original description of Solanum persicifolium; Martius 255 appears to be the more widely distributed of the syntypes cited in the protologue and as it is a numbered collection is more reliable for tracing duplicates. The duplicate in M here selected as the lectotype has annotations in Martius’s hand. Morton annotated his photograph [Morton neg. 8668] of Martius s.n. (São Paulo, the other syntype) at M as holotype, but there is no evidence on the specimen or in the protologue that this is the case.

Morong (Morong and Britton 1893) clearly intended Solanum handelianum as a replacement name for Solanum angustifolium Lam.; he cites “S. angustifolium Lam., Illus. no. 2343, not Miller” and although he cites a collection of his own from near Asunción in Central Paraguay (Morong 818), he does not append “n.sp.” to the epithet as he does for all new taxa he described in 1893 from his own collections.

Specimens examined.

Argentina. Buenos Aires: Buenos Aires, 1831, Bacle 55 (F, G, G-DC); La Plata, Berisso, 15 Dec 1941, Beffon s.n. (SI); Barrancas de Belgrano, Apr 1946, Castellanos 822 (CORD); Buenos Aires, 10 Apr 1900, Debeaux 70 (F, GH, US); Río Negro, Allen, Feb 1939, Hunziker 161 (CORD); Martín Coronado, 8 Apr 1942, Hunziker 3616 (CORD); Río de la Plata, Reserva Natural Costanera Sur, 9 Jan 1993, Liede & Conrad 3003 (MO); Avellaneda, Feb 1918, Molfino 25707 (US); Isla Martín García, Apr 1935, Pastore 326 (SI); Quilmes, Rodríguez 179b (A, SI); Barrancas al Sud, 12 Mar 1902, Venturi 40 (CORD); Catamarca: Ancasti, Las Palomas, 5 Mar 1950, Brizuela 884 (CORD); Pomán, Dec 1910, Spegazzini s.n. (SI); Chaco: La Fidelidad, 1918, Jörgensen 2827 (SI); 1 de Mayo, Colonia Benítez, 30 Sep 1971, Martínez et al. s.n. (SI); Colonia Benítez, orilla Río Tragadero, 24 Mar 1948, Schulz 7251 (F, MO); Corrientes: Bella Vista, Paraje Rincón del Ambrosio, 12 Oct 1975, Irigoyen 248 (MO); Esquina, Islas frente a Esquina, 30 Nov 1974, Krapovickas et al. 26860 (G, MO); Esquina, Colonia Libertador, arroyo Barrancas, 15 Mar 1975, Krapovickas et al. 27773 (G, MO); Bella Vista, Bella Vista, nameless island in Río Paraná, some 2 km above Bella Vista, 28 Jan 1956, Pedersen 3716 (G, GH, US); Concepción, Carambola, 29 Feb 1972, Pedersen 10075 (A, CORD, L, MO, S); Empedrado, Estancia La Yela, 2 Jun 1984, Pedersen 13917 (G, MO); Córdoba: San Justo, San Francisco, 7 Dec 1946, Baliguo 944 (B); Capital, Córdoba, estación de ferrocarril Manuel Belgrano, 15 Mar 2000, Chiarini 316 (CORD); Capital, Córdoba, Apr 1916, Stuckert 23259 (CORD); Río Segundo, Pilar, en un cerco sobre calle 25 de Mayo, 20 Oct 1991, Subils 4520 (CORD); Entre Ríos: Uruguay, Concepción del Uruguay, alrededores del Banco Pelay, 2005, Barboza et al. 1566 (CORD); Gualeguaychú, alrededores, Dec 1936, Cabrera 3966 (F); Victoria, Isla del Pillo, Río Paraná, 20 Apr 1984, Oberti s.n. (CORD); Victoria, Isla del Pillo, casa de Sr. Tomas A. Nuñez, 7 Apr 1996, Oberti s.n. (CORD); Delta de Paraná y Medanos (Sud de Entre Rios), costas del Río Paraná-Mirri, 7 Jan 1904, Pennington 128 (CORD); Concepción del Uruguay, Banco Pelay, en camino a Banco Pelay, 5 Apr 1994, Solís Neffa et al. 59 (GH); La Paz, Isla Curuzú-Chalí, 30 Jan 1981, Troncoso & Bacigalupo 3127 (MO); Formosa: Pirané, Palo Santo, NE de Palo Santo, entre Palo Santo et le Riacho Pilagá, 85 m, 8 Nov 1986, Charpin & Eskuche 20253 (G); Los Matacos, 15 km east, 600 m, 11 Oct 1938, Eyerdam & Beetle 22940 (GH); Pilcomayo, Paraíso, 8 Oct 1948, Morel 6155 (B); Laishí, Herradura, 12 Oct 1950, Pedersen 1231 (G, US); La Rioja: Estación Cebollar, 13 Jan 1910, Spegazzini s.n. (SI); Mendoza: Guaymallén, El Borbollón, 28 Feb 1947, Villafaño 828 (B); Misiones: Posadas, in ripa limosa fluminis Alto Paraná, 16 Nov 1907, Ekman 816 (G, S, US); Capital, Nemesio Parma, 30 Feb 1994, Guillén 393 (CORD); San Pedro, El Alcázar, 6 Apr 1949, Schwindt 1499 (CORD); Salta: Anta, Puerto Verde, 19 Dec 1947, Luna 551 (B); Rosario de Lerma, Campo Quijano, 1200 m, Feb 1941, Meyer 3654 (F, GH); Capital, Ciudad de Salta, Av. Chile, proximo al Río Arenales, 12 Jun 1977, Novara et al. 443 (CORD); Capital, Ciudad de Salta, vias del FFCC entre la estación y calle Olavarria, 1190 m, 25 Oct 1989, Novara 9053 (CORD, G, S); General Güemes, Ojo de Agua, ruta 10, 6-9 km al NE de Gral. Guemes, 650 m, 12 May 1990, Novara & Bruno . 9873 (CORD, G, S); San Juan: Rawson, Médano de Oro, 650 m, 16 Jan 1987, Kiesling & Maglioli 6455 (SI); Santa Fe: General Obligado, Arroyo Malabrigo, entre Reconquista y Fortín Olmos, ca. 25 km de Reconquista, 10 Nov 1954, Hunziker 10372 (CORD); Canal Viejo de Santa Fé a Colastiné, Jan 1936, Job 734 (F, GH); Isla Mascota, La Reconquista, Río Paraná, Jan 1936, Job 948 (F); General Obligado, Villa Ocampo, (Isla), 29 Jan 1964, Panigatti 483 (CORD); Las Colonias, entre Esperanza y Santa Fé, 11 Apr 1984, Penseiro 1510 (SI); San Cristóbal, Arroyo Las Conchas, Ruta 13, 4 Apr 1984, Prado 625 (CORD); D.P.Malabrigo, F.C. [ferrocarril] a Reconquista, 13 m de Chico Malabrigo, 20 Nov 1906, Schroeter 164 (SI); Santiago del Estero: Rivadavia, Ruta Nac. 34, 100 m, 17 Jul 1964, Hunziker & Cocucci 17773 (CORD); Quebracho, Paso de la Cina, a orillas de Río Dulce, a unos pocos km al norte de Limache, 6 Jul 1991, Hunziker 25443 (CORD); Robles, Colonia Jaímez, 12 Sep 1948, Luna 1354 (CORD); Tucumán: Villa Luján, 11 Feb 1919, Venturi 272 (SI, US).

Bolivia. Santa Cruz: Cordillera, Bañados de Izozog, Cachari, ‘ACARARENDA’, 340 m, 11 Mar 1991, Navarro et al. 294 (MO); Florida, Samaipata, 1700 m, 17 Jul 2000, Wood 16448 (K).

Brazil. Mato Grosso do Sul: margens do Rio Miranda, 2 Jul 1983, Conceicao 1441 (MO); Rio Grande do Sul: Guaíba, 24 Jan 1949, Rambo 40159 (B, F); Porto Alegre, Rio Guahyba, Mar 1899, Reineck & Czermack 379 (G); Morro do Côco, Viamao, 22 Nov 1979, Soares 175 (F); Rio de Janeiro: environs de Rio de Janeiro, Glaziou 13087 (K, LE); Botafogo, Guillot s.n. (P); Itaipu, near Rio de Janeiro, 28 Oct 1828, Lund s.n. (BM); Rio de Janeiro, Macrae s.n. (GOET); Eugenho Velho, Rio de Janeiro, Miers 3885 (K); Rio de Janeiro, May 1832, Riedel 407 (US); Rio de Janeiro, 1844, Widgren s.n. (S).

Paraguay. Alto Paraguay: Parque Nacional Defensores de Chaco, alrededores del Madrejòn, 17 Jul 1985, Brunner 1218 (G); Madrejón, 20 Jun 1983, Hahn 1474 (BH, G); Boquerón: Estancia Toro Mocho, 17 Feb 2006, Egea et al. 886 (BM); Central: Asunción, Jun 1874, Balansa 2012 (G); Ypacaray, Cuervo Cui (Cordillera de Altos), Sep 1913, Hassler 11756 (BM, E, G, GH, L); Lago Ypacaraí, Feb 1903, Fiebrig 939 (E, F, G, GH, K, L); Asunción, Bahia, Jun 1988, Mereles 1193 (G); Río Salado, cruce en el camino Linguio-Embocada, Jul 1971, Schinini 4030 (G); Limpio, Rincón El Peñon, 27 May 1987, Zardini 2672 (G); Estero de Ypoá, Puerto Guyrati on Río Paraguay, 22 Jun 1993, Zardini & Guerrero 36385 (G); Concepción: Puerto Risso, on Rio Paraguay, between Rio Apa & Rio Aquidaban, 1908, Fiebrig 4079 (G, K); prope Concepción, Aug 1901, Hassler 7193 (BM, G, GH, K, P, S); Río Napegue, Aug 1988, Mereles 1311 (G); Concepción, S. Chaco, 91 m, Jun 1944, Sandeman 4834 (K); Cordillera: Río Salado [Palado], Aug 1901, Hassler 3207 (BM, G, GH, K, S); San Bernardino, Feb 1915, Rojas 1624 (S); San Bernardino, ufer des Río Salado, Feb 1916, Rojas 8563 (K); western side of Río Piribebuy basin, 27 km W of Arroyos y Esteros, 19 May 1990, Zardini & Velázquez 20317 (G); Guairá: Yataití, river Tebacuary, Mar 1933, Jörgensen 4657 (GH); Itapúa: Trinidad, 1914, Chodat 39 (G); Misiones: Llanuras de Santa Ana, Anonymous 76 (G); Villa Florida, Río Tebicuary, en banco de arena frente a la cuidad, 20 Jul 2000, Mereles & González Parini 7990 (G); Paraguarí: Estero de Ypoá, northern part, 10 km above Nueva Italia on Arroyo Cañabe, 23 Jun 1990, Zardini & Velázquez 21657 (G); Presidente Hayes: Carretera trans-Chaco, km 320, 15 May 1984, Billiet & Jadin 3063 (MO); Estancia Vanguardia, 29 Mar 2004, Egea & Centrón 376 (BM); Estancia La Golondrina, Villa Hayes, 9 Sep 1982, Hahn 673 (BH, G); Estancia Santa Asunción, 25 Oct 2004, Peña-Chocarro & Egea 1940 (BM); Ruta Ñ, frente a la entrada a Vanguardia, 3 Feb 2005, Peña-Chocarro & Egea 2404 (BM).

Uruguay. Colonia: Colonia Valdense, near Gallant, Feb 1957, Dubugnon 213 (G); Montevideo: Montevideo, Courbon s.n. (G); Montevideo, Fruchard s.n. (F); Atahualpa, 25 m, Apr 1925, Herter 190 (BH, F, G, GH, MO, S, SI); Tacuarembó: Rincón de la Laguna, 15 Feb 1947, Castellanos 17776 (CORD); ruta 26 km 300, pasando el puente sobre el arroyo Yaguarí, 28 Nov 2001, Seijo et al. 2541 (MO).

5. Solanum angustifidum Bitter, Repert. Spec. Nov. Regni Veg. 12: 544. 1913

http://species-id.net/wiki/Solanum_angustifidum

Figure 16
Type.

Argentina. Córdoba: San Vicente, T. Stuckert 4021 (lectotype, designated by Morton 1976, pg. 62: CORD [CORD00004111]; isolectotypes: G [G00070138], GOET [GOET-5966]).

Description.

Erect or lax, scandent shrubs, 1–2 m. Stems erect, glabrous; new growth glabrous. Bark of older stems reddish brown to grey. Sympodial units plurifoliate, not geminate. Leaves deeply pinnatifid (pinnate) with 1–4 pairs of leaflets, 2–6.5 cm long, 2–4 cm wide, elliptic in outline, membranous, both surfaces glabrous; primary veins 4–6 pairs; base attenuate onto the petiole; margins deeply pinnatifid, the leaflets (0.5-)2–2.5 long, 0.3–0.4 cm wide at their widest point; apex rounded; petioles 0.5–1.5 cm long, glabrous, sometimes twining. Inflorescences terminal or occasionally lateral, 2–6 cm long, several times branched, with 20–50 flowers, glabrous but with a few simple uniseriate trichomes at the point of pedicel insertion; peduncle 1–3 cm; pedicels 0.7–1(-1.7) cm long, slender, ca. 0.5 mm in diameter at base and apex, glabrous, spreading at anthesis, articulated at the base in a short sleeve, leaving a tiny peg on the inflorescence axis; pedicel scars irregularly spaces 1–4 mm apart. Buds ellipsoid, the corolla exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 1–1.5 mm long, cup-shaped, the lobes 0.5–1 mm long, deltate, glabrous or minutely papillate at the tips. Corolla 1.6–2 cm in diameter, lilac, purple or occasionally white, stellate to deeply stellate, lobed 2/3 to 3/4 of the way to the base, the lobes 6–9 mm long, 3–4 mm wide, spreading at anthesis, densely papillate on the margins and midvein abaxially, glabrous adaxially. Filament tube minute, the free portion of the filaments variable, with one filament longer than the rest 2–2.5 mm long, the other four 1–1.5 mm long, all glabrous; anthers 5–6 mm long, 1–1.5 mm wide, ellipsoid, yellow, poricidal at the tips, the pores opening to slits with age. Ovary glabrous; style 9–11 mm long, glabrous; stigma capitate, minutely papillose. Fruit a globose berry, 0.7–1 cm in diameter, black or violet-black when ripe, the pericarp thin and shiny, glabrous; fruiting pedicels 1–1.5 cm long, ca. 1.5 mm in diameter at the base, woody, pendent; fruiting calyx tightly investing the lower portion of the berry and slightly accrescent. Seeds 10–20 per berry, ca. 3 mm long, 2.5 mm wide, flattened reniform, pale brown, the surfaces minutely pitted, the testal cells square or circular. Chromosome number: not known.

Figure 16.

Solanum angustifidum Bitter. (A drawn from Tressens et al. 560 B–E drawn from Pedersen 7150 F drawn from Cristóbal et al. 1618). Illustration by Bobbi Angell.

Distribution

(Figure 17). Central and northern Argentina, with a few collections from Bolivia; 45–2000 m. Often cultivated (see Figure 1) and found escaped; Bolivian specimens are thought to be escapes from cultivation (Morton 1976).

Figure 17.

Distribution of Solanum angustifidum Bitter.

Ecology.

Widely distributed in many vegetation types, from chaco habitats to high elevation dry forests.

Common names.

Argentina. jazmín, jazmín de Córdoba, jazmín de cielo (Morton 1976); Córdoba: yerba vaca (Stuckert 2416); Jujuy: jazmín (Cabrera 1983).

Conservation status.

Least Concern (LC); EOO >50, 000 km2 (LC) and AOO >5, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum angustifidum is distinctive amongst members of the Dulcamaroid clade in having consistently pinnatifid leaves that are divided nearly to the midrib into usually three pairs of linear lobes; in general in the group pinnatifid and simple leaves both occur on individual plants. Although normally a shrubby plant, the petioles of leaves on terminal branches occasionally are elongate and twine around other vegetation. This led Morton (1976) to suggest that this characteristic made Solanum angustifidum a linking species between the sections Dulcamara and Jasminosolanum (both now recognised as part of the larger more inclusive Dulcamaroid clade, Weese and Bohs 2007).

This species is not easily confused with any other in the region; Solanum salicifolium has simple or only shallowly pinnatifid leaves and simple inflorescences of usually white flowers, and Solanum endoadenium has densely glandular pubescent leaves and orange (rather than black) fruits. Solanum viscosissimum of southeastern Brazil has deeply pinnatifid leaves, but is strictly vining and also always has simple leaves on the same stems. Solanum seaforthianum is the only other member of the Dulcamaroid clade to have leaves that are almost always pinnate and anthers on unequal filaments, but that species has much broader leaf divisions, rather than the narrow lobes of Solanum angustifidum, and is always a vine.

Cabrera (1983) states that Solanum angustifidum is commonly cultivated in Argentina for its abundant flowers (see Figure 1), and material from Bolivia appears to have been in association with habitations (Morton 1976; M. Nee, pers. comm.), suggesting that Solanum angustifidum is not native there, but is cultivated for its showy purple flowers.

Specimens examined.

Argentina. Buenos Aires: Jardín Botánico, 25 Nov 1922, Anon. s.n. (SI); Distrito Federal, Buenos Aires, en el jardin de la estancia Prack, Hunziker 2267 (CORD); Pergamino, Parodi 8832 (GH). Catamarca: Santa María, La Soledad, 12 Feb 2012, Barboza et al. 3489 (BM, CORD). Chaco: Resistencia, Margarita Belén, 15 Oct 1946, Aguilar 920 (G); Barranqueras, 35 m, 12 Nov 1913, Curran 31 (US); Tapenagá, Enrique Urien, campo Bonazzola, lote 9, Nov 1940, Rodrigo 2405 (GH); San Fernando, 15 km W de Ruta 12 por Ruta 89, 29 Dec 1976, Schinini 13864 (G, MO); Colonia Benítez, Apr 1932, Schulz 2066 (MO); Colonia Benítez, 23 Sep 1941, Schulz 3906 (MO); bords du Río Las Garzas, 100 m, 17 Nov 1902, Wagner s.n. (P); Corrientes: Paso López, ruta 12, 1.2 km NW de Paso Lopez, 22 Nov 1969, Carnevali 1746 (MO); Sauce, Río Guayquiraró, 18 km S de Sauce, 9 Oct 1977, Cristóbal et al. 1618 (MO); Emechado, El Pallo, 2 leguas al Este, 29 Aug 1945, Ibarrola 3188 (G); Esquina, Ruta 27, 27 km S de Esquina, 1 Dec 1974, Krapovickas et al. 27062 (G, MO); Esquina, Arroyo Saturno, 14 Mar 1975, Krapovickas et al. 27684 (G, MO); Empedrado, on the road from San Luis de Palmar to Mburucuyá, just north of the ford across the Río Empedrado, 20 Sep 1952, Pedersen 1834 (G, K, MO, US); Empedrado, Estancia La Yala, 20 May 1956, Pedersen 3916 (G, LE, US); Curuzú Cuatiá, Ruta 12 km 716, 20 Dec 1957, Pedersen 4785 (G, US); Saladas, Santo Domingo, 21 Jan 1950, Schwarz 9313 (CORD); Saladas, Estancia Pancho, 14 Feb 1950, Schwarz 9702 (CORD); Curuzú Cuatiá, Paso López, 17 km E, ruta 12, 29 Oct 1974, Tressens et al. 560 (MO); Córdoba: Capital, Córdoba, camino que une la Evenida Colon al 6000 con la ruta provincial 28 (ex. nac. 20), por detras de la Escuela de Aviación, 2 Dec 1973, Ariza Espinar 2857 (CORD); Punilla, Valle Hermoso, Valle de Punilla, Vaquerias, en camino de tierra que va de Casa Grande a Valle Hermoso, despues del segundo paso a nivel (a la derecha), 14 Dec 1972, Barrera et al. 8244 (CORD); Sierra Chica, Cerro Negro cerca de Cerro Azul, 9 Mar 1955, Castellanos 3353 (CORD); Ischilín, Deán Funes, 5 Nov 1945, Cuezzo 873 (B, CORD); Punilla, Salsipuedes, 6 Apr 1947, Dawson 1744 (K); Colón, entre Salsipuedes y Bello Horizonte, en el camino a La Falda, 29 Oct 1974, Hunziker 22611 (CORD); La Falda, Jan 1936, Job 548 (S, US); Alta Gracia, 600 m, 30 Mar 1930, King 631 (BM); Capital, Córdoba, 1 Jan 1885, Kurtz 964 (CORD); San Alberto, Villa Cura Brochero, Sierra Grande, falda O, 2 Feb 1948, Meyer 13518 (CORD); Colón, Jesús María, 530 m, 23 Dec 1947, Meyer 13602 (CORD); Colón, Salsipuedes, 26 Mar 1944, O’Donell & Rodríguez 690 (A); San Alberto, Nono, 22 Mar 1944, O’Donell & Rodríguez 721 (A, CORD); Colón, Ascochinga, 14 Mar 1944, O’Donell & Rodríguez 884 (F); Hendiolaza, 16 Mar 1944, Ruiz de Huidrobo 69 (GH); Río Segundo, Pilar, 2 May 1999, Subils 4662 (CORD); Entre Ríos: Capital, Río Paraná, 1891, Anetto s.n. (CORD); ruta Villaguaey a C. del Uruguay, bajos del Río Gualaguaey, 10 Nov 1973, Burkart et al. 30139 (MO, SI); La Paz, Paso Yunque, Río Guayquiraró, 7 Nov 1973, Burkart et al. 30151 (SI); Río Gualeguay, ruta Villaguay a C. del Uruguay, 10 Nov 1973, Burkart 30319 (GH, K); Las Barrancas, Doering s.n. (CORD); Puerto Nuevo, barranca del Paraná, 1 Dec 1912, Hicken s.n. (SI); Feliciano, San José de Feliciano, El Caraya, Feb 1948, Martínez Crovetto 4850 (K); Villaguay, Estancia Santa Martha, 1937, Museh s.n. (SI); Federación, Estancia Buena Esperanza, 15 Nov 1964, Pedersen 7150 (A, K); N of La Paz, Estancia Santa Cruz Cué, 7 Nov 1965, Walter & Walter 331 (B); Formosa: Jujuy, Pilagá, 5 Oct 1945, Pierotti 4201 (B); Pilagá, Pilagá, 3 Oct 1945, Pierotti 4271 (BM); La Rioja: Sanagasta, Sanagasta, 1500 m, 18 Feb 1944, Hunziker 4752 (CORD); Famatina, Famatina, Ruto 40, 9 Jan 1947, Hunziker 1825 (CORD); Famatima, Aguaditas, 7 Apr 1949, Toscani 51 (K); Salta: La Caldera, ruta 9, 18 km camino a Jujuy, 15 Nov 1947, Dawson 2032 (K); La Caldera, Dique Campo Alegre, ruta 9, km 1232, proximo a ruinas abandonadas de antiguo camping, 10 Jan 2005, Novara 12197 (CORD); San Juan: Iglesia, Arrequintín, 15 Mar 1989, Pedersen 15243 (G); San Luis: San Martín, San Martín, 1000 m, 13 Feb 1944, Varela 519 (CORD, G); Santa Fe: San Martín, Piamonte, alrededores, 24 Nov 1974, Astegiano 35 (CORD); General Obligado, Berna, ruta 11, km 759, 25 Jan 1992, Bernadello & Galetto 790 (CORD); San Cristóbal, Arroyo Las Conchas, Ruta 13, 26 Nov 1983, D’Angelo & Penseiro 611 (CORD); Cuty Lai, 6 Apr 1917, Hosseus 75 (CORD); Vera, entre Margarita y Vera, 8 Nov 1954, Hunziker 10357 (CORD); Capital, Santa Fe, en el Río Paraná, 7 Apr 1901, Kurtz 11857.5 (CORD); Sorrento, 7 Apr 1901, Kurtz 11857 bis (CORD); Villa Guillermina, Nov 1939, Meyer 3067 (US); General Obligado, camino entre Villa Guillermina y El Rabón, 18 Feb 1988, Penseiro & Tivano 3262 (CORD); San Justo, Fives Lille, 4 Jan 1937, Ragonese 2680 (US); Gral. Obligado, Mocoví, 1904, Venturi 51 (CORD, K); Tucumán: Capital, Tucumán, 450 m, 17 Oct 1897, Lillo 2030 (A); Camino Madillal, road Tucumán to Racas, 760 m, 27 Dec 1935, Mexia 4333 (MO).

Bolivia. Potosí: Suipacha, 2800 m, Aug 1946, Cárdenas 3731 (US). Tarija: Tarija, Jul 1932, Cárdenas 188 (GH); Mendez, Tomatitas, ca. 5 km N of Tarija, 2050 m, 21 Jan 2000, Wood et al. 15824 (K).

6. Solanum aspersum S.Knapp, PLoS ONE 5(5): e10502. 2010

http://species-id.net/wiki/Solanum_aspersum

Figure 18
Type.

Colombia. Putumayo: vertiente oriental de la Cordillera, entre Sachamates y San Francisco de Sibundoy, 1600-1750 m, 30 Dec 1940, J. Cuatrecasas 11471 (holotype: COL; isotypes: F [F-1335119], US [US-1799731]).

Description.

Woody vine, of unspecified length or height. Stems densely and evenly pubescent with antrorsely curved simple uniseriate trichomes 0.5–1.5 mm long, these few-celled with a large basal cell, arising from expanded bases and eventually deciduous; new growth densely pubescent with simple uniseriate trichomes to 1.5 mm long, these pale straw-colored in herbarium specimens; bark of older stems greenish brown, minutely tuberculate from the bases of the deciduous trichomes. Sympodial units plurifoliate. Leaves simple, (1-)3.5–9 cm long, (0.6-)1.5–4.6 cm wide, ovate to narrowly ovate, widest in the basal third, membranous or chartaceous, strongly discolorous, the upper surfaces evenly pubescent on veins and lamina, the trichomes to 2 mm long, simple, uniseriate, arising from expanded bases giving the lamina surface a tuberculate appearance, the lower surfaces densely and evenly pubescent with simple uniseriate trichomes to 2 mm long, these 2–3-celled with the basal cell largest, denser on the veins; primary veins 7–9 pairs, impressed above in herbarium specimens; base truncate or shallowly cordate; margins entire, not revolute; apex acute to acuminate; petioles 0.7–2 cm long, densely pubescent with simple trichomes like those of the stems and leaves. Inflorescences terminal on leafy short shoots, 3–15 cm long, globose to ellipsoid in outline, branching many times, with 2 principal basal branches, with 12–60 flowers, densely pubescent with simple trichomes; peduncle 0.5–3 cm long, the branching very near the junction with the stem; pedicels 0.5–0.8 cm long, < 0.5 mm in diameter at the base and apex, pubescent with 1–2-celled simple trichomes to 1.5 mm long, spreading at anthesis, articulated near the base from a small sleeve, leaving a small peg on the axis; pedicel scars irregularly spaced 1–10 mm apart, the inflorescence rachis bending at almost right angles at articulation points. Buds narrowly ellipsoid, the corolla strongly exserted from the calyx tube. Flowers all perfect, 5-merous. Calyx tube ca. 2 mm long, conical, the lobes 0.5–1 mm long, deltate to broadly deltate, pubescent with simple trichomes, these sparser than on the rest of the inflorescence. Corolla 1.2–1.7 cm in diameter, white, pink or “pale blue” (violet?), narrowly stellate, lobed nearly to the base, the lobes 6–7 mm long, 1.5–2 mm wide, reflexed at anthesis, densely pubescent abaxially with weak simple papillate trichomes to 0.5 mm long, these denser on tips and margins, glabrous adaxially. Filament tube < 0.5 mm, the free portion of the filaments ca. 0.5 mm long, glabrous; anthers 4–4.5 mm long, ca. 1 mm wide, ellipsoid, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 5–6 mm long, pubescent with weak simple trichomes to 0.5 mm, more densely pubescent in the basal half; stigma capitate-truncate, the surface minutely papillose. Fruit a globose berry, ca. 1.3 cm in diameter (immature?), green or yellowish green, the pericarp thin and shiny, glabrous; fruiting pedicels 0.9–1 cm long, 1–1.5 mm in diameter at the base, woody and spreading. Seeds not seen from mature berries, apparently 10+ per berry and flattened reniform. Chromosome number: not known.

Figure 18.

Solanum aspersum S. Knapp. (A–C drawn from MacDougal et al. 4251 D–E from Zak 1857A). Originally published in Knapp (2010). Illustration by Bobbi Angell.

Distribution

(Figure 19). Solanum aspersum occurs in widely separated and isolated populations along the Andes from central Ecuador into Colombia, from 1600 to 2500 m.

Figure 19.

Distribution of Solanum aspersum S.Knapp

Ecology.

Montane forests and forest margins.

Conservation status.

Vulnerable/Near Threatened (VU/NT); EOO >50, 000 km2 (LC) and AOO >5, 000 km2 (LC); the widely separated, isolated populations and few collections, however, mean that this species is of conservation concern. See Moat (2007) for explanation of measurements.

Discussion.

The few specimens of Solanum aspersum have usually been annotated as the more common and widely distributed Solanum aureum, also from Andean Ecuador. Solanum aspersum differs from that species in its simple uniseriate, rather than congested-dendritic, pubescence and in the elongate buds that open to deeply stellate flowers. Specimens of Solanum aureum from Azuay province in Ecuador have shiny adaxial leaf surfaces like those of Solanum aspersum, but always have the characteristic golden dendritic pubescence of that species rather than the simple pubescence of Solanum aspersum. The leaves of Solanum aspersum are usually more cordate than those of Solanum aureum, but some populations of Solanum aureum approach Solanum aspersum in overall leaf morphology at first glance. Solanum aspersum has a very scattered distribution all along the Andes from northern Colombia to central Ecuador and is likely to be found in more of the intervening parts of the cordilleras, but it is apparently rare and easily overlooked.

A single collection from Cajamarca in northern Peru (Diaz et al. 9717, NY) with simple trichomes on the inflorescence and completely glabrous leaves may also be this species; it shares the elongate buds and simple trichomes, but may represent a different taxon.

Specimens examined.

Colombia. Antioquia: Frontino, Nutibara, region Muri, camino hasta La Blanquita, 1440 m, 10 Jul 1986, Acevedo et al. 1213 (US); Urrao, between Urrao and caicedo, 21 km E of Urrao, near high point on road, 2710 m, 27 Feb 1989, MacDougal et al. 4251 (MO). Cauca: between La Cumbre and Quebrada La Isla, headwaters of Río Dinde, 28 Aug 1944, Core 1111 (US); Cundinamarca: Chiquinquirá, 2500 m, 17 Feb 1950, Sneidern 5825 (S).

Ecuador. Napo: Parroquia Cosanga, 6 kms de la carretera Cosanga-El Aliso, 2200 m, 23 Aug 1990, Jaramillo et al. 12110 (MO); Cantón Quijos, Reserva Ecológica Antisana, Río Aliso, 8 km al suroeste de Cosanga, 2500 m, 15 Nov 1998, Vargas et al. 3043 (MO). Pichincha: Reserva Florística-Ecológica “Río Guajalito”, km 59 de la carretera antigua Quito-Santo Domingo de los Colorados, a 3.5 km al NE de la carretera, 2200 m, 28 Mar 1987, Zak 1857A (F, MO).

7. Solanum aureum Dunal, Solan. Syn. 16. 1816.

http://species-id.net/wiki/Solanum_aureum

Figure 20
Solanum loxense Dunal, Solan. Syn. 16. 1816.
Type: Ecuador. Loja: Loja, A. Humboldt & A. Bonpland s.n. (holotype: P [P00136336]).
Solanum clematideum Bitter, Repert. Spec. Nov. Regni Veg. 16: 86. 1919.
Type: Ecuador. Azuay: Cuenca, Pindilic [Pindilig], 2600–3000 m, 11 Oct 1888, F. Lehmann 4948 (holotype: B [F neg. 2654], destroyed; lectotype, designated here: K [K000438685]).
Solanum aureum Dunal var. riobambense Werd., Notizbl. Bot. Gart. Berlin-Dahlem 12: 379. 1935.
Type: Ecuador. Chimborazo: Ríobamba, N of Sanancajas, 3300 m, 28 Jul 1933, L. Diels 355 (holotype: B, destroyed; no duplicates traced).
Type.

Ecuador. Base of Chimborazo, June 1802, A. Humboldt & A. Bonpland 3142 (holotype: P [P00136318, Morton neg. 8151]; isotypes: B-W [B-W-4318, IDC microfiche 271-315.297:5, F neg. 2890], P [P00136319]).

Description.

Woody vines or lax shrubs to 5 m tall, often in open areas. Stems densely pubescent with stiff, golden dendritic trichomes to 1 mm long, the trichome branches numerous and very short; new growth densely golden-pubescent with congested dendritic trichomes. Bark of older stems pale brownish yellow, glabrescent. Sympodial units plurifoliate, not geminate. Leaves simple, 1.5–7(-11) cm long, 0.9–3(-6) cm wide, elliptic to narrowly elliptic, membranous to slightly chartaceous or fleshy, discolorous, the upper surfaces shiny, sparsely and evenly pubescent with erect dendritic trichomes to 0.5 mm long, these with short and congested branches, occasionally glabrous with only a few trichomes along the midvein, the lower surfaces densely pubescent on veins and lamina with erect, golden, dendritic trichomes to 1 mm long, with short, congested branches, the surface appearing golden; primary veins 7–10 pairs, paler than the lamina, golden beneath; base truncate to acute, occasionally oblique; margins entire, not markedly revolute; apex acute; petioles 0.5–2 (3.5) cm long, densely golden pubescent with dendritic trichomes like those of the stems, occasionally twining to aid in climbing. Inflorescences terminal, occasionally becoming lateral and internodal, 5–10 cm long, globose in outline, many times branched, leafy near the basal branches, with 20–40 (+) flowers, densely pubescent with golden dendritic trichomes like those of the stems; peduncle very short, the transition between stem and inflorescence not always clear; pedicels 0.5–0.6 cm long, stout or appearing so from dense pubescence, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading at anthesis, densely golden-pubescent with stiff dendritic trichomes to 1 mm long, articulated at the base leaving a small sleeve or peg 0.5–1 mm long on the axis; pedicel scars irregularly spaced 1–10 mm apart, obscured by the pubescence. Buds globose to broadly ellipsoid, the corolla strongly exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 2–2.5 mm long, conical, the lobes 1.5–2 mm long, deltate to narrowly deltate, the apex blunt, densely to sparsely pubescent with golden dendritic trichomes to 1 mm, the pubescence usually sparser than that of the pedicel. Corolla 1.5–1.7 cm in diameter, violet or purple to deep purple, stellate, lobed ca. 2/3 of the way to the base, the lobes 5–7 mm long, 3.5–5.5 mm wide, planar at anthesis, densely pubescent abaxially with a mixture of simple and dendritic trichomes to 0.5 mm long, these weak and tangled, denser on the tips and margins, the interpetalar sinuses glabrous, glabrous adaxially. Filament tube ca. 0.5 mm long, the free portion of the filaments ca. 1 mm long, glabrous; anthers 4–5 mm long, 1.2–1.5 mm wide, glabrous, ellipsoid, loosely connivent, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 7–8 mm long, glabrous or densely and minutely papillate in the basal 1.3; stigma capitate, minutely papillate. Fruit a globose berry, ca. 1 cm in diameter, purplish black and shiny when ripe, glabrous, the pericarp thin; fruiting pedicels 1–1.2 cm long, ca. 1 mm in diameter at the base, woody and spreading, the calyx lobes woody and persistent around the base of the berry. Seeds 20–30 per berry, ca. 3 mm long, 2.5–3 mm wide, flattened-reniform, the margins undulate, reddish brown, the testal surface minutely pitted, the cells pentagonal in outline. Chromosome number: not known.

Figure 20.

Solanum aureum Dunal. (A drawn from Juncosa 2335, NY B drawn from Zak 1845 C–G drawn from Rose 22963 H drawn from Cerón et al. 11842). Illustration by Bobbi Angell.

Distribution

(Figure 21). In Andean Ecuador and Peru, from 2500 to 3700 m elevation, most commonly collected around 3000 m.

Figure 21.

Distribution of Solanum aureum Dunal.

Ecology.

Most commonly collected in montane forests, páramos and páramo margins (“ceja de selva” or “jalca”).

Conservation status.

Least Concern (LC); EOO >50, 000 km2 (LC) and AOO >5, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum aureum is widespread in Ecuador and has been considered to include several species here recognised as distinct: Solanum sanchez-vegae of northern Peru, Solanum dichroandrum of northern Colombia and Venezuela and Solanum aspersum of Ecuador and Colombia, with which Solanum aureum is sympatric. Solanum aureum is almost always described as a vine, but occasionally as shrubby, a habit variously common in the Dulcamaroid clade. Solanum sanchez-vegae and Solanum dichroandrum differ from Solanum aureum in their looser, less dense leaf pubescence and somewhat larger flowers. All three taxa have dark purplish black berries, but those of Solanum aureum are somewhat smaller. Solanum aureum and Solanum sanchez-vegae may overlap in distribution in northern Peru in the Huancabamba depression; Solanum dichroandrum is only known Colombia and Venezuela. Specimens now recognised as Solanum aspersum had long been considered as conspecific with Solanum aureum, but differ from it in their simple trichomes from bullate bases, elliptic buds and flowers with more lanceolate corolla lobes. These two species are sympatric in northern Ecuador. Solanum cutervanum is also sympatric with Solanum aureum through much of its range and is easily confused with it. The most obvious difference in the two taxa is habit, with Solanum aureum being a vine and Solanum cutervanum a shrub, but also the leaf and stem trichomes of Solanum cutervanum are beige, more elongate and the branches shorter than those of Solanum aureum, in which the trichomes are golden and more classically dendritic.

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

In describing both Solanum aureum and Solanum loxense, Dunal cited specimens in the Humboldt and Bonpland herbarium (“v.s.h. H. et B.”); no sheets of either are presently in P-Bonpl., but material collected by Humboldt and Bonpland is housed in the general collection and is certainly that used by Dunal in describing these taxa. Many specimens from the original Humboldt and Bonpland collection were apparently re-integrated into the general collections after use in the 19th century rather than being returned to the special collection (see Knapp and Spooner 1999; Knapp 2007b).

Specimens examined.

Colombia. Cauca: Pitayó, Prov. of Popayán, Anonymous s.n. (K); Putumayo: road from Sibundoy to Pasto, between La Maria and Páramo de San Antonio, 2900 m, 1 Jun 1946, Schultes & Villarreal 7536 (GH, K, US); Valle del Cauca: Cuchilla de Barragán, Cordillera Central, vertiente occidental, hoya del Río Bugalagrande, 3250 m, 12 Apr 1946, Cuatrecasas 20605 (F, US).

Ecuador. Azuay: Cantón Cuenca, Yanasacha, Parroquia Baños, 2925 m, 26 Dec 1976, Boeke 595 (GH, MO); Cruz Pamba region above Baños, ca. 15 km southwest of Cuenca, 2743 m, 29 Jun 1945, Camp E-3940 (MO); Campamento Molón, road in construction Sigsig-Gualaquiza, 2900 m, 11 Apr 1968, Harling et al. 8218 (MO x2); Cumbe, 24 Sep 1918, Rose et al. 22963 (GH, US); Bulán, Padrehurco-Amapala, en las riberas del río Chanín, 2776 m, 18 Oct 2000, Verdugo et al. 285 (HA); Cantón Girón, Girón, El Chorro, 2700 m, 22 Jun 2001, Verdugo et al. 353 (HA); Bolívar: Río Tataguazó, carretera Santiago-San Vicente-Alizo “San Agustin”, 2500 m, 24 Feb 1987, Zak 1741 (F); Carchi: Páramo del Ángel, 3300 m, 19 Jul 1945, Acosta Solís 10571 (F); El Ángel, 1 Jan 1931, Benoist 3630 (P); La Rincónada, ranch between Ibarra and Tulcán, 3000 m, 10 Aug 1923, Hitchcock 20947 (GH, US); ca. 2 km along the road El Angel-Tulcán, 3150 m, 14 May 1973, Holm-Nielsen et al. 5207 (F, MO); Road Julio Andrade-Palestina, 3300 m, 27 Dec 1980, Holm-Nielsen et al. 29707 (BM); near El Angel on old road to Tulcán, 3060 m, 23 Feb 1984, Juncosa 2335 (MO); Cantón Espejo, Reserva Ecológica El Ángel, Libertad-Morán, El Salado, 3300 m, 30 Oct 1993, Palacios 11547 (MO); Quebrada del Río Angel, 3 km NE of San Isidro on road to El Angel, 28 Jan 1967, Sparre 14280 (S); Cañar: north of Biblian, 2900 m, 6 Apr 1974, Harling & Andersson 13239 (MO); carretera Alausi-Cañar, desvio a Oyashig-Hacda. El Carmen al N de Cañar, 3270 m, 12 Aug 1987, Zak 2400 (F, MO); carretera Alausi-Cañar, desvio a Oyashig-Hacda. El Carmen al N de Cañar, 3270 m, 12 Aug 1987, Zak 2403 (F, S); Chimborazo: Cubillín, Cordillera Oriental, 3300 m, 1 Mar 1944, Acosta-Solís 7575 A (F); Cantón Riobamba, Parque Nacional Sangay, páramo de Pinlilligue, entre Alao y La Tranca, 3300 m, 18 Aug 1990, Cerón et al. 11842 (MO); 3 km E of Alao, above Río Alao, on road to Huamboya (Morona-Santiago), 2 Dec 1988, Dorr & Barnett 6207 (MO); Cantón Riobamba, Riobamba, 3300 m, 22 Mar 1934, Schimpff 896 (G); Bayushig, carretera Riobamba-Penipe-Bayushig, 3400 m, 18 Feb 1986, Zak 1635 (F); Cantón Riobamba, Parque Nacional Sangay, carretera Chimborazo-Chambo-Pungalá-Alao, 3150 m, 9 Aug 1987, Zak 2360 (F, F, MO); Cotopaxi: Pilalo, 2400 m, 9 Jul 1968, Holm-Nielsen, & Jeppesen 1541 (S); Road Pilaló-Zumbagua, 10 km above Pilaló, 3150 m, 28 Jul 1980, Holm-Nielsen & Quintana 24654 (BM); Imbabura: Shanshipamba-La Esperanza, 2950 m, 16 Nov 1949, Acosta-Solís 14422 (F); Laguna Cuicocha, crater lake 30 W of Ibarra, steep slopes around lake and on the smaller island in the lake, 3100 m, 24 May 1973, Holm-Nielsen et al. 6321 (S); Cantón Otavalo, Otovalo, Páramo de Mojanda, 3200 m, Lehmann K-222 (K); Lake Cuicocha, island in lake, 3100 m, 29 May 1939, Penland & Summers 761 (F, GH, US); Cantón Cotacachi, Reserva Ecológica Cotacachi-Cayapas, islote Teodoro Wolff, laguna de Cuicocha, 3100 m, 30 Aug 1991, Peñafiel et al. 313 (BM, MO); Carretera Pimampiro-Chuga-Palmar Chico; entre Chuga y Palmar Chico, 2800 m, 16 Jan 1988, Zak & Jaramillo 3353 (BM, F x2, GH, MO, MO); Loja: between Loja and San Lucas, 2100 m, 6 Sep 1923, Hitchcock 21497 (GH, US); Cerro de Celica. Celica-Guachanamá, Km 14-18, 2700 m, 13 Apr 1994, Jørgensen et al. 157 (BM); El Cisne-Zaruma, unfinished road km 2.9, 2340 m, 12 Dec 1994, Jørgensen et al. 1397 (MO); Catamayo-Catacocha, km 25, turnoff at Las Chinchas towards Piñas, km 2.3, 2250 m, 13 Dec 1994, Jørgensen et al. 1463 (MO); Pichincha: Cantón Cayambe, Oyacachi, Cordillera Oriental, 3200 m, 26 Sep 1945, Acosta-Solís 11148 (F); entre Oyacachi y Comenia, Cordillera Oriental, 3000 m, 27 Sep 1945, Acosta-Solís 11204 (F); El Llalo, 3201 m, 29 Dec 1938, Balls B-5826 (BM, E, K, US); base du Pichincha, 23 Mar 1930, Benoist 2211 (P); Paluguillo, 19 Feb 1931, Benoist 3908 (P); Cantón Cayambe, Reserva Cayambe-Coca, zona de amortiguamiento, 3200 m, 2 Jan 2000, Cuamacás et al. 579 (MO); Cantón Quito, Pifo, Hacienda Los Andes, 3500 m, 24 Jan 1991, Palacios 6906 (MO); Lloa, Sodiro s.n. (P); Hac. Paluguillo, on road Pifo-Papallacta, 3200 m, 19 Jul 1967, Sparre 17727 (S); carretera Quito-Guantopugro-Yanacocha, 3400 m, 22 Mar 1987, Zak 1845 (F, MO); Cantón Mejía, Parroquia El Chaupi, faldas del Volcán El Corazón, 3300 m, 23 May 1988, Zak & Jaramillo 3698 (BM, MO); Tungurahua: entre Leito y La Cima, Cordillera Oriental, 2700 m, 15 Nov 1944, Acosta-Solís 8994 (F); Cantón Mocha, Mocha, 6 Jul 1896, André 3922 (K); road between El Triunfo and Patate, summit of road that descends to Patate, 3140 m, 26 Feb 2003, Bohs et al. 3132 (UT); Cantón Mocha, Mocha, base of Chimborazo, 2743 m, Jameson 46 (K); Cantón Ambato, Ambato, 3048 m, Jul 1939, Sandeman 37 (BM, K); Cantón Mocha, Mocha, alrededores del pueblo, 2900 m, 30 Sep 1995, Villacres 373 (F).

Peru. [perhaps Ayabaca?], 1909, Weberbauer 6411 (F); Cajamarca: San Ignacio, San José de Lourdes. Base del Cerro Picorana, 25 Aug 1999, Díaz et al. 10743 (MO); Piura: Huancabamba, carretera entre Canchaque y Huancabamba, 2800 m, 14 Jan 1988, Díaz et al. 2719 (MO); Huancabamba, Cruz Blanca-Turmalina, 2600 m, 15 Sep 1981, López M. et al. 8926 (BM, MO); Huancabamba, Huancabamba, 2591 m, Aug 1943, Sandeman 4262 (K).

8. Solanum boldoense Dunal & A.DC., Prodr. [A.P. de Candolle] 13(1): 679 1852

http://species-id.net/wiki/Solanum_boldoense

Figure 22
Solanum cardiophyllum Dunal, Prodr. [A.P. de Candolle] 13(1): 89. 1852, non Solanum cardiophyllum Lindl., 1848.
Type: Cuba. Havana: “in Havana”, B.M. Boldo s.n. (holotype: MPU).
Solanum scandens Sessé & Moc., Fl. Mexic., ed. 2, 53. 1894.
Type: Cuba. Sin. loc., Sessé & Mociño s.n. (lectotype, designated by Knapp 2008b, pg. 20: MA [MA-604676, F neg. 48318]; isolectotype: MA [MA-604675, F neg. 48343, 5381, upper frag. in photo, sheet now divided]).
Type:

Based on Solanum cardiophyllum Dunal, non Solanum cardiophyllum Lindl., 1848.

Description.

Woody vining shrub or liana. Stems flexuous, glabrous; new growth minutely pubescent, the trichomes ca. 0.4 mm, simple, uniseriate, later glabrous. Bark of older stems dark reddish brown. Sympodial units plurifoliate. Leaves simple or occasionally very shallowly 3-lobed, 4.5–7 cm long, 3–5 cm wide, ovate to cordate or narrowly cordate, glabrous on both surfaces or with a few minute uniseriate simple trichomes along the midrib above; primary veins 5–7 pairs, drying reddish; base cordate or truncate and oblique; margins entire; apex acuminate; petioles 2.3–3.5 cm long, glabrous, twisting to aid in climbing up supports. Inflorescences leaf-opposed or terminal, 10–30 cm long, ovoid to ellipsoid in overall shape, branching to 5 times, with 50–100 flowers, glabrous; peduncle 1.5–9 cm long, glabrous; pedicels slender, 1.2–1.7 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, nodding, glabrous, articulated in distal quarter just below the calyx tube leaving an elongate peg, occasionally articulated in the basal half of the pedicel, but always leaving a distinct remnant; pedicel scars widely spaced ca. 1 cm apart, appearing as a series of elongate pegs due to articulation point. Buds globose and somewhat inflated, the corolla strongly exserted from the calyx tube. Flowers all perfect, 5-merous. Calyx tube 3.5–4 mm long (to articulation), the upper part 2–2.5 mm long atop a conical receptacle-like structure, the lobes absent or mere undulations on the rim of the tube, glabrous. Corolla 2–2.3 cm in diameter, purple or violet, stellate, lobed ca. 3/4 of the way to the base, the lobes 0.8–1 mm wide, 0.4–0.6 cm wide, planar (or slightly cupped?) at anthesis, glabrous, the margins and cucullate tips densely papillose. Filament tube < 0.1 mm, glabrous; free portion of the filaments ca. 0.75 mm long; anthers 4–4.5 mm long, 2–2.5 mm wide, stout, poricidal at the tips, the pores lengthening to slits with age. Ovary conical, glabrous; style 0.8–0.9 cm long, glabrous, the stigma capitate, the surface minutely papillose. Fruit a globose berry, 1–1.2 cm in diameter, red when ripe, the pericarp thin and shiny; fruiting pedicels 1.1–1.3 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, not particularly woody, deflexed, the basal portion of the calyx tube expanding in fruit to be clearly differentiated above the articulation point, appearing somewhat swollen. Seeds ca. 10 per berry, 3–3.5 mm long, 1.5–2 mm wide, flattened reniform, pale brown, the surfaces minutely pitted, the testal cells sinuate in outline. Chromosome number: not known.

Figure 22.

Solanum boldoense Dunal & A.DC. (A–G drawn from Clark 10597 and J. Clark photographs taken in the field H drawn from Wright 381, NY). Illustration by Bobbi Angell.

Distribution

(Figure 23). Known only from Cuba and a single collection from Haiti, at low to middle elevations. Several very old single collections seen from Jamaica (K000196458) and Puerto Rico (Plée s.n., P00549340) may mislabelled as to locality or from cultivated plants.

Figure 23.

Distribution of Solanum boldoense Dunal & A.DC.

Ecology.

A relatively rare plant in forests and along forest edges.

Conservation status.

Possible Near Threatened (possible NT); EOO <100, 000 km2 (possible NT) and AOO <100, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum boldoense is very similar to, and has been confused with (see below), Solanum dulcamaroides of Mexico. The pedicel articulation point serves to easily distinguish the two species: in Solanum boldoense it is the distal quarter of the length and long pegs are left when flowers fall, while in Solanum dulcamaroides the pedicel articulates at or very near the base.

Dunal used the name Solanum cardiophyllum for this species, but before publication realised it had already been used by Lindley for a species of potato. His replacement name is based on the same type, collected by B.M. Boldo near Havana.

Considerable confusion exists over the use of the epithet “scandens” in Sessé and Mociño’s works, but it is clear that on page 53 of Flora Mexicana, they were using Solanum scandens in the sense of a new name, different from that of Solanum scandens L., which they re-named Solanum nutans. The specimen (MA-604676) labelled “S.scandens IC.” in the Sessé and Mociño herbarium at MA with the locality “Hava. et Queretaro” demonstrates the confusion over the identity of all these plants. MA-604676 is a specimen of Solanum boldoense, suggesting [confirming?] the localities were added to the labels and the labels to the specimens after the fact, and that botanists in Madrid were not treating these two species are different. Another sheet of the same plant, MA-604675, appears to be from the same gathering. The “IC.” referred to on the label of MA-604676 may be plate 6331.1503 of the Torner Collection, that was used by Dunal to describe Solanum dulcamaroides (see discussion below). The plate is only partly finished and could be either species, but the pedicel articulation near the rhachis suggests it is Solanum dulcamaroides; epitype material is selected in the treatment of Solanum dulcamaroides to fix the usage this name.

Specimens examined.

Cuba. Havana: Havana, Delessert s.n. (G); Havana, 1841, Karwinski s.n. (LE); Las Tunas: In Cuba Orientali [note on US sheet - “fl. purple, Cliffs, Rio Sta. Cruz, July”], 1856, Wright 381 (G, G, GH, GOET, K, LE, MO, S, US); Las Villas: Gavinas, Trinidad Mountains, San Blas-Buenos Aires, 18 Sep 1941, Gonzales 177 (GH); Pinar del Río: San Diego de los Baños, 31 Aug 1910, Britton et al. 6674 (F, US); La Palma, Pan de Guajaibón, trail on or near summit of Pan de Guajaibón, 700 m, 20 Dec 2008, Clark et al. 10597 (BM); San Cristobal, Loma del Pimiento, 29 Sep 1920, Ekman s.n. (S); on the road between Mameyar and El Toro, 14 Sep 1923, Ekman 17544 (S); Santa Cruz de los Pinos, vicinity of Taco-Taco River, 29 Oct 1925, Brother León 12543 (GH); Santa Clara: Las Lagunas, Buenos Aires, 762 m, 5 Dec 1928, Jack 6823 (A).

Haiti. La Hotte, between La Cueva and Placer Bonita, 1067 m, 1 Aug 1950, Howard 12255 (GH).

Jamaica. Sin. loc, Anonymous s.n. (K, LE).

Puerto Rico. “Antilles. Porto Rico”, Plée s.n. (P).

9. Solanum calileguae Cabrera, Hickenia 1: 162. 1978

http://species-id.net/wiki/Solanum_calileguae

Figure 24
Type.

Argentina. Jujuy: Dpto. Valle Grande, road to Altos de Calilegua, 31 Oct 1974, A.L. Cabrera, N.B. Deginani, A. Giaioti, R. Kiesling, E. Zardini & F.O. Zuloaga 25639 (holotype: SI; isotype: LP [n.v.]).

Description.

Woody vine. Stems scandent, glabrous to pubescent with a mixture of transparent simple and dendritic uniseriate trichomes to 1 mm long; new growth densely pubescent with simple and mostly dendritic trichomes. Bark of older stems yellowish brown. Sympodial units plurifoliate. Leaves simple to deeply 5-lobed, the lower lobes complete and the leaves apparently pinnate, 5.5–9 cm long, 2.5–3.5 cm wide, narrowly elliptic, the divided leaves to 10 cm long, 10 cm wide, elliptic in outline, membranous, the upper surfaces sparsely pubescent on the veins and lamina with dendritic 3–4-celled trichomes to 0.5 mm long, the lower surfaces densely pubescent along the veins with dendritic trichomes to 0.5 mm long like those of the upper surface, these sometimes extending to the lamina; primary veins 7–9 pairs; base truncate, oblique and asymmetric; margins entire or lobed, the lobes to 5 cm long, 2 cm wide; apex acute; petioles (1-)1.5–3(-6+) cm long, sparsely to densely pubescent with dendritic trichomes, especially adaxially, twining. Inflorescences terminal, later lateral, to 13 cm long, many times branched, with up to 30+ flowers, sparsely to densely pubescent with a mixture of simple and dendritic uniseriate trichomes to 0.5 mm long; peduncle to 8 cm long; pedicels 0.9–1.1 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, slender, spreading at anthesis, sparsely to densely pubescent with a mixture of simple and dendritic trichomes like those of the inflorescence axis, articulated at the base from a small sleeve, leaving a small peg on the inflorescence axis; pedicel scars irregularly spaced up to 9 mm apart. Buds broadly ellipsoid, the corolla about halfway exerted from the calyx before anthesis. Flowers all perfect, 5-merous. Calyx tube 2–3 mm long, conical, irregularly splitting into lobes 2–4 mm long, densely pubescent with dendritic trichomes, these denser on the tube. Corolla 2–3 cm in diameter, white, stellate-rotate, lobed 1/2 to 2/3 of the way to the base, the lobes 9–12 mm long, 4–5 mm wide, planar at anthesis, densely pubescent abaxially with minute simple trichomes to 0.5 mm long, pubescent adaxially along the distal half of the midvein with minute simple trichomes to 0.5 mm long, the tips cucullate. Filament tube minute, the free portion of the filaments 0.5–1 mm long, minutely puberulent within with tangled, weak simple uniseriate trichomes; anthers ca. 5 mm long, 2 mm wide, ellipsoid, yellow, loosely connivent, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style ca. 9 mm long, densely pubescent in the basal half with weak, tangled simple uniseriate trichomes; stigma minutely clavate, the surface minutely papillose. Fruit a globose berry, ca. 1 cm in diameter (fide Cabrera 1983). Seeds not seen from mature berries.

Figure 24.

Solanum calileguae Cabrera.(A–D drawn from Nee & Bohs 50809 E drawn from Nee & Bohs 50768). Illustration by Bobbi Angell.

Distribution

(Figure 25). Known only from the foothills of the Andes in the Provinces of Jujuy and Salta, Argentina, 1600–1850 m; possibly also occurring (sterile voucher only) in adjacent Bolivia.

Figure 25.

Distribution of Solanum calileguae Cabrera.

Ecology.

In premontane forests dominated by Juglans L. (Juglandaceae) and Podocarpus Pers. (Podocarpaceae).

Conservation status.

Endangered (EN); EOO <5, 000 km2 (EN) and AOO <2, 000 km2 (VU). See Moat (2007) for explanation of measurements.

Discussion.

Solanum calileguae is a distinctive species with its dense dendritic pubescence and large, white flowers. It has a restricted Andean distribution, and is easily distinguished from other members of the group potentially occurring in the area. It differs from Solanum uncinellum, which also has dendritic pubescence on the stems and leaves, in its ellipsoid (rather than long pointed) buds, its rotate-stellate rather than deeply stellate flowers and in its anthers borne on equal filaments. Solanum uncinellum, although widespread over most of South America, has only once been collected in the Andean foothills of Argentina, but not from the same area where Solanum calileguae occurs. Solanum calileguae is most similar and perhaps most closely related to Solanum flaccidum, from which it differs in the dendritic pubescence and anthers borne on equal filaments. Solanum flaccidum occurs in southeastern Brazil, and the two species are unlikely to be found together in the field.

Specimens examined.

Argentina. Jujuy: Ledesma, Abra de las Cañas, camino a Velle Grande, 24 km NW de Libertador Gral. S. Martín, 1600 m, 8 Nov 1974, Krapovickas et al. 26583 (G, MO); Palpalá, Cerro Zapla, above Quebrada Los Tomates, on road to Cerro Zapla, 1850 m, 13 Apr 2000, Nee et al. 50756 (CORD, NY); Valle Grande, 2 km N of Abra de Cañas on road from Calilegua to Valle Grande, 14 km by road S of San Francisco, 1600 m, 14 Apr 2000, Nee & Bohs 50768 (CORD, MO, NY); Valle Grande, ca. 1 km (by air) E on trail from San Francisco towards Altos de Calilegua, 1700 m, 18 Apr 2000, Nee & Bohs 50809 (BM, MO); Valle Grande, ca. 1 km (by air) E on trail from San Francisco towards Altos de Calilegua, 1700 m, 18 Apr 2000, Nee 50809 (MO); Salta: Santa Victoria, camino al Río San José, desde el desvio del camino de Los Toldos a Lipeo, 1823 m, 28 Sep 1998, Ahumada & Agüero 8178 (CORD).

10. Solanum coalitum S.Knapp, Novon 17: 212. 2007

http://species-id.net/wiki/Solanum_coalitum

Figure 26
Type.

Ecuador. Loja: Yangana-Valladolid, km 1.1, track to Sierra Toledo, km 18.5, 3250 m, c. 4°23'S, 79°06'W, 14 Nov 1997, G. Lewis & B. Klitgaard 3719 (holotype: LOJA; isotypes: AAU, BM [BM000846493], K [K000585495], NY [NY00888405], QCA, QCNE [QCNE-740]).

Description.

Subshrubs to 1 m tall, sometimes scandent and trailing. Stems glabrous and shining, usually appearing warty from the prominent leaf scars; young stems and leaves completely glabrous or sometimes with a few scattered loose white branched trichomes to 0.5 mm long. Bark of older stems dark brown, shining. Sympodial units plurifoliate. Leaves simple, 2.5–10.4 cm long, 0.7–3.5 cm wide, narrowly elliptic to less commonly elliptic, the upper surfaces glabrous and shiny, sometimes with scattered branched trichomes at the edge where the margin is revolute, the lower surfaces glabrous or sparsely papillate, the papillae drying reddish brown, perhaps glandular; primary veins 5–10 pairs, drying darker than the lamina; base acute to attenuate; margins strongly revolute, pubescent on the upper surfaces where turned under; apex acute; petiole 0.3–1.6 cm long, glabrous and shiny. Inflorescences terminal, 2.5–6 (-10) cm long, branched 4–6 times, with 3–15(-20) flowers, glabrous and shining, or with scattered loosely branched trichomes along the axes; peduncle 2–5.5 cm long; pedicels in flower 0.8–1.3 cm long, stoutish, ca. 1 mm in diameter at the base, ca. 2 mm in diameter at the apex, nodding, glabrous, minutely papillate or sparsely pubescent with loosely branched white trichomes ca. 0.3 mm long, articulated at the base and inserted in a short sleeve ca. 1 mm long; pedicel scars closely spaced and clustered at inflorescence branch tips. Buds globose when very young, soon elliptic and strongly exerted from the calyx tube. Flowers all perfect, 5-merous. Calyx tube 2.5–4 mm long, cup-shaped, strongly constricted at the base, thick and coriaceous, glabrous or with a few branched trichomes like those of the inflorescence axis, the lobes 1–1.5 mm long, broadly deltate or minute, glabrous, with the margins glabrous or densely pubescent with branched trichomes ca. 0.3 mm long. Corolla 2–2.6 cm in diameter, violet to dark mauve-purple, lobed 3/4 of the way to the base, stellate, the lobes 0.9–1.3 mm long, 0.5–0.8 cm wide, slightly campanulate or planar at anthesis, densely pubescent with simple or dendritic trichomes ca. 0.5 mm long on the margins and tips, sometimes with scattered simple trichomes on the abaxial lobe surface, these denser on the petal midvein, the adaxial surface glabrous. Filament tube less than 0.5 mm long; free portion of the filaments 1–1.5 mm long, glabrous; anthers 5–6 mm long, 1.5–2 mm wide, loosely connivent, poricidal at the tips, the pores lengthening to slits with age. Ovary conical, glabrous; style 1–1.2 cm long, straight, glabrous; stigma clavate or 2-lobed, bright green (fide Lewis & Klitgaard 3719), the surface minutely papillate. Fruit a globose berry, 1.2–1.5 cm in diameter, shiny and black when mature, the pericarp thin; fruiting pedicels 2–2.2 cm long, 2–2.5 mm in diameter at the base, 2–3 mm in diameter at the apex, erect, thick and woody. Seeds ca. 10 per berry, 4–5 mm long, 3–4 mm wide, flattened reniform, reddish brown, the margins not enlarged, the surfaces minutely pitted, the testal cells sinuate in outline. Chromosome number unknown.

Figure 26.

Solanum coalitum S. Knapp. (A, B drawn from Jorgensen 2188 C D drawn from G. Lewis photographs of Lewis 3179). Illustration by Bobbi Angell.

Distribution

(Figure 27). Endemic to Ecuador, only known from the páramo of Cerro Toledo S of Loja along the road leading to the Peruvian border, on ridges between the towns of Yangana and Valladolid, at 3150-3460 m in southwestern corner of Parque Nacional Podocarpus.

Figure 27.

Distribution of Solanum coalitum S.Knapp.

Ecology.

Páramos and forest edges.

Conservation status.

Critically Endangered (CR); EOO <100 km2 (CR) and AOO <10 km2 (LC). See Moat (2007) for explanation of measurements. Knapp (2007c) gave a preliminary conservation status of EN (endangered) or critically endangered for this species. The fact that Solanum coalitum occurs only within the boundaries of the Parque Nacional Podocarpus is good news for its ultimate protection and conservation, but its very restricted distribution in an isolated habitat means it certainly is of concern.

Discussion.

Solanum coalitum is a striking species with its large, fleshy bright purple flowers and black fruits. Some specimens of Solanum coalitum have been identified as Solanum stenophyllum, with which it is very similar. Solanum coalitum differs from Solanum stenophyllum in its subshrubby, sometimes trailing habit, its glabrous stems and leaves (except for the peculiar marginal trichome band), its cup-shaped (rather than conical) calyx and its slightly larger flowers that are glabrous abaxially. Trichomes of Solanum coalitum when they occur are looser and more openly dendritic than the almost echinoid trichomes that are distinctly yellowish of Solanum stenophyllum. The fruiting pedicels of Solanum stenophyllum appear to be nodding when fruit are mature, while those of Solanum coalitum are erect. Specimens of Solanum stenophyllum have been collected from the province of Loja (i.e., Jørgensen et al. 477, 1068, BM) from further north and at slightly lower (2600-3000 m) elevations than Solanum coalitum and in drier and/or degraded forests. Solanum stenophyllum grows as a shrub or small treelet, usually in disturbed situations. Some individuals of Solanum stenophyllum in southern Ecuador are very sparsely pubescent, but the conical calyx and yellowish closely branched trichomes serve to distinguish these plants.

Solanum coalitum is distinguished from the very similar Solanum imbaburense by its broadly deltate, rather than long triangular calyx lobes, and its leaves with sparsely papillate undersides.

The sole locality in which Solanum coalitum has been encountered is the páramo of Cerro Toledo in the extreme southwestern corner of Parque Nacional Podocarpus, one of the largest protected areas in Ecuador. Cerro Toledo is a mixed páramo of tussock grasses and shrubby vegetation on the divide of the Cordillera de Sabanillas; the area is a pathway used by local people to take cattle from one drainage to another, and as such has a medium level of disturbance (Hofstede et al. 2002). Roads constructed by the military to allow access to radio towers have opened the area to others. Hofstede et al. (2002) suggest that the inhospitable nature of the climate in the region (wet, cold and windy) will limit human incursion on a large scale. Cerro Toledo is isolated from other páramo regions of southern Ecuador, and represents one of the southernmost extensions of the páramo habitat in the Andes (Luteyn 1999).

Specimens examined.

Ecuador. Loja: Yangana, Cerro Toledo, ascending road from Yangana to Numbala toward the antennas. Area next to antennas at summit of cerro, 3400 m, 27 Mar 2005, Bohs et al. 3320 (BM, LOJA, QCNE); carretera Yangana-Toledo, 3420 m, 28 Dec 1988, Jaramillo 10606 (AAU); road from Yangana to Cerro Toledo, km 18-22 to antennas, 3460 m, 3 Nov 2000, Jørgensen et al. 2188 (BM, NY); Yangana-Cerro Toledo, páramo of Cerro Toledo, 3420 m, 28 Dec 1988, Jørgensen et al. (AAU, BM); Cerro Toledo, Parque Nacional Podocarpus, 3350 m, 1 Dec 1988, Madsen et al. 75641 (BM); Cerro Toledo, 2500 m, 30 Oct 1989, Madsen 86333 (BM); Cerro Toledo, E of Yangana, Parque Nacional Podocarpus, 3400 m, 26 Feb 1985, Øllgaard et al. 58162 (BM).

11. Solanum crispum Ruiz & Pav., Fl. Peruv. 2: 31, t. 158a. 1799

http://species-id.net/wiki/Solanum_crispum

Figure 28
Solanum ligustrinum Lodd., Bot. Cab. 1963. 1833.
Type: Chile. Sin. loc., seeds sent by H. Cuming in 1831 (lectotype, designated here: Loddiges, Bot. Cab. No. 1963. 1833).
Solanum concavum Lindl., Edwards’s Bot. Reg. 28(Misc.): 57. 1842.
Type: Chile. sin. loc., H. Cuming 263 (lectotype, designated here: CGE; isolectotypes: BM [BM000935822], E [E00057570], GH [GH00077603], K [K000585719]).
Solanum laetum Kunze, Linnaea 16: 352. 1842.
Type: Germany. Cultivated in Leipzig, 1837, Anon. s.n. (no specimens cited or found; synonymy ex descr.).
Solanum syringaefolium Kunth & C.D.Bouché, Ind. Sem. Hort. Berol. 10. 1845.
Type: Chile. sin. loc., T.C. Bridges s.n. (holotype: B, destroyed [F neg. 2746]).
Witheringia berteroana J.Rémy, in Gay, Fl. Chil. 5: 65. 1849, as “berterianum
Type: Chile. Tagua-Tagua, C.G.L. Bertero s.n. (lectotype, designated by Knapp 1989, pg. 74: P [P00324728]).
Witheringia crispa (Ruiz & Pav.) J.Rémy, in Gay, Fl. Chil. 5: 63. 1849.
Type: Based on Solanum crispum Ruiz & Pav.
Witheringia gayana J.Rémy, in Gay, Fl. Chil. 5: 67. 1849.
Type: Chile. sin. loc., C. Gay s.n. (lectotype, designated by Knapp 1989, pg. 74: P [P00335390]; isolectotype: K [K000585718]).
Witheringia tomatillo J.Rémy, in Gay, Fl. Chil. 5: 64. 1849.
Type: Chile. sin. loc., C. Gay s.n. (lectotype, designated by Knapp 1989, pg. 74: P [P00325721]; isolectotype: K [K000585722]).
Solanum angustifolium Lam. var. brevifolium Dunal, Prodr. [A.P. de Candolle] 13(1): 90. 1852.
Type: Chile. Sin. loc., 1828, E. Poeppig 74 [diar. n. 34] (holotype: G-DC [G00144942, Morton neg. 8402, IDC microfiche 800-61.2068:II.7]; isotypes: BM [BM000935826], LE, P [P00319643, Morton neg. 8145]).
Solanum congestiflorum Dunal, Prodr. [A.P. de Candolle] 13(1): 92. 1852.
Type: Chile. Aviluco, C.G.L. Bertero 634 (lectotype, designated by Knapp 1989, pg. 74: P [P00325722]; isolectotype: MO [MO-3461608]).
Solanum congestiflorum Dunal var. longifolium Dunal, Prodr. [A.P. de Candolle] 13(1): 92. 1852.
Type: Chile. Región × (Los Lagos): Valdivia: “in Chile australis provincia Valdivia”, C. Gay herb. 3e envoi 211 (lectotype, designated by Knapp 1989, pg. 74: P [P00325724]).
Solanum crispum Ruiz & Pav. var. elaeagnifolium Dunal, Prodr. [A.P. de Candolle] 13(1): 92. 1852.
Type: Chile. Región VI (Liberador): Prov. O’Higgins: circa Rancagua, 1833, C.G.L. Bertero 640 (lectotype, designated by Knapp 1989, pg. 74: G-DC [G00144948, Morton neg. 8406]; isolectotype: P [P00325727, Morton neg. 8175]).
Solanum crispum Ruiz & Pav. var. ligustrinum (Lodd.) Dunal, Prodr. [A.P. de Candolle] 13(1): 92. 1852.
Type: Based on Solanum ligustrinum Lodd.
Solanum pyrrhocarpum Phil., Anales Univ. Chile 21(2): 383. 1862.
Type: Chile. Región VIII (Bío-Bío): Prov. Ñuble: Chillan, sin. coll. (lectotype, designated by Knapp 1989, pg. 74: SGO [SGO-55460, barcode SGO000004551]).
Solanum sadae Phil., Linnaea 33: 203. 1864–1865.
Type: Chile. Región VI (O’Higgins): Colchagua, sin. coll. (lectotype, designated by Knapp 1989, pg. 74: SGO [SGO-55450, barcode SGO000004595]; isotype: E).
Solanum landbeckii Phil., Linnaea 33: 204. 1864–1865.
Type: Chile. Región VI (O’Higgins): Colchagua, C.L. Landbeck s.n. (lectotype, designated by Knapp 1989, pg. 74: SGO).
Solanum izquierdii Phil., Anales Univ. Chile 43: 522. 1873. as “izquierdi
Type: Chile. Región Metropolitana: Prov. Santiago, Aculeo (ca. 33°50'S, 70°55'W), V. Izquierdo s.n. (lectotype, designated by Knapp 1989, pg. 74: SGO [SGO-55455, barcode SGO000004573]; isolectotype: K [K000585727]).
Solanum gayanum (J.Rémy) F.Phil., Cat. Pl. Vasc. Chil. 228. 1881.
Type: Based on Witheringia gayana J.Rémy
Solanum tomatillo (J.Rémy) F.Phil., Cat. Pl. Vasc. Chil. 229. 1881.
Type: Based on Witheringia tomatillo J.Rémy
Solanum pugae Phil., Anales Univ. Chile 91: 7. 1895.
Type: Chile. Región VIII (Bío-Bío): Prov. Ñuble, Cerro Centinela, F. Puga s.n. (holotype: SGO, specimens not located).
Solanum pannosum Phil., Anales Univ. Chile 91: 9. 1895.
Type: Chile. Región VII (Maule): Prov. Curicó, Los Maquis, ca. 35°43'S, 70°47'W, M. Vidal s.n. (holotype: SGO, specimens not located; isotype: W [W-1908-10271]).
Solanum berteroanum (J.Rémy) Phil., Anales Univ. Chile 91: 8. 1896.
Type: Based on Witheringia berteroana J.Rémy
Solanum tagua Kuntze, Revis. Gen. Pl. 3(2): 226. 1898.
Type: Based on Witheringia berteroana J.Rémy
Solanum congestiflorum Dunal var. syringaefolium (Kunth & Bouché) Reiche, Anales Univ. Chile 123: 721. 1908.
Type: Based on Solanum syringaefolium Kunth & C.D.Bouché
Solanum congestiflorum var. pannosum (Phil.) Reiche, Anales Univ. Chile 123: 722. 1908.
Type: Based on Solanum pannosum Phil.
Type.

Chile. Región VIII (Bío-Bío): Concepción: “in Chile ruderatis copiosé in Conceptionis urbis sepibus, et ad Carcamo et Palomares tractus”, H. Ruiz & J. Pavón s.n. (lectotype, designated by Knapp 2008c, pg. 312: MA [MA-747012]; isolectotype: MA [MA-747101]).

Description.

Shrubs or small trees, often lax and scrambling, 0.4–5 m tall. Stems glabrous or pubescent with tiny dendritic trichomes; leaf scars somewhat prominent; new growth glabrous to densely pubescent with fine, dendritic trichomes. Bark of older stems pale brownish-yellow, glabrous, and shiny. Sympodial units plurifoliate. Leaves simple, 2.7–7.5 (10) cm long, 1–3 (7) cm wide, ovate to narrowly ovate, occasionally somewhat elliptic, larger and broader in plants growing in shade and in juvenile plants (see discussion), the adaxial surfaces glabrous or with a few dendritic trichomes along the main veins, the abaxial surfaces glabrous or puberulent with dendritic trichomes, these usually denser along the veins; primary veins 6–12 pairs, pubescent below; base truncate or somewhat cordate, not winged on to the petiole; margins entire, undulate or crispate; apex acute to acuminate; petiole 0.5–1 (2.2) cm long. Inflorescences terminal, later appearing lateral from overtopping of shoots, 2–10 cm long, flat-topped or pyramidal, branching 5–7 times, with 10–20 flowers, glabrous or sparsely pubescent with dendritic trichomes like those of the stems and leaves; peduncle 1–5 cm long; pedicels 1–1.3 cm long, tapering from a basal diameter of ca. 0.5 mm to an apical diameter of ca. 1 mm, nodding at anthesis, glabrous or with a few scattered dendritic trichomes, articulated at the base and inserted in a sleeve ca. 0.5 mm long; pedicel scars closely spaced in clusters. Buds globose when young, later elliptic, the corolla strongly exserted from the calyx tube. Flowers all perfect, 5-merous. Calyx tube 1–1.5 (2) cm long, conical, the lobes 0.5–1 mm long, deltate to long-triangular, glabrous or with a few scattered dendritic trichomes abaxially, adaxially glabrous. Corolla 1.2–2.5 cm in diameter, violet or occasionally white, lobed 3/4 to 7/8 of the way to the base, the lobes 5–9 mm long, 3–5 mm wide, planar or somewhat reflexed at anthesis, densely pubescent with simple (in glabrous plants) or dendritic (in pubescent plants) trichomes abaxially, the trichomes denser at the tips of the lobes, glabrous adaxially. Filament tube less than 0.5 mm long; free portion of the filaments 1–1.5 mm long, glabrous; anthers 3.5–5 mm long, 1–2 mm wide, loosely connivent, poridical at the tips, the pores becoming slit-like with age. Ovary glabrous or with a few dendritic and simple trichomes at the apex, especially in otherwise pubescent plants; style 0.6–1 cm long, pubescent with dendritic or simple trichomes along the entire length; stigma clavate or capitate, the surface minutely papillose. Fruit a globose or ellipsoid berry, 0.8–1 cm in diameter, bright red when ripe, changing from green to yellow or orange during ripening, with thin pericarp; fruiting pedicels 1.2–1.6 cm long, ca. 1 mm in diameter at the base, woody, deflexed. Seeds ca. 11 per berry, 2–3 mm long, 1.5–2 mm wide, flattened lenticular, reddish-brown, the surfaces minutely pitted. Chromosome number: n = 12 (vouchers: Knapp 8632, Knapp 8633 cultivated material).

Figure 28.

Solanum crispum Ruiz & Pav. (A drawn from Taylor 10235 B drawn from Nee 54654 C drawn from Landrum 8216 D drawn from Biese 78 E–H drawn from Landrum 4459). Illustration by Bobbi Angell.

Distribution

(Figure 29). Chile from Quillota south to the island of Chiloé, from 10–2500 m elevation. Solanum crispum is also known from scattered collections in Argentina along the border with Chile; in the Neuquén area these plants are usually found in association with villages and it is suspected that rather than being native, they have been brought from Chile and cultivated for their medicinal properties (C. Ezcurra, pers. comm.).

Figure 29.

Distribution of Solanum crispum Ruiz & Pav.

Ecology.

Solanum crispum grows in Nothofagus (Nothofagaceae) forest, often in second growth, and in a wide variety of moist microsites in otherwise dry habitats.

Common names.

Chile: tomatillo, natre, natrien, natri, tomatilla (see Knapp 1989).

Conservation status.

Least Concern (LC); EOO >50, 000 km2 (LC) and AOO >5, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum crispum is one of the most variable and is the most southerly species of the Solanum nitidum species group, occurring to 43 degrees S latitude. It also has a huge elevational range, occurring from sea level to nearly 3000 m in a wide variety of habitats. Two pubescence forms occur throughout the range of Solanum crispum: glabrous plants were traditionally called Solanum crispum and pubescent ones Solanum congestiflorum (see Knapp 1989). Specimens of intermediate pubescence are rare, but the new growth of glabrous plants is always dendritic-pubescent. In a cladistic analysis (Knapp 1989) the two forms were treated as separate, but were strongly resolved as sister taxa. Pubescence may be related to habitat, but this effect has not been studied in any detail; polymorphism in pubescence is extremely common in members of the Dulcamaroid clade and elsewhere in Solanum. The pubescent form often has larger, more repand leaves than does the glabrous form. This raises the intriguing possibility that the pubescent form is paedomorphic, retaining the shape and indument of juvenile leaves.

Medicinal uses of Solanum crispum have been recorded for over two centuries, beginning with Ruiz and Pavón in Flora Peruviana (1797), where the plant was reported to be used as a febrifuge. It has been reported as used against the fevers called ‘congo’ and ‘chavalongo’.

Solanum crispum has been cultivated in the United Kingdom since the early part of the 19th century. Specimens were introduced to Kew Gardens from the island of Chiloé (Chile) by Mr. Anderson (Hooker 1844) and a variety still in cultivation today was developed at the Glasnevin Botanic Gardens in Dublin. Solanum crispum is usually classed as a climber, but this is due to its lanky habit in the British climate. It does not possess the twining stems, petioles or tendrils of a true climber, but it is not a robust, erect plant.

Many monographers in Solanum have stated that holotypes or lectotypes for Ruiz and Pavón names were in the Madrid herbarium (MA), but without specifying a particular sheet. In a few cases, only one sheet exists, thus making lectotypification relatively straightforward, but in others multiple sheets exist in the Ruiz and Pavón herbarium at MA, meaning that previous type designations are not sufficiently precise. It is unlikely that any of these specimens are actually holotypes; the dispersal of specimens at the time of the expedition and subsequently through sale and loss means lectotypification is essential even if only a single sheet is present at MA. Knapp (1989) lectotypified Solanum crispum citing only a sheet in MA; this was rectified in 2008 by citation of the particular sheet (Knapp 2008c) as the lectotype.

Knapp (1989) erroneously lectotypified Solanum ligustrinum using one of the specimens cited by Dunal (1852) when making the new combination rather than the plate from Loddiges’ (1833) original description (in which seeds from an un-numbered collection sent by H. Cuming are cited). The epithet ligustrinum is correctly lectotypified here using Loddiges’ plate (Figure 30), as it is the only extant element unambiguously related to the protologue.

Figure 30.

Lectotype of Solanum ligustrum Lodd. (Loddiges 1833: tab. 1963). Reproduced with permission of the Natural History Museum Botany Library.

Specimens examined.

Argentina. Mendoza: Santa Rosa de los Andes to Uspallato Pass, Moseley s.n. (BM, LE); Neuquén: Minas, a 18 km de las Ovejas camino a las lagunas Epu-Lauquén, mallin de la Culebra, 1450 m, 14 Jan 1964, Boelcke et al. 10786 (SI).

Chile. Región IV (Coquimbo): Elqui, sector bordering road to Pachon, 38 km SE of Guard Control Post at Cerro Tololo, 2416 m, 16 Jan 2004, Acosta-Solís & León BB 181 (K); Illapel, Dec 1862, Landbeck s.n. (W); Elqui, Baño de Pangue, 1800 m, 20 Sep 1947, Sparre 2628 (S); Ovalle, Ovalle, ca. 86 km from Ovalle on road Caren to Río Mostazal, 1800 m, 11 Nov 1938, Worth Morrison 16448 (G); Región Metropolitana: Rio Colorado, Paso Uspallata, 6 Jan 1886, Philippi & Borchers s.n. (BM); Santiago, Oct 1933, Grandjot s.n. (GOET); Melipilla, Las Vizcachas, ca. 10 km from La Dormida, 1920 m, 7 Dec 1938, Morrison 16760 (G); Santiago, 1876, Philippi s.n. (G); Santiago, 1862, Philippi s.n. (G); Farallones, 2500 m, 15 Apr 1969, Plowman 2683 (LE); ); Santiago, Cerro San Cristobal, Nov 1869, Reed 1869 (BM); Santiago, Prov. Santiago, Cord de Santiago, Rio San Francisco, 2200 m, Dec 1924, Werdermann 439 (BM); Río San Francisco, Cordillera de Santiago, 2200 m, Dec 1927, Werdermann 479 (G); Región V (Valparaíso): Santa Rosa de los Andes, May 1882, Ball s.n. (LE); Los Señales, Dec 1829, Bertero 1323 (G-DC); Quillota, Aug 1829, Bertero 1327 (G-DC); Llaillai, “Atacama”, 9 Oct 1884, Borchers s.n. (F); Región VI (O’Higgins): prov. Colchagua, Cerro Echaurrina, San Fernando, 13 Oct 1926, Montero 33 A (F); Colchagua, 1862, Philippi s.n. (G); Región VII (Maule): Talca, Cordillera de Los Andes: 18 km on road to Laguna del Maule from first border post, 4 Feb 1998, Baxter et al. 26 (BM); Cordillera de Curicó, Valle del Toro, 1500 m, 1903, Bürger 48 (GOET); Región VIII (Bío-Bío): Baños de Chillán, Jan 1878, Anonymous s.n. (W); Concepción, 26 Sep 1939, Bailey 941 (BH); Concepción, Quebrada Honda, entre Liriquén y Tomé, 50 m, 16 Oct 1986, Basualto et al. 37 (MA); Ñuble, Termas de Chillán, Refugio El Aserradero, 1240 m, 13 Nov 1986, Basualto et al. 156 (MA); Talcahuano, 12 Nov 1950, Brooke 6951 (F); Biobío, Antuco, Cordillera de los Andes, Fundo Los Ciervos, passing into El Toro hydroelectric central and crossing the Río Polcura, 860 m, 28 Jan 2004, Brownless et al. DCI-1045 (BM); Ñuble, Chillán, road to Termas Chillán at Puente Torrealba, 1514 m, 25 Dec 2003, Gardner & Knees 6776 (BM); Ñuble, Chillán, 1856, Germains.n. (G); Prov. Ñuble, Chillán, 1856, Germain s.n. (F); Concepción, 1825, Macrae s.n. (G); Coronel, 1864, Oschenius s.n. (GOET); Chillán, Philippi s.n. (K); Ñuble, Termas de Chillán, 1450 m, 11 Dec 1987, Rechinger & Rechinger 64317 (B); Ñuble, Río Ñuble, 40-50 km desde San Fabian hacia la Cordillera, entre rio Nuble y rio Los Sauces, 26 Feb 1968, Zalensky III 81-82 (LE); Región IX (Araucanía): Malleco, Victoria, Hotel El Bosque on Ruta 5 south-bound from Victoria, 338 m, 24 Jan 2004, Brownless et al. DCI-903 (BM); Cautín, Villarrica, road from Meseta San Judas to western edge of Lago Colico, 500 m, 20 Dec 2003, Gardner & Knees 6734 (BM); Malleco, road from Victoria to Termas de Tolhuaca before Parque Nacional Tolhuaca near to Puente Tacadero, 942 m, 30 Dec 2003, Gardner & Knees 6870 (BM); Cautín, Río Zuerpe, 28 Sep 1905, Middleton s.n. (G x2); Región X (Los Lagos): Island of Chilóe, Anonymous 7966 (BM); Island of Chilóe, Anonymous 7967 (BM); Archipelago de Chiloe, Downton 6 (BM); Región XIV (Los Ríos): Valdivia, 20 Oct 1904, Buchtien s.n. (G); Valdivia, 22 Oct 1905, Buchtien s.n. (G); Valdivia, 28 Sep 1896, Buchtien s.n. (G); Chiloé, Chiloé, Caldeleugh s.n. (G); Valdivia, Valdivia, coastal road from Curiñanco to Niebla, 19 Dec 2003, Gardner & Knees 6723 (BM); Valdivia, Panguipulli, 150 m, Oct 1924, Hollermayer 324 (BM); Cordillera de Ranco, Lechler 827 (GOET); Valdivia, road from La Union to El Mirador, 700 m, 30 Mar 1969, Plowman 2640 (GH); Valdivia, Corral, Amargos, San Carlos, 20 m, 6 Jan 1954, Sparre & Smith 399 (G); Valdivia, Panguipulli, 150 m, Oct 1924, Werdermann 324 (G);

12. Solanum cutervanum Zahlbr., Ann. K. K. Naturhist. Hofmus. 7: 7. 1892

http://species-id.net/wiki/Solanum_cutervanum

Figure 31
Solanum angustifolium Ruiz & Pav., Fl. Peruv. 2: 33, t. 163b. 1799, non Solanum angustifolium Miller, 1768, nec Solanum angustifolium Lam., 1793.
Type: Peru. Huánuco: Acomayo, H. Ruiz & J. Pavón s.n. (lectotype, designated by Knapp 2008c, pg. 310: MA [MA-747093]; isolectotypes: F, MA [MA-747094, MA-747095]).
Solanum pulverulentum Pers., Syn. 1: 223. 1805.
Type: Based on Solanum angustifolium Ruiz & Pav., Fl. Peruv. 2: 33, t. 163b. 1799, non Solanum angustifolium Miller, 1768, nec Solanum angustifolium Lam., 1793.
Solanum aureum Dunal var. angustelanceolatum Bitter, Bot. Jahrb. Syst. 54, Beibl. 119: 13. 1916.
Type: Peru. Huánuco: Chaglla, 3100-3200 m, c. 9°46'S, 1909-1914, A. Weberbauer 6700 (holotype: B, destroyed; lectotype, designated by Knapp 1989: 90: F [F-628476, F neg. 69667]; isolectotypes: GH [GH00077582], MOL, US [US-1444961]).
Solanum aureum Dunal var. latelanceolatum Bitter, Bot. Jahrb. Syst. 54, Beibl. 119: 13. 1916.
Type: Peru. Huánuco: Chaglla, 3100-3200 m, c. 9°46'S, 1909-1914, A. Weberbauer 6700 (holotype: B, destroyed; lectotype, designated here: US [US-1444961]; isolectotypes: F, MOL, US).
Type.

Peru. Cajamarca: Cutervo, C. von Jelski 30 (holotype: W [W 1891-0004325, F neg. 33065]).

Description.

Shrubs to small trees, 1–7 m tall. Stems and leaves densely covered with loosely branching golden tree-like trichomes; leaf scars somewhat raised, the stem not winged; new growth densely pubescent with golden tree-like trichomes above and below. Bark of older stems dark reddish-brown, sparsely pubescent with the tree-like trichomes of the young stems. Sympodial units plurifoliate. Leaves simple, 6.5–13 cm long, 1.7–5 cm wide, elliptic or occasionally narrowly elliptic (type), the upper surfaces of the blades drying dark, sparsely pubescent with golden tree-like trichomes, these mostly along the veins, the lower surfaces pubescent with golden trichomes like those of the upper surfaces, the pubescence denser than that above; primary veins 8–12 pairs, sparsely pubescent; base acute, not winged on to the petiole; margins entire, not markedly revolute; apex acute; petiole 0.5–2 cm long, densely golden pubescent. Inflorescences terminal, later appearing lateral or in the fork of the branches, 4–10 cm long, pyramidal, branching ca. 10 times, with 10–20 flowers densely pubescent with loose golden tree-like trichomes like those of the young stems; peduncle 1–4 cm long; pedicels 0.7–1.3 cm long, ca. 0.5 mm in diameter at the base tapering to an apical diameter of 1 mm, slightly nodding at anthesis, densely pubescent with golden tree-like trichomes, articulated at the base and inserted in a sleeve ca. 0.5 mm long; pedicel scars closely spaced and congested at the inflorescence branch tips. Buds ellipsoid, the corolla strongly exserted from the calyx tube. Flowers all perfect, 5-merous. Calyx tube 1.5–2 mm long, conical, the lobes deltate, 1.5–2 mm long, densely pubescent abaxially with golden tree-like trichomes, densely pubescent adaxially with dendritic and simple trichomes. Corolla 1.5–1.8 cm in diameter, violet or occasionally white, lobed 3/4 of the way to the base, the lobes 7–8 mm long, 4–5 mm wide, planar at anthesis, densely pubescent abaxially with tiny dendritic trichomes, these denser at the tips of the lobes, adaxial surfaces glabrous. Filament tube absent; free portion of the filaments 1–1.5 mm long, occasionally slightly pubescent near the base; anthers 3.4–4 mm long, 1–1.5 mm wide, loosely connivent, poricidal at the tips, the pores becoming slit-like with age. Ovary glabrous or with a few dendritic trichomes at the apex, glabrate in fruit; style 5–7 mm long, sparsely to densely pubescent at the base or along its entire length with golden dendritic trichomes; stigma bilobed, the surface minutely papillose. Fruit a globose, purplish-black berry, with thin pericarp, 1–1.2 cm in diameter; fruiting pedicels 1–1.5 cm long, woody, nodding to more or less erect, ca. 1 mm in diameter at the base. Seeds ca. 8–10 per fruit, 3–4 mm × 2.5–3.5 mm, flattened lenticular, reddish-brown, the surfaces minutely pitted. Chromosome number: not known.

Figure 31.

Solanum cutervanum Zahlbr. (drawn from Hutchison & von Bismarck 6571). Reproduced from Knapp (1989) with permission of the Natural History Museum Botany Library. Illustration by Margaret Tebbs.

Distribution

(Figure 32). Andean Peru from Piura to Puno with a single collection known from Bolivia; 2500–3300 m.

Figure 32.

Distribution of Solanum cutervanum Zahlbr.

Ecology.

In rocky uplands, cloud forests and along trails in forest, usually growing in the open.

Common names:

Peru: rama de serrano (Knapp 1989).

Conservation status.

Least Concern (LC); EOO >100, 000 km2 (LC) and AOO >10, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum cutervanum had long been confused with and placed in the synonymy of S. nitidum. Specimens of both these species were previously annotated as Solanum pulverulentum Pers., perhaps due to their superficial similarity and sympatric distribution. The two species, however, are very different. Solanum cutervanum has golden or brownish tree-like trichomes on stems, leaves and inflorescences and black berries, while Solanum nitidum has more delicate, greyish, strictly dendritic trichomes and red ripe berries.

Solanum cutervanum is closely related to Solanum ruizii, also from central Peru. It differs from that species in its often dichasial branching, rounded leaf bases, smaller flowers and deltate calyx lobes. These sister taxa are sympatric and flower size may contribute to reproductive isolation. Some populations of Solanum cutervanum from N Peru have broadly elliptical leaves superficially reminiscent of Solanum aureum, a vining species with golden strictly dendritic pubescence.

Unfortunately the species name Solanum pulverulentum Pers., so long in use for this species, is a later homonym of Solanum pulverulentum L., itself a superfluous name for Solanum tomentosum L., an African prickly species (see Vorontsova and Knapp, in review). Linneaus only used the epithet pulverulentum in the 1759 edition of his Systemae Naturae, and in later publications reverted to the use of his original epithet tomentosum for the South African species.

Many monographers in Solanum have stated that holotypes or lectotypes for Ruiz and Pavón names were in the Madrid herbarium (MA), but without specifying a particular sheet. This is incorrect when several sheets are present in MA. Knapp (1989) lectotypified Solanum angustifolium citing only a sheet in MA; this was rectified in 2008 by citation of the particular sheet (Knapp 2008c).

In describing Solanum aureum vars. angustelanceolatum and latelanceolatum Bitter (1916) used sheets of Weberbauer 6700 at B for both names; he stated that within a branch leaf morphology was relatively uniform, but that between branches (probably sheets, as he never saw these or any other South American Solanum in the field) there was a great deal of variation. He compared Weberbauer 6700 to a sheet at B collected by Humboldt (an isotype of Solanum aureum) and clearly stated (Bitter 1916) that the Humboldt sheet represented the typical variety, and that various sheets of Weberbauer 6700 were his new taxa. I have selected the F duplicate as the lectotype of var. angustelanceolatum (Knapp 1989) and here selected the US sheet of the same number with slightly wider leaves as the lectotype of var. latelanceolatum as no duplicate material annotated by Bitter has been found. It is likely he was using material at Berlin (now destroyed) for his description.

Specimens examined.

Bolivia. La Paz: Franz Tamayo, Parque Nacional Madidi, Puina Viejo, ca. 3 km río abajo por camino al E del río, 3345 m, 20 Jun 2005, Fuentes 8507 (NY).

Ecuador. Azuay: Cantón Sevilla de Oro, Sevilla de Oro, eastern Cordillera, 4-6 km N of village, 2743 m, 14 Aug 1945, Camp E-4698 (MO); Cantón Cuenca, Cuenca, Cuenca, Parroquia Cumbe, 2682 m, 3 Jul 1991, Cerón 15542 (MO); Cantón Sevilla de Oro, Sevilla de Oro, 2950 m, 18 Apr 1968, Harling et al. 8454 (MO); Azuay/Morona Santiago: road Gualaceo-Limón, at the pass point, La Virgen, 3500 m, 26 Feb 1993, Harling & Ståhl 26713 (MO); Loja: Las Chinchas, region central, 2250 m, 12 Apr 1944, Acosta-Solís 7786 (F); Cordillera de Las Lagunitas. Amaluza-Jimbura-Zumba, Km 36, 3390 m, 22 Nov 1994, Jørgensen et al. 747 (BM, MO); Parque Nacional Podocarpus, above Cajanuma, trail from ‘Centro de Información’ toward ‘Lagunas de Compadre’, 3100 m, 19 Jan 1989, Madsen 85561 (BM).

Peru. Amazonas: Luya, Camporedondo, Tullanya, base Cerro Huicsocunga, 3075 m, 7 Dec 1996, Díaz & Peña 8853 (MO); Chachapoyas, Cerros de Calla-calla, uppermost slopes and summit, near kms 403-407 of Balsas-Leimebamba road, 3400 m, 18 Aug 1962, Wurdack 1704 (USM); Cajamarca: Jaén, Sallique, 3300 m, 26 Jun 1998, Campos et al. 5102 (MO); Jaén, Sallique. Quebrada Grande, camino entre La Cocha y Tablón, 2750 m, 2 Jul 1998, Campos et al. 5183 (BM, MO); Jaén, Lanchal, La Concha, Dist. Sallique, 2960 m, 16 Jun 1998, Díaz et al. 9597 (MO, USM); Jaén, Sallique, Quebrada grande, ruta entre La Cocha y Tablón, 2770 m, 30 Jul 1998, Díaz et al. 9779 (BM, MO, MOL); Jaén, Paramillo de Pomahuaca, antes del pajonal, 3200 m, 8 Nov 1999, Díaz & Campos 10908 (USM); El Pargo, 42 km E of Llama, ca. 14 km SE of Tunas Pampas, 3000 m, 8 Sep 1991, Gentry et al. 74570 (MO, USM); El Pargo, 16 km E of Tunas Pampa, ca. 42 km E of Llama on road to Huambos, 3000 m, 18 Sep 1991, Gentry et al. 74897 (MO, USM); Chota, Bosque El Pargo, entre Llama y Huambos, 3010 m, 12 Aug 1994, Leiva G. et al. 1485 (BM); Cusco: Calca, Choquecancha, Azulcocha, Dist. Lares, 3832 m, 18 Feb 2005, Valenzuela et al. 4978 (NY); Huánuco: Huamalíes, Monzón, cerros al sudoeste de Monzón, 3500 m, Weberbauer 3310 (MOL); Lambayeque: Ferrañafe, Cañariaco, Distrito Cañaris, sector Cañariaco, 3083 m, 16 Aug 2008, Marcelo Peña et al. 3688 (MOL); Pasco: Oxapampa, Distrito Huancabamba, Lanturachi, sector Santa Barbara, camino a Milpo, 2824 m, 10 Oct 2003, Perea et al. 702 (BM); Piura: Huancabamba, El Tambo, 3 Jun 1961, Acleto 294 (USM); Huancabamba, Carmen de la Frontera, alturas de Nueva York, 3280 m, 27 Jul 2006, Cano et al. 16766 (USM); Huancabamba, carretera entre Canchaque y Huancabamba, 2800 m, 14 Jan 1988, Díaz et al. 2719 (MO); Huancabamba, Mitopampa (Huancabamba-Cuello del Indio), 2650 m, 22 Jul 1975, Sagástegui et al. 8251 (MO); San Martín: Mariscal Caceres, Parque Nacional Rio Abiseo, cerro al sur de campamento Chochas, 3500 m, 30 Jun 1996, Cano et al. 7439 (USM); Distrito de Huicongo, valle de Ruibarbos, 3650 m, 12 Jun 2001, León & Ramírez 5200 (BM); Mariscal Cáceres, Parque Nacional Rio Abiseo, Puerta del Monte, 3200 m, Young 1573 (K); Mariscal Cáceres, Parque Nacional Rio Abiseo, Chochos, 3400 m, 14 Feb 1986, Young 2809 (K, MOL, USM); Mariscal Cáceres, Parque Nacional Rio Abiseo, Chochos, 3450 m, 13 Jun 1986, Young 3763 (K); Mariscal Cáceres, Parque Nacional Rio Abiseo, 3400 m, 3 Jul 1986, Young 3866 (K, MOL, USM); Mariscal Cáceres, Parque Nacional Rio Abiseo, along trail to El Mirador, Puerta del Monte, NW corner of Park, 3100 m, 11 Jul 1987, Young & León 4475 (USM).

13. Solanum dichroandrum Dunal, Prodr. [A.P. de Candolle] 13(1): 86. 1852

http://species-id.net/wiki/Solanum_dichroandrum

Figure 33
Solanum sanctaenevadae Dunal, Prodr. [A.P. de Candolle] 13(1): 678. 1852.
Type: Colombia. “Venezuela, Mérida, Sa. Nevada”, 8000, 1847, N. Funck & L.J. Schlim 1621 (holotype: G-DC [G00144896]; isotypes: BM [BM000849512], G [G00070229, F neg. 6742, IDC microfiche 800-61.2067:III.6], P [P00325803, Morton neg. 8183]).
Solanum dichroandrum Dunal var. glabrisculum Dunal, Prodr. [A.P. de Candolle] 13(1): 679. 1852.
Type: Colombia. “Venezuela, Prov. de Mérida, Sa. [Sierra] Nevada”, 8000 ft, Feb 1846, N. Funck & L.J. Schlim 1126 (holotype: G-DC [G00144978, IDC microfiche 800-61.2068:I.5]; isotypes: BM [BM000778190], G [G00070156], P [P00325804, Morton neg. 8181], P [P00325805], W).
Solanum endotrichum Bitter, Repert. Spec. Nov. Regni Veg. 12: 161. 1913.
Type: Colombia. Cundinamarca: Miquinquirá near Bogotá, Jul 1909, Bro. Idinaël 57 (holotype: MPU).
Solanum schlimii Bitter, Repert. Spec. Nov. Regni Veg. 16: 85. 1919.
Type: Colombia. Magdalena or Guajira: Prov. Río Hacha, Sierra Nevada de Santa Marta, 3100 m, L. Schlim [“Linden”] 831 (lectotype, designated here: G [G0070139, Morton neg. 8553]; isolectotypes: BM [BM000887368], BR, G [G00070140, F neg. 23165; G00070216], P [P00371281, Morton neg. 8322]).
Type.

Venezuela. “Caracas”, 1842, J. Linden 433 (holotype: G [G00070155, F neg. 8583]; isotypes: BM [BM000778189], G [G00104266], K [K000545358, K000545359], P [P000326802, Morton neg. 8182]).

Description.

Woody vine to 8 m long. Stems flexuous and appearing somewhat warty from prominent leaf scars, almost glabrous to densely pubescent with loose dendritic trichomes 1–1.5 mm long, these with few, long branches; new growth sparsely to densely pubescent with loose transparent dendritic trichomes to 1.5 mm long. Bark of older stems pale brown or pale reddish brown, glabrescent. Sympodial units plurifoliate. Leaves simple, 3–7(-11.5) cm long, 1–3.5 cm wide, elliptic to narrowly elliptic, membranous, the upper surfaces glabrous with a few scattered dendritic trichomes along the veins to sparsely pubescent with loose dendritic trichomes to 0.5 mm long, these occasionally mixed with a few simple uniseriate trichomes of the same size, the lower surfaces almost glabrous to densely pubescent with loose dendritic trichomes on the veins and lamina, usually also with sparse glandular papillae on the lamina; primary veins 8–9 pairs, usually reddish brown beneath; base attenuate to acute; margins entire, not revolute; apex acute to acuminate; petioles 0.5–2 cm long, pubescent like the stems, in herbarium specimens usually drying darker and more pubescent adaxially, occasionally twining. Inflorescences terminal, 4–8 cm long, globose to depressed-elliptic in outline, many times branched, with 10–50 flowers, glabrous to pubescent with loose, dendritic trichomes like those of the stems; peduncle 0.3–1.2 cm long, usually branching very near the base; pedicels 1–1.5 cm long, slender, ca. 0.3 mm in diameter at the base, ca. 1 mm in diameter at the apex, glabrous to loosely pubescent with dendritic trichomes, spreading at anthesis, articulated at the base, leaving a swollen area on the axis; pedicel scars irregularly spaced 1–10 mm apart, the inflorescence rachis bent at the articulation points. Buds ellipsoid, the corolla strongly exserted from the calyx tube in early bud. Flowers apparently all perfect (although the type has no long-styled flowers), 5-merous. Calyx tube 1–2.5 mm long, conical, the lobes 1–1.5 mm long, deltate with thickened margins and an elongate apex, sparsely to densely pubescent with loose dendritic trichomes. Corolla (1.2)1.5–2.5 cm in diameter, white or white tinged with lilac, stellate, lobed ca. 3/4 of the way to the base, the lobes 7–10 mm long, 4–6 mm wide, planar at anthesis, densely papillate on the tips and margins, the papillae sometimes extending to the lobes abaxially. Filament tube minute, the free portion of the filaments 1–1.5 mm long, glabrous or minutely pubescent with simple uniseriate trichomes less than 0.2 mm long; anthers ca. 4 mm long, 1 mm wide, glabrous, ellipsoid, loosely connivent, yellow, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 6–9 mm long, minutely puberulent with tiny simple uniseriate trichomes, these denser in the basal half; stigma capitate, the surface minutely papillose. Fruit a globose berry, ca. 2 cm in diameter, black when ripe, green when immature, dull and matte, glabrous, the pericarp thin; fruiting pedicels 1.5–2 cm long, ca. 1 mm in diameter at the base tapering to an apical diameter of 1.5–2 mm, woody and somewhat deflexed. Seeds 10–12 per berry, ca. 5 mm long, 4.5 mm wide, flattened-reniform, dark brown, the surfaces minutely pitted, the testal cells small and rectangular. Chromosome number: not known.

Figure 33.

Solanum dichroandrum Dunal. (A–F drawn from Riina et al. 784 G–I drawn from Killip & Smith 17916). Illustration by Bobbi Angell.

Distribution

(Figure 34). In northern South America in the coastal and western ranges of northern Colombia and Venezuela, extending to the Andes in Venezuela; 2200 to 3300 m.

Figure 34.

Distribution of Solanumdichroandrum Dunal.

Ecology.

Growing in cloud forests, probably in open areas.

Conservation status.

Least Concern (LC); EOO >100, 000 km2 (LC) and AOO >10, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum dichroandrum is very similar morphologically to a number of other Andean members of the Dulcamaroid clade such as Solanum sanchez-vegae, Solanum aureum and Solanum luculentum. It is most similar to Solanum sanchez-vegae, sharing with that species large flowers and loose dendritic pubescence. The two taxa differ in their seed number (with Solanum dichroandrum having twice as many of seeds), style pubescence (glabrous in Solanum sanchez-vegae, minutely puberulent in Solanum dichroandrum), and in flower size, with Solanum dichroandrum having somewhat smaller flowers. The two taxa are not sympatric, and neither is Solanum dichroandrum sympatric with Solanum aureum, from which it differs in having larger flowers, looser leaf and stem pubescence and fewer seeded berries. Solanum dichroandrum is sympatric with Solanum luculentum, from which it differs in leaf and stem pubescence (Solanum luculentum is completely glabrous), flower size and sex expression (Solanum dichroandrum has perfect flowers, whereas Solanum luculentum is almost certainly dioecious).

Specimens collected in the Colombian Department of Norte de Santander around Vetas (Killip & Smith 17267, 17313, 17388, 17910, 17916) are very pubescent and have slightly smaller flowers than the type, but otherwise fall within the range of variation seen in Solanum dichroandrum.

In describing Solanum schlimii, Bitter cited both BR and the herbarium of Barbey-Boissier (now part of the general collections at G) in the protologue; the lectotype selected (G0070139) is the cited sheet from the Barbey-Boissier herbarium and is annotated by Bitter.

Specimens examined.

Colombia. Antioquia: San José de San Andrés, 1 May 1948, Correa V.& Velásquez V. 35 (US); Boyacá: Pauna, carretera a Muxo, Las Curcubitas, km 114-117, 2850 m, 12 Nov 1948, García-Barriga 13236 (US); Cesar: Páramo de Sabana Rubia, 3250 m, 22 Jul 1987, Cuadros 3726 (MO); Cundinamarca: Bogotá, 1919, Brother Ariste-Joseph A-418 (US); La Guajira: Cerro del Espejo, N slopes, Serrania de Perijá, Venezuela border, 2560 m, 28 Apr 1987, Gentry & Cuadros 57183 (MO); Magdalena: Quebrada de Floridablanca, east of Manaure, Sierra de Perijá, 2700 m, 10 Nov 1959, Cuatrecasas & Romero-Castañeda 25199 (F); Río Garaban, headwaters of Río Aracataca, Sierra Nevada de Santa Marta, 2860 m, 23 Jul 1944, Kernan 159 (US); Mamancanaca, and vicinity, 3300 m, 27 May 1977, White & Alverson 611 (MO); Norte de Santander: Cerro de Oroque, limites entre los Departmentos Norte de Santander y Cesar, Cordillera Oriental, 3700 m, 22 Jul 1974, García-Barriga & Jaramillo 20721 (MA, US); Santander: Vetas, 3100 m, 16 Jan 1927, Killip & Smith 17267 (GH, US); Vetas, 3100 m, 16 Jan 1927, Killip & Smith 17388 (A, GH, US); Vetas, 3100 m, 16 Jan 1927, Killip & Smith 17916 (A, GH, US).

Venezuela. Lara: Parque Nacional Dinira, Páramo de Jabón, laderas nororientales, 3000 m, 28 Dec 1999, Riina et al. 867 (BM); Mérida: near Laguna de Coromoto, Sierra Nevada, 3200 m, 7 Aug 1958, Dennis 2185 (K); Pueblo Hondo, carretera a Mérida km 115-117, 2350 m, 24 Nov 1948, García-Barriga 13295 (US); Laguna de Coromoto, 3200 m, Jun 1958, Schwabe s.n. (B); Táchira: La Grita, Páramo el Rosal, 2800 m, 8 Oct 1965, Bernardi 10898 (K, LE); Zumbador, hacia Queniquea, 2500 m, 31 Jul 1984, Bono 4066 (MO); Páramo el Pantano, 2500 m, 16 Nov 1976, Charpin & Jacquemond 13451 (MO); Páramo de la Negra, slopes below páramo, above La Grita, 2430 m, 7 Jul 1944, Steyermark 57101 (F).

14. Solanum dulcamara L., Sp. Pl. 185. 1753

http://species-id.net/wiki/Solanum_dulcamara

Figure 35
Solanum scandens Neck., Delic. Gallo-Belg. 1: 119. 1768.
Type: Based on Solanum dulcamara L.
Dulcamara lignosa Gilib., Fl. Lit. Inch. 1: 37. 1782.
Type: Based on Solanum dulcamara L.
Lycopersicon dulcamara (L.) Medik., Beobacht. 245. 1783.
Type: Based on Solanum dulcamara L.
Solanum rupestre F.W.Schmidt, Fl. Boem. 2: 96, tab. 241. 1793.
Type: Czech Republic. “Termas Carolinas, Dorotheenau” [Karoly Vary] (no specimens found, plates unpublished; synonymy ex descr.).
Dulcamara flexuosa Moench, Meth. 514. 1794, nom. illeg. superfl.
Type: Based on Solanum dulcamara L.
Solanum ruderale Salisb., Prodr. Stirp. Chap. Allerton 133. 1796, nom. illeg. superfl.
Type: Based on Solanum dulcamara L.
Solanum dulcamara L. var. villosissimum Desv., Observ. Fl. Angers 111. 1818.
Type: France. Anjou: environs de Brissac, N.A. Desvaux s.n. (possible type specimen: P [P00582490]).
Solanum dulcamara L. var. rupestre (H.W.Schmidt) Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 581. 1819.
Type: Based on Solanum rupestre F.W.Schmidt
Solanum persicum Willd., Syst. Veg., ed. 15 bis [Roemer & Schultes] 4: 662. 1819.
Type: “In Persia, Pallas”(neotype, designated here: BM [BM000942806]).
Solanum littorale Raab, Flora 2: 414. 1819.
Type: Switzerland. Vaud: Lac Leman “inter Vidy et Pully; prope Ouchy et Champlande”, C.W. Raab s.n. (no specimens found; synonymy ex descr.).
Solanum kieseritzkii C.A.Mey., Verz. Pfl. Cauc. 113. 1831.
Type: Azerbaijan. Lenkoran, 23 May 1830, C.A. Meyer s.n. (holotype: LE; isotypes: LE (3 sheets), OXF).
Solanum assimile Friv., Flora 19: 439. 1836.
Type: Greece. “Turkey in Rumelia”, I. Frivaldszky s.n. (holotype: BP [n.v.]; isotype: NY [NY00169744]).
Solanum dulcamara L. var. tomentosum W.D.J.Koch, Syn. Fl. Germ. Helv. 2: 508. 1837.
Type: Based on Solanum littorale Raab
Solanum dulcamara L. var. album G.Don, Gen. Hist. 4: 409. 1838.
Type: “Dulcis amara flor albo”, Besler, Hortus Eystettensis, fol. 46, t. ii. 1613. (lectotype, designated here: Besler, Hortus Eystettensis, fol. 46, t. ii. 1613 [illustration on p. 180 in reprints]).
Solanum dulcamara L. var. carneum G.Don, Gen. Hist. 4: 409. 1838.
Type: Sweden. “Celsius, Ups. 32, Linnaeus, Fl. Suec. p. 66” (no specimens found).
Solanum dulcamara L. var. hirsutum G.Don, Gen. Hist. 4: 409. 1838.
Type: “Plant hairy, or downy. Flowers violaceous. On the sea-coast.” (no specimens cited).
Solanum dulcamara L. var. violaceum G.Don, Gen. Hist. 4: 409. 1838.
Type: “Dulcis amara flor coeruleo vulgatica”, Besler, Hortus Eystettensis, fol. 46, t. iii. (lectotype, designated here: Besler, Hortus Eystettensis, fol. 46, t. iii on folio 46 t. iii. 1613 [illustration on p. 180 in reprints]).
Solanum dulcamara L. var. cordifolium Peterm., Fl. Lips. Excurs. 173. 1838.
Type: Germany. Sin. loc. (no specimens found).
Solanum dulcamara L. var. ovatum Peterm., Fl. Lips. Excurs. 173. 1838. as ovata.
Type: Germany. Sin. loc. (no specimens found).
Solanum dulcamara L. var. marinum Bab., Man. Brit. Bot. 210. 1843.
Type: Ireland. County Galway: Connemara, Renvyle [“Renville”] (lectotype, designated here: CGE; isolectotype: BM [BM000941486]).
Dulcamara flexuosa Moench var. cordata Opiz, in Bercht. & Opiz, Oekon.-techn. Fl. Böhm. 3: xv. 1843.
Type: Based on Solanum dulcamara L. var. cordifolium Peterm.
Dulcamara flexuosa Moench var. ovata Opiz, in Bercht. & Opiz, Oekon.-techn. Fl. Böhm. 3: xv. 1843.
Type: Based on Solanum dulcamara L. var. ovatum Peterm.
Dulcamara flexuosa Moench var. hastiifolia Opiz, in Bercht. & Opiz, Oekon.-techn. Fl. Böhm. 3: xvi. 1843. as hastaefolia.
Type: Czech Republic [?]. sin. loc., P.M. Opiz s.n. (holotype: PR?).
Solanum lignosum Sloboda, Rostlinnictví 358. 1852.
Type: “Dulcamara Marina, Solanum lignosum Rayi” (no specimens found).
Solanum dulcamara L. subsp. cordatum (Opiz) Dunal, Prodr. [A.P. de Candolle] 13(1): 78. 1852.
Type: Based on Solanum dulcamara L. var. cordifolium Peterm.
Solanum dulcamara subsp. ovatum (Peterm.) Dunal, Prodr. [A.P. de Candolle] 13(1): 78. 1852.
Type: Based on Solanum dulcamara L. var. ovatum Peterm.
Solanum dulcamara L. var. rupestre (F.W.Schmidt) Dunal, Prodr. [A.P. de Candolle] 13(1): 79. 1852.
Type: Based on Solanum rupestre F.W.Schmidt
Solanum dulcamara L. var. palustre Dunal, Prodr. [A.P. de Candolle] 13(1): 79. 1852.
Type: Greece. “Lernacicis”, J. Sartori s.n. (no specimens found, see discussion).
Solanum dulcamara L. var. hirsutum Dunal, Prodr. [A.P. de Candolle] 13(1): 79. 1852.
Type: Greece. “Lernacicis”, J. Sartori 84 (lectotype, designated here: G-DC [G00144899]).
Solanum dulcamara L. var. laciniatum Dunal, Prodr. [A.P. de Candolle] 13(1): 78. 1852.
Type: United States of America.Massachusetts: Boston, 1827, Mrs. Dutton s.n. (lectotype, designated here: G-DC [G00144856]).
Solanum dulcamara L. subsp. hastiifolia (Bercht. & Opiz) Dunal, Prodr. [A.P. de Candolle] 13(1): 78. 1852.
Type: Based on Dulcamara flexuosa Moench var. hastiifolia Opiz in Bercht. & Opiz
Solanum dulcamara L. var. alpinum Schur, Enum. Pl. Transsilv. 478. 1866.
Type: Romania. (Transylvania) Siebenbürgen [Sibiu], ”Arpaser-Kerzesor Alpen” [Arpas], Jul-Aug, J.F. Schur s.n. (holotype: GOET?, not found).
Solanum dulcamara L. var. macrocarpum Maxim., Index Seminum [St. Petersburg] suppl. 1869: 26. 1870.
Type: japan. Jezo Island: Hakodate, 1861, C. Maximowicz s.n. (lectotype, designated here: LE; isolectotypes: G [G00301664], LE).
Solanum dulcamara L. var. integrifolia Willk. in Willk. & Lange, Prodr. Fl. Hispan. 2: 526. 1870.
Type: Spain. Aragón: Jaca, Jun 1850, H.M. Willkomm 313 (lectotype, designated here: BM [BM000847923]; isolectotype: G [G00357859]).
Solanum dulcamara L. var. indivisum Boiss., Fl. Orient. 4: 285. 1879.
Type: Based on Solanum persicum Willd.
Solanum dulcamarum St.-Lag., Ann. Soc. Bot. Lyon, 7: 135. 1880, nom. illeg. superfl.
Type: Based on Solanum dulcamara L.
Solanum dulcamara L. forma subglabrum Kuntze, Trudy Imp. S.-Peterburgsk. Bot. Sada 10: 222. 1887.
Type: Turkmenistan. Asgabat [As’chabad], May 1886, O. Kuntze [Gen. Komarov] s.n. (holotype: B, destroyed; lectotype, designated here: NY [NY00172263]).
Solanum rupestre Waisb., Oesterr. Bot. Z. 45: 143. 1895, non Solanum rupestre F.W.Schmidt, 1793.
Type: Austria. “ad pagum Rödlschlag solo serpentino”, 700-750m, Jun-Jul 1894, A. Waisbecker s.n. (no specimens found, holotype: BP?).
Solanum dulcamara subvar. villosissimum (Desv.) Bornm., Beih. Bot. Centralbl. 33: 305. 1915.
Type: Based on Solanum dulcamara L. var. villosissimum Desv.
Solanum macrocarpum (Maxim.) Kudô, in Kudô & Susaki, Hokkaido-Yakuyo-Shokub t. 84. 1922.
Type: Based on Solanum dulcamara L. var. macrocarpum Maxim.
Solanum dulcamara L. var. canescens Farw., Papers Mich. Acad. Sci. 2: 39. 1923.
Type: United States of America. Michigan: Rochester, 15 Aug 1909, O.A. Farwell 2105 (holotype: BLH [n.v.]).
Solanum dulcamara L. forma albiflora Farw., Papers Mich. Acad. Sci. 2: 39. 1923.
Type: United States of America. Michigan: Trenton, 26 Jul 1921, O.A. Farwell 5944 (holotype: BLH [n.v.]).
Solanum dulcamara L. forma albiflorum House, Bull. New York State Mus. 254: 613. 1924.
Type: United States of America. New York: Livingston County, Caledonia, F. Beckwith s.n. (holotype: NYS [n.v.]).
Solanum maximowiczii Koidz., in Mayebara, Fl. Austrohigo 50, 85. 1931.
Type: Based on Solanum dulcamara L. var. ovatum Peterm.
Solanum macrocarpum (Maxim.) Koidz., Acta Phytotax. Geobot. 1: 23. 1932, nom. illeg., non Solanum macrocarpum (Maxim.) Kudô
Type: Based on Solanum dulcamara L. var. macrocarpum Maxim.
Solanum megacarpum Koidz., Acta Phytotax. Geobot. 4: 159. 1935.
Type: Based on Solanum dulcamara L. var. macrocarpum Maxim.
Solanum depilatum Kitag., Lin. Fl. Manshur. 390. 1939.
Type: China. [Heilongjiang]: “Chéjché, Manchur. bor.”, M. Kitagawa s.n. (type not located).
Solanum asiaemediae Pojark., Not. Syst. Herb. Inst. Bot. Acad. Sci. URSS 17: 330. 1955.
Type: Uzbekistan. Kokaidskii region, Tamariskovii Krai, [lower Najman-Saj River], 21 Jul 1934, A.V. Prozorovskii 187 (holotype: LE).
Solanum pseudopersicum Pojark., Not. Syst. Herb. Inst. Bot. Acad. Sci. URSS 17: 328. 1955.
Type: Russian Federation. Starvropolskii Krai: Pyatigorsk, Mashukh [protologue Ciscaucasia], 3 Jul 1896, D. Litvinov s.n. (holotype: LE).
Solanum marinum (Bab.) Pojark., in Komarov, Fl. URSS, 22: 16. 1955.
Type: Based on Solanum dulcamara L. var. marinum Bab.
Solanum persicum Roem. & Schult. subsp. pseudopersicum (Pojark.) Schönb.-Tem., Fl. Iranica 100: 14: 1972.
Type: Based on Solanum pseudopersicum Pojark.
Solanum kitagawae Schönb.-Tem., Fl. Iranica 100: 15. 1972.
Type: Based on Solanum depilatum Kitag.
Solanum dulcamara L. luc. atroviolaceum Máthé, Bot. Közlem. 59(2): 129. 1972.
Type: Hungary. Nógród: “ad pagum Nótines”, I. Máthé s.n. (holotype: VBI [n.v.]).
Solanum dulcamara L. forma lucidum Máthé, Bot. Közlem. 59(2): 129. 1972.
Type: Hungary. Pest: “ad viam pagi Vácrátot”, I. Máthé s.n. (holotype: VBI [n.v.]).
Solanum dulcamara L. var. pusztarum Soó, Acta Bot. Acad. Sci. Hung. 18(1-2): 172. 1973.
Type: Hungary. Mittelungarn, Kom. Bács-Kiskun, Bugacpuszta, R. Sóo s.n. (holotype: BP [n.v.]).
Solanum dulcamara subsp. pusztarum (Soó) Soó, Acta Bot. Acad. Sci. Hung. 23(3-4) : 382. 1978 [“1977”].
Type: Based on Solanum dulcamara L. var. pusztarum Soó
Solanum borealisinense C.Y.Wu & S.C.Huang, Fl. Reipubl. Popul. Sin. 67(1): 84, tab. 21, Figure 1–7. 1978.
Type: Based on Solanum depilatum Kitag.
Type.

“Europa”, Anon. (lectotype, designated by Knapp and Jarvis 1990, pg. 335: LINN 248.7).

Description.

Herbaceous or woody vine, above ground stems trailing to 8–10 m long and with spreading or creeping underground stems. Stems very rarely glabrous, more often pubescent with simple uniseriate or dendritic trichomes with short branches, or a mixture of the two types, these often tangled, 4–8-celled, to 1.5 mm long, usually white, pubescence density extremely variable; new growth usually white pubescent. Bark of older stems grey to yellowish grey. Sympodial units plurifoliate. Leaves simple to ternately pinnatifid, extremely variable in shape and size, even along a single stem, 2.5–7 cm long, 1.2–6 cm wide, elliptic or ovate to cordate in outline, membraneous, the upper surfaces glabrous to moderately pubescent with simple uniseriate or dendritic trichomes to 1.5 mm long on the veins and lamina, the lower surfaces sparsely to densely pubescent with trichomes like those of the upper surfaces, but usually denser; primary veins 6–9 pairs, usually pubescent; base truncate or cordate; margins entire or the leaves lobed, the lobes most commonly 2, rarely more, basal, narrowing near the sinuses; apex acute to acuminate; petiole 0.5–2 (+) cm long, pubescent like the stems. Inflorescences terminal or lateral, not leaf-opposed, (1-)4–15 cm long, many times branched, with up to 40 flowers, only a few open at a time, glabrous to moderately pubescent, the rachis often purplish in hue; peduncle (0.5-)1–7 cm long; pedicels 6–12 mm long at anthesis, ca. 1 mm in diameter, slender, spreading, often purplish green, glabrous to sparsely pubescent with simple unieriate or more rarely dendritic trichomes to 0.5 mm long, articulate at the base in a small sleeve leaving a prominent swollen peg on the axis; pedicel scars irregularly spaced 1–5 (-10) mm apart, the axis zig-zag. Buds turbinate, the corolla long-exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 1–1.5 mm long, broadly conical, the lobes < 0.5 mm long, broadly triangular, glabrous or pubescent with uniseriate white trichomes, the apex pointed, the margins papillate. Corolla 1.5–2 cm in diameter, purple, violet or white, with green and white shiny spots at each lobe base, deeply stellate, lobed 3/4 of the way to the base, the lobes 6–8 mm long, 2.5–3 mm wide, strongly reflexed at anthesis, glabrous or minutely papillate on tips and margins, occasionally densely pubescent with simple uniseriate trichomes abaxially, glabrous adaxially. Filament tube minute, the free portion of the filaments to 0.5 mm long, glabrous; anthers 4.5–6 mm long, ca. 1 mm wide, fused into a single column and tightly connivent, poricidal at the tips, the pores not lengthening to slits with age. Ovary glabrous; style 5–9 mm long, glabrous; stigma minutely capitate, the surface papillose. Fruit a globose to ellipsoid berry, 0.6–1.1 cm long, 0.6–1.5 cm wide, bright red when ripe, the pericarp thin and shiny; fruiting pedicels to 1.3 cm long, 1–1.5 mm in diameter, not markedly woody, spreading. Seeds >30 per berry, ca. 3 mm long, ca. 2 mm wide, flattened reniform, pale yellow or tan, the surfaces minutely pitted, the cells of the testa pentagonal. Chromosome number: n=12 (many counts available at ICPN, http://mobot.mobot.org/W3T/Search/ipcn.html)

Figure 35.

Solanum dulcamara L. (All drawn from live plants in Battleboro, Vermont, USA). Illustration by Bobbi Angell.

Distribution

(Figure 36). Solanum dulcamara is widely distributed across Eurasia and northern North America, where it is also common; sea level to ca. 2000 m. The North American populations are thought to be introductions, but it is possible that the species has a truly circumboreal distribution.

Figure 36.

Distribution of Solanum dulcamara L.

Ecology.

Solanum dulcamara is a weedy species and grows in a wide variety of temperate habitats, often associated with water and open places with abundant light. Although somewhat woody, it rarely reaches into the canopy but is more often found in thickets and sprawling in other low vegetation.

Common names.

Douce-amére (French); woody nightshade, bittersweet (English speaking Europe)

Conservation status.

Least Concern (LC); EOO >100, 000 km2 (LC) and AOO >10, 000 km2 (LC). See Moat (2007) for explanation of measurements. Solanum dulcamara has a circumboreal distribution and is very common so it is not of conservation concern. Peripheral populations such as those in eastern Russia, however, may harbour interesting genetic variation (see Knapp 2011).

Discussion.

Solanum dulcamara is one of the most variable species in the Dulcamaroid group, all of which show extremes of phenotypic plasticity. It is also the most widespread, and is found throughout the northern hemisphere, in Europe and Asia from Spain to Siberia and into extreme northern Japan, and across the northern United States (reported from Florida on the PLANTS database [http://plants.usda.gov/java/profile?symbol=SODU], from where I have seen no specimens), where it is presumed to have been introduced from Europe. I can find no documentary evidence for the introduction of Solanum dulcamara into the New World, but early collections from North America held in BM (e.g., Bartram, Clayton) do not contain specimens of this species. The earliest collections I have seen are from the Boston area in the early 19th century (e.g., Dutton s.n.); it is possible that Solanum dulcamara was introduced to North America for its medicinal properties.

Some of the variation in Solanum dulcamara appears to be related to habitat (e.g., coastal forms with succulent leaves that have been recognised as var. marinum and Solanum littorale) but much of the variation is in leaf size and division, pubescence and flower color; I have been able to find no pattern related to either geography or habitat in this variation. Turesson (1922) suggested that differences in morphology related to habitat were genetic, despite his finding that in non-coastal environments thick-leaved coastal forms developed thinner, less pubescent leaves. Sun and shade leaves of Solanum dulcamara have been studied for physiological and some morphological properties (Clough et al. 1979a, b). Differences in morphology can occur along a single stem depending upon the shade environment. Unlike some other species in the Dulcmaroid clade (e.g., Solanum umbelliferum) variation does easily sort into distinguishable units, but the sheer range and complexity of the variation has led to the description of many taxa within what I recognise here as a single highly polymorphic species, especially in floristic treatments from Central Asia (Pojarkova 1955; Schönbeck-Temesy 1972), where much of the variation exists.

Chemical variation in Solanum dulcamara has also been intensively studied in eastern Europe (Máthé and Máthé 1972, 1979); populations with pubescent individuals from the northen part of the eastern European range tended to have higher percentages of the alkaloid aglycone tomatidinol while more glabrous populations had higher concentrations of soladulcinine. The concentrations of various alkaloids were affected by the environmental conditions under which plants were grown (Máthé et al. 1975) as well as by what were presumed to be genetic (ecotypic) factors (Bernáth and Tétényi 1977). Clough et al. (1979a) and Pegtel (1985) both suggested that ecotypic differentiation did not explain the variation observed, but instead that the variability was the result of short-term phenotypic acclimatization to differing environments. A study using AFLP (Associated Fragment Length Polyomorphisms) data from Europe-wide accessions of Solanum dulcamara found almost no genetic variation or population structuring either geographically or in association with habitat (Golas et al. 2010).

A few of the more extreme forms have recognised at the species level, but the variation in the diagnostic features of these forms is continuous across the range of the species. Identifications of individual specimens by botanists describing these variants were occasionally inconsistent from duplicate to duplicate.

Plants occurring in the swampy areas around the hot springs at Lenkoran in what is now Azerbaijan were described as Solanum kieseritzkii; all of the collections from this locality are composed of small erect shoots connected with creeping stems and have very small inflorescences of only a few flowers.

Glabrous plants with simple leaves have been recognised as Solanum pseudopersicum; glabrous plants with divided leaves as S. kitagawae.

Pubescent plants with simple leaves were described as Solanum persicum; these tended to be from the southern part of the species range in Eurasia, but occur throughout the species’ range.

Large-fruited plants from the eastern margins of the Asian range have been called var. macrocarpum; these fruits also tend to be more ellipsoid than globose, but as with other chracters, this occurs sporadically throughout the range of the species.

White-flowered plants have been given varietal or subspecific status both in Europe and the United States; polymorphism in flower color is common in Solanum in general.

Solanum dulcamara has a long history of use in medicine in Europe and in the United States. Gerard (1597:350) says “The leaves and fruit of the Bitter-sweet are in temperature hot and dry, clensing and wasting away” and cites as its “Vertues” effects on liver and spleen, against jaundice and bruising, and of its use to help breathing difficulties and as a restorative after childbirth. The common name bittersweet (or douce-amère) comes from the Latin ‘Amaradulcis’, and refers to the fact that when chewed, the twigs taste at first bitter, then sweet (Grieve 1931). Bittersweet had many uses ranging from inflammatory diseases of all kinds (such as asthma and rheumatism) to nymphomania and syphilis (Dunal 1813; Felter and Lloyd 1898); Grieve (1931: 590) states “there are few complaints for which it has not at some time been recommended.” As ‘Dulcamara’ or ‘Dulc’ (van Zandvoort 1996), the stems are used still in homeopathic medicine (Bharatan et al. 2002). It is today listed as a poisonous plant in many countries and US states (see for example the FDA at http://www.accessdata.fda.gov ). Michel-Félix Dunal treated the uses and superstitions associated with this species (Dunal 1813) and concluded that the many properties attributed to bittersweet were mostly not substantiated in fact. The steroidal alkaloids responsible for the active properties of Solanum dulcamara were not discovered until the late 19th century, they are principally dulcamarine and solanine, the first of which is responsible for the bittersweet taste and the second for the narcotic effects (Grieve 1931).

Solanum dulcamara has been thought to be an intermediate host and possible source of primary infections for several agronomically important potato diseases such as late blight (caused by the oomycete Phytophthora infestans (Mont.) de Bary) and brown rot/bacterial wilt (caused by the bacterium Ralstonia solanacearum (Smith) Smith). Eradication of Solanum dulcamara has been attempted as a control of brown rot (Persson 1998), and use of water from ditches in which Solanum dulcamara grows is a known cause of brown rot spread in irrigated potato fields. The role of Solanum dulcamara in the spread of late blight, however, has been shown to be minimal (Golas et al. 2010b), as most plants are resistant, and those that are susceptible to the disease do not harbour it over winter. Even in very favourable conditions only sporadic infections occur (Cooke et al. 2002; Flier et al. 2003; Dandurand et al. 2006). Solanum dulcamara has novel genes for resistance to late blight, and there is scope for its use in new potato breeding programs in Europe (Golas et al. 2010b). Little work has been done to investigate the mechanisms or genetics of resistance to the major diseases of potato in species outside the Potato clade, but new breeding methods have stimulated the investigation of disease in wild species such as Solanum dulcamara (Golas et al. 2010b) and Solanum nigrum L., a European hexaploid member of the Morelloid clade (Lebecka 2008, 2009).

Typification of the many synonyms of Solanum dulcamara has been very difficult. Many of the early names are found in early 19th century floristic accounts, and while validly and effectively published, usually do not cite specimens. I suspect many of these were not necessarily based on specimens, but rather on field observations. I have not neotypified any of the synonyms for which I could not find specimens directly linked to the original description. In addition, many of the European floras coined complex series of replacement names in the early parts of the 19th century, before the codes of nomenclature became firmly established. Where I have been able to trace the identity and derivation of epithets at varying ranks through citation of specimens I recognised these names as homotypic, but for many of the early names (e.g., those coined by George Don in 1838) the links are very tenuous and I have thus recognised them as heterotypic (and often have not designated types).

The type of Solanum persicum was a Pallas specimen held in Berlin, now destroyed. A Pallas specimen in BM that matches the description perfectly has been selected as a neotype, although it may not be a duplicate of the original.

The many varietal names coined by George Don (1838) were based not on specimens, but on literature references (e.g., var. album) or personal observations (var. hirsutum). His var. hirsutum referred to plants that were “Hairy or downy. .. On the sea coast”, with no specimens or plates cited. I have not neotypified this name. Varieties album, violaceum and plenum were based on one plate (plate 46) in Hortus Eystettensis (Besler 1613) which is composed of three plants (see Figure 37): one white-flowered Solanum dulcamara that I have designated as the lectotype of var. album, one violet flowered Solanum dulcamara that I have designated the lectotype of var. violaceum, and a central plant, used by Don as the basis for var. plenum, that is a drawing of Nigella sativa L. (Ranunculaceae) (see Figure 37). He cannot have been looking very carefully!

Figure 37.

“Dulcis amara” from Hortus Eystettensis (Besler 1613). The centre plant is clearly Nigella (Ranunculaceae), but was described as a variety of Solanum dulcamara by David Don. Reproduced with permission of the Natural History Museum Botany Library.

Petermann (1838) coined several new infraspecific taxa in his local flora of the Leipzig region; I have found no material related to these names. Solanum maximowiczii was based on “Solanum dulcamara var. ovatum Maxim.” and was coined at the specific level to represent that variety. In his discussion of Solanum dulcamara, however, Maximowicz (Regel and Maximowicz 1870) refered explicity to Dunal’s variety ovatum and did not coin a new name. Thus, this specific epithet is homotypic with the chain of epithets linked to the variety ovatum treated by Maximowicz, ultimately traceable to the (presumably) first instance of this varietal name being used in Petermann’s floristic treatment (Petermann 1838). Dunal (1852), cited by Maximowicz, cites Opiz (Berchtold and Opiz 1843) and Opiz cites Petermann as the origin of the varietal name. I have found no specimens associated with these names.

Dunal’s (1852) treatment of Solanum dulcamara for the Prodromus has a broad range of variation recognised as a single taxon; he recognised 14 infraspecific entities at two levels, mostly based on literature citations, his final words in the description - “Multum variat.” - couldn’t be more true. He recognised as A, B and C forms previously recognised by Opiz (Berchtold and Opiz 1843) and Petermann (1838)cordatum, ovatum and hastiifolia (as hastaefolia). Below them on the page he recognised a series of 11 taxa (“Hae tres formae vicissum variant”) that he listed with Greek letters (α to ξ, leaving out ι, κ and λ), some of which had binomial and others polynomial epithets. Infraspecific taxa denominated with Greek letters in Dunal (1852) have always been recognised at the varietal rank, and no where else in the treatment of Solanum in the Prodromus does Dunal use an apparently nested set of ranks as he does with Solanum dulcamara. I am therefore assuming that he intended his A, B and C ranks to be hierarchically above the varietal epithets and have treated them as subspecies in the synonymy. It is possible that var. hastiifolia of Opiz is in fact an illegitimate name for var. flexuosa, as it seems from the text that Opiz (Berchtold and Opiz 1843) treated this variety as the typical one, but I cannot be certain and so have recognised it as legitimate. The polynomial varietal epithets (e.g., “corollis violaceis”, “corollis albis”, “corollis carneis”, “flore pleno”, corollis variegatis”) are not legitimate names; I have lectotypified the rest based on either herbarium material or literature citations cited by Dunal. For var. laciniatum, Dunal cited specimens in both P and G-DC; I have found only the G-DC sheet, so have selected it as the lectotype.

The holotype of Solanum dulcamara forma subglabrum was destroyed in Berlin and a duplicate of Kuntze’s collection held in NY has been selected as the lectotype.

In describing Solanum dulcamara var. macrocarpum, Maximowicz (Regel and Maximowicz 1870) cites as a locality “Circa Hakodate insulae Jezo pluribus locis”. Several collections of Maximowicz’s from Hakodate dated 1861 are held at LE; others are in BM, GH and NY. It is very unclear whether any of these are true duplicates, so I have selected one of the flowering and fruiting sheets at LE as the lectotype and another at LE that is almost identical in morphology as a possible isolectotype.

Wasibecker’s (1895) Solanum rupestre is a later homonym of Solanum rupestre H.W.Schmidt. It is possible that the name Solanum serpentini Borbas & Waisb., apparently coined in 1897 (Soó 1968: 153) is a replacement name for the homonym, as the epithet indicates an origin on serpentine soils (like the type of Solanum rupestre Waisb.). I have been unable to locate the place of publication of Solanum serpentini, and so treat Solanum rupestre Waisb. as a synonym of Solanum dulcamara and Solanum serpentini as a doubtful name (see Doubtful and excluded names and Names not validly published).

Máthé and Máthé (1972) undertook an exhaustive study of the variation of Solanum dulcamara in Hungary and described two new variants, one at the form level (forma lucidum) and the other (luc. atroviolaceum) at the rank ‘lucus’ that was used in by Hungarian botanists between variety and forma. He followed Soó (1968) who transfered many earlier epithets to the rank of lucus, however, none of these are validly published, as no explicit reference to the basionym’s place of publication was made as is required by the Code.

I here cite specimens I have examined; these do not cover all European countries, although Solanum dulcamara is recorded from throughout Europe. I have selectively cited material from the range edges, as that is where I anticipated taxonomic problems might lie. Solanum dulcamara is very commonly collected and the specimens cited here do not completely represent the range.

Specimens examined.

Afghanistan. Kurrum valley, at Zabardastkalla and Alikhél, Dec 1879, Aitchison 861 (BM, GH, K, LE, S); Sere Tange, south of Marshad valley, Keshan, 1250 m, 30 Jun 1969, Carter 375 (K); Parwan, 33 miles from main Salang road, by side of Gharband River, 1920 m, 13 Jun 1969, Hewer 1274 (K, LE); Gulbahar, 14 May 1937, Koelz 11428 (LE, US).

Albania. Shkoder, North Albania, environs of the city of Shkoder near the river Umri, 14 Jul 1959, Kuvaev 95 4 (LE).

Algeria. Clemcen, vers les Cascades, 800 m, 24 Aug 1932, Faure s.n. (S); Elenicens?, Aug 1849, Romain s.n. (BM).

Andorra. Santa Julia de Loria, de borda del Sabater a Tolse, 840 m, 27 Aug 2002, Aedo et al. 8428 (MA).

Armenia. Kafanskii Region, near Ang, 24 Aug 1980, Avetisyan s.n. (MO); Karassan, 1837, Koch s.n. (LE); Arpa, between Iahichevan and Yerevan River Arpa to the right of the bridge, 21 Jun 1974, Mamakli s.n. (LE); Kirovakan station, 26 May 1953, Tamamschian, S., s.n. (LE); Megra, Megrinskii Region, 21 Jul 1954, Tamamshek & Deiisova s.n. (LE); Kirovakai, 26 Aug 1953, Tamaschira s.n. (LE); Alaverdskii reg., S. Sanain, Uzunar, 25 Jul 1960, Avegisyan & Gabrielian s.n. (LE).

Austria. Tyrol: Tirolia centralis, in valle Gschnitz prope pagum Trins, Kerner 3290 (BM, LE); Vienna: Wien, erster Bezirk, an der Grundmauer des Naturhistorischen Museums auf der Seite der Ballariastrasse, 200 m, 2 Oct 1992, Wallnöfer 4212 (BM).

Azerbaijan. Distr. Schemacha, Fagraüksch, Prope st. Scharodilskaja, ad lacum Fagraüksch, 792 m, 8 Aug 1900, Alexeenko 11304 (LE); Bakinskaya Oblast’, Kusary, Kubinskii region, 20 Jul 1900, Grigoreev 20 (LE); Lenkoran, 1834, Hohenacker s.n. (G-DC, K, LE, MO); Distr. Kuba, Seput, 24 Jul 1930, Kasumova s.n. (LE); Eshakchi, 3 Jul 1909, Kirichenko s.n. (LE); Shirvan steppe, inter Kerzi-Kent et Kotoran, 24 Aug 1930, Kolakovskii s.n. (LE); Gandzha Oblast’, District Kazakh, near the road to Poyly, 13 Jun 1928, Kolakovskii s.n. (LE); Lenkoranskiy Okrug, [near village of Nipolaevna, on river Vilyazh-chai], 30 Jul 1931, Matveeva 791 (LE); Gandzha Oblast’, Bardy, Agdam District, on the Karabkh steppe, near town, by the River Terter, 17 Aug 1927, Prilipko s.n. (LE); distr. Saljany, Saljany, prope st. Dzhafarchan, 6 Jul 1927, Prilipko s.n. (LE); distr. Lenkoran, Tangovan, near village Tangovan, 26 Jun 1931, Shipchinskii 275 (LE); Talysh, Lenkoranskii, 16th km on the road from Lenkoran to Leric, left bank of the river Lenkoran-chai, 100 m, 1 Jul 1952, Vasiliev & Vasilieva s.n. (LE); 18 km from Sheka, 26 Jun 1972, Vlasov s.n. (LE).

Belgium. Brussels: garden on Chaussee Pas in Auderghem, near a forest east of Brussels area, Jul 1987, Geerinck-Coutrez 4343 (BM); West Flanders: Knokke, Le Zwin, 21 Jun 1971, Auquier et al. s.n. (LE); Knokke, Zrvin, 8 Aug 1955, Lonvalrée 6781 (BM).

Bosnia-Herzegovina. Mostar, 1886, Bornmüller 1405 (B).

Bulgaria. Varna, Aug 1886, Bornmüller s.n. (B); 8-9 km W of Velingrad, along the stream Lukovica, 980 m, 8 Jul 1982, Frost-Olsen 4401 (MA); Vitosa, 10 Jun 1927, Georgieff s.n. (LE); Malkija Kozuh, Struma valley by the hill Malkija Kozuh, 9 Jun 1980, Kuzmanov 801405 (B); Samokor, 30 Aug 1912, Post 277 (G); Sliven, 16 Jul 1907, Schneider 710 (BM); Prilap, May 1917, Stribrny s.n. (LE).

Canada. British Columbia: Vancouver, Vancouver 8: Alma. Roadside, 18 Jul 1954, Bird 43 (BM);, S. Okanagon, White Lake, Oliver road, 480 m, 6 Jul 1961, Vrugtman & Campbell 610348 (BM); New Brunswick: Kent County, Newcastle, rocky shore of Miramichi River, 17 Jul 1945, Dore & Gorham 45555 (G); St. John County, Fairville, near St. John River, 19 Sep 1926, Fernald et al. 394 (GH); Bass River, Kent County, Fowler s.n. (LE); Newfoundland: Salmonier, 1931, Ayre s.n. (GH); Nova Scotia: Annapolis County, Granville, North Mtn, 18 Jul 1921, Bartram & Long 24435 (GH, LE); Yarmouth County, Markland, Cape Forchu, 13 Jul 1921, Fernald et al. 24434 (K, LE); Shelburne County, East Jordan, 4 Aug 1921, Fernald & Long 24436 (GH); Halifax County, Halifax, 30 Sep 1924, Jack 3644 (GH); Guysborough County, Guysborough, 14 Aug 1930, Rousseau 35472 (K); Ontario: Plevna, 26 Jul 1902, Fowler s.n. (GH); Hastings County, Marmora, Concession 3-4, Twin Sister Lakes, 8.5 miles NNE of Marmora, 14 Aug 1952, Gillett & Hammond . 6922 (G); Bruce County, Tobermory, 2 Jul 1933, Krotkov 7758 (GH); Waterloo County, Westmount Woods, 1 mi. W of Kitchener, 16 Jun 1939, Montgomery 338 (GH); Brittania, 18 Jun 1921, Rolland 15680 (GH); Halton County, Great Lakes Region, Lake Medad, 25 Jun 1940, Soper & Burcher 2011 (BM, GH); Prince Edward Island: Queens County, Charlottetown, 28 Aug 1912, Fernald et al. 7789 (GH); Quebec: Iberville County, Ile aux Noix, 12 Aug 1928, Adrien 2098 (K, LE); Missisquoi, Noyan, 23 Jun 1959, Gervais & Lavigne s.n. (G); Pontiac County, Beech Grove, 19 Aug 1943, Lamarre s.n. (MA).

Croatia. Dalmatia, Spalolo, Mossor, 1886, Bornmüller s.n. (B); Golubevec coal district, 1924, Leathes s.n. (BM); island of Veglia (Crk-Venica), Sep 1924, Leathes s.n. (BM).

Denmark. Copenhagen, Lyngby Mose, 14 Jul 1941, Dahl D-45 a (BM); Odder, Sondrup Plantage, ca. 11km S of Odder, W part of the plantation, 25 Jul 1978, Frost-Olsen 1748 (LE); Lake Tissø, D42, Jul 1968, Hjorih-Olsen 10 (BM); Jutland, Silkeborg, Borre Lake, 1 Aug 1968, Jensen et al. 464 (BM, MA); North Jutland, Stenbjerg, 9 Jul 1964, Larsen & Laegaard s.n. (LE); Skagen, 9 Aug 1973, Larsen et al. S.L. 6847 (BM, MA).

Egypt. Sin. loc, Anon., s.n. (LE).

Estonia. At village Oiu, 12 Sep 1967, Aasamaa 11616 (MO); Dorpet [Dorpat = Tartu], 12 Jul 1860, Gruner s.n. (BM);Vijlandi County, Tipu, River Halliste, 14 Jul 1994, Reier s.n. (B).

Finland. Pyhäranta, Rihtniemi point and village, 21 Aug 1968, Alava et al. s.n. (LE); Nauvo, Seili, near church, 4 Aug 1976, Arkkola et al. s.n. (MA); Åland, Nafsby, Prov. Hammarland, 28 Jul 1963, Haakana s.n. (LE); Kupittaa, Turku, Varsinais-Suomi, 14 Aug 1924, Kari s.n. (LE); South Häme, Heinola, Tommola, Maitiaislahti, 7 Aug 1980, Kemppainen s.n. (MA); Kärkölä, Järvelä, middle part of the NE shore of Lake Hähkäjärvi, just NW of the man-made cape, 97 m, 29 Jun 1992, Lampinen 14764 (MA); Kuitia, Parainen, Lemlaxö, 7 Aug 1971, Kukkonen 9106 (LE); Sauvo, Karona, near the church, 13 Jul 1976, Lempiäinen & Ravanko s.n. (MA); Broby, Nylandia, par. Pyttis, 28 Jun 1963, Majander & Nordström s.n. (LE); Norra byn, par. Sottunga, 11 Jul 1973, Nordström s.n. (LE); Uusimaa, Jätkäsaari, Helsinki, harbour, 28 Jul 1968, Oinonen s.n. (LE); Versinais-Suomi, Iniö, Norrby, 13 Jul 1929, Valle s.n. (MA); Jyddo-ojen, Foglo, Ahvenanmaa (Åland), 29 Aug 1907, Wahlberg s.n. (LE).

France. Alsace: at the edge of the river at Haguenau, 24 Jul 1846, Billot 160 (BM); Auvergne: Auliac, 26 Sep 1835, Anonymous s.n. (BM); chemin de Surat à Maringues; chemin de Bussière à St.Clement, Jun 1906, Caumel 334 (BM); Brittany: chemin de Chantenay à Rochemaurice, 12 Aug 1901, Gadeceau 5106 (BM); Corsica: zwischen Pte d’Abatesco und Mignattaja S von Ghisonaccia, 10 Aug 1933, Aellen 4601 (LE); Languedoc-Rousillon: Bouches du Rhone, Camargue, Tour du Valat, SE of Mas, 11 Jun 1968, Kendrick & Moyes 231 (BM); Bouche du Rhone, Camargue, Etang de la Dame, 2 miles SE of Salin de Giraud, 21 Jun 1968, Kendrick & Moyes 346 (BM); Loire-Atlantique: Ouest de France, environs de Vertou, Jun 1880, Gadeceau s.n. (BM); Nord Pas de Calais: Beuvrequent, ruisselet de Slack, 24 Jul 1986, Coutrez 3780 (MA); Pays-de-la Loire: Nantes, loin le Fevre, 18 Jul 1847, Genevier s.n. (BM); Picardy: Aisne, St. Quentin, Jun 1871, Moignier s.n. (BM); Poiteau Charente: Rochel, Plaisance en Oléron (Charente-Inferieure), 20 Oct 1892, Reau s.n. (BM); Rhône-Alpes: Bonneville, bords d’Arve, Depierre s.n. (BM); Haute Savoie, route d’Etrumbiern a Mornex, 14 Jun 1859, Déséglise 281 (BM); Hirault, Mont Blanc, Station Crenugua, 7 Jul 1954, Witte 14547 (MA).

Georgia. Dusheti District, south and east of Fudauri, north of Pasanauri, slopes along Georgian M3 Road (Georgian Military Highway), 1784 m, 5 Aug 2005, Atha et al. 5005 (US); Chevsurya, between Barshak and Reshka, 1 Jul 1903, Busch s.n. (LE); Akmolinsk prov., Kokczetav District, 4 May 1913, Drobov 203 (S); South of the Krestovyy Pereval, Georgian Military Highway, Belaya Aragvi valley, between Ordzhonikidze and Tbilisi, Gruzinskaya S.S.R. along walls in the village of Mleti, 4 Jul 1971, Jenkins 2946 (BM); Somchetya, Teleti, near Tiblisi, Oct 1923, Juzepczuk s.n. (LE); Klukhorskii, between villages of [illegible] and Madniekheva, 13 Aug 1946, Kemularia-Nathadze & Chinthibidze s.n. (LE); Kutaisi, Ozur, 15 Jun 1914, Kikodse s.n. (LE); Abkhasia, Teberdinskii, Protected Area [Terbeda], 1938, Komarov & Komarova s.n. (LE); Kazbek, Dushetskii region, near the 31st field, Tifliskaya Province, 1916, Krylov & Steinberg s.n. (LE); Megrelia, near village Inghuri towards city Tobashi, 14 Aug 1952, Madenova & Kuthatheladze s.n. (LE); along the Georgian Military road in the area of Krestovi Pass N of Tibilisi, 31 Jul 1984, McNeal et al. 332 (MO); Atskhur, Akhaltsikhskii region, vicinity of village Atskhur, 16 Jul 1926, Meffert 314 (LE); Kazbegi, Aug 1971, Menitskii s.n. (LE); Abkhasia, Sukhumsk, District of Tsebeld, in the Peukirskoe gorge on open slopes of the mountain Agysh near the road, 23 Aug 1937, Pobedimova 383 (LE); Abkhasia, Gagra, on the lower road towards Zhoevarskii waterfall, 4 Nov 1915, Sakharov s.n. (LE); Mtiuleti, Kasbegi Region, Dariali Gorge, Greater Caucasus Mountain Range; ca. 12 km N of Kasbegi village, on footpath to Gveleti village to local waterfalls, 1600 m, 24 Aug 1998, Schmidt et al. 2891 (G); Tiblisi, by the River Vera, 26 May 1917, Shishkin s.n. (LE); Adzharskaya, Nizovaya Natanebi, 4 Aug 1948, Sochava & Semenova s.n. (LE); Adzharskaya, gorge of the river Bakhvis-tskali, 9 May 1941, Tatishvili s.n. (LE); valley of the River Terek to the south of the Station Lars, 25 Aug 1949, Vasilchenko et al. 926 (LE); Sukhum, on the right bank of the River Besleta, near the famous factory, 29 Jul 1951, Vasiliev s.n. (LE).

Germany. Bayern: Oberbayern, München, obhere Speilbahn des Golfplatzes Thalkirchen 1 km sudlich des Asam-Schlössels, 550 m, 12 Jul 1992, Förther 48 (B); Lkr. Ostallgau, NW Trauchgau, 845 m, 24 Jul 2001, Herb. Willing [Willing & Willing] 13418 D (B); Lkr. Garmisch-Partenkirchen, SW Krün, 911 m, 5 Aug 2001, Herb. Willing [Willing & Willing] 15543 D (B); Berlin: Pichelswerder, Bez. Spandau, 35 m, 4 Jun 1998, Herb. Willing [Eisenblatter & Willing] 4624 (B); North Rhine-Westphalia: Münster in Westfalen, Wilms s.n. (BM); Rheinland-Pfalz: Westerwaldkreis, Seeburg, westerwalder Seenplatte, Haidenwalder bei Seeburg, 421 m, 2 Jul 1978, Kalheber 78-424 (B); Schleswig-Holstein: Rendsburg, Larsen et al. 106 (BM, LE, MA); Thüringen: Am Ufer des Altenberger Teiches bei Eisenach, 28 Jul 1947, Launert s.n. (BM).

Greece. Thracia, prov. Xanthi, in valle infra pagum Dhamari, 4 km infra junctionem torrentium, 500 m, 16 Aug 1978, Greuter 16361 (B); Dhrama, prope phylacion nom. Praseki, a pago Ano Vrondous, ca. 6 km boreo-occidentum versus, 1320 m, 22 Aug 1978, Greuter 16703 (B); Mt. Olympe, Thess, Jun 1929, Guiol 642 (BM); Etolia-Akarnania, Arahova (Ar 358), 900 m, 8 Jul 1996, Herb. Willing [Eisenblatter & Willing] 48029 (B); Thesprotia, Ep. Souliou, NW Paramithia (The 209), 190 m, 26 Jun 1997, Herb. Willing [Eisenblatter & Willing] 55434 (B); Trikala, Ep. Kalambakas, Notia Pindhos, NW Kalambaka (Tri 200), 330 m, 18 Jul 1997, Herb. Willing [Eisenblatter & Willing] 60744 (B); Nom. Ionnanina, Ep. Dhodhonis, Tsepelovo (Ioa 574), 1000 m, 17 Jul 1998, Herb. Willing [Eisenblatter & Willing] 67904 (B); Nom. Kastoria, Ep. Kastorias, SW Vyssinia (KAS 164), 820 m, 31 Jul 1998, Herb. Willing [Eisenblatter & Willing] 71725 (B); Pella, Ep. Edhessis, W Loutraki (Pel 115), 260 m, 27 Jul 2000, Herb. Willing [Eisenblatter & Willing] 86148 (B); Arkadhia, Elati, 1180 m, 27 Sep 2003, Herb. Willing [Willing & Willing] 118384 (B); Paranestion, Nomos Drama, Dimos Paranestion, river Nestos ca. 1 km SW of Paranestion, 100 m, Aug 1996, Liebfritz & Schuler s.n. (B); Nomos Kavala, Dimos Keramoti, mouth of river Nestos, 1 m, 31 Aug 1998, Schuler 98/447 (B); Ipiros, prob. Ioannina, distr. Konitsa, Konitsa, on both sides of the Aoos River above the bridge, 460 m, 23 Jul 1971, Stamatiadou 13382 (BM).

Hungary. Sin. loc., Jul 1877, Anonymous s.n. (BM); Kozragohid, Budapest, 26 Aug 1904, Degen s.n. (LE).

India. Jammu and Kashmir. Kashmir, Kishtwar, 1829 m, 17 Sep 1876, Clarke 31361 (BM); above Barangalla, 2000 m, 25 Jun 1902, Drummond 13886 (E); Gandarbal, Sind valley, 1585 m, 1 Jun 40, Ludlow & Sherriff 8101 (BH, BM, E); Hajipir pass, Poonch to Uri, 2134 m, 4 Jul 1934, Stewart 14000 (A); Naranag, Wangat valley, 2134 m, 8 Aug 1939, Stewart & Stewart 18104 (GH); Poonch & Azad Kashmir, below Hajiperi, 2134 m, 3 Jul 52, Stewart & Nasir 24062 (BM).

Iran. Ghilan, Rescht, 13 Jul 1902, Alexeenko 47 (LE); Fars, Tang-e-Sorkh, between Ardakan and Yasuj, 2450 m, 18 Jun 1977, Bokhari & Edmondson 2159 (G); Meshed, 1941, Mrs Donaldson s.n. (K); Amol, 25 miles S of Amol, Central Elburz N. slopes, 1219 m, 21 Aug 1966, Furse 9071 (K); . Keredj, 29 Sep 1934, Gauba 829 (B); Kurdestan, 31 km N of Sanandaj, 2000 m, 8 Jul 1964, Grant 16064 (MO); Dozein, 5 km N of Dozein, which is a village 55 km SE of Gonbad-e-Kabus, 950 m, 19 Jun 1977, Hewer 3961 (K); Khairat, Mazenderan, 2439 m, 25 Jul 1940, Koelz 16569 (US); Kuhi, near Teheran, Pidjehek, Jul, Kotschy s.n. (BM); Fars, 36 km NW Ordekam, 14 Jul 1959, Pabot 2446 (G); Qotur River, W Khvoy towards the Turkish border, 1800 m, 10 Jul 1971, Rechinger 41735 (B, G); Keredj, Elburs Mountains, Keredj River, 6 Jun 1937, Rechinger 733 (S); Hamadan, 2134 m, 7 Sep 1929, Rogers 538 (K); Joistan, Talighan Valley, 1829 m, Jul 1953, Trott 24 (K); Astava, shoreline of Caspian Sea, 21 Aug 1966, Wright 65 (K).

Iraq. Baghdad, Pig Island, 10 May 1960, Agnew et al. W-1679 (K); Tigris Plain. near Mosul, 200 m, 5 May 1936, Low 284 (BM); Penjiwin, 1200 m, 8 Jun 1948, al Rawi 12159 (K, US); Pushtashan, 15 km NE of Rania, lower slope of Qandil Range, 1150 m, 29 Jul 1957, al Rawi & Serhang 23865 (K); Erbil, Pushtashan, Montes Qandil, (Kurdistan), 1000 m, 28 Jul 1957, Rechinger 11027 (G).

Ireland: West Galway, Connemara, Renville, Babington 299 (BM); Cork, Bishopsbrook, 1849, Herb. J. Carroll s.n. (BM); South Kerry, Dooks, 15 Jul 1903, Marshall s.n. (BM); Meath, Drogheda, Aug 1956, McClintock s.n. (BM); North Kerry, Walls between Killarney and Mucross, 24 Aug 1883, Ridley s.n. (BM); South Kerry, Rossbeigh Dunes, 5 Aug 1935, Simpson 35620 (BM); South-East Galway, near Dalystown, 24 Jun 1952, Simpson 52108 (BM).

Israel. Gilead, Burmah to Gerashi, 4 May 1886, Anonymous s.n. (K).

Italy. Abruzzo: Teramo, cerca Castelli, Findo de al Salsa, 1100 m, 2 Jul 2002, Navarro et al. 4348 (MA); Apulia: Puglia, farm about 7 km NE of Taranto, 1881, Vitantonio 1675 (BM); Emilia-Romagna: Parma, Anonymous s.n. (BM); Friuli-Venezia Guilia: Trieste, Sistiana, 25 May 1980, Greuter et al. 17348 (B); Sicily: Nebroda Mtns., ad pedes montium Madoniarum prope Polizzi, 600 m, 6 Jul 1874, Strobl s.n. (BM); Catania, Randazzo, Lago de Gurrida, 820 m, 5 Jun 2000, Ávarez et al. 1490 (MA); Tuscany: Agro Lucchese, Lucca country side, Jun 1835, Puccinelli s.n. (BM).

Japan. Hokkaido: Sapporo, 4 Jul 1903, Arimoto s.n. (GH, MO); Iburi, Tomakomai, 30 Jul 1930, Akiyama s.n. (S); between Kotoni and Sapporo, 22 Aug 1929, Dorsett & Morse 1128 (US); Yezo, prov. Nemuro, to Attoko-too from Attoko-Nemuro-gun, 8 Jul 1959, Furuse s.n. (A, S); on the coasts vicinity and Oikamanai-numa Bansei, Taiki-choo, Hirowo-gun, Province Tokachi, 19 Jul 1974, Furuse 6434 (K); Wakkanai-Park. Wakkanai-shi. Province Kitami. Hokkaido ((Yezo)), 18 Aug 1975, Furuse 9465 (K); Jyuusam-mura. Nishi-tsugaru-gun. Province Mutsu. Hondo, 20 Sep 1955, Furuse 29993 (K); Watsakinai. Toyotomi-choo. Teshio-gun. Province Teshio. Hokkaido (Yezo), 2 Jul 1975, Furuse 8980 (K); prov. Iburi, between Tomakomai-shi and Lake Shikotsu, 15 Aug 1981, Takahashi 1650 -A (NY); Honshu: Shimura, Itabashi-ku, Tokyo Prefecture, Jun 1907, Makino 75314 (LE); Shiga, Yotsugawa, Adogawa Cho, Takashima gun, Lake Biwa, 86 m, 17 Sep 1991, Kanda & Kuribayashi 1 (A); Hondo, Oze-ga-hara, prov. Kozuke, 26 Aug 1950, Mizushima 417 (A); Aomori-shi, Arakawa, below Jogakura Spa, 810 m, 8 Aug 2000, Yonekura 5960 (A).

Kazakhstan. West Kazakhstan Prov., Right side of river Ural, 2 km downstream of Kopseharov, 15 Feb 1955, Grubov & Lynbarskii 32 (MO); South Kazakhstan Oblast, Shymkent, prov. Syr-Darja, Distr. Tschimkent, Tchiluk, 1908, Knorring 604 (S); Tselinogradskaya Oblast’, near village of Kurgaljinskii, on the lower eastern shore of the Lake Kuraljin, in the surroundings of the settlement of Karazhar, 20 Jun 1980, Sidorova 155 (LE).

Lebanon. Ain Zahatallah, Mt. Lebanon, Aug 1912, Letchworth s.n. (K).

Lithuania. Holonety, dist. Wilkomierz, 30 Jul 1895, Rudiminowna s.n. (B, BM).

Luxembourg. Jardin Botanique, 20 Aug 1837, Gay s.n. (K).

Macedonia. Popovo, NW foot of Krusa Balkan, 3 m E of Lake Doiran [Doirani], 1918, Gooding s.n. (BM); Kojnsko, distr. Duditza-et Saharupa-planina, 550 m, Jun 1917, Schultze-Jena 138 (B).

Mongolia. southwestern Mongolia, Khovdinskii Aymak, southern slope of the Mongolian Altai, 45 km to the E of the villageg of Bulgan, along the River Bulgan, on the border with KNR [China], 1979, Gubanof 7321 (LE); “ad Mongolia borealis, Altai australis ad Totyask nigrum”, 1876, Potanin s.n. (K).

Morocco. Xauen, 650 m, 28 May 1928, Font Quer 354 (BM); Tazzeka, 13 km from Taza on minor road, Ras-El-Ma, 990 m, 5 Jul 1993, Jury et al. 11718 (K).

Norway. Oslo, 24 km hinter, am fjord, 8 Sep 1980, Meyer s.n. (B); Stjordal, near Hagra, 1926, Trethewy s.n. (BM); Astre Biarum, 6 Sep 1902, Triatz s.n. (BM).

Poland. Wolka wysoka prope Sanniki, distr. Gostyn, 22 Jul 1895, Drymmer s.n. (BM); Orawicza, in Banat, 6 Aug 1840, Wierzbicki s.n. (GOET).

Portugal. Salvador, as margenes do rio Sever, Sera de S. Maccede, 12 Jun 1959, Beliz & Raimundo 5046 (MA); inter Garvas et Panvies, Jun 1875, Daneau 1184 (BM); Alto Alentejo, Elvas, Frequisia de Varche, Horto do Carlos Ranita, margem da Ribereira do Cancão, 30 Sep 1984, Guerra 1518 (MA); Beira Alta, Vizeu, Santa Comba-Dão, Pinheiro de Azer, descida para a capela de Sra. do Pranto, 11 Jun 1982, Marques 2087 (MA); Rangel prope a Coimbra, Jun 1889, Moller s.n. (BM); Oliveira, M.P. de, s.n. (BM); Entremadura, Obidos nas margens dos vales na La Goa de Obidos, 14 Jun 1917, Rainha 377 (MA); Idanha a Nova, Ribeira do Poasul, Jul 1883, Ricardo da Cunha s.n. (BM); Alijó, Pinhão, beira-Doura, foz do Rio Pinhão, 30 Jun 1070, Rozeira et al. s.n. (MA); Douro Litoral, near Oporto, 2 Jul 1887, Murray s.n. (BM); Madeira, sin. loc, 1804, Turner s.n. (K); Tagum, 1848, Welwitsch 202 (LE).

Romania. Comm. Balcani, prope pagum Frumoasa, 550 m, 12 Jul 1972, Barabas 487 (BM).

Russian Federation. Central Federal District: Moscow, 30 Aug 1992, Croat 73991 (MO); Far Eastern Federal District: Amurskaya Oblast’, Radde, around the village, on the River Amur, 25 Sep 1926, Selivanova s.n. (LE); Amurskaya Oblast’, Zea, Zeyskii region, around the town of Zea, 30 Jul 2000, Shaligin s.n. (LE); Dalnevostochnii Krai, near Valdivostock, 12 Jul 1929, Transhel 284 (LE); Kurilskii Islands, Iturup [illegible], 914 m, 29 Aug 1946, Vorobeev s.n. (LE); Archangelsk Oblast, Archangelsk, Beresnik, 1919, Grantham s.n. (BM); around Vladivostock, on the bank of Lake Khanka, 10 Aug 1929, Shishkin 1306 (LE); island of Sakhalin, village of Vladimirovskoe, 26 Jun 1899, Shestunov 53 (LE); Primorskaya Oblast’, around 25 verst [measure of distance] to the west of Vladivostock, on the slope of a mountain, 29 Jun 1926, Ivanova s.n. (LE); Primorskaya Oblast’, Khabarovsk, left bank of the River Amur, near Khabarovsk, 25 Jul 1926, Solokhin s.n. (LE); Primorskaya Oblast’, Petrovolikova Bay, island of Furugelma, 21 Sep 1922, Kozlov 483 (LE); Primorskaya Oblast’, the village of Chernshovekoe [Chernyshevka], 30 Jul 1902, Paltzevskii s.n. (LE); Sakhalin Oblast, on the sea slope of southern Sakhalin, between the settlements of Khvostovo and Altasovo, 13 Aug 1959, Pobedinova & Konovalova 1347 (LE); North Caucasian Federal District: Tegenekli, basin of the River Baksan, right side of the gorge Prielbrusye, 23 Aug 2000, Menitskii & Popova s.n. (LE); Nalchik, to the south of the settlement of the village Belaya Rechka, 2 Aug 2000, Menitskii & Popova s.n. (LE); [Chechnya] Itun-kale, Argun, left side of river Argun, above the village, 23 Aug 1988, Menitskii & Nikolaev 38 (LE); [Dagestan] Machachkala, by the river Terstagay, 26 Sep 1926, Orlov s.n. (LE); [Dagestan] Gergebilskii region, between villages of Arakani and Mogok, 14 Jun 1961, Tsvelev et al. 914 (LE); [Dagestan] Levashinskii region, between villages of Gersibil and Levashy, 27 Jul 1961, Tsvelev et al. 4175 (LE); Chechyna, Terekaya Oblast’, Beslan, May 1901, Alexeenko 14374 (LE); Kabardino-Balkar, Bezeniyskoe gorge, on the slope of the mountain Ushtensk, 12 Aug 1995, Shkhagapsoev & Korchevskaya 675-2000 (LE); Ordzhonikidzevskii, “Chechnya-Ingushetia”, 22 Jul 1966, Vlasov & Kuvas s.n. (LE); slope of the left bank of the River Bizhgon 10 km S of the station Storozhevaya, 31 Aug 1947, Borisova 369 (LE); Stavropolskii Krai, bank of River Podkushok in the region of Pyatigorsk, 8 Aug 1990, Yagushevskii s.n. (LE); Terekaya Oblast’, Nalchik, near town, 30 Jun 1917, Paltseva s.n. (LE); Northwestern Federal District: Ladozhskoe Lake, town Lakhdempokhye, 6 Jul 1961, Pobedimova & Gladkova 97 (LE); near St. Petersburg, pag. Olgino, in litore sinus Fennici, Jun 1919, Czernjakovskaja s.n. (BM); Siberian Federal District: Siberia, regio transbaicalensis, Utotschkina, Selenga River, 500 m, Jul 1900, Ehnberg s.n. (G); Irkutsk Oblast, Irkutsk, Besser s.n. (K); Krasnoyarsk Krai, Yeniseysk, “Siberia, Jenisei, Jeniseisk”, 23 Jun 1876, Arnell s.n. (S); Central Siberia, region of Novosibirsk, near railway station Seyatel’ ca. 3 km Nw of Akademgorodok near river Ob’, 2-3 km below hydroelectric dam, ca. 20 km SSE of Novosibirsk, 140 m, 24 Jul 1979, Iltis et al. 1115 (G, S); Podkova, Yekaterinava, 21 Jun 1906, Yálovaya s.n. (LE); Southern Federal District: Mineralnie Vody, eastern slope of the mountain Zmeyka, 7 Aug 1996, Menitskii & Popova 122 (LE); River Mzymty [Myzmyta] near the mouth of River Agipse, 7 Sep 1928, Leskov s.n. (LE); Tuapsinsk, Black Sea Region, 8 Jul 1915, Litvinov s.n. (LE); [Adygea] Kubanskii Oblast’, Maykopsk, between the stations of Dondukovkskaya and Sergieskaya, in the valley of the River Fars, May 1915, Kozo-Polyanski & Preobrasheiski s.n. (LE); Krasnodavskii Krai, Novorossika, Black Sea region, near the road on the high shore of the sea, 25 Jun 1923, Pojarkova 156 (LE); Urals Federal District: Goygol Rayon, Goygol, territory of Elizabethapolis [Yekaterinburg], 8 Jun 1844, Kolenati 1753 (LE).

Serbia. mountain above Jviato Petka, 22 Aug 1921, Yermoloff s.n. (BM).

Slovenia. within 6 miles of Bled (in Slovenja, at the borders with Austria), which includes the lower slopes of Mt.Triglav, 304 m, 1923, Leathes s.n. (BM).

Spain. Alcabete: Reolid, 800 m, 19 Aug 1985, Figuerola s.n. (MA); Andalucia: Granada, Castril, pr. Cortijo del Nacimiento, 1100 m, 21 Jul 1978, Charpin et al. 15109 (MA Asturias: Villaviciosa, playa de España, 5 m, 5 Oct 1992, Aedo 2469 (MA); Avila: Conabeltrán (?), Jul 1918, Cogolludo s.n. (MA); Baleares: Mahón, San Sancho, 18 Apr 1899, Pons, Gereau s.n. (MA); Burgos: Carazo, junto a la Fuente de la Mora, Peñas de Cervera y aledaños, 1200 m, 24 Jul 1979, Pons Sorolla & Susanna 616 (MA); Caceres: Baños de Montemayor, 15 May 1944, Cabellero s.n. (MA); Cadiz: collected in a farm named ‘Marcellino’ in a rough country between Los Barrios and Algeciras, 60 m, 14 Jun 1973, Brinton-Lee 1346 (BM); Cantabria: Santander, Cueto, El Panteón del Inglés, 22 May 1992, Busqué s.n. (MA); Castellón: Villafarnés, on same sheet several specimens, also from Castielfavib and Vallanca (Valencia, 1792, 1793), and Jardin de la Chevette (France, 1781), 1792, Anonymous s.n. (MA); Castilla: Bujedo, 28 Jul 1920, Elías s.n. (BM); Catalunya: Castello, Castillo de Villamalefa, Río Villahermosa, 540 m, 22 Oct 1998, Riera et al. 19505 (MA); Cerdagne, Llivia, val de l’Estahuja, 1300 m, 10 Aug 1918, Sennen s.n. (BM); Cuenca: Hoz de Cañizares, 10 Jul 1932, Cabellero s.n. (MA); Tarancón, 7 Apr 1977, López 8 (MA); Cuidad Real: Frencaliente, margenes del Río Pradillo, 600 m, 7 Jul 1997, García Río s.n. (MA); Galicia: Lugo, Riberas de Lea, 25 Jul 1956, Carreira s.n. (MA); Guadalajara: Zaorejas, Hoz del Tajo, piscifactoria del Campillo, 900 m, 18 Jul 1981, Muñoz 512 (MA); Huelva: Almonte, Coto de Donaña, Reserva de La Rocina, 28 Jun 1977, Castroviejo & Porta 733 (MA); Madrid: San Agustín de Guadalquix, riberas del Río Guadalquix, 3 Aug 1982, Moreno s.n. (MA); Navarro: Pamplona, 16 Jun 1971, Moriyóu s.n. (MA); Salamanca: alrededores de Bejar, Aug 1914, Cogolludo s.n. (MA); Segovia: Granja de San Ildefonso, desde la finca La Sauca hacia la carretera a Torrecabelleros, 30TVL1529, 1150 m, 9 Sep 1985, García Adá & Egido 1816 (MA); Soria: Cañon del Río Lobos, 14 Jul 1980, Buades s.n. (MA); Tarragona: 20 km SW of Tarragona, between Cambrils and Hospitalet, 29 May 1962, Brummitt et al. 298 (BM); Toledo: ribera del Río Tajo, 500 m, 18 Jun 1983, Egido 947 (MA); Valencia: Bco. Bocairente, 510 m, 7 Jul 1980, Palasí s.n. (MA); Valladolid: Encinos de Esqueva, en Arroyo de los Frios, 11 Jul 1983, Fernández Alonso 2071 (MA); Zamora: Parque Natural Lago de Sanabria, Forcadura, 1100 m, 1 Jul 1987, Anonymous s.n. (MA).

Sweden. Uppland, ad sopes Upsalia passim, Aug 1870, Ahlberg s.n. (BM); Uppland, Funbo Parish, southern shore of Lake Goren, 6 Jul 1944, Alm & Smith 272 (BM); Bohnslam, Väderöarna, 10 Jul 1909, Almquist s.n. (BM); Brohuslan, Fjalibacha, 3 Jul 1910, Almquist s.n. (BM); Uppsala, Andersson s.n. (MA); Skafto, Gaso, 20 Jun 1948, Ankarsward s.n. (BM); Södermanland, Toiosocken, Stora Grasskar, 6 Aug 1922, Asplund s.n. (BM); Västergötland, Goteborg, 14 Aug 1959, Blom s.n. (BM); Skåne, Buutofha, Jul 1893, Johansson s.n. (BM); Scania, Kivik, Aug 1911, Johansson s.n. (BM); Malmo, Aug 1886, Johansson s.n. (BM); Östergötland, Linköping, Sep 1912, Johansson s.n. (BM); Sm. Oscasthaven, 4 Aug 1909, Köhler s.n. (BM); Yonstrup, Jul 1861, Mortensen s.n. (BM); Södermanland, Paroecia Strängnäs, Sundby, 27 Jun 1925, Samuelsson 1371 (BM); Bohuslän, Högås Parish, the innermost part of the bay Svältekilen, 14 Aug 1960, Santesson 13962 (BM).

Switzerland. Vaud: Montherod pres d’Aubonne, Anonymous s.n. (BM); inter Nyon et Lausanne ad lacum Lemanni, Jul 1838, Sander s.n. (BM); Zürich: Rümlang, Gemeinde Rümlang, Näherer Einschlag, 420 m, 20 Jun 1947, Bührer s.n. (MA).

Syrian Arab Republic. Monts Arnanis, 762 m, Aug 1906, Haradjian 481 (K); Zahleh, Cacle [coll. Prof. Post, ex. Herb. J.T. Rothrock], Post s.n. (F).

Tajikistan. Giesar, Pamiro-alay, near the village of Giesar on the river Khanaka, 31 Aug 1942, Grigoriev 757 (LE).

Turkey. Ouchak, Phrygie, Bords du ruisseau d’Ouchak (Phrygie), 910 m, 9 Jul 1858, Balansa 1137 (A, B, BM, K); Cumasia, 24 Apr 1889, Bornmüller 344 (B); Prov. Konya distr., Ermeneh (Cilican tracks), Sorivandt, 1200 m, 15 Aug 1949, Davis 16221 (K); Ankara prov., Hacikacum Valley, 11 Jun 1952, Davis & Dodds 18783 (BM, K); Cankiri-Kalecik, 9 Jun 1954, Davis & Polunin 21737 (BM); B9 Agri, Murst Valley, 3-5 km from Tutak to Hamur, 1600 m, 2 Jun 1966, Davis 44044 (K); Suluçem, (Musum) S edge of Balik G, 2300 m, 23 Jul 1966, Davis 47185 (K); C10 Hakkari, Nehil çayi, 48-55 km from Hakkari to Yuksekova, 1600 m, 14 Jun 1966, Davis 44905 (K); Cankiri-Kalecik, 9 Jun 1954, Davis 21737 (K); Tokat-Artova, 4 Sep 1954, Davis & Polunin 24854 (K); Abant Gol, Prov. Bolu, 1400 m, 11 Jul 1962, Davis & Coode D-37291 (K); Prov. Zonguldak, Bartin-Amasra, 25 Aug 1960, Khan et al. 796 (K); Çibuktal, bei Ankara, 10 Jun 1932, Kotte s.n. (K); Adapazari, Kuntay 8735 (MO); Dorukhan geçidi, Nord-seite (A4 Zonguldak), 800 m, 8 Aug 1982, Raus 6964 (B); Takiansi, 304 m, 29 Aug 1878, Regel s.n. (LE); Yagan, Erzerumskii sandzhak, river Arakas above Yagan, 9 Jul 1916, Sapozhnikov & Shishkin s.n. (LE); River Fol-chai, Trapezundskii sandzhak, 28 Jun 1917, Shishkin s.n. (LE);Vitse, Lazistan, 12 Aug 1912, Shishkin s.n. (LE); Pamphylia, Antalya, 23 Jun 1936, Tengwall 48 (GH, K, S); Bolu, Koru Motel, 850 m, 30 Aug 1972, Uotila 20137 (G); Ismid, (Nicaea), 1834, Wiedemann s.n. (LE); vicinity of Tortum Lake. East Turkey, 30 Jul 1961, Winter 278 (BM).

Turkmenistan. Germad, 8 Jun 1899, Antonov s.n. (LE); gorge Verkhanyaya Firyuza, central Kopet Dag, 31 Aug 1945, Blinoskii s.n. (LE); going up the river Dyuynee near the spring Autet, 1200 m, 25 May 1928, Bobrov & Yarmelenko 168 (LE); Karasashinskii region, river Gebe-Zvud, 14 Jul 1931, Borieova 472 (LE); Zakaspiiskaya Oblast, Shorkala [Shor-Kala], Alta-Yaba, aul Shorkala, 23 Jun 1924, Bukinich 45 (LE); Kopet Dag, road Utsch-Kuyu Nukhur, 28 Jun 1925, Fedchenko et al. 790 (LE); Germad, on the banks of the river Saki[l]yad, near village of Germad, 13 Jul 1934, Gnezdillo 97 (LE); Zakaspiiskaya Oblast’, Askabadskii area, mountains of Kopet-Dag, gorge of Chuli, 4 May 1912, Lipskii 1807 (LE); Zakaspiiskaya Oblast’, Askabadskii area, mountains of Kopet-dag, Chuli, 17 Jun 1911, Mickelson 487 (LE); Zakaspiiskaya Oblast’, [Finshchi], Jun 1895, Minkevich s.n. (LE); Tilutschi, 4000 m, 23 Jul 1879, Regel s.n. (K); Zakaspiiskaya Oblast’, Chuli Mountains, 26 Jun 1912, Samokish 1373 (LE); Zakaspiiskaya Oblast’, Chuli Mountains, near Askabad, 9 Jun 1911, Sevdturabov s.n. (LE); Kisil-arwat, Karakala, by a little river the valley of [Foltere], 10 Jul 1901, Sintenis 2035 (B, LE).

Ukraine. Kiev, 28 Aug 1906, Lonaczewsky s.n. [7272a](BM, MO); Zakarpatskaya Oblast’, Uzhgorodsky Raion, road to Imina (Transcarpatia Province, Uzhgorod District), 26 Jul 1967, Apentyeva s.n. (K).

United Kingdom. Channel Isles: Guernsey, Petit Bo, Jul 1883, Fawcett s.n. (BM); Jetou, 26 Jun 1932, Meinertzhagen s.n. (BM); Sark, 8 Jul 1953, Sowerby 4 (BM); England: Huntingdonshire, Woodwalton, 7 Jun 1914, Adamson s.n. (BM); Leicestershire, Scraptoft, 15 Aug 1949, Bangerter s.n. (BM); Warwickshire, Between Pillerton Priors and Eatington; Ettington parish, 9 Jul 1961, Bangerter 1058 (BM); North Wiltshire, Calne, 27 Aug 1914, Barton s.n. (BM); Nottinghamshire, Cresswell Gorge, east of Cresswell, 60 m, 22 Jul 1963, Bowden & Hillman 137 (BM); Yorkshire, Spern Point, 18 Aug 1070, Castroviejo s.n. (MA); Buckinghamshire, Aston Clinton parish, Dancers End Nature Reserve, 16 Aug 1942, Dandy, J.E., 943 (BM); Herefordshire, Startop’s End Reservoir. Tring Rural parish, 20 Jun 1943, Dandy 974 (BM); East Suffolk, Corporation Marshes, Walberswick; Walberswick parish, 6 Jun 1961, Elword 19 (BM); Kent, Wrotham, 21 Aug 1913, Fox 1200 (BM); West Cornwall & Scillies, Tresco harbour, Isles of Scilly, 9 Jun 1963, Gerrans 1098 (BM); Cornwall, Daymer Bay, Trebetharick, near Weybridge, 8 Jul 1967, Halliday 163 /67 (LE); East Gloucestershire, Clapton, 9 Aug 1884, Hanbury s.n. (BM); West Kent, Woolwich Arsenal, 23 Aug 1894, Herb. A.H. Wolley-Dod s.n. (BM); Oxfordshire, Banbury, 3 Aug 1863, Herb. Alfred French s.n. (BM); South Essex, Romford, Jun 1914, Herb. H. Stanley Redgrove s.n. (BM); East Cornwall, Tintagel, 26 Jun 1907, Herb. L.B. Hall 306-1134 (BM); West Cornwall & Scillies, Lizard, 27 May 1886, Herb. R.P. Murray s.n. (BM); West Gloucestershire, over Sapperton Tunnel, 18 Jun 1908, Herb. W.C. Barton s.n. (BM); Lancashire, road to Newland, Hodgson 738 (BM); Berkshire, Speen Moors near Newbury, 8 Sep 1894, Jackson s.n. (BM); East Norfolk, Lopham Middle Fen, 3 Jul 1973, Jones & Vickery 20 (BM); Hampshire, Southampton, edge of The Avenue, Southampton Common, 17 Jul 1964, Kerr 220 (LE); South Lancashire, Nr. Freshfield, Ainsdale Dunes Reserve, 26 Jul 1966, Kendrick 30 (BM); Suffolk, Brome, 19 Jul 1832, Kirby s.n. (BM); Durham, Hartlepool, North sands, Sep 1872, Lees s.n. (BM); West Lancashire, Lancaster, Shingle at Bare, Morecambe Bay, 27 May 1912, Lees s.n. (BM); Lincolnshire, Skegness, 24 Aug 1904, Linton s.n. (BM); North Essex, Frinton to Clacton, 14 Sep 1889, Linton s.n. (BM); South Hampshire, Lymington, Pilewell, 24 Aug 1893, Linton s.n. (BM); East Kent, Wittersham, May 1927, Meinertzhagen s.n. (BM); Norfolk, Sutton Broad, Sep 1928, Meinertzhagen s.n. (BM); North Devon, Woolacombe, 14 Sep 1931, Meinertzhagen s.n. (BM); South Somerset, Knowle, Jul 1881, Painter s.n. (BM); Staffordshire, Knypersley, Jul 1886, Painter s.n. (BM); Hertfordshire, Hadley Wood, 16 Jul 1882, Parker s.n. (BM); West Norfolk, Thompson, 12 Aug 1917, Robinson s.n. (BM); South Devon, Dartmouth, 30 Jul 1952, Robson 350 (BM); Dorset, Pentridge, Cranborne Chase, 19 Jun 1934, Simpson 34161 (BM); West Sussex, Chichester, Kingley Vale, West Stoke, 12 Aug 1905, Standen s.n. (BM); Cheshire, Parkgate, 12 Oct 1965, Valdés s.n. (MA); Surrey, Farncombe near Godalming, edge of R. Wey, 14 Aug 1952, Welch 4613 (BM); North Somerset, Bristol, Clifton Down, 15 Jul 1929, White s.n. (BM); Derbyshire, Monsal Dale, near the station, 21 Jun 1916, Wilmott 708 (BM); Middlesex, Buckingham Palace garden, 9 Jun 1995, Wiltshire s.n. (BM); Isle of Man: Andreas, 22 Jun 1929, Paton s.n. (BM); Scotland: East Lothian, Anonymous s.n. (BM); Fife, Burntisland, 12 Jun 1847, Anonymous s.n. (BM); East Sutherland, Creich, road between Invershin and Bonarbridge, 20 Aug 1958, Groves 2838 (BM); East Lothian, Haddington, Longniddry, 1850, Herb. J.T.I. Boswell-Syme s.n. (BM); West Perthshire, Clackmannanshire, Dollar [v.c.87], 1851, Herb. J.T.I. Boswell-Syme s.n. (BM); Mid Ebudes, Mull, near Nun’s Pass, west of Carsaig, 24 Sep 1967, Kenneth & Stirling s.n. (BM); Renfrewshire, Crossmyloof, 14 Aug 1953, Mackechnie s.n. (BM); Easterness, Beauly, 5 Aug 1892, Marshall s.n. (BM); near Mull of Kintyre, 2 Sep 1961, McClintock s.n. (BM); South Aberdeenshire, Jul 1947, Robson 349 (BM); Wales: Pembrokeshire, Pembroke, Lydstep, 28 Aug 1907, Bickham 1134 (BM); Radnorshire, Gladestry, Jul 1951, Edwards 5 (BM); Glamorgan, Sully Island, Jul 1905, Herb. H.J. Riddelsdell s.n. (BM); Merionethshire, Dolgelley, [v.c.48], 15 Aug 1923, Herb. W.C. Barton s.n. (BM); Radnorshire, Between Erwood and Aberedw, 17 Aug 1885, Ley s.n. (BM); Holyhead, Anglesey, 4 Aug 1955, Robertson D-11 (LE).

United States of America. California: Shasta County, Fall River Hills, 3 miles SW, 4 Jul 1934, Howell 12411 (GH); District of Columbia: Washington DC, National Arboretum, 15 Jul 1965, Mazzeo & Meyer 1090 (G); Idaho: Boise, 18 Jul 1911, Clark 345 (G, GH); Power County, Meader’s Trout Farm, 28 May 1961, Thiel s.n. (LE); Illinois: Kane County, Elgin, north part, 24 Jul 1928, Benke 4878 (GH); McHenry County, Fox River Grove, 18 Jun 1934, Benke 5661 (GH); Lake County, Lake Villa, 3 Aug 1906, Gleason & Shore 123 (GHDu Page County, Morton Arboretum, near Lisle, 26 Oct 1922, Palmer 22347 (A); Indiana: Wells County, Harrison TP, 31 May 1903, Deam s.n. (GH); Noble County, Kendalle, Martin Farm about 2 miles southeast of Kendalle, 23 Aug 1928, Dean 46131 (GH); Berrien County, Niles, 17 Jun 1933, Palmer 40391 (A); Putnam County, Greencastle, East Seminary Street, 1 Sep 1940, Welch 6120 (GH); Maine: Washington County, Lake View Campground, Moose Island, along western passage into Passamaquoddy Bay, N of East Port, 10 m, 30 Aug 2000, Nee & Atha 50937 (BM); Maryland: Cumberland, Jul 1849, Prior s.n. (K); Massachusetts: Hampden County, Knightville Dam, vicinity of overlook on Hwy. 112, about 1.3 mi N of Knightville, 23 Jun 1984, Bates & Elsik 83 (LE); Berkshire County, Tyringham Valley, 10 Sep 1926, Meredith s.n. (LE); Suffolk County, Franklin Park, 7 Jul 1883, Seymour s.n. (USM); Michigan: Mason County, Hamlin Lake, Ludington, 25 Jul 1910, Chaney 91 (GH); Flint, Aug 1877, Clarke 2195 (K); Cheboygan County, Douglas Lake, northern end of Lower Peninsula, Gleason 269 (K); Clinton County, East Lansing, 7 miles northeast near route 78, 9 Sep 1955, Perdue 2016 (GH); Montana: Powell County, Grant Kohrs Ranch National Historic Site, Deerlodge on US Hwy 10, 1373 m, 12 Aug 1983, Ray 194 (GH); Nevada: Washoe County, Truckee River, 3-7 miles W of Sparks, 1219 m, 6 Aug 1937, Moore & Franklin 929 (GH); New Jersey: Union County, Summit, Passiac River, 5 Jun 1969, Hellquist s.n. (GH); Cape May County, Five-mile Beach, 3 Oct 1899, MacElwee 1448 (GH); New York: Tompkins County, Ithaca, Dryden Road, near Dwyer’s Bridge, 12 Jun 1916, Bechtel 7126 (GH); Washington County, Lake George, N. Beaver Creek, Vaughn’s, north of Hudson Lake, 12 Jun 1913, Burnham s.n. (GH); Westchester County, Yonkers, Oatskill Aqueduct, 10 Jul 1934, Gleason et al. 1404 (LE, MA); Tompkins County, in creekbed of Taughannock Creek upstream from the falls, 24 Sep 1983, Kearney 132 (BM); Bronx, New York Botanical Garden, in the Swale, 28 m, 9 Jun 2006, Nee & McClelland 54448 (G); Cayuga County, Lake Como, 15 Jun 1919, Randolph & Wiegand 12864 (GH); Orange County, Aleck Meadow Reservoir, Black Rock Forest, 6 Jul 1936, Raup 7502 (GH); Yates County, Canandaigua Lake, south end, 19 Aug 1933, Swartley 66 (A); Delaware County, North Harpersfield, 11 Jun 1906, Topping 137 (A); North Carolina: Haywood County, Waynesville, US 276 near city limits, W side of highway, 823 m, 16 Aug 1969, Pittillo & Hensley 3102 (GH); Ohio: Butler County, Oxford, south of Oxford, 13 Jul 1958, Cobbe m-52 (G); Portage County, Garrettsville, 15 Jun 1895, Webb 68 (GH); Oregon: Multnomah County, St. Johns, near slough below St. Johns, 23 Jun 1915, Gorman 3575 (K); Polk County, vacant lot back of Taylor’s, 30 May 1960, Murata 71 (GH); Willamette County, Salem, 16 Jul 1917, Nelson 1684 (GH); Washington County, Hillsboro, 23 Aug 1911, Smith 4093 (GH); Pennsylvania: Northumberland County, Turbotville, 4 mi NNW of town, 13 Aug 1940, Adams 5416 (A); Berks County, McKnight’s Gap, 3/4 mile SW of town, 164 m, 12 Jun 1941, Berkheimer 2565 (GH); Buchs County, Riegelsville, 3 mi below (Kintnerville), 27 Jun 1925, Dreisbach 3573 (LE); Pike County, Kimble, along Lackawanna River, 24 Jun 1936, Fogg 10784 (A); Montgomery County, Jenkintown, 28 Jun 1938, Frazier s.n. (A); Somerset County, Tire Hill, beside Pa. Highway 53, 0.9 km SSE of Tire Hill, 427 m, 9 Aug 1956, Shetler 266 (LE); Fulton County, Needmore, 15 Jun 1947, Wahl 2514 (A); Utah: Tooele County, Open Reventment Area, zone 12, 23 Jul 2000, Long 1090 (GH); Vermont: Bellamy Falls, Anonymous s.n. (K); Leicester, 18 Aug 1907, Dutton s.n. (G); Virginia: Arlington County, Clarendon, Allard’s garden, 10 May 1938, Allard 4653 (GH); Frederick County, Belle Grove, near Middletown, 23 May 1943, Hunnewell 17854 (GH); Rappahannock County, Piney River, 27 Jun 1949, Hunnewell 19218 (GH); Washington: Yakima County, Silah Valley, 29 Aug 1902, Cotton 882 (GH); Skamania County, White Salmon, along Hwy 830 just W of White Salmon, , 10 Aug 1961, Dennis 2323 (A, G, GH); King County, Seattle, 9 Aug 1935, Eyerdam s.n. (BM, G); Spokane County, Chattaroy, 12 Jul 1923, Lackey s.n. (G); Thurston County, Mima Prairie, along the Black River 3 miles south of Littlerock, 22 Jun 1939, Meyer 1650 (GH); Whatcom County, Northwood Swamp, 26 Jun 1937, Muenscher 8394 (GH); Whitman County, Bonnie Lake, north of Rock Lake, 29 May 1938, Sharsmith 3570 (GH); West Virginia: Pendleteon County, Snowy Mountains, east slope, 13 Aug 1931, Core s.n. (GH); Smyth County, Marion, 29 Jun 1892, Small s.n. (K); Sweet Spring, 8 Sep 1903, Steele & Steele 246 (GH); Wisconsin: Dane County, Rice Lake, 19 Jun 1926, Fassett 2783 (GH); Iron County, Mercer, near outlet of Grand Portage Lake, 17 Sep 2000, Nee & Atha 50993 (BM); Brown County, Kellogg’s Work Farm, 6 Aug 1902, Schuette 96 (GH); Wyoming: Fremont County, Lander, along the Popo Agie River just south of city limits; overflow channel of river, 1615 m, 2 Aug 1965, Scott 588 (GH).

Uzbekistan. on the right bank of the Amu-dari [Amu-Darya River] between two rocks, near [Kumksntau], 15 Jul 1873, Karolkov & Krause s.n. (LE); Karakalpakiya, near Amu-dari [Amu-Darya River], 22 Aug 1874, Smirnov s.n. (LE).

15. Solanum dulcamaroides Poir., Encycl. Suppl. 3: 751. 1814

http://species-id.net/wiki/Solanum_dulcamaroides

Figure 38
Solanum macrantherum Dunal, Solan. Syn. 16. 1816, nom. superfl.
Type: Based on Solanum dulcamaroides Poir.
Solanum nutans Sessé & Moc., Fl. Mex., ed. 2, 50. 1894.
Type: Mexico. “urbe Queretaro”, M. Sessé & J. Mociño s.n. (neotype, designated by Knapp, 2008b, pg.18: MA [MA-604674, F neg. 48343]).
Solanum sarmentosum Sessé & Moc., Fl. Mex., ed. 2, 51. 1894.
Type: Mexico. “Habitat in Queretari et Temescaltepec hortis” M. Sessé & J. Mociño s.n. (lectotype, designated by Knapp, 2008b, pg. 18: MA [MA-604621, F neg. 48321]; isolectotypes: G [G00071088, F neg. 34123], MA [MA-604620, F neg. 48317], LE).
Solanum megalospermum J.L.Gentry & Child, Brenesia 16: 147. 1979.
Type: Guatemala. Baja Verapaz: Sierra de las Minas, 5 km S of Purulhá, 1600 m, 2 Jan 1973 [“1963” on label at MO], L.O. Williams, A. Molina R. & T.P. Williams 41964 (holotype: F [F-1986736, F neg. 62207]; isotypes: K [K000488211], MO [MO-2270289], NY [NY00139007]).
Type.

Mexico. Based on an unpublished illustration in the Sessé and Mociño collection (lectotype, designated here: Hunt Botanical Institute 6331.1503); Mexico. Sin. loc., M. Sessé & J. Mociño s.n.. (epitype, designated here: MA [MA-602624]).

Description.

Woody vine, often reaching into the canopy. Stems when young sparsely to densely pubescent with tangled, very weak simple uniseriate trichomes to 0.5 mm long, some trichomes furcate or dendritic in more pubescent individuals; new growth sparsely to densely pubescent with simple and furcate uniseriate trichomes, these tangled and weak, ca. 0.5 mm long. Bark of older stems yellowish brown, glabrate, corky on very large stems (fide Nee 23735). Sympodial units plurifoliate. Leaves usually simple, occasionally pinnatifid, especially on younger stems, (2.5-)4–10 cm long, (1-)2–8 cm wide, elliptic to obovate, usually widest in the basal third, slightly thick and fleshy, the upper surfaces glabrous and shiny with the trichomes confined to the veins or uniformly pubescent on the veins and lamina with simple uniseriate trichomes to 0.5 mm long, the lower surfaces glabrous or the pubescence similar to that of the upper surfaces, but the trichomes denser along the veins; primary veins 6–7 pairs, connected by a prominent marginal vein ca. 0.5 cm from the margin; base acute to truncate or very occasionally somewhat cordate; margins entire or with 1–2 pairs of basal lobes, the lobes 0.5–0.7 cm long, each with a petiolule to 0.4 cm long; apex acuminate, often with an elongate tip; petioles to 5 cm long, glabrous or pubescent like the stems and leaves, the trichomes denser on the channelled adaxial groove, twining to aid in climbing. Inflorescences terminal or lateral, 7–20(+) cm long, longer in fruit, many times branched, with up to 80 flowers, glabrous or pubescent with simple uniseriate and occasionally furcate trichomes like the stems; peduncle 1.5–5 cm long; pedicels 1–1.6 cm long, slender, ca. 1 mm in diameter at the base, 1–1.5 mm in diameter at the apex, deflexed or nodding at anthesis, glabrous or sparsely pubescent with simple uniseriate trichomes adaxially, minutely papillate abaxially, articulated at the base from a tiny sleeve, leaving a small peg on the inflorescence axis; pedicel scars irregularly spaced 5–10 mm apart. Buds globose and inflated with prominent angles at the petal margins, the corolla strongly exserted from the minute calyx tube long before anthesis. Flowers all perfect, 5-merous. Calyx tube 1.5–2 mm long, flattened to somewhat conical, the lobes < 0.5 mm long, forming mere teeth on the rim of the tube, glabrous to sparsely pubescent with simple trichomes, these denser on the minute apices. Corolla (2-)2.5–4 cm in diameter, violet to deep purple, very showy, stellate, lobed ca. 2/3 of the way to the base, the lobes 0.9–1.5 mm long, 0.6–0.7 cm wide, spreading to slightly cupped at anthesis, densely papillate all long the margins and on the cucullate tips, otherwise glabrous. Filament tube ca. 0.5 mm, the free portion of the filaments 1.5–3 mm long, glabrous; anthers 4.5–5 mm long, 3–3.5 mm wide, ellipsoid to almost globose, yellow, loosely connivent, the abaxial surfaces thickened and enlarged and the thecae not visible, drying papillate, poricidal at the tips, the pores not lengthening to slits with age. Ovary glabrous; style 14–16 mm long, glabrous; stigma small capitate, the surface minutely papillose. Fruit a globose berry, 2–2.5 cm in diameter, bright red and juicy when ripe, the pericarp thin and shiny; fruiting pedicels 1.5–2.5 cm long, 1–1.5 mm in diameter at the base, hanging from the weight of the fruit; fruiting calyx a flattened plate. Seeds ca. 20 per berry, 5–6 mm long, 4–5 mm wide, flattened reniform, reddish brown or pale brown, in immature fruit the surfaces minutely pitted, in mature fruits the lateral testal cell walls prominent and giving the seed a hairy appearance, these to 1 mm long and creating a prominent wing around the seed, the testa cells rectangular in outline. Chromosome number: not known.

Figure 38.

Solanum dulcamaroides Poir. (A drawn from Saunders 8220 B drawn from Nee 23837 C drawn from Nee 23735 D–F drawn from Cortés 29 G drawn from Lopez 340). Illustration by Bobbi Angell.

Distribution

(Figure 39). From central Mexico (Estado Colima) to Nicaragua; from almost sea level to 2100 m.

Figure 39.

Distribution of Solanum dulcamaroides Poir.

Ecology.

In deciduous and evergreen forests, often a canopy vine.

Conservation status.

Least Concern (LC); EOO >100, 000 km2 (LC) and AOO >10, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum dulcamaroides is a beautiful plant, with large, fleshy flowers and bright red fruits. It is one of the larger vining members of the Dulcamaroid clade, often growing well into the canopy of rainforests. Although it in some ways resembles Solanum dulcamara (as mentioned by Dunal 1813), it can be easily distinguished from that species by its inflated, globose angular buds (rather than the turbinate buds of Solanum dulcamara) and the much larger flowers (to 4 cm in diameter) without green spots at the corolla lobe bases. It also is somewhat similar morphologically to Solanum boldoense of Cuba (see Knapp 2008b for a discussion of the mix-up associated with these two species in the collections of Sessé and Mociño) and the more widespread and commonly cultivated Solanum seaforthianum. Solanum dulcamaroides differs from both those species in its distinctive anther morphology, with the abaxial surface thickened so that the thecae are not visible and the anthers appear as small football-shaped structures, and with pores that do not markedly lengthen to slits with age. The filaments of Solanum dulcamaroides and Solanum boldoense are equal, while in Solanum seaforthianum one filament is longer than the rest. Solanum boldoense has the articulation point of the pedicels just below the base of the calyx tube, while the pedicels of both Solanum dulcamaroides and Solanum seaforthianum are articulated at the base from a small sleeve. Pinnatifid leaves are rare in flowering or fruiting specimens of Solanum dulcamaroides (see below) and Solanum boldoense, while they are common in Solanum seaforthianum.

Although very few specimens of Solanum dulcamaroides have pinnatifid leaves, Nee (1993) records that divided leaves are borne on lower parts of the plant, and that plants cultivated from seed first produced profoundly pinnatifid leaves and eventually produced only simple leaves once stems became flowering. This developmental transition from pinnatifid juvenile to simple adult leaves appears to be relatively common in the Dulcamaroid clade (see Solanum flaccidum), but is often not noticed as herbarium specimens are usually made only from the terminal portions of reproductive stems. From herbarium specimens, it is apparent that leaf size decreases as the stem makes the transition to the inflorescence.

Leaf pubescence varies considerably in Solanum dulcamaroides, as it does in many other members of the group. Some specimens (especially those from the west of Mexico) are densely pubescent, with some of the trichomes furcate or with a few branches, but the majority of the trichomes are simple. Other plants, among them the type of Solanum megalospermum (Williams et al. 41964), are almost entirely glabrous. Plants from Nicaragua have smaller flowers (to 2.5 cm in diameter) than do plants from Mexico, flowering collections of Solanum dulcamaroides from Guatemala are a priority to see if this is a simple cline or due instead to environmental differences.

The epithet “dulcamaroides” has long been attributed to M.-F. Dunal rather than J.L.M. Poiret, the editor of the supplement to Lamarck’s encyclopaedia (see Nee 1982). While most of the 41 Solanum epithets published in the supplement are indeed attributable to Dunal, Solanum dulcamaroides has the diagnosis ascribed to Dunal, but the epithet was not, and was therefore probably coined by Poiret himself. Dunal later cites “Solanum dulcamaroides Poir.” in synonymy with his Solanum macrantherum, attributing the epithet “dulcamaroides” to Poiret only, thus indicating that Poiret did not use the epithet Dunal had wanted for this plant.

Solanum dulcamaroides (and its later homonym Solanum macrantherum) was described based on an illustration seen by Dunal in the collection of original drawings done for the Sessé and Mociño expedition to Mexico and Central America (see McVaugh 2000) and on his own, unpublished illustration, now held in the collections at MPU. Dunal is likely to have seen this illustration when Mocino was in Montpellier between 1812 and 1817 (see McVaugh 2000: 12); this set of original illustrations was then copied for Candolle, but no images of Solanum species are held in the set of copies at G. The original held in the Hunt Botanical Institute is thus the only material associated with the name. Material linked to this name was described by Sessé and Mociño as Solanum sarmentosum and Solanum nutans, and herbarium sheets linked with those names can be found in MA (see Knapp 2008b), duplicates of these are also held in G. An illustration in the Torner Collection of the Hunt Botanical Institute (accession number 6331.1503, Figure 40) that clearly shows the distinctive bud morphology of this species is here chosen as the lectotype of Solanum dulcamaroides. Epitype material is held in MA amongst the collections of the Sessé and Mociño expedition: I have selected as the epitype the sheet best matching the leaf morphology in the watercolour. Considerable confusion over the identities of Solanum dulcamaroides and the very similar Cuban endemic Solanum boldoense is apparent in the labelling of sheets at MA; this is discussed under Solanum boldoense above and in Knapp (2008b).

Figure 40.

Lectotype of Solanum dulcamaroides Poir. (Torner Collection plate 6331.1503). Reproduced with permission of the Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh, PA. Torner Collection of Sessé and Mociño Biological Illustrations. Rights reserved.

Specimens examined.

Guatemala. Baja Verapaz: Reserva Municipal Los Cerritos, Baja Verapaz, Salamá. Cerro el Portezuelo, Reserva Municipal Los Cerritos, 1020 m, 10 Mar 2009, Christenhusz et al. 5662 (BM); El Quiché: Sacapulas, 1600 m, 12 Jan 1974, Molina R. et al. 30376 (F); Huehuetenango: Sierra de los Cuchumatanes, 15 Sep 1941, Johnston 1978 (F).

Mexico. Chiapas: Ocozocoautla de Espinosa, 32 km N of Ocozocoautla along road to Mal Paso, 762 m, 19 Oct 1965, Breedlove & Raven 13575 (F, MO, US); Motozintla de Mendoza, Huixtla, 45-50 km NE along road to Motozintla, 1900 m, 28 Dec 1972, Breedlove & Thorne 31051 (MO); Siltepec, El Porvenir, 10 km NW, por camino a Siltepec, 2750 m, 17 Aug 1985, Hernéndez & Ramírez H. 901 (MEXU); Chiapa de Corzo, El Chorreadero, 5.6 miles east of Chiapa de Corzo along Mexican Highway 190, 762 m, 15 Jul 1966, Laughlin 1285 (US); Ocozocoautla de Espinosa, Chapapote to Viente Casas, 13 Nov 1964, MacDougall 449s (MEXU); Unión Juárez, en el camino Talquina-Cima de Volcán Tacana, rumbo a la linea divisoria con Guatemala, 1800 m, 20 Oct 1985, Martínez S. 14166 (MEXU); bewteen Mazapa and Motozintla, 19 Jul 1941, Matuda 4812 (F, GH, MEXU); Ocuilapa, 1036 m, 21 Aug 1895, Nelson 3031 (GH, US); Colima: Comala, Rancho El Jabalí, 20 km (airline) N of Colima in the SW foothills of the Volcán de Colima, on main ranch road near the stables, 1400 m, 21 Sep 1991, Sanders et al. 11609 (MEXU); Comala, Rancho El Jabalí, 22 km (ariline) N of Colima in the SW foothills of the Volcán de Colima, Colima/Jalisco line passes through ranch; 100 m from Lago El Epazote on the road from the ranch headquarters, 1400 m, 3 Aug 1991, Vázquez V. 1001 (MO); Guanajuato: Cerro Capulín, western slopes, just E of Mexico 43, ca. 8 km NNE of Uriangato on road to Salamanca, 1900 m, 17 Sep 1977, Iltis & Doebley 111 (F, MEXU); Puerto Nieto, 13 Aug 1947, Kenoyer 2032 (GH); San Miguel de Allende, 28 Nov 1967, Lape 1 (GH); Allende, Microondas Calderón, en el cerro de Alcoer, 2180 m, 12 Aug 1987, Mora Benítez 827 (MEXU); Moroleón, Piñícuaro, 2120 m, 26 Jul 1986, Zamudio 4162 (MEXU); Guerrero: Taxco, 14 Jun 1937, Abbott 210 (GH); Taxco, 12 Aug 1937, Abbott 358 (ECON); Mina, Manchón, 6 Jul 1937, Hinton 10516 (GH, K, US); Hidalgo: Ajacuba, Ejido Santiago Texontlale, camino que va rumbo a la Mesa Chat, cerro al NW del poblado Santiago Textontlale, sierra de Mexe, 2440 m, 14 Sep 1988, Díaz V. 207 (MEXU); Ajacuba, La Peñitas Blancas, NE del panteón de Emiliano Zapata, ladera S de la sierra Chicavasco, ejido San Nicolas Tecomatlán, 2260 m, 23 Aug 1988, Díaz V. & Valverde G. 91207 (MEXU); Metzquititlán, Barranca de Venados, 1800 m, 6 Aug 1979, Hernández M. 3610 (MEXU); Carretera Federal a Pachuca 3 km de la caseta, 9 Jul 1977, Martínez Marino 44 (MEXU); Pueblo Viejo, 9 Jul 1977, Martínez Marino 50 (MEXU); Jalisco: Autlán de Navarro, El Chante to Guisar, Sierra de Manantlán, 2160 m, 17 Aug 1980, Breedlove & Almeda 45756 (MEXU); Zapopan, Arroyo Las Coronillas, 6 km al ONO de Nexitipac, sobre el cauce del arroyo, 1500 m, 9 Aug 2001, Carrilo-Reyes 2479 (MEXU); Tuxpán, Atenquique, 1 km S por carr. a Colima, luego 9 km al O por brecha al Volcán de Colima, 1700 m, 23 Aug 1988, Fuentes O. 546 (MO); Santa Cruz de Astillero, Santa Cruz de Astillero, 1500 m, 12 Aug 1984, Hernández M. & Hernández V. 9537 (MEXU); Tonila, Cerro Alto, en al microondas, 2000 m, 12 Jul 1992, Huerta M. et al. 230 (MEXU); 1.5 km S of Rincón de Manantlán, at very base of Sierra de Manantlán, exactly 13 km due S of El Chante, 1560 m, 9 Oct 1980, Iltis & Guzmén M. 3164 (F, MEXU); Tepatitlán, Aug 1882, Kerber s.n. (A); Cihuatlán, adelante del aserradero, camino a La Cumbre, 1460 m, 15 Jun 1988, López F. et al. 733 (MEXU); Atenquique, along logging road 5 km S of Atenquique, 1829 m, 4 Aug 1965, Mertz 191 (MEXU); brecha Chalapa-Mezcala, 1590 m, 14 Jul 1974, Puga 6739 (MEXU); Tapalpa, km 22 de la brecha Tapalpa-Ve. Carranza, 2100 m, 24 Aug 1986, Rodríguez C. & Suárez J. 591 (MEXU); Michoacán: Morelia, N of Zapote, 4 Aug 1910, Arsène s.n. [5970] (K, L, US); Punguato, vicinity of Morelia, 2100 m, 16 Jul 1909, Arsène 2894 (US); Lago de Pátzcuaro, 1 km al W de San Andres Tizorondaro, 2060 m, 27 Nov 1977, Cabellero & Mapes 40 (MEXU); Cuitzeo, Jéruco, ca. 3 km al WNW, 1890 m, 14 Sep 1999, Carranza & Silva 5824 (MEXU); Lagunillas, Cerro El Águila, 3 km del poblado de Huatzanguio, 2415 m, 15 Aug 2008, Cornejo Tenorio & González Castañeda 2986 (MEXU); Jiquilpan, in and around a stone quarry on Route 110 between Km 25 and ca. 25.5 km W of Jiguilpan, 29 Aug 1966, Cruden 1231 (GH); Erongarícuaro, Oponguio, 2100 m, 5 Jul 1990, Díaz Barriga 6201 (MEXU); 2 km de Pomoca, carretera a Maravatio, 2100 m, 21 Mar 1991, Díaz Barriga et al. 6854 (MA, MEXU); Quiroga, Santa Fe de Laguna, 2200 m, 23 Oct 1985, Escobedo 465 (MEXU); Zitácuaro, Zitácuaro-Loma Larga, 15 May 1938, Hinton 11857 (GH, K, US); Coalcomán, S. Naranjillo, 15 Jul 1939, Hinton et al. 13951 (F, GH, US); Morelia, La Concepción, approx. 500 m al N, 2200 m, 10 Sep 1985, Huerta B. 160 (MEXU); Ruta 15 Morelia a Zitácuaro, ca. 10 km al E de Morelia, 2100 m, 22 Oct 1981, Lorence & Ramamoorthy 3742 (F, MEXU); Lake Cuitzeo, 9 Aug 1892, Pringle 4188 (BM, F, G, GH, GOET, K, LE, MEX, US); Tlazazalca, Tlazazalca, camino a Cerro La Cruz, 1800 m, 5 Jul 1990, Pérez & García 1344 (MEXU); Coeneo, 3 km al NE de Azajo, 2110 m, 14 Jul 1988, Ramos 142 (MEXU); Susupato, La Ziranda, 1650 m, 26 Sep 1989, Rzedowski 49012 (MEXU); Indaparapeo, 4 km al S, sobre el camino a Las Peras, 2100 m, 27 Sep 1989, Rzedowski 49037 (MO); Huaniqueo, Tendeparacua, 3 km al S, 2150 m, 21 Oct 1990, Rzedowski 50418 (MEXU); Alvaro Obregón, El Cerrito, cerca de Tzintzimeo, 1900 m, 21 Jul 1986, Santos Martínez 1579 (MEXU); Huaniqueo, Coeperio, SE del pedregal grande, 1.4 km al NE de Coeperio, 2150 m, 14 Oct 1992, Silva Saenz 406 (MEXU); 17 km al E de Morelia, en la desv. a Union Progreso, 1850 m, 24 Sep 1979, Soto N. & Ramírez S. 1748 (MEXU); Tuxpán, La Providencia, 5.3 km al SO de Malacate, 1750 m, 1 Nov 1989, Torres & Ramírez 13501 (MEXU); Contepec, El Tambor, aprox. 3 km al E de Tuxtepec, 2350 m, 25 Oct 1986, Zamudio R. & Murillo 4971 (MEXU); Morelos: Tepoztlán, bajada en el camino del Parque a Texpotlán, 21 Jul 1940, Miranda 507 (MEXU); Cuernavaca, 10 Oct 1840, Moore s.n. (K); Cerro Tepozteco, 20 km NO de Cuernavaca, 4 Jun 1987, Torres C. & Miller 9690 (F, MEXU); Tepozteco, 1800 m, 1 Oct 1971, Vázquez 3437 (MEXU); México: Temascaltepec, Real de Arriba, 1930 m, 4 Jun 1932, Hinton 835 (K, MA); Temascaltepec, Cerro Muñeca, 2300 m, 15 Aug 1932, Hinton 1360 (BM, F, G, K, MEXU, US); Temascaltepec, Temascaltepec, 1750 m, 18 Sep 1932, Hinton 1697 (BM, G, GH, K, K); Ixtapán de la Sal, Valle de Mexico, 1800 m, 2 Sep 1951, Matuda 20864 (MEXU); 3 km N of Ixtapan de la Sal along Highway 55, 1650 m, 28 Jul 1964, Mick & Roe 344 (BM); Puerta del Gato, 7 km al NW de Zitácuaro, 1950 m, 20 Jul 1964, Rzedowski 18351 (MEXU); Malinalco, carretera Malinalco-Tenancingo, poco despues de Malinalco, cima de cerros, 2180 m, 15 Jul 2001, Vibrans 7366 (MEXU); Oaxaca: San Bartolo, Aug, Andrieux 185 (G-DC, GH, K); Soyaltepec, Presa de Temazcal, a 2 km al N de los vertederos de la presa, 23 Sep 1984, Cabrera & Torres 7255 (MEXU); San Lucas Ojitlán, del poblado de El Zapotal a Mata de Caña, 60 m, 23 Jan 1989, Calzada 14291 (MEXU, MO); San Felipe Usila, Poblado Pas Escalera, senda para la cima del Cerro Paso Escalera, 450 m, 19 Oct 1989, Calzada 14997 (MEXU); Suchixtepec, 19 km al S de San José Pacifico, carretera Miahuatlán-Pochutla, Dto. Miahuatlán, 2180 m, 1 Oct 1988, Campos V. 2457 (MEXU); Presa Miguel Alemán, lado SO de la presa, 13 Feb 1982, Cedillo Trigos & Torres 1031 (MEXU); Cerro San Antonio, 1550 m, 26 Jun 1906, Conzatti 1410 (BH, GH, MEXU); San Miguel Soyaltepec, a 4 km de Temazcal camino a vertador, Dto. Tuxtepec, 60 m, 8 Mar 1986, Cortés et al. 266 (MEXU); San Pedro Teutila, El Faro, 670 m, 19 Jan 2005, Cruz Espinosa 2431 (MEXU); Tuxtepec, 21 miles S on Hwy. 175 to Oaxaca, 40 m, 30 Oct 1980, Fryxell & Lott 3224 (BH, MEXU, MO); San Juan Mixtepec, Dto. Miahuatlán, 2050 m, 13 Jul 1997, Hunn OAX-1493 (MEXU); Santa María Jacatepec, La Joya, Cerro Quemado, 500 m, 6 Nov 1986, López Vargas 363 (MEXU); San Miguel del Puerto, Llano de Horno, limite de los terrenos comunales, Dto. Pochutla, 360 m, 20 Jul 2000, López 340 (MEXU); Quiechapa, cercanias, 23 Oct 1953, MacDougal s.n. (MEXU); Chiltepec, and vicinity, Dto. Tuxtepec, 20 m, Jul 1940, Martínez Calderón 333 (A, MEXU, US); San Miguel Soyaltepec, Cerro Tepezcuintle, Torre 38 de la L.T. Temascal II-Oaxaca Potencia, 203 m, 11 Nov 2004, Martínez Feria & Juárez García 78 (MEXU); San Miguel del Puerto, 30 m de la desviación camino a Santa Catarina Janmixtepec, orilla Río Zimatán, Dto. Pochutla, 450 m, 29 Sep 2001, Pascual 16 (MEXU); San Miguel del Puerto, 3.4 km al oeste del pueste San Lorenzo, sobre la brecha a Monte Carlo, Dto. Pochutla, 690 m, 25 Sep 2001, Salas M. et al. 4001 (MEXU); San Juan del Estado, 18 Jun 1894, Smith s.n. (MEXU); San Pablo, Santa Cruz, 10-12 km al NW de Oaxaca, Distrito de Etla, 1680 m, 2 Nov 1978, Solano & Vara 305 (MEXU, MO); Temascal, 13 Dec 1961, Sousa 1093 (MEXU); Santa María Jacatepec, El Águila, al O de San Agustin, entrando por La Reforma, 28 km al SO de Tuxtepec, 550 m, 19 Jan 1988, Torres C. & Martínez S. 11034 (MEXU); San Pedro Teutila, a 400 m de la brecha a El Faro, 1 km hacia el este de Faro, 571 m, 7 Oct 2002, Velasco Gutiérrez & Juárez 83 (MEXU); Santa Cruz Itundujia, Dto. Putla, La Laguna a 2.88 km en LR (SW) de Santa Cruz Itundujia, 2181 m, 20 Jun 2008, Velasco Gutiérrez et al. 2837 (MEXU); Puebla: Puebla, Jardín del Calvarío, 2175 m, 1 Sep 1907, Arsène 2330 (GH, US); Morelia, environs, 2073 m, Aug 1840, Galeotti 1183 (G); Tehuacán, 29 Jul 1897, Pringle 7496 (BH, F, GH, GOET, K, L, US); El Riego, Jul 1905, Purpus 1285 (F, GH); Cerro de Chicamole, Aug 1909, Purpus 3969 (BM, F, GH, US); Querétaro: km 16-17 camino entre carretera a Mexico y Amealco, 2300 m, 2 Sep 1977, Argüelles 802 (MEXU); Los Cues, 2200 m, 20 Jul 1981, Argüelles 1666 (MEXU); Hacienda Ciervo, near Cadereyta, 26 Aug 1905, Rose et al. 9846 (US); Amealco, Quiotillos, 2200 m, 18 Jul 1989, Rzedowski 48594 (MEXU); Veracruz: Catemaco, ca. 3 km al E de Laguna Catemaco en el camino al Bastonal, 550 m, 11 Aug 1972, Beaman 6436 (F); Orizaba, 10 Sep 1866, Bourgeau 3054 (G, GH, K, L, LE, US); San Andrés Tuxtla, Laguna Encantada, 8 km N de San Andres Tuxtla, 300 m, 26 Jan 1978, Calzada 4223 (F, MEXU); San Andrés Tuxtla, 2 km al norte, 20 Jan 1987, Cedillo Trigos 3780 (MEXU); La Luz près Córdoba, 27 Sep 1882, Kerber 68 (BM, G, GOET, K, LE, US); Catemaco, at highest point on road from Catemaco to Sontecomapán, 5 km N of junction with road around Laguna de Catemaco, 8 km (by air) NE of Catemaco, 510 m, 6 Dec 1981, Nee 23735 (F, MO); Ixtaczoquiatlán, Orizaba, ca. 2 km N of town, south of old (non-cuota) road between Fortín and Orizaba, 1000 m, 7 Dec 1981, Nee 23837 (BH, F, GH, MEXU, MO); Jilotepec, Vista Hermosa, 1200 m, 20 Oct 1977, Ortega O. 726 (MEXU); San Andrés Tuxtla, Estación Biológica Las Tuxtlas, camino Corte Ruiz Cortinez, lote 71, 160 m, 3 Oct 1986, Sinaca Colin 1016 (MEXU); Catemaco, Cerro de Buenavista, N de Catemaco, 490 m, 3 Oct 1998, Torres R. & Campos V. 46 (MEXU); Jilotepec, Rincón del Muerto, 1400 m, 26 Apr 1971, Ventura A. 3505 (F); La Concepción, 4 km de la desviacion carretera Concepción, 1050 m, 22 Jul 1976, Zola B. 552 (F, MEXU).

Nicaragua. Estelí: Mesas Moropotente, ca 16.0 km (by road) NE of Hwy. 1 at Estelí, 1310 m, 11 Jun 1981, Henrich & Stevens 454 (BM); Estelí, km 144.5, Fundacion CECALII, 782 m, 21 Jun 2001, Rueda et al. 16497 (MO).

16. Solanum endoadenium Bitter, Repert. Spec. Nov. Regni Veg. 12: 546. 1913

http://species-id.net/wiki/Solanum_endoadenium

Figure 41
Solanum endoadenium Bitter var. robustius Bitter, Repert. Spec. Nov. Regni Veg. 12: 547. 1913.
Type: Argentina. Catamarca: Cuesta de Muschaca, F. Schickendantz 285 (holotype: B, destroyed; lectotype, designated here: CORD [CORD00004206]).
Solanum incurvipilum Bitter, Repert. Spec. Nov. Regni Veg. 12: 158. 1913.
Type: Argentina. Salta: El Paso “Al Alizar” [Alizal?], 2400-2600 m, between Pampa Grande, 1740 m and Cerro Cristal, 2610-1700 m, E. Nelson in F. Kurtz 12512 (holotype: S [S04-2919]; isotype: CORD [CORD00004229]).
Type.

Argentina. La Rioja: Pie de Cuesta, above Vallecito, Sierra Famatina, G.H.E.W. Hieronymus & G. Niederlein 746 (lectotype, designated by Morton 1976, pg. 88: G [G00070235]; isolectotypes: CORD [CORD00004199, CORD00004200], LE, P [P00335096]).

Description.

Shrubs 1–1.5 m tall. Stems erect, sparsely to densely pubescent with patent simple uniseriate trichomes ca. 0.5 mm long, often gland-tipped and the plants viscous; new growth densely pubescent. Bark of older stems yellowish brown, the leaf bases prominent. Sympodial units plurifoliate. Leaves simple, 1.8–6 cm long, 0.5–2 cm wide, lanceolate to more or less linear, slightly fleshy, the upper surfaces uniformly pubescent with sparse to dense simple uniseriate trichomes ca. 0.5 mm long, sometimes gland-tipped, the lower surfaces variable from almost glabrous with the simple trichomes confined to the veins to densely and uniformly pubescent with simple uniseriate trichomes ca. 0.5 mm long; primary veins 6–9 pairs, drying yellowish red; base attenuate; margins entire, densely pubescent; apex acute to acuminate, the ultimate tip rounded; petioles 0.5–1 cm long, sparsely to densely pubescent with simple trichomes like those of the stems, usually not twining. Inflorescences terminal or becoming lateral, 2–4 cm long, simple to several times branched, with 5–30 flowers, glabrous to densely pubescent with simple uniseriate trichomes like the stems; peduncle 1–3 cm long; pedicels 0.6–0.8 cm long, filiform, < 0.5 mm in diameter at the base and apex, erect to somewhat spreading, glabrous to pubescent with simple uniseriate trichomes, articulated at the base from a small sleeve, leaving a small peg on the inflorescence axis; pedicel scars irregularly spaced 1–4 mm apart. Buds globose to slightly ellipsoid, the corolla exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 1–1.5 mm long, conical, the lobes 1–1.5 mm long, triangular, sparsely to densely pubescent with simple uniseriate trichomes. Corolla 0.7–1 cm in diameter, dark bluish purple to pale violet with green spots at the base of the lobes, stellate, lobed 1/2 to 2/3 of the way to the base, the lobes 3–3.5 mm long, 2–2.5 mm wide, spreading or perhaps cupped, densely pubescent all along the tips and margins with simple trichomes < 0.2 mm long. Filament tube minute, the free portion of the filaments ca. 0.5 mm long, glabrous or minutely pubescent with glandular trichomes; anthers 1.5–2 mm long, ca. 1 mm wide, ellipsoid, loosely connivent, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 4–5 mm long, glabrous or occasionally glandular in the lower half; stigma minutely capitate, the surface minutely papillose. Fruit a globose berry, 0.6–0.8 cm in diameter, orange when ripe, the pericarp thin and shiny, glabrous; fruiting pedicels 1.2–2 cm long, ca. 1 mm in diameter at the base, not markedly woody, nodding or spreading. Seeds 12–14 per berry, 3.5–4 mm long, 3–3.5 mm wide, flattened reniform, reddish brown, the surfaces minutely pitted, the testal cells very small, rectangular to square. Chromosome number: n=12 (Moscone 1992).

Figure 41.

Solanum endoadenium Bitter. (A, C–G drawn from Hunziker et al. 25550 B Legname & Vervoorst 154). Illustration by Bobbi Angell.

Distribution

(Figure 42). Eastern Andean slopes in central to northern Argentina and adjacent Bolivia, from 1500–3000 m.

Figure 42.

Distribution of Solanum endoadenium Bitter.

Ecology.

Forests and open areas above treeline; sometimes growing in sandy soil in puna vegetation.

Conservation status.

Least Concern (LC); EOO >100, 000 km2 (LC) and AOO >10, 000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Unlike many of the members of the Dulcamaroid clade, Solanum endoadenium is a shrubby species and is apparently never a vine or even scandent. The leaves are consistently simple, and apparently never lobed or pinnatifid. Solanum endoadenium is easily recognised by its purple flowers, orange fruits and dense, often glandular-viscid pubescence of simple trichomes. The older stems are warty from the persistent leaf bases; this distinctive character distinguishes Solanum endoadenium from the somewhat similar Solanum salicifolium from the same region. Solanum endoadenium is further easily distinguished from Solanum salicifolium by its larger, usually branched inflorescences and more spreading pubescence (rather than appressed-ascending).

Leaf size in Solanum endoadenium is quite variable, and varies within as well as between individuals. In general leaves near the stem tips are smaller than those lower down. Bitter (1913) stated that the epithet was derived from the glandular pubescence on the inside of the filaments. The berry of Solanum endoadenium stays on the plant a long time, and apparently cracks to release the seeds (see Figure 6D); whether this is always the case is not clear.

Morton (1976) lectotypified Solanum endoadenium with the only syntype (of seven in Bitter’s original description) from La Rioja province, as Bitter (1913) stated he was basing his description on material from La Rioja. In his lectotypification, Morton cites the collection as Hieronymus & Neiderlein 746, but the only specimen at G [G00070235] has no collection number, nor do the duplicates at LE and P. Morton’s annotation slip on this G sheet states “isosyntypus” and is dated 1961; he may have made a transcription error later, as this is the only specimen at G that bears this locality and collector information. I have therefore assumed he meant this sheet in his lectotypification and have accepted his designation of the G sheet as the lectotype. I have found no duplicates of Schickendantz 285 (the type collection cited in the protologue of var. robustius) in any of the herbaria where other Schickendantz duplicates have been found (GOET, SI); Bitter cited “herb. Hieronymous” in the original description (probably at B and now destroyed).

Specimens examined.

Argentina. Catamarca: Belén, al incio de la Cuesta de Randolfo, 3009 m, 31 Jan 2008, Barboza et al. 1997 (CORD); Belén, próximo a Quebrada de Randolfo rumbo a Nacimientos de San Antonio, 12 Feb 2012, Barboza et al. 3476 (BM, CORD); Pomán, Sierra de Ambato, falda oeste, Mutquin, en Valle Muerto, 4 Feb 1910, Castillón 1613 (CORD); Andalgalá, Quebrada del Río Pisavil, 26 Nov 1948, Filipovich 66 (CORD); Ambato, Cumbres de Narváez, falda oeste, Ruta 62, km inmediaciones de Las Chacritas, entre Narváez y Singuil, 1850 m, 10 Dec 1965, Hunziker et al. 18555 (CORD); Ambato, Sierra de Ambato, falda E, subiendo desde El Rodeo hacia el Cerro Manchado, 2300 m, 23 Feb 1967, Hunziker 19075 (CORD); Pomán, Sierra de Ambato, falda oeste, subiendo desde El Rincón hacia Las Casitas, rumbo al Cerro Manchado, 2300 m, 18 Feb 1970, Hunziker & Ariza 20349 (CORD); Ambato, Cu