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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. 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. Full descriptions and synonymies including designations of lectotypes or neotypespreliminary conservation assessments, illustrations, distribution maps, and an extensive list of Guararema brusque online dating are provided for all species.
With approximately species J. 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. 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. This treatment is part of this collaborative effort.
The major clades of Solanum Guararema brusque online dating Bohs For revisions of individual species published on-line see Solanaceae Source http: Dunaldivided the genus into two major groups, Inermia unarmed solanums and Aculeata armed solanumsbased 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. Dunal maintained as separate the genera Lycopersicon Mill. Danertused 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 but recognised them at different ranks.
He recognised 21 sections with numerous subsections, and listed component taxa in each. Child and Lester provided a synopsis of infrageneric classification of Solanumbased largely on the work of Bitter and Seithe They did not indicate component species of any of their recognised sections, which comprise fewer species than those of Nee Child and Andropedas Rusby plus a variety of other taxa see Bohs for discussion.
Most current 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. 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. The Dulcamaroid clade as recognised by Bohs and treated here is comprised of elements from several previously recognised subgenera and sections of Solanum.
The common European woody nightshade or bittersweet, Solanum dulcamara L. It was the only member of this group described by Linnaeuswho recognised its extreme variability but did not explicitly name any of these variants, although he did explicitly name variants of other highly variable taxa.
This polynomial refers to the South African species Solanum africanum Mill. Dunal had only seen herbarium specimens or live plants of three of these taxa Solanum dulcamaraSolanum 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 were scattered over three groups based on leaf division.
It is clear that the species now recognised here as members of the Dulcamaroid clade were in Guararema brusque online dating groups due to their highly variable leaf morphology that can be pinnatifid to simple on a single stem see below.
He included Solanum viscosissimum Sendtn. Difficulty in grouping these taxa which today we recognise as closely related is perhaps understandable as label data from 19 th century herbarium specimens is sparse and usually does not include any mention of habit. Seithe 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 a section; Seithe was unsure as to the affinities of these taxa. The rest of the species were included in Guararema brusque online dating sections Dulcamara e. Those taxa included in Anthoresis are all the members of the Solanum nitidum species group sensu Knapp and were grouped together based on the possession of highly branched trichomes.
He pointed out the extreme morphological similarity between members of his section Holophyllum and Brevantherum Seithe; species in both these groups have dichasial branching, but they differ in their trichome morphology. Species such as Solanum pulverulentum Pers. Knapp studied the members of 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. Child established subsection Nitidum A. Child based on this monograph Knappbut did not suggest further relationships or examine material in detail. Child also segregated the Californian species Solanum xanti A. In his treatment Solanum in the New World Nee 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 His circumscription of section Holophylla also includes all of the members of the Geminata clade as treated by Knapp All the vining species plus other species previously recognised as section Parasolanum Bitter, e.
Although the two groups are morphologically quite similar, in that both contain
Guararema brusque online dating that climb with the aid of petioles see belowsequence 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 ; Weese and Bohs 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. The unexpected relationship of the Solanum species group Knapp with the vining dulcamaroid solanums was revealed in analyses of DNA sequence data Bohs and Olmstead ; Bohs ; Weese and Bohs The monophyly of the Dulcamaroid clade as recognised here had previously never been suggested.
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.
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 Although the structure of relationships within the Dulcamaroid clade is very poorly resolved and the taxa sampled relatively few, a
Guararema brusque online dating sister relationships are apparent within the clade.
In the ndhF analysis Bohs five groups form a polytomy: 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 nitidumSolanum pubigerum Guararema brusque online dating Solanum aligerumbut not Solanum Guararema brusque online dating. Relationships amongst the vining dulcamaroid species are less clear, but additional sampling will certainly help with the delimitation of smaller groups within the clade.
Guararema brusque online dating of these relationships are Guararema brusque online dating poorly supported in both analyses, so any sister taxon relationships should be regarded as preliminary. Members of the Dulcamaroid clade are all woody plants and vary from shrubs or lax shrubs to vines see Figure 1. 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. Habit of members of the Dulcamaroid clade. 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 ; T.
Darwin 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. Danert, and Child and Lester documented sympodial units and anthoclades patterns of foliar lateral branches and associated inflorescences in the Solanaceae. The Guararema brusque online dating of leaves per sympodium is very regular and of major taxonomic significance in the entire genus Solanum see Knapp 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.
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 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 Members of the Solanum nitidum are either monochasial or some species have dichasial branching see Knapp ; in species with dichasial branching two axillary shoots develop and the branching is dichotomous e.
Most of these species have a mixture of the two branching types but are predominantly one or the other. The leaves of members of the Dulcamaroid clade are extremely variable, both across the group Guararema brusque online dating within individual plants see Figure 2 and individual species illustrations. 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.
Leaf variation in members of the Dulcamaroid clade. A Leaf variation along a single short stem, Solanum salicifolium Barboza Guararema brusque online dating al.
Trichome types found in members of the Dulcamaroid clade. Leaf polymorphism is common in the group and in many species individual stems bear both simple and pinnatifid leaves e. 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 viscosissimumlateral 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 ; 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 pinnatifidbut 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.
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