Corresponding author: Alexander P. Sukhorukov (
Academic editor: E. Roalson
The former
Sukhorukov AP, Nilova MV, Krinitsina AA, Zaika MA, Erst AS, Shepherd KA (2018) Molecular phylogenetic data and seed coat anatomy resolve the generic position of some critical Chenopodioideae (Chenopodiaceae – Amaranthaceae) with reduced perianth segments. PhytoKeys 109: 103–128.
The family
Some of the most recent and drastic taxonomic changes have been proposed by
Further changes were subsequently proposed by
The recent split of
It has become apparent in recent years that fruit and seed characters are also useful in distinguishing members of the former
Amongst the species of the former
To resolve this issue, we have included these two species, in addition to several accessions of taxa sampled for the first time [
Several new taxa were included in the phylogenetic analysis for the first time:
Voucher information and GenBank accession numbers for the species of
Species | Old names (if applicable) | GenBank accession number | |||
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Total genomic DNA was extracted from herbarium samples according to
PCRs for two chloroplast markers (
Primers and cycler programmes used for the molecular analysis.
Marker | Primer sequences and combination | Reference | Cycler programmer |
---|---|---|---|
ITS | ITS5 5'-GGA AGT AAA AGT CGT AAC AAG G-3' |
|
95 °C for 5 min, 33 cycles of amplification (95 °C for 15 s, 55 °C for 30 s, 72 °C for 40 s), 72 °C for 5 min |
ITS4 5'-TCC TCC GCT TAT TGA TAT GC-3' | |||
rbcLaF 5'- ATG TCA CCA CAA ACA GAG ACT AAA GC-3' |
|
95 °C for 5 min, 35 cycles of amplification (95 °C for 10 s, 55 °C for 30 s, 72 °C for 40 s), 72 °C for 5 min | |
rbcLaR 5'-GTA AAA TCA AGT CCA CCR CG-3' |
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atpB-rbcL F 5'-GAA GTA GTA GGA TTG ATT CTC-3' |
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95 °C for 5 min, 35 cycles of amplification (95 °C for 20 s, 56 °C for 30 s, 72 °C for 60 s), 95 °C for 20 s, 56 °C for 80 s, 72 °C for 8 min | |
atpB-rbcL R 5'-CAA CAC TTG CTT TAG TCT CTG-3' | |||
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Tab C 5'-CGA AAT CGG TAG ACG CTA CG-3' |
|
95 °C for 5 min, 35 cycles of amplification (95 C for 1 min, 50 °C – 65 °C (increasing in 0.3 C per cycle) for 1 min, 72 °C for 4 min), 72 °C for 5 min |
Tab D 5'-GGG GAT AGA GGG ACT TGA AC-3' | |||
Tab E 5'- GGT TCA AGT CCC TCT ATC CCC-3' | |||
Tab F 5'ATI' TGA ACT GGT GAC ACG AG 3' |
Sequencing was performed following Sanger methods on an Applied Biosystems 3730 DNA Analyser using ABI PRISM BigDye Terminator v. 3.1 (Center of Collective Use “Genome”, Institute of Molecular Biology, Moscow, Russia). The sequencing primers were the same as the amplification primers.
The raw forward and reverse sequences were checked and combined in BioEdit sequence alignment editor v. 7.0.5.3 (
To show the relationships between taxa, we reconstructed various phylogenies using Bayesian analysis, maximum likelihood (
The carpology of the tribe
The cross-sections of the seeds were prepared using a rotary microtome Microm HM 355S (Thermo Fisher Scientific, USA) and then examined using a Nikon Eclipse Ci (Nikon Corporation, Japan) light microscope and photographed using a Nikon DS-Vi1 camera (Nikon Corporation, Japan) at the Department of Higher Plants, Lomonosov Moscow State University. Before sectioning, the seeds were soaked in water:alcohol:glycerine (1:1:1) solution, dehydrated in ethanol dilution series and embedded in the Technovit 7100 resin (Heraeus Kulzer, Germany).
The phylogenetic analysis based on nrDNA (ITS) and combined cpDNA analyses (
Best tree from the BEAST analysis of the ITS
In the ITS analysis (Figure
Like the ITS phylogenetic analysis, the combined
Best tree from the BEAST analysis of the combined
This study highlighted the fact that these species, with the exception of
Pericarp of
Cross-section of the seed of
Cross-section of the seed of
Additional noteworthy characters evolved in
Taxon/Character | Life history | Perianth segments | Cells of the outer pericarp layer | Pericarp adherence to the seed coat | Seed shape and colour | Seed surface | Seed keel | Thickness of seed-coat testa (µm) | Acicular outgrowths of the testa cells | Presence of spatial heterospermy | Seed embryo position |
---|---|---|---|---|---|---|---|---|---|---|---|
|
short-lived perennial herb | basally connate | spongy | scraped off the seed | roundish, red | alveolate | – | 12–20 | – | – | vertical |
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annual | free | not spongy | easily ruptured | roundish, red | undulate | + | 7–10 | – | – | vertical |
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annual or short-lived perennial herb | basally connate | not spongy | hardly removed | roundish, red | alveolate, with hairy-like outgrowths | – | 17–25 | + | – | vertical |
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perennial herb | basally connate | spongy | scraped off the seed | roundish, red | smooth | – | 37–45 | – | + | vertical, rarely horizontal |
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perennial herb | basally connate | spongy | scraped off the seed | roundish, red | alveolate | – | 25–30 and 37–45 (heterospermous) | – | + | vertical |
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annual or short-lived perennial herb | basally connate | not spongy | hardly removed | ovate, red | undulate | + (two keels and a groove between them) | 12–15 | – | + | vertical |
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annual or short-lived perennial herb | connate to 1/3 | not spongy | hardly removed | ovate, red | undulate | + (two keels and a groove between them | 15–18 | – | + | vertical |
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annual or short-lived perennial herb | almost free | not spongy | hardly removed | ovate, red | undulate | + (two keels and a groove between them) | 10–12 | – | – | vertical |
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annual or short-lived perennial herb | basally connate | not spongy | hardly removed | ovate, red | alveolate | + (two keels and a groove between them) | 10–12 | – | – | vertical |
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annual | free, or perianth absent | not spongy | hardly removed | roundish, red | alveolate, with hairy-like outgrowths | – | 8–10 | + | – | vertical |
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annual or short-lived perennial herb | basally connate | not spongy | hardly removed | ovate, red | alveolate | + (two keels and a groove between them) | 15–17 | – | – | vertical |
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annual or short-lived perennial herb | basally connate | not spongy | hardly removed | ovate, red | undulate | + (two keels and a groove between them) | 10–12 | – | + | vertical |
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annual | fused in almost all flowers, free only in some flowers | not spongy | easily ruptured | roundish, red | minutely pitted | – | 10–15 | + | vertical and horizontal | |
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annual | basally connate | not spongy | easily ruptured | roundish, red | minutely pitted | – | 10–15 and 17–25 (heterospermous) | + | + | vertical and horizontal |
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annual | basally connate | not spongy | hardly removed | roundish, red | smooth (minutely pitted) | + (one keel) | 12–15 | + | – | vertical |
|
annual | connate to the middle or almost to the top | spongy | scraped off the seed | roundish, red | reticulate with minutely pitted dots | – | 12–20 | + | – | vertical and horizontal |
|
annual | basally connate | not spongy | easily ruptured | roundish, red | reticulate with minutely pitted dots | – | 20–25 | + | + | vertical and horizontal |
|
annual | basally connate | not spongy | scraped off the seed | roundish, red | minutely pitted | + (one keel) | 12–15 | + | – | horizontal, rarely vertical |
|
annual | basally connate | not spongy | easily ruptured | roundish, red | reticulate with minutely pitted dots | – | 10–15 | + | vertical and horizontal | |
|
annual | basally connate | papillate | scraped off the seed | roundish, black | minutely pitted | – | 42–50 | + | – | horizontal |
The phylogenetic position of
The importance of morphological characters used to delineate species within the genus
Habit of
In the absence of molecular phylogenetic data, it is clear that carpological characters must be taken into consideration when determining the generic placement of taxa in either
This species was initially described as
Shoot of
Recently,
Annual, glabrous, branched or not; lateral branches if present ascending; leaves cauline (rosulate leaves absent), densely located, spatulate-oblong, with a short petiole up to 1 cm or sessile, entire; inflorescence leafy (bracts similar to stem leaves); flowers sessile or shortly pedicellate, unisexual intermixed in small glomerules (Figure
As
The type specimen lodged at GH contains several plants collected from different areas in California and almost all of them were collected after the description of
South-western North America (USA, North Mexico).
The new generic name is composed by the prefix “neo” (new) and the core name
In the
We thank Eric H. Roalson and anonymous reviewers for the comments on the previous draft of the paper and Igor Pospelov and Steve Matson for the excellent images of