ACTA BOT. CROAT. 81 (2), 2022 121 Acta Bot. Croat. 81 (2), 121–128, 2022 CODEN: ABCRA 25 DOI: 10.37427/botcro-2022-009 ISSN 0365-0588 eISSN 1847-8476 Blitum venetum (Chenopodiaceae), a new species from the north-eastern Dolomites (Italian Eastern Alps) Duilio Iamonico1*, Katia Sciuto2,3, Adriano Sfriso2, Carlo Argenti4, Marion A. Wolf2,3 1 University of Rome Sapienza, Ce.R.S.I.Te.S., via XXIV Maggio 7, 04100-Latina, Italy 2 Ca’Foscari University of Venice, Department of Environmental Sciences, Informatics and Statistics, via Torino 155, 30172-Venezia Mestre, Italy 3 University of Ferrara, Department of Chemical, Pharmaceutical and Agricultural Sciences, via Luigi Borsari 46, 44121-Ferrara, Italy 4 Pietriboni street 7, 32100-Belluno, Italy Abstract – A new species, Blitum venetum Iamonico, Argenti, Sciuto & Wolf is described from the Dolomites Massif (North-Eastern Italy) on the basis of molecular analyses (nuclear ITS and plastid trnL-F regions) and morphological investigation. The new species is similar to B. bonus-henricus and B. californicum, but it differs from these taxa by characters of leaves (width, pubescence, and margins), inflorescence (presence of bracts), perianth segments (colour), and seeds (diameter, colour, and seed testa ornamentation). Moreover, in phylogenetic reconstructions B. venetum results as clearly separated from the other species of the genus; this finding was further strengthened by the nucleotide divergences calculated between the sequence of the type specimen and other Blitum taxa, which are comparable with the interspecific divergences calculated inside this genus. Keywords: Blitum, Blitum bonus-henricus, Blitum californicum, ITS, taxonomy, tnrL-F. Introduction The genus Blitum L. [tribe Anserineae Dumort., Cheno- podiaceae Vent. sensu Hernández-Ledesma et al. 2015; some authors, e.g. APG IV (2016) consider Chenopodiaceae as included in Amaranthaceae Juss.], traditionally included in Chenopodium L. s.l. during the last centuries (with a few exceptions), was recently taxonomically resurrected by Fuentes-Bazan et al. (2012a, b) based on phylogenetic stud- ies of Chenopodioideae Burnett, where a reclassification of the whole subfamily was proposed recognizing four tribes and ten genera. The generic classification outlined in Fuentes- Bazan et al. (2012b) has been updated in many recent pub- lications (e.g., Iamonico 2012, Mosyakin 2013, Uotila 2011, 2017, Sukhorukov et al. 2013, Sukhorukov and Kushunina 2014) and generally accepted (e.g., Hernández-Ledesma et al. 2015; some online floras and databases, e.g., French flora; POWO 2021-onward). Blitum (incl. Agathophytum Moq., Monolepis Schrad., Scleroblitum Ulbr., Carocarpidium S.C.Sand. & G.L.Chu) is currently known to include about 12 species and nothospe- cies (see Fuentes-Bazan et al. 2012b, Mosyakin 2013, Hernández-Ledesma et al. 2015, Sukhorukov et al. 2018), ten of which occur in the Northern Hemisphere [B. asiaticum (Fisch. & C.A.Mey) S.Fuentes, Uotila & Borsch (= Monolepis asiatica Fisch. & C.A.Mey.), B. bonus-henricus (L.) Rchb., B. californicum S.Watson, B. capitatum L. s.l., B. korshinskyi Litv., B. litwinowii (Paulsen) S.Fuentes, Uotila & Borsch, B. nuttalianum Schult. (= Monolepis nuttalliana (Schult.) Greene), B. petiolare Link (= Chenopodium exsuccum (C.Loscos) Uotila), B. ×tkalcsicsii (H.Melzer) Mosyakin, B. virgatum L. (= Chenopodium foliosum Asch.)], one in South America (B. antarcticum Hook. f.) and one [B. atriplicinum F.Muell. (= Scleroblitum atriplicinum (F.Muell.) Ulbr.)] in Australia. As part of the ongoing study on Chenopodium s.l. (e.g., Iamonico 2010, 2011, 2012, 2013, 2014, Iamonico and Jarvis 2012, Mosyakin and Iamonico 2017), a population belong- ing to the genus Blitum was found in the Dolomites Massif (North-Eastern Italy). After a detailed analysis of pertinent literature, examination of specimens in herbaria and in vir- tual herbaria online (type material included), and molecular investigation, we reached the conclusion that the population * Corresponding author e-mail: d.iamonico@yahoo.it IAMONICO D., SCIUTO K., SFRISO A., ARGENTI C., WOLF M. A. 122 ACTA BOT. CROAT. 81 (2), 2022 found cannot be identified as belonging to any known spe- cies of Blitum. Consequently, we describe here a species new for science and compare it with the related taxa. Materials and methods Morphological and nomenclatural study The work is based on field surveys, analysis of relevant literature (protologues of the names involved in the present study are included), and examination of specimens pre- served in the following herbaria: BR, GH, K, LINN, NY, P, RO, SD, UC, and US (codes according to Thiers 2022). Pho- tographs of inflorescence and fruits were made using an op- tical stereoscope Wild Heerbrugg M8. Molecular study Genomic DNA was extracted using the Genomic DNA purification kit (Thermo Scientific™, Waltham, MA, USA). The nuclear ITS region and the plastid trnL-F region were amplified following Fuentes-Bazan et al. (2012a). The ob- tained PCR products were cleaned using the HT ExoSAP-IT (Applied Biosystems™, Waltham, MA, USA) and sequencing was carried out at the Eurofins Genomics Sequencing Ser- vice (Germany), with the same primers employed in the am- plification reactions. For sequencing of the trnL-F locus, two internal primers were also employed as reported in Fuentes-Bazan et al. (2012a). The GeneStudio sequence analysis software (http://genestudio.com/) was used to as- semble the final consensus sequences. The new sequences were deposited in the International Nucleotide Sequence Database Collaboration (INSDC) repositories, through the European Nucleotide Archive (ENA) platform, with the fol- lowing GenBank accession numbers: OU753550 (ITS re- gion) and OU753551 (trnL-F region). The obtained sequences were compared with those pres- ent in the INSDC archives using the BLAST program avail- able at the USA National Center for Biotechnology Infor- mation (NCBI) web server (http://www.ncbi.nlm.nih.gov). Two distinct datasets of ITS and trnL-F sequences were cre- ated, including the newly obtained sequences and other suitable sequences available in the INSDC repositories, fol- lowing the most recent classifications for the genus Blitum (Fuentes-Bazan et al. 2012b, Sukhorukov et al. 2018). For each analysis, a sequence of Chenopodium pallidicaule Aellen (INSDC accession number: HE577438 for the ITS locus; INSDC accession number: HE577573 for the trnL-F locus) was used as outgroup to orient the tree. Multiple se- quence alignments were generated with MUSCLE (Edgar 2004) and used for phylogenetic analyses. The ITS multi- alignment included 15 sequences for a total of 507 aligned positions and the trnL-F multialignment included 13 se- quences for a total of 987 aligned positions. Phylogenetic analyses were performed with MEGA v. X program (Kumar et al. 2018) using the Maximum Parsimo- ny (MP) and the Maximum Likelihood (ML) methods. For ML, the model that best fit the data, found with the “Find best DNA Models” tool implemented in MEGA v. X under the BIC criterion (Schwarz 1978), was T92 + G for both the molecular markers. Nonparametric bootstrap re-sampling (Felsenstein 1985) was performed to test the robustness of the obtained topologies (1000 replicates). Bayesian Inference (BI) analyses were carried out with MrBayes version 3.1.2 (Ronquist and Huelsenbeck 2003). The analyses included two independent MCMC runs, each composed of four chains (three heated and one cold); each MCMC ran for 1 × 106 generations, sampling trees every 100 generations. The sampling of the posterior distribution was considered to be adequate if the average standard deviation of split frequen- cies was ≤ 0.01. The first 2500 trees were discarded as burn- in, as determined by stationarity of log likelihood assessed using Tracer version 1.5 (Rambaut and Drummond 2007). The consensus topology and posterior probability values were then calculated from the remaining trees. The final tree pictures were created with Inkscape v. 0.92 (Figs. 1a, 1b). Alignments of the ITS and trnL-F sequences of different Blitum species were also obtained with MUSCLE (Edgar 2004) to calculate the percent identities within the genus (Figs. 1c, 1d). Results Morphology The genus Blitum is morphologically characterized by having stems often unbranched or nearly so, or mainly branched at or near the base, leaves often forming basal ro- settes, stigma lobes 2–4, and seed vertical with the seed coat displaying a visible protoplast (Fuentes-Bazan et al. 2012b, Iamonico 2013, Sukhorukov et al. 2018). The genus is rath- er heterogeneous morphologically (Clemants and Mosyakin 2003, Zhu et al. 2003, Sukhorukov et al. 2018), the plants we found in the Dolomites Massif (North-Eastern Italy, Veneto region) appear to resemble both B. bonus-henricus L. and B. californicum S.Watson. In particular, they show the follow- ing combination of characters: perennial (other Blitum s pecies, except perennial B. bonus-henricus and B. californicum, are usually annual), erect to ascending [B. asiaticum (Fisch. & C.A.Mey.) S.Fuentes, Uotila & Borsch, B. litwinowii (Paulsen) S.Fuentes, Uotila & Borsch, and B. nuttalianum Schult. have stems prostrate], basal rosette with many leaves [B. asiaticum and B. nuttalianum have just 1 or 2 leaves, that may wither away completely by anthesis], large and more or less triangular leaves with their base hastate or cordate, more than 5–6 cm long [the other species have leaves more or less rhomboid, with base cuneate, and blades usually up to 5–6 cm long (B. virgatum L. and B. capitatum L. have leaves triangular and up to 8 cm long, but they are annual species)], perianth with 5 segments (other species of the genus usually have 1–3 segments) not becoming red and succulent at the fruiting stage (in B. virgatum and B. capitatum perianth segments are fleshy and red at maturity, forming a berry-like structure). BLITUM VENETUM SP. NOV. ACTA BOT. CROAT. 81 (2), 2022 123 In comparison with Blitum bonus-henricus (native in Europe, introduced in North America) and B. californicum [endemic to California and Mexico (Baja California)], the Italian population differs morphologically by both vegeta- tive and generative characters, i.e. size, pubescence and leaf margins, colour of perianths segments, structure of the in- florescence, and size, colour, and surface of seeds (Tab. 1; see also the diagnosis under the taxonomic treatment). Molecular data In both phylogenetic reconstructions, those based on the ITS and trnL-F markers (Figs. 1a, 1b), the sequences ob- tained from the type specimen collected in the Belluno Do- lomites (Italy) resulted clearly separated from the other spe- cies of Blitum. In the ITS tree (Fig. 1a) our sequence was included in a well-supported clade (100/100/1.00) with a se- Fig. 1. Phylogenetic reconstructions obtained with ML method. Molecular analyses based on the ITS region (a) and molecular analyses based on the trnL-F region (b). For each node, the support values from ML bootstrap, MP bootstrap and BI posterior probabilities are reported, repectively. Only bootstrap supports ≥ 50% and posterior probabilities ≥ 0.70 are shown. Values for nodes that obtained sup- port in only one of the phylogenetic analyses were omitted. For each of the downloaded sequences, the species name, followed by the INSDC accession number between parentheses, is reported. The new species is highlighted with a grey box. Scale bar represents ex- pected number of nucleotide substitutions per site. (c) Percent identity matrix (507 aligned positions) of the taxa belonging to the genus Blitum based on analysis of ITS region. (d) Percent identity matrix (987 aligned positions) of the taxa belonging to the genus Blitum based on analysis of trnL-F region. a) b) c) d) IAMONICO D., SCIUTO K., SFRISO A., ARGENTI C., WOLF M. A. 124 ACTA BOT. CROAT. 81 (2), 2022 quence of B. bonus-henricus from Austria (HE577372). This phylogenetic relationship was not confirmed by the phylo- genetic analysis based on the plastid spacer trnL-F (Fig. 1b), where the Belluno specimen and B. bonus-henricus were placed again as sister taxa, but without any statistical sup- port. On the basis of the percent identity matrices obtained with MUSCLE (Figs. 1c, 1d), the nucleotide divergences cal- culated between the Blitum sp. specimen from Belluno and each of the other recognized Blitum species ranged from 0.40% (Blitum sp. vs. B. bonus-henricus) to 4.37% (Blitum sp. vs. B. californicum) for the nucleotide ITS marker and from 0.86% (Blitum sp. vs. B. bonus-henricus) to 3.70% ( Blitum sp. vs. B. petiolare) for the plastid trnL-F marker. The interspecific divergences calculated between the ITS sequences of the currently taxonomically accepted species of this genus ranged from 0.20% [Blitum nuttalianum vs. B. asiaticum; B. capitatum subsp. capitatum vs. B. capitatum subsp. hastatum (Rydb.) Mosyakin; B. capitatum vs. B. litwinowii; B. capitatum vs. B. virgatum subsp. montanum (Uotila) S.Fuentes, Uotila & Borsch] to 3.98% (B. californicum vs. B. petiolare; B. californicum vs. B. antarcticum). The ITS sequence of B. litwinowii and that of the subspecies B. virgatum subsp. montanum did not show divergences. For the plastid trnL-F spacer, the interspecific divergences ranged from 0.21% (B. litwinowii vs. B. virgatum subsp. virgatum) to 4.47% (B. bonus-henricus vs. B. petiolare). Therefore, for both analysed markers, the nucleotide di- vergences calculated between the sequence of Blitum sp. from Belluno and other Blitum species are comparable with the interspecific divergences calculated inside this genus. Discussion The results obtained in the present study, both morpho- logical and molecular data, support the description of the population from the Belluno Dolomites as a new species, Blitum venetum, which is formally described below. Blitum venetum can be reliably distinguished from the closely related species B. bonus-henricus and B. californicus by various vegetative and generative morphological charac- ters. All diagnostic characters found for B. venetum have high taxonomic value in the classification of both Blitum and the intrageneric taxa of Chenopodium s.l. (Mosyakin and Clemants 1996, Clemants and Mosyakin 2003, Zhu et al. 2003, Sukhorukov et al. 2018). Its recognition as a dis- Tab. 1. Morphological comparison among Blitum bonus-henricus, B. californicum, and B. venetum. Differential characters are in bold. Blitum bonus-henricus Blitum californicum Blitum venetum Stem Erect to ascending, often unbranched, 20–70 cm, glabrous (sometimes farinose) Erect to ascending, often unbranched, 25–80 cm, glabrous (sometimes farinose) Erect to ascending, branched, 40–80(–90) cm, glabrous and sparsely pubescent in the inflorescence part Leaves Green, adaxially sparsely farinose, non-aromatic, petioled (petiole 5–20 cm), blade triangular, 5–15 × 3–10 cm (size decreasing along the stem upwards), base truncate to hastate or cordate; entire and undulate margins, apex usually acute Green, adaxially sparsely farinose, non-aromatic, petioled (petiole 1–12 cm), blade triangular, 4–10 × 3–9 cm, (size decreasing along the stem upwards), base truncate to cordate; margins dentate (in larger leaves up to 10 teeth, teeth acute to acuminate, 0.3–1.0 cm long), apex acute to acuminate Green, glabrous, non-aromatic, petioled (petiole 1–10 cm), blade triangular, 10–13 × 11–13 cm (size decreasing along the stem upwards), base truncate to hastate or cordate; margins dentate (in larger leaves 6–8 teeth, teeth acuminate, 1.0–2.5 cm long), apex acute to acuminate Inflorescence Terminal and lateral spiciform inflorescences ebracteate; spikes 5–20 cm long, the terminal one usually longer than the lateral ones; glomerules about 3–5 mm in diameter Terminal spiciform inflorescence ebracteate; spikes 5–20 cm long; glomerules about 3–5 mm in diameter Terminal and lateral spiciform structures; each glomerule subtended by a linear bract (1.0–2.5 mm long); some lower glomerules subtendend by the upper cauline leaves; spikes 6–8 cm long, the terminal one longer than the lateral ones; glomerules about 2 mm in diameter Flowers Perianth segments connate up to the half, lobes 5, oblong to elliptic, 0.5–1.1 × 0.8–1.5 mm, scarcely keeled through- out, apex obtuse to rounded, glabrous, green throughout; stamens 4–5; stigmas 2(–4) Perianth segments connate up 1/2–2/3, lobes 5, oblong to elliptic, 0.6–1.0 × 0.4–1.2 mm, scarcely keeled throughout, apex obtuse to rounded, glabrous, green throughout; stamens 4–5; stigmas 2 Perianth segments connate up to the 1/3 of the total length, lobes 4–5, ovate, about 0.5 × 1.0 mm, scarcely keeled throughout, apex rounded, glabrous, green with reddish apical part; stamens 4–5; stigmas 2 Fruits Achenes obovoid, 1.5–2.2 mm; pericarp adherent. Achenes obovoid, 1.5–2.0 mm; pericarp adherent. Achenes obovoid, 1.5–2.0 mm; pericarp adherent. Seeds Seeds vertical, obovoid, 1.5–2.0 mm in diameter, margins usually not rounded; seed coat red, smooth Seeds vertical, obovoid, 1.5–2.0 mm in diameter, margins rounded; seed coat red Seeds vertical, obovoid, about 1 mm in diameter, margins rounded; seed coat blackish, alveolate. BLITUM VENETUM SP. NOV. ACTA BOT. CROAT. 81 (2), 2022 125 tinct species is further supported by its phylogenetic posi- tion in both nuclear ITS and plastid trnL-F trees, as well as considering, for both of the analysed markers, the nucleo- tide divergences among various Blitum species. Blitum venetum Iamonico, Argenti, Sciuto & Wolf, sp. nov. Holotype. Italy, Veneto region, Belluno administrative Province, Vigo di Cadore Municipality, locality Casera Razzo, 46°28’41.8”N, 12°36’33.3”E, 1740 m a.s.l., nitrophi- lous grasslands, 11 August 2020, C. Argenti s.n.; holotype: RO! (Fig. 2). Diagnosis (Fig. 3) – Blitum venetum differs from B. bonus- henricus and B. californicum in having the leaves wider (11–13 cm vs. 3–10 cm and 3–9 cm, respectively), gla- brous (vs. adaxially sparsely farinose), with dentate and not undulate margins (vs. entire and/or undulate margins), the perianth segments with red or reddish terminal parts (vs. green throughout), the spike-like inflorescences bracteate, each floral glomerule subtended by a linear bract (vs. spici- form inflorescences ebracteate or nearly so), and the seed smaller (about 1 mm in diameter vs. usually 1.5–2.0 mm). In comparison with B. bonus-henricus, B. venetum also dif- fers by the colour and surface of the seeds (blackish and al- Fig. 2. Holotype of Blitum venetum (RO). IAMONICO D., SCIUTO K., SFRISO A., ARGENTI C., WOLF M. A. 126 ACTA BOT. CROAT. 81 (2), 2022 veolate vs. red and smooth; also B. californicum has reddish seeds). Molecular vouchers. OU753550 (ITS region) and OU753551 (trnL-F region). Description. Stems erect to ascending, branched, 40– 80(–90) cm, glabrous; sparsely inflated hairs in the inflores- cence part. Leaves green, non-aromatic, blades glabrous; petiole 1–10 cm; blade triangular, 10–13 × 11–13 cm (size decreasing along the stem upwards), base truncate to has- tate or cordate; margins dentate (in larger leaves with 6–8 teeth, teeth acuminate, 1.0–2.5 cm long), apex acute to acu- minate. Inflorescences: glomerules mostly densely arranged in terminal and lateral spiciform structures, each subtended by a glabrous leaf-like bract (0.5–3.0 × 2–5 cm); the upper- most bracts with attenuate bases (edges of the leaf base forming an angle of about 90°), with two basal teeth (trian- gular) or with entire margins (lanceolate); each glomerule subtended by a linear bract (1.0–2.5 mm long), with mem- branous hyaline borders; some lower glomerules solitary and sessile, subtendend by the upper cauline leaves; spikes 6–8 cm long, the terminal one longer than the lateral ones; glomerules about 2 mm in diameter. Flowers: perianth seg- ments connate up to the 1/3 of the total length, lobes 4 or 5, ovate, about 0.5 × 1.0 mm, scarcely keeled throughout, apex rounded, glabrous, green with reddish apical part; stamens 4 or 5; stigmas 2. Achenes obovoid, 1.5–2.0 mm; pericarp adherent. Seeds vertical, obovoid, about 1 mm in diameter, margins rounded; seed coat blackish, alveolate. Etymology. The specific epiteth is dedicated to the Ital- ian region (Veneto) in which the new species occurs. Proposed vernacular name. Blito del Veneto (Italian), Veneto’s Goosefoot (English). Flowering time. August to September. Distribution area and habitat. Blitum venetum is known only from locus classicus, i.e., locality Casera Razzo (Bel- luno administrative Province, Veneto region, North-East- ern Italy), where it grows on nitrophilous grasslands at 1740 m a.s.l. The locus classicus is part of the territory of the Do- lomiti Bellunesi National Park (date of establishment: 1988) and of the UNESCO World Heritage Site “The Dolomites” (date of inscription: 2009; see http://whc.unesco.org/en/ list/1237). IUCN Conservation assessment. Only one population was found (locus classicus) and it occupies a small area (less than 50 m2). On the basis of criterion B3 of IUCN (2019), Blitum venetum is here assessed as Critically Endangered (CE). Additional specimens examined Blitum bonus-henricus. Habitat in Europae ruderatis, Herbarium Linnaeus no. 313.1 (LINN, lectotype). Belgium: Bastogne, 1884, A. Verheggen s.n. (P01089431); Theux, 18 Fig. 3. Blitum venetum. A – habitat, B – young individual, C – details of the terminal inflorescence, D – seed. Scale bars: B – 3 cm, C – 3 mm, D – 0.5 mm. Photo: A-B – C. Argenti, C-D – D. Iamonico. BLITUM VENETUM SP. NOV. ACTA BOT. CROAT. 81 (2), 2022 127 June 1959, J. Pelgrims 753 (BR0000027710821V). France: Or- cières, village Merlette, 1830 m a.s.l. 13 July 1980, G. Sag 986 (P00036524). Italy: Lazio region, Montagna d. Duchessa (Velino), June 1946, B. Anzalone 10739 (RO); Lazio region, Piani di Livata al M. Autore, 21 July 1966, B. Anzalone 10737 (RO); Lazio region, Monti della Laga (Amatrice – RI), Fos- so di Selva Grande, 1350-1500 m a.s.l., 6 July 1997, B. Anza- lone 10734 (RO!); Trentino-Alto Adige, San Martino di Cas- trozza, August 1958, B. Anzalone s.n. (RO). Netherlands: Epen, July 1954, W. Voss s.n. (BR0000026765815V). Slove- nia, Jesenice, 1200 m a.s.l., 18 August 1976, G. Sag 986 (P00041309). Sweden: Uppland, Bondkyrka parish, about 1½ km southwest of Berthåga, 18 September 1956, C. G. Alm 2972 (US03654843). Switzerland: Wegkant to Davos Platz, July 1975, W. Verbeke s.n. (BR0000025468366V). U.S.A.: New York, Long Island, East Hampton, 10 June 1877, E. S. Miller 321 (US03652612). Blitum californicum. Mexico: Baja California, Ensena- da, Sierra San Pedro Martir. Colony under live oak, Los Encinos, 10 August 1969, R. Moran 16539 (UC1361814). U.S.A.: California, San Diego county, in rocks at base of cliff ½ mile a southeast of Eel Point, 28 January 1967, R. M. Beauchamp 134 (SD00017034); San Diego county, west side of San Felipe Wildlife Area, under shrubs and mesquite, 26 March 1997, S. Bell 248 (SD); San Buena Ventura, March 1861, W. H. Brewer 244 (GH00936570, syntype, image available at https://plants.jstor.org/stable/10.5555/al. ap.specimen.gh00936570); Fort Tejon, 1857-1859, J. Xantus de Vesey 104 (GH00936572, syntype, image available at https://plants.jstor.org/stable/10.5555/al.ap.specimen. gh00936572); s.d., J. C. Fremont s.n. (NY00008484, syn- type, image available at http://sweetgum.nybg.org/science/ vh/specimen-details/?irn=217758); s.d., D. Douglas s.n. (K000898461, syntype, image available at http://apps.kew. org/herbcat/getImage.do?imageBarcode=K000898461). Acknowledgements Thanks are due to the Directors and Curators of the her- baria cited. References Angiosperm Phylogeny Group, Chase, M.W., Christenhusz, M.J.M, Fay, M.F., Byng, J.W., Judd, W.S., Soltis, D.E., Mabberley, D.J., Sennikov, A.N., Soltis, P.S., Stevens, P.F. (APG IV), 2016: An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of Linnean Society 181, 1–20. Clemants, S.E., Mosyakin, S.L., 2003: Chenopodium L. 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