Chenopodium betaceum, Andrzejowski, 1862

Mosyakin, Sergei L., 2017, Notes on taxonomy and nomenclature of Chenopodium acerifolium and C. betaceum (C. strictum auct.) (Chenopodiaceae), Phytotaxa 324 (2), pp. 139-154 : 144-146

Chenopodium betaceum and the problem of C. strictum

It was commonly assumed by most authors (except Beaugé 1974, Gervais 1979, and a few others) that the name Chenopodium strictum was validly published in 1821 by Roth (1821: 180). However, this name was first validated in the volume no. 6 (published August–December 1920: see IPNI 2017 –onward) of Caroli a Linné ... Systema Vegetabilium … (ed. 15 bis, volumes 1–4 by Roemer & Schultes) (Schultes 1920: 264). Schultes ascribed the name to Roth, provided the Latin description, the provenance and collector (“ H. [Habitat] in India orientali. B. Heyne ”), highlighting that the description was provided in Roth’s manuscript (“ Roth nov. pl. Spec. MSS ”). Finally he compared the new species with C. lanceolatum Muehlenberg ex Willdenow (1809: 291) by characters of leaf shape and with C. album by the inflorescence structure (“Quoad foliorum figuram proxime accedit ad Ch. lanceolatum Mühlenb. ; quoad inflorescentiam vero ad Ch. album ”). Roth (1821) provided an updated and expanded description of the species. It seems that the earlier date and place of valid publication of the name C. strictum (1820, not 1821) was first emphasized, at least in the 20 th century, by Beaugé (1974) in his thorough historical overview of C. album and related taxa.

The name Chenopodium strictum Roth (in Schultes 1820: 264) is usually applied now to a tetraploid species (or, in a wide sense, to a group of species and/or infraspecific taxa) of the C. album (sensu latissimo) aggregate ( Aellen & Just 1943, Aellen 1960–1961, Uotila & Suominen 1976, Uotila 1977, 1997, 2001a, 2001b, Skripnik 1987, Lomonosova 1992, Mosyakin 1996, 2003, 2012, Mosyakin & Fedoronchuk 1999, Clemants & Mosyakin 2003, Zhu et al. 2003, Iamonico 2010, Feodorova 2014, Sukhorukov 2014 etc.). Before 1929, if and when this species was accepted, it was usually mentioned in European publications under the name C. striatum ( Krašan 1894: 255) Murr (1896: 32) . However, Aellen (1929a) suggested that C. striatum is conspecific with C. strictum . He provided a few arguments in favor of his decision, but cited the opinion of Murr (1904), who commented that one of the species names listed by Moquin-Tandon (1849), for example, C. strictum or a doubtful species C. virgatum 2 Thunberg (1815: 143), might be “an older synonym” of his taxon (recognized by Murr in 1904 as a subspecies of C. album ): “Es wird sich übrigens noch darum handeln festzustellen, ob Ch. striatum (Kraš.) nicht noch in einer der von Moquin-Tandon aufgeführten Arten, z. B. in sp. 6 Ch. strictum Roth oder in der zweifelharten sp. 62 Ch. virgatum Thunb. ein älteres Synonym besitzt” ( Murr 1904: 223). However, on the same page (in a footnote) Murr also compared his C. album subsp. striatum to a North American taxon now usually accepted as a variety of C. berlandieri ( Clemants & Mosyakin 2003) : “Von den nordamerikanischen Formen kommt dem Ch. striatum das Ch. Boscianum Moq. mit länglich eiförmigen, grünen Blattern, stark rotstreifigen Stengeln und sehr glänzenden Samen nahe, teilt aber die Neigung zur Mikrophyllie mit der folgenden ssp.” [the next subspecies was “ssp. glomerulosum Rchb. ” sensu Murr]; translation: “Of the North American forms, Ch. striatum is approached by Ch. Boscianum Moq. with oblong-ovate green leaves, strongly redstriped stems and very shining seeds, but [the latter] shares the tendency to microphylly with the following ssp.” [ “ssp. glomerulosum Rchb. ” sensu Murr—S.M.]).

Aellen’s decision to synonymize C. striatum with C. strictum was soon accepted in many standard European floras and other publications (see e.g., Iljin & Aellen 1936: 64–65, Mansfeld 1939: 102), and the name C. strictum is now widely applied (in a wide or strict sense) to the Eurasian tetraploids related to the hexaploid C. album (see references above).

Several segregate tetraploid species are also sometimes recognized in this group, such as C. striatiforme Murr (1901: 51) [≡ C. strictum Roth subsp. striatiforme (Murr) Uotila (1977: 199) ], C. glaucophyllum Aellen (1929b: 155) [≡ C. strictum subsp. glaucophyllum (Aellen) Aellen in Aellen & Just (1943: 67) ≡ C. strictum var. glaucophyllum (Aellen) Wahl (1954: 38) ], and C. novopokrovskyanum (Aellen) Uotila (1993: 192) [≡ C. album subsp. novopokrovskyanum Aellen (1938: 3) ]. Dvořák (1984a, 1984b, 1989, 1993 etc.) recognized in Europe several other segregate species and supposed hybrids, but his extreme “splitter’s” approach was followed by only a few botanists.

However, the application of the name C. strictum to a rather widespread Eurasian species (and probably to its relatives in North America, if they are indeed native there) was soon considered at least questionable. Iljin & Aellen (1936: 65) already noted that the plants of C. strictum occurring in the former USSR differ from typical Indian plants in having terminal spicate partial inflorescences arranged in a pyramidal general inflorescence. Because of that they proposed the new combination C. strictum subsp. striatum (Krašan) Aellen & Iljin (in Iljin & Aellen 1936: 65) [see also Brenan (1964: 94), who accepted this taxon as C. album subsp. striatum (Krašan) Murr (1904: 222) ].

2 The taxonomic identity of C. virgatum Thunb. (described from Japan) still remains insufficiently understood. Judging from the digital images (including close-ups of the inflorescence provided by Mats Hjertson) of the holotype (UPS, Herb. Thunberg 6444), the plant is definitely not a member of Chenopodium sect. Acuminata Ignatov (1988: 18) and is not conspecific with C. vachellii Hooker & Arnott (1838: 269) because it lacks the characteristic multicellular tubular hairs on inflorescence branches (see also discussion in Mosyakin & Iamonico 2017). Instead, it belongs to the Chenopodium album aggregate, as it was suggested by Aellen (1960–1961), but is morphologically different from the species accepted here as C. betaceum . In recent publications (e.g., Uotila 2001b, Sukhorukov 2014) C. virgatum is recognized as a separate species distinct from both C. album and “ C. strictum ”.

Mosyakin (1996: 41–42), judging from the available type fragment and the protologue, emphasized that the name C. strictum may in fact belong to one of the taxa or hybrids of the C. album aggregate and because of that it is quite possible that the correct name for the East and Central European species is C. betaceum , which is definitely applicable to the European plants. Sukhorukov (2014) noted that the morphological diversity of Indian taxa belonging to the C. album aggregate is very different from the patterns observed in Europe and also concluded that it is quite possible that “for European populations the priority name might be C. betaceum ” [in Russian: “Не исключено, что длЯ европейских популЯций приоритетным наЗванием может быть C. betaceum ” (Sukhorukov 2014: 229) ]. Dvořák (1989, 1993), who preferred to treat C. strictum , C. striatum , C. striatiforme and several other taxa as separate species and interspecific hybrids, assumed that C. strictum “is a hybridogenous species in whose genotype there participate C. album L. <…>, C. diversifolium (Aellen) Dvořák , C. eustriatum Dvořák and C. striatiforme J. Murr ” ( Dvořák 1989: 198). Dvořák (1986) added a note on the herbarium specimen GZU-000273164 (see below): “ Chenopodium striatum (Krašan) J. Murr ist nicht identisch mit Chenopodium strictum Roth ” (see JSTOR Global Plants 2017–onward).

The type of Chenopodium strictum (the standing lectotype, designated by Dvořák 1989: 201; a fragment of a specimen that was originally in B but was destroyed in 1943 during World War II) has a much-branched cymosepaniculate inflorescence superficially similar to inflorescences of C. pedunculare Bertoloni (1837: 32 ; see also Dvořák 1984c, Walter 1995, Paśnik 1999) or C. lanceolatum , which are hexaploid taxa (“microspecies”, or infraspecific entities, or just morphotypes?) of the C. album aggregate. A very similar much-branched cymose inflorescence is also peculiar to the original Linnaean specimen (LINN 313.9) of C. viride Linnaeus (1753: 219) ; this Linnaean name is also considered to be applicable to a form of the C. album aggregate (see Uotila 1978) but in the past it was often misapplied to the diploid species now known as C. suecicum Murr (1902: 341) . In contrast, type specimens of C. betaceum and C. striatum , as well as most of other European and western Asian specimens usually identified as “ C. strictum ”, have linear (“spicate”) inflorescences, normally with moniliform arrangement of glomerules.

Inflorescences similar to those of the type of C. strictum also occur in some other Indian plants, e.g. in C. purpurascens Jacquin [1776–1777: 43, tab. 80; lectotype designated by Dvořák 1994: 127 (Fig. 10), 128]. It is worth noticing that Murr (1904: 223) cited the name “ C. purpurascens β [var.] lanceolatum Moq. -Tand.” ( Moquin-Tandon 1849: 67) in synomymy of his C. album subsp. striatum . In turn, Chenopodium purpurascens was sometimes considered (most probably erroneously) as conspecific with C. giganteum D. Don ( Don 1825: 75). However, Sukhorukov & Kushunina (2014: 10, 18) claimed that “ C. bengalense seems to be an older name for the taxa known as C. giganteum or C. album subsp. amaranticolor ”. The lectotype of C. bengalense (designated by Sukhorukov & Kushunina 2014: 18; Fig. 5 on page 19) also has upper leaves very similar in shape to those of the type of C. strictum . The standing lectotype of C. strictum in its inflorescence branching habit is also more similar to the lectotype of C. giganteum (designated by Sukhorukov & Kushunina 2014: 18) than to European plants. At least two additional species described from India ( Pandeya et al. 1998, Pandeya & Pandeya 2003) may be also related to C. strictum , but their real identity remains obscure because their published descriptions and illustrations are insufficient for positive identification.

Sukhorukov & Kushunina (2014: 18) cited the combination C. bengalense as validated in 1821. However, in 1821 the name “ Chenopodium bengalense . Spielm.” ( Steudel 1821: 92) was cited only in synonymy of the accepted name Atriplex bengalensis : the name in Chenopodium was thus not validly published (Art. 36.1 of ICN, McNeill et al. 2012). Steudel (1840) accepted the name C. bengalense (as “ benghalense ”) in the 2 nd edition of his Nomenclator, so the name should be cited as C. bengalense (Lam.) Spielm. ex Steudel (1840: 348) [ Atriplex bengalensis Lamarck (1783: 276) ].

Despite several relevant publications (e.g., Aellen 1929c, Beaugé 1974, Dvořák 1992, 1994, Sukhorukov & Kushunina 2014), the taxonomy and identity of the mentioned presumably native Indian and/or SoutheastAsian taxa of Chenopodium remain highly problematic, and further studies are necessary. The real identity of C. strictum (which at present remains nomen dubium) also remains obscure and will be probably clarified after in-depth studies of Chenopodium from India and adjacent regions.

Thus, the inflorescence branching pattern and leaf shape observed in the type of Chenopodium strictum are not typical for C. betaceum or most other plants of Eurasian “ C. strictum ”. It should be also emphasized that most of Eurasian plants of C. betaceum (incl. C. striatum ) have striate stems and branches, with stripes of deep beet-red color (in old herbarium specimens the stripes often become dark olive green). This prominent feature, which is reflected in the epithets “ betaceum ” (similar to Beta in its color) and “ striatum ” (striped), is not well manifested in the plant fragments of the standing lectotype of C. strictum . The absence of mature fruits in that lectotype also hampers its precise identification. In contrast, the original specimen (lectotype) of C. betaceum reported below leaves no doubt that it is conspecific with C. striatum and not conspecific with C. strictum , as it has been already concluded by Iljin.

In his treatment of Chenopodium for the Flora of the Ukrainian SSR, Iljin (1952: 306–308) accepted C. betaceum and placed C. striatum in its synonymy. He commented that C. betaceum is a “good southern species of goosefoot” [in Ukrainian: “Добрий південний вид лободи…” ( Iljin 1952: 308)] that was only recently recognized in Western Europe under the name C. striatum ; he further commented that “now, however, as demonstrated by the study of Andrzejowski’s herbarium kept in Kiev, the priority in establishing and recognizing this species should be credited to that author, who described it under the name C. betaceum Andrz. already in 1862” [In Ukrainian: “…тепер, Як покаЗало вивченнЯ гербарію АндржіЄвського, що ЗберігаЄтьсЯ в КиЄві, пріоритет на встановленнЯ і виділеннЯ цього виду треба Залишити За цим автором, Який описав його під наЗвою C. betaceum Andrz. ще в 1862 р.” ( Iljin 1952: 308)]. He also mentioned that European plants differ from Indian C. strictum mainly in inflorescence characters. Iljin further suggested that C. betaceum is probably not a native European plant and hypothesized that it may have originated in the eastern parts of the “Ancient Mediterranean region”. In Soviet historical biogeography that term, as outlined by Popov (1963, reprinted in 1983) (ОбЛасть ДревнегО СредиЗемьЯ in Russian, which can be also translated as the “Ancient Middle-Earth” because the term СредиЗемье but not СредиЗемнОмОрье was used), covers not only the Mediterranean area but also the Irano-Turanian region, the zone that can be also called the Tethyan area (of the ancient Tethys and Paratethys oceans). It means that Iljin (l.c.) placed the hypothetical native (or ancestral?) range of C. betaceum somewhere in a wide zone stretching from the Eastern Mediterranean to Central Asia.

Historical and geographical considerations also indicate that the Indian plant originally described by Roth as C. strictum is not identical with the European and western and Central Asian plants since the time of publication of Aellen’s article ( Aellen 1929a) and until recently commonly called “ C. strictum ”. Benjamin Heyne (1770–1819) worked and collected plants mainly in the southern regions of India ( Heyne 1814, Burkill 1953, Stewart 1982, Kochhar 2013). He arrived to Tranquebar ( Tharangambadi, Tamil Nadu state) in 1793 and then visited and/or stayed in Madras (Chennai, Tamil Nadu), Bangalore (Bengaluru, Karnataka state), Mysore (Mysuru, Karnataka) and some other places ( Kochhar 2013: 2), traveled “from Cuddapa [Kadapa (formerly known as Cuddapah), Andhra Pradesh state] to Hydrabad [Hyderabad, Andhra Pradesh]” and “from Bengalore [Bangalore or Bengaluru, Karnataka] to Trichinopoly [Tiruchirappalli, Tamil Nadu]” (see Heyne 1814: 461–462), and died in Madras in 1819. It is documented that in 1813 Heyne “passed on a considerable number of plant specimens to the German botanist Albrecht William [Wilhelm —S.M.] Roth (1757–1834), who published an account of 200 of them in Novae plantarum species praesertium Indiae Orientalis ” ( Kochhar 2013: 12). For almost all new taxa described by Roth (1821), no geographical information has been provided except “ India Orientalis” (meaning India in the sense of the [British] East India Company, as opposed to the West Indies of the New World, and not to be interpreted as “eastern parts” of India): “there is scarcely a single precise record of locality from back to back of the book” ( Burkill 1953: 859). We anyway can thus safely assume that the specimens of Chenopodium provided by Heyne to Roth were collected somewhere in southern India. However, I am not aware of any reliable records or herbarium specimens from southern India that are referable to Eurasian plants now usually called “ C. strictum ” (following Aellen 1929a). If such plants (“ C. strictum ” sensu Aellen and many other authors who followed him) indeed occur in India, they are most probably restricted to the northern regions of the country, closer to the border with Pakistan (see Uotila 2001a). It is worth noticing that C. strictum (in either sense) is not reported in recent treatments of Chenopodiaceae of Nepal ( Sukhorukov & Kushunina 2014) and India ( Paul 2012); however, in the latter publication it is definitely one of several glaring omissions. Thus, historical and geographical data also testify (although indirectly) that the type of C. strictum (in the strict sense) collected by Heyne and provided by him to Roth most probably represents some local southern Indian taxon of the C. album aggregate, but not the plant called “ C. strictum ” in recent literature.

In my opinion, the best solution would be to stop the misapplication of the name C. strictum to European and some Asian tetraploid plants and to restore the name C. betaceum , as it was proposed by Iljin (1952; see above), similar to what happened with the restored (also by Iljin) and now almost universally accepted name C. acerifolium .Alternatively, one might argue that C. strictum was a name used widely since 1929 and until recently, and because of that its recent usage might deserve to be preserved, e.g. through its conservation with a conserved neotype. However, cases of similar abandonment of previously widely accepted names happened in Chenopodium many times. For example, the Linnaean names C. viride and C. serotinum Linnaeus (1756: 12) were commonly misapplied before for the taxa now properly known as C. suecicum Murr and C. ficifolium Smith (1800: 276) , respectively. Thus, the name C. strictum should again join a long list of obscure names of uncertain taxonomic identity that were proposed in Chenopodium in the 18 th and 19 th centuries and which are now not in current use, such as C. paganum Reichenbach (1832: 579) , C. patulum Roth (1821: 181) , and many others (for more examples and a historical overview of some of those names, see Beaugé 1974).