Taxonomic & Nomenclatural History
History
Both Cyperus and Cyperus papyrus were published in the first edition of Species Plantarum. Linnaeus typically did not include a description for C. papyrus, however he subsequently added one in the second edition.
Willdenow in 1812 published Papyrus antiquorum Willd., a species he differentiated from C. papyrus. This was significant in the recognition of variation within C. papyrus sensu L. because subsequent authors have recognised this taxon to fall within the overall delineation of C. papyrus, and it has persisted through all the subsequent monographs of this species, until 1999 when the WCSP view that it is a synonym of the type ssp. C. papyrus L. ssp. papyrus.
Parlatore was the first to explicitly subdivide C. papyrus, creating in addition C. syriacus Parl. in 1854, to accommodate plants in Palestine and Sicily, on morphological characters. His publication was not accessed for this study and subsequent authors do not report the characters, and nor do they record the weight given to geographic location in the delineation. From this point, therefore, the relationship between taxonomic delineation and spatial distribution is unresolved. All subsequent treatments of the infraspecies rely upon morphology, but also reflect geographic separation.
Chiovenda in Il papiro in Italia (1931), describes new species of Cyperus, later to become, in large part the basionyms for the infraspecies recognised by Kükenthal and others. That he was subdividing Linnaeus’ C. papyrus was clearly known to him, since, he places C. mossambicensis Klotzsch in the synonymy of his new C. nyassicus (albeit with a question mark), along with synonymising other previously published names of C. papyrus pro parte. Therefore, the choice of species as the appropriate taxonomic rank was a deliberate one. His taxa are those later recombined by Kükenthal, with the exception of C. papyrus ssp. madagascariensis (Willd.) Kük (C. madagascariensis Willd.), which he omits. This is odd, since he cites Willdenow’s work, when he, for example, synonymises Papyrus antiquorum (Willd.), and therefore could be expected to have read Willdenow’s account of P. madagascariensis, published in the same work.
The characters on which he divides C. papyrus s.l. are several, and summarised here:
i) Anther connective tissue – exserted or not;
ii) Anther connective tissue – shape of exsertion;
iii) Glume outline shape;
iv) Spikelet length;
v) Rhachilla wings blunt or acute;
vi) Glume apex acute or rounded;
vii) Raceme condensed or lax;
viii) Racemes sessile, sub-sessile or not;
ix) Floret shape; and
x) Style length compared to ovary.
Two species he published from single herbarium specimens, C. elapsus and C. panormitanus. Both are dismissed by later authors (Kükenthal, 1936, Tournay, 1950) as being insufficiently known, and possibly extinct. C. panormitanus is certainly of Sicilian distribution (Palermo), the second species from this island, with C. papyrus. He is not clear about C. elapsus, however the specimen is from the herbarium in Florence and may well have been a third Sicilian segregate species.
Kükenthal, in his monograph of 1936, still the most recent one, largely accepts Chiovenda’s species and descriptions, while commenting both that Chiovenda had limited African material available - a significant proportion of the specimens Chiovenda cites are Italian - and that some of his characters are idiosyncratic. The shape of the spikelet wings relied upon by Chiovenda was considered particularly inconsistent by Kükenthal. Kükenthal emended Linnaeus’ brief and sparse species description. Kükenthal cites Kunth as the author responsible for recombining Papyrus madagascariensis Willd. as C. madagascariensis, and relies on Kunth’s description for this taxon.
Notably, Kükenthal wrote that C. papyrus s.l is introduced in Europe, he describes its centre of origin as the area of the While Nile (Sudan) from where it spread west through central Africa and south to Madagascar and Zimbabwe. He gives 13° N as the Northern limit of its natural range, and 26° S as the Southern. Therefore, he states, it was introduced to Egypt and Italy for the purposes of paper production. This casts doubt on the emphasis Chiovenda and others have placed upon the Italian material of papyrus and the belief that it represented one, two or more distinct taxa.
Kükenthal, like Tournay later, refers to C. antiquorum var. palaestinae Chiov. Chiovenda clearly separates this variety from ssp. antiquorum, giving it separate diagrams, however, he appears to neglect to formally publish it, it is not in his list of described species, nor is it described nor included in his key. He lists specimens from Palestine under C. antiquorum. Kükenthal refers to the list of figures given by Chiovenda, (Chiovenda 1931, 119), and the illustrations it refers to, as the protologue for this variety.
Tournay (1950) accepts Chiovenda’s and Kükenthal’s treatments largely, but adds his own ssp. niliacus var. niliacus. He established this name to replace Chiovenda’s C. antiquorum (Willd.), as adopted and recombined to C. papyrus ssp. antiquorum (Willd.) Chiov. by Kükenthal. His view was that Willdenow only had Sicilian material and, therefore, was not describing an Egyptian-Sudanese taxon, furthermore that this name was synonymous with the type. However, Tournay does not provide a description of his own, and given that he places Willdenow’s P. antiquorum pro parte under the synonymy of both C. papyrus L. and his own ssp. niliacus, there is not enough information to be certain of his delineation, therefore I have here followed the Plant List in rejecting this name. Further investigation is needed to determine if he published a description elsewhere.
Tournay l.c. refers C. antiquorum Chiov. var. palaestinae Chiov. to ssp. niliacus var. palaestinae, the consequence of having synonymised C. antiquorum, and this name is here treated as a synonym for the same reasons.
Tournay refers to all type specimens as lectotypes, whereas Chiovenda appears to nominate one or more specimens to accompany his descriptions, which may, therefore, be syntypes. The specimens are the same in both publications.
Kew’s acceptance or rejection of infraspecies, as published in the on-line Plant List (2010) is cited as based on Govaerts (1999), Govaerts and Simpson (2007) or Govaerts (2011). Govaerts 1999 relegates some infraspecies to C. papyrus L. s. l. whereas Govaerts & Simpson, 2007 and Govaerts, 2011 treat them as synonyms of C. papyrus L. ssp. papyrus. It appears to be the case, therefore, that C. papyrus ssp. papyrus, is a “lowest common denominator” for the synonymy, no attempt has been made to place synonyms under specific taxa. It is not clear why these authors have accepted some of Kükenthal’s infraspecies, but rejected others. C. papyrus ssp. ugandensis (Chiov) Kük. is synonymised, as is ssp. antiquorum (Willd.) Kuk. (and therefore ssp. antiquorum var. palaestinae Chiov (Kuk)), without explanation, others including, for example, ssp. zairensis (Chiov.) Kük are accepted. This may be due to the sparcity of specimens.
The Plant List records confidence in its treatment of the infraspecies as uniformly “Low”, likely to mean they arise from conflicting and unresolved sources. Because of this, the lack of published explanation for the piecemeal treatment of Kükenthal’s names, and because the synonymised names from Kükenthal and Chiovenda are all referred without distinction to synonymy, the accepted taxa for this study are based upon Kükenthal’s monograph.
Hoenselaar, for Flora of Tropical East Africa (Hoenselaar, 2010), only accepts C. papyrus s. l. However, no explanation is given for this, and, therefore, it is likely that this work did not attempt to distinguish below species level. Similarly, the account of C. papyrus in Flora Zambesiaca (unpubl.) will disregard infraspecies taxa, for want of a usable treatment (M. Lock, pers. comm. 22/01/2013). N. B., the herbarium material at Kew is arranged into Kew Herbarium areas, but not specifically into infraspecies, nor, given the only partial acceptance of taxa in Kükenthal’s monograph, is there an up-to-date key to infraspecies available to Flora writers.
Following the work of Tournay (1950), the situation remained unchanged until 1977, when a new subspecies was described for Egypt. This followed the discovery, in 1968, by Professor Dr. N. el Hadidi, of a stand of papyrus growing in an isolated salt lake between Cairo and Alexandria, at Umm Risha Lake in the Wadi Natroun. The species had been considered extinct in Egypt (Serag, 2003). The Umm Risha Lake material was considered unlike that of any other subspecies because of its geographical isolation, “indistinctly triangular to terete” stem (p. 183) and the shape of the anther connective appendage (Chrtek and Slavíková, 2003). C. papyrus ssp. hadidii Chrtek & Slavíková is detailed below, however, it is recorded as a synonym of C. papyrus ssp. papyrus in the Kew List.
Cyperus; Cyperus papyrus
Cyperus; Cyperus papyrus
Cyperus; Cyperus papyrus
Cyperus; Cyperus papyrus
Cyperus; Cyperus papyrus
Cyperus; Cyperus papyrus
Cyperus; Cyperus papyrus