Conspectus subspecierum

 

Cyperus L., Sp. Pl. ed. 1, p. 44, 1753

Lectotype Cyperus esculentus L. (Britton, 1907: 6).  (Huygh et al., 2010).

Cyperus papyrus L., Sp. Pl. ed. 1, p. 47, 1753 and ed. 2, p. 70, 1762, emend Kük., in Engler (ed.), Pflanzenreich, IV 20, booklet 101: p. 57, (1936).

Description:   Large, emergent, aquatic perennial.  Rhizomes short, covered in thick, black scales.  Roots tough, extending 1m or more in suitable conditions.  Rootlets numerous.  Culms erect to 5m, 5 (-6)cm across at the base (widest point), smooth, trigonous (angles very rounded, particularly on lower parts), green (photosynthetic); spreading when old.  Pith solid, white-light brown, vascular bundles prominent.  Leaves reduced, sheathing and restricted to basal ±50cm of culms; reddish-blackish brown when young; expanding with age and colours restricted to lateral edges sub-marginally; tough, margins scarious; becoming papery, colours fade and sheaths split ventrally with age.  Inflorescence large (30-) 50 (-60)cm diameter, compound, umbel-like, hemispherical when young, becoming sub-globose with age.  Some inflorescences smaller, entirely sterile, or minutely fertile.  Bracts (B1) variable, 10-15 (-17)cm long, (10-) 12-25mm wide at base; broadly triangular to triangular-attenuate; reddish dorsally in bud, becoming green then papery and brown with age; patent to slightly recurved.  Rays “branches”, (R1) 50-150, sub-equal, smooth, green, (15-) 20-30 (-35)cm long, thin (2) 3-4 (5)mm wide, trigonous to triquetrous apically, flexible; with basal, sheathing, dark brown tipped, red-veined prophylls (P1) 2-2.5cm.  Spikes (S1) terminal on rays, in small umbel-like clusters (1-) 3 (-4), length 2 – 3cm, elongate-cylindrical; subtended by  3 (-4) bracteoles (B2), (5-) 10-15 (-16)cm long, thin, wiry, green, trigonous; rough-scabrid in one direction, apex to base.  Rachillas (R2), unequal 0-1cm to first spikelet; winged; with basal, short, sheathing prophylls (P2).  Spikelets (S2) arranged spirally, sessile, oblong to oblanceolate, flattened to sub-terete; minute basal bract (B3), 0.3-0.4cm, lanceolate to sub-ovate; ovate, with basal, semi-sheathing prophyll (P3) 0.4-0.6cm.  Florets (F1), bisexual, arranged distichously, 4 – 8 (-10). Glumes 1-2mm, blunt to mucronulate.  Scales “rachilla wings” 1mm, chaff-like, brown, lanceolate, curling on drying.  Style long; stigmas 3, long, protruding, persistent.  Ovary superior, one-celled.  Stamens 3, filaments fine, elongating; anthers (0.8-1.2mm), basifixed, bilocular, caducous; connective tissue distinct, red, with an apical extension ± 0.1mm.  Nutlets 3-angled, 0.8-1.3mm, brownish-black, oblong, sometimes concave ventrally, shortly apiculate, with short-lived, translucent.

Vernacular names:    Papyrus, Papyrus reed.

Lectotype:      "Ɵ Papyrus 15", Material from Linn. (UPS). This specimen is listed by Juel (1931: 13).  Designated by Simpson in Cafferty & Jarvis (2004) for C. papyrus s.l..

Notes:             Since C. papyrus ssp. Cyperus is the autonym of C. papyrus, and was lectotypified by Clarke, this type appears to be superfluous.

_____ ssp. papyrus

C. Papyrus L., Sp. Pl. ed. 1, p. 47 (1753) and Sp. Pl. ed. 2 (1763) p. p. type with regard to herb. material and synonymy Van Royen p. p., Bauhin p. p., Monti, Scheuchzer, Micheli p. p.; Morison p. p.;  L., Syst. Nat. ed. 12, 2, p. 82 (1767);  Lam., Ill. Genres, 1, p. 147 (1791) p. p. type with regard to herb. material from Paris bot. garden;  Roem. and Schultes in L., Syst. Veg., (ed. 18), 2, p. 218 and 876 (1817) p. p. type with regard to  herb. material; from Sicily, and to synonymy of Micheli p. p. and  Willdenow p. p.;  Boeck., Linnaea, 36, p. 303 (1869-1870) p. p. type, with regard to Sicily and synonymy of L. p. p., Willd. p. p., and Parl (1851); Thistleton-Dyer, Gard. Chron., ser. 2, 3, p. 78 and 81, fig 14 (1875) p. p. type with regard to Sicily;  Chiov., Lav. R. Ist. Bot. Modena, 1, p.78, tab. 3, fig 8 and tab. 4, fig. 8 (1931).

Papyrus Jussieu in Caylus, Mem. Litt. Acad. R. Inscr. B.-L., Paris, 26, p. 280, fig. 1-2, (1759) p. p. with regard to Sicily.

C. papyraceus Crantz, Inst. Rei Herb., 1, p. 411 (1766) p. p. with regard to synonymy L. p. p.

 P. antiquorum Willdenow, Abhandl. Phys. Klasse K.-Preuss. Akad. Wiss., 1812-1813, p. 70 (1816) p. p. with regard to herb. material and synonymy of L. p. p.

C. Papyrus L. var. α Roem. and Schultes, in L., Syst. Veg., (ed. 18), 2, p. 218 (1817).

P. domesticus Poiret In Bory, Dict. Class. Hist. Nat., 37, p. 447 (1825) p. p. with regard to synonymy of L. p. p.

P. sicula Parlatore, Hooker’s Journ. Bot., ser. 4, 3, p. 189 (1851). Nom. nud.

C. syriacus Parlatore, FI. Ital., 2, p. 43 (1852) and Mém.  Acad. Sc. Inst. Imp. France, Sc. Math, and Phys., 12, p. 498, tab. 1 (1854) p. p. L. type with regard to herb. material from Sicily.

Chlorocyperus Papyrus Rikli, Pringsh. Jahrb. Wiss. Bot., 27, p. 563 (1895) p. p. type with regard to synonymy L. p. p.

C. siculus (Parlatore) Chiov., Delect. Sem. Spor. Horti Bot. Catinensis, 1927, p. 2 (1927).

C. panormitanus Chiov., Lav. R. Ist. Bot. Modena, 1, p. 79, tab. 3, fig. 10 and tab. 4, fig. 10, (1931).

C. elapsus Chiov., Lav. R. Ist. Bot. Modena, 1, p. 79, tab. 3, fig. 10 and tab. 4, fig. 10, (1931)

C. papyrus L. ssp. siculus (Parlatore) Kük.. in Engl., Pflanzenreich, 101, p. 48 (1935).

Lectotype:      “One sheet marked “Papyrus” in Linnaeus’s hand, Linn.” (UPS), Clarke (1894: 308).  Bullo et al. (2011) state that Clarke was designating LINN 70.38, which they saw.  Kunth (1837: 64) cites “Linn. Spec. 70”, presumably the same specimen.  Neither Chiovenda nor Kükenthal list specimens from Linnaeus under C. papyrus L. or C. papyrus ssp. siculus (Parl.) Kük. respectively.

N.B. Tournay (1950) and others note that Linnaeus only had access to material from Sicily, and so the type is of Italian, not directly African, origin.

Specimens:    Sizilien: Bei Syrakus am Flusse Anapo nahe bei der Quelle Ciana, heute Pisma genannt und 4-5 km entfernt bei La Maddalena (Hohenacker n. 549!, Ross n. 2301, Todaro n. 1330! u. a. §).

Description:   Anther connectives beyond locule apices 1-3 X anther breadth.  Glumes narrowly elliptical.  Spikelets 1-3cm.  (Translated after Chiovenda, 1931). 

Notes:             Linneaus augmented his publication of the name with additional descriptive material in Species Plantarum ed. 2.  His name is based on a specimen of Van Royen, and he cites, by way of description, the predecessor authors that Van Royen cited, namely Bauhin etc., with the addition of Morison.  Chiovenda published C. elapsus to accommodate one specimen material from Micheli (Tournay, 1950), Tournay does not list it as a synonym.  C. panormitanus Chiov. is regarded as a synonym of C. papyrus s. l. in the Plant List, but not by Tournay, it may, therefore be better placed, following investigation under another subspecies.

____ ssp. antiquorum (Willd.) Kük., in Engler (ed.), Pflanzenreich, IV 20, booklet 101: p. 47, (1936).

C. Papyrus L., Sp. Pl. ed. 1, p. 47 (1753) and Sp. Pl. ed. 2, p. 70 (1763) p. p. non-type with regard to herbarium and synonymy Morison p. p.;  Roem and Schultes in L., Syst. Veg., (ed. 18), 2, p. 218 and 876 (1817) p. p. non-type with regard to synonymy Bruce p. p. and Willdenow p. p.  Delile, Descr. Egypte, Hist. Nat., 2, p. 50 (1813) ; Hooker, f. and Bentham in Hook., Nig. FL, p. 551 (1849); Parlatore, Mém. Acad. Sc. Inst. Imp. France, Sc. Math, et Phys., 12, p. 500, tab. 1 (1854);  Boeck., Linnaea, 36, p. 303 (1869-1870) p. p. non-type with regard to synonymy L. p. p. and Willdenow p. p.; Thistleton-Dyer, Gard. Chron., ser. 2, 3, p. 78 (1875) p. p. with regard to Abyssinia;  Boeck., Flora, 62, p. 555 (1879) non-type with regard to Schweinfurth 1154 and 3749 ; Clarke in Dur. et Schinz, Consp. Fl. Afr., 5, p. 571 (1894) p. p. non-type with regard to Lagos, Vieux Calabar, Kameroun and Ogôoué and in Thistleton-Dyer, Fl. Trop. Afr., 8, p. 374 (1901) p. p. non-type with regard to from Upper Guinea.

C. papyraceus Crantz, Inst. Rei Herb., 1, p. 411 (1766) p. p. type with respect to synonymy. L. p. p.

Papyrus Savary, Lettr. Egypte, 1, p. 322 (1788); Bruce, Trav. Discov. Source Nile, 5, App., p. 14, tab. 1 (1790) p. p. with regard to Egypt and Abyssinia.

P. antiquorum Willdenow, Abhandl. Phys. Klasse K.-Preuss. Akad. Wiss., 1812-1813, p. 70 (1816) p. p. non-type with regard to synonymy of L. p. p.

C. Papyrus L. var. β Roem. and Schultes, L., Syst. Veg., (ed. 18), 2, p. 218 (1817).

P. domesticus Poiret. in Bory, Dict. Class. Hist. Nat., 37, p. 447 (1825) p. p. non-type with regard to synonymy of L. p. p.

C. Papyrus L. var. antiquorum (Willdenow p. p. non-type) Clarke, Journ. Linn. Soc, Lond., Bot., 30, p. 571 (1894) p. p. non-type with regard to Bassin de la Gazelle, Pays des Niam-Niam and Bassin du Nil Blanc and loc. cit. p. 374 (1901) p. p. non-type with regard to Nile Land.

Chlorocyperus Papyrus (L. p. p. non-type) Rikli, Pringsh. Jahrb. Wiss. Bot., 27, p. 563 (1895), p. p. non-type with regard to synonymy L. p. p. 

C. antiquorum (Willdenow p. p. non type) Chiov., Lav. R. Ist. Bot. Modena, 1, p. 73, tab. 2, fig. 4 and tab. 4, fig. 4, (1931).

C. papyrus L. ssp. niliacus Tournay var. niliacus, Bulletin de la Société Royale de Botanique de Belgique/Bulletin van de Koninklijke Belgische Botanische Vereniging, 82(2), 349, (1950).  Nom. nud.

Lectotype:      Soudan Anglo-Egyptian, Nil Blanc, 1862, Petherick (K).

Specimens:    Anglo-agyptischwer  Sudan: Am Weissen Nil (Petherick!, Werne!); am Gazellenfluß (Schweinferth n.1154! §); Land der Niam-Niam (Schweinferth n3749!) am Blauen Nil bis Abessinien, Tanesee 1850 m (F. Rosen !); Gallahochland, am Awalasee (Ellenbeck n. 1703!).   Westwärts bis zum Tschadsee; Französ Kongo, Ogowefluß (Buchholz!); Kamerun, Njong-Upa (H. Winkler n. 835!); Süd-Nigeria; Lagos (Barter!).

Description:   Anther connective not or scarcely extending beyond tips of anther locules; and shortly arched.  Spikes lax, shortly pedunculate.   (Translated after Kükenthal, 1935).   Anther connectives not or scarcely beyond locule apices.  Spikelets 5-10mm.  6-10 florets.  Glumes elliptical-rotund.  (Translated after Chiov., 1931)

Notes:             This synonymy, selectively from Tournay (1950) is placed by him under ssp. niliacus Tournay, here regarded as a synonym itself.  The lectotype cited here, is, therefore the one given by Tournay to ssp. niliacus.

_____ ssp. antiquorum var. palaestinae (Chiov.) Kük., in Engler (ed.), Pflanzenreich, IV 20, booklet 101: p. 47, (1936).

C. Papyrus L., Sp. Pl. ed. 1, p. 47 (1753) and Sp. Pl. ed. 2, p. 70 (1763) p. p. non-type with regard to herbarium and synonymy Bauhin p. p., Morison, p. p., Micheli p. p. and Van Royen p. p.; Roem and Schultes in L., Syst. Veg., (ed. 18), 2, p. 218 and 876 (1817) p. p. non-type with regard to synonymy Micheli p. p., Bruce p. p. and Willdenow p. p.; Thistleton-Dyer, Gard. Chron., ser. 2, 3, p. 78 (1875) p. p. with regard to Palestine.

C. papyraceus Inst. Rei Herb., 1, p. 411 (1766) p. p. type with respect to synonymy L. p. p.

Papyrus Bruce, Trav. Discov. Source Nile, 5, App., p. 14, tab. 1 (1790) p. p. with regard to Syria.

P. antiquorum Willdenow, Abhandl. Phys. Klasse K.-Preuss. Akad. Wiss., 1812-1813, p. 70 (1816) p. p. non-type with respect to synonymy of L. p. p.

P. domesticus Poiret in Bory, Dict. Class. Hist. Nat., 37, p. 447 (1825) p. p. non-type with regard to synonymy of L. p. p.

C. syriacus Parlatore, Mém. Acad. Sc. Inst. Imp. France, Sc. Math, et Phys., 12, p. 498, tab. 1 (1854);  p. p. non-type with regard to herb. material from Syria and synonymy Bauhin p. p., Micheli p. p., L. p. p. and Willdenow p. p.

C. Papyrus L. var. syriacus (Parl. p. p. non-type) Boeck., Linnaea, 36, p. 304 (1869-1870)

Chlorocyperus Papyrus (L. p. p. non-type.) Rikli, Pringsh. Jahrb. Wiss. Bot., 27, p. 563 (1895), p. p. non-type with regard to synonymy L. p. p. 

C. antiquorum (Willdenow p. p. non-type) Chiov. var. palaestinae Chiov., Lav. R. Ist. Bot. Modena, 1, p. 119, tab. 3, fig. 5-7 and tab. 4, fig. 5-7,  (1931).

C. Papyrus L. ssp. niliacus Tournay var. palaestinae (Chiov.) Kük. in Engl., Pflanzenreich, IV 20, booklet 101: p. 47, (1935).

Lectotype:      Palestine, lac de Galiliée, 1872, Hayne (K).

Specimens:    Palastina: Ras es Ain; Meromtal (W.A. Hayne); Galilaischer See (W. A. Hayne, O. Kersten!); am Strande bei Jaffa (Michon, Letourneux!)

Description:   Anther connective not or scarcely extending beyond tips of anther locules, truncate across apices of locules.  Spikes short, thick.  (Translated after Kükenthal, 1935) .

Notes:             The Flora of Palestine (Feinbrun-Dothan, 1981) considers the issues of subspecies in respect to their native populations, concluding that the anther connective character first used by Chiovenda is unreliable and therefore persisting with the species sensu lato.

_____ ssp. hadidii Chrtek and Slavíková Preslia vol. 49: p. 183-185, (1977).

Holotype:       Lacus Umm Risha, Wadi Natroun, Aegyptus, Nabil el Hadidi 1968, (CAI).

Specimens:   

Description:   Culms smooth, rounded to inconspicuously triangular.  Anthela composite,  many rays. Anthelulae dense, with 3 spikes, spikelets shortly pedunculate or subsessile, 6-8 (-10) mm long, pitch-black to brown.  Glumes densely imbricate, adpressed, elliptic-ovate to elliptic, apex bluntly acute, violet-striped. Anther connective extending beyond anther apices, three times or four times the length of the shorter side of the anther apex.  (Translated after Chrtek & Slavíková, 1977) 

Notes:             Boulos (2005) treats this name as a synonym of C. papyrus.  Since he does not explain, nor retain any infraspecies as accepted names, here the name is retained as accepted.

 

_____ ssp. madagascariensis (Willd.) Kük., in Engler (ed.), Pflanzenreich, IV 20, booklet 101: p. 48, (1936).

Papyrus Jussieu in Caylus, Mem. Litt. Acad. R. Inscr. B.-L., Paris, 26, p. 280, fig. 3(1), (1759) p. p. with regard to Madagascar.

P. madagascariensis Willdenow, Abhandl. Phys. Klasse K.-Preuss. Akad. Wiss., 1812-1813, p. 72 (1816).

C. Papyrus L. var. γ Roem. and Schultes, L., Syst. Veg., (ed. 18), 2, p. 218 (1817).

C. madagascariensis (Willd.) Roem. and Schultes, L., Syst. Veg., (ed. 18), 2, p. 876 (1817);  Clarke, Journ. Linn. Soc, Lond., Bot., 30, p. 568 (1894) and in Thistleton-Dyer, FI. Cap., 7, p. 184 (1897).  Kunth.

C. Papyrus L. var. madagascariensis (Willdenow) Boeck., Linnaea, 36, p. 304 (1869-1870); ? Thistleton-Dyer, Gard. Chron., ser. 2, 3, p. 78 (1875)

C. imerinensis Boeck., Engl. Bot. Jahrb., 5, p. 500 (1884).

C. madagascariensis (Willdenow) Roem. and Schultes var. imerinensis (Boeck.) Clarke, Journ. Linn. Soc, Lond., Bot., 30, p. 568 (1894).

C. madagascariensis (Willdenow) Kunth, Enum Plant. 2, p. 64 (1837).

Lectotype:      Madagascar, s. l. n. d. Aubert du Petit-Thouarts in Willdenow, 1414. (B).

Specimens:    Malagassisches Gebiet: Madagaskar (Thouars!, Baron n. 552 §, 3274, 5110); östliche Region (Poivre, Geay n. 7778);  Centralregion, Imerina (Hildebrandt n. 3798!, Perrier de la Bäthie n. 1483); Tananarive; Nanisana; Moramanga (K. Afzelius!) u. a. St. Mascarenen: Reunion. Mauritius (N. J. Andersson!). Trop. Ostafrika: Delagoabay (Forbes!). Deutsch-Ostafrika: Kihmandjaro, im Sumpfe Mboloti zwischen Sanjafluß und Kwarefluß (A. Peter n. 2719!)

Lectotype:      Nyasaland, fleuve Shire, 1887, L. Scott (K)

Description:   Anther connective not or scarcely extending beyond tips of anther locules.  Culms triquetrous. Bracts narrow.  Bracteoles anthelulas barely exceeding [spikes] usually much shorter.  Glumes often intensely brick red.  Spikelet wings acute.  Translated after Kükenthal, 1935).

Notes:             Tournay (1950) omitted the work of Kunth (Kunth, 1837), despite Kükenthal relying on it for the description of this subspecies.  Here, therefore it is included, but separately, as it has not been subject to scrutiny of the description or specimens, as clearly undertaken by Tournay.

_____ ssp. nyassicus (Chiov.) Kük., in Engler (ed.), Pflanzenreich, IV 20, booklet 101: p. 47, (1936).

C. nyassicus Chiov., Lav. R. Ist. Bot. Modena, 1, p. 76, tab. 2, fig. 1-2 and tab. 4, fig. 1-2,  (1931).

Lectotype:      Nyasaland, fleuve Shire, 1887, L. Scott (K).

Specimens:    Nyassaland:  Kondowe gegen Karonga 600-1800m (Whyte); Ufer des Shire bei Shamo (Scott); im Stromgeveit des Sambesi (G. Adanson n. 51§).

Description:   Anther connective not or scarcely extending beyond tips of anther locules, linear-cylindrical.  Glume apices subacute.  (Translated after Kükenthal, 1935).   Anther connectives beyond locule apices cylindrical-narrow.  Glumes more acute.  Raceme lax, peduncled.  Spikelets often elongate 1-3cm, many flowered.  (Translated after Chiov. 1931).

_____ ssp. ugandensis (Chiov.) Kük., in Engler (ed.), Pflanzenreich, IV 20, booklet 101: p. 47, (1936).

C. mossambicensis Klotzsch with Boeck. in Peters, Reise Mossamb., Bot., p. 541 and 575, in syn. (1864) nom. illegit.

P. mossambicensis Parlatore with Boeck. In Peters, Reise Mossamb., Bot., p. 575, in syn., tab. 59 (1864) nom. illegit.1. c, p. 575, in syn., tab. 59 (1864) nom. illegit

C. ugandensis Chiov., Lav. R. Ist. Bot. Modena, 1, p. 77, tab. 2, fig. 3-3b and tab. 4, fig. 3, (1931).

Lectotype:      Uganda, Naivasha, 1900, H. Johnston 249 (K).

Specimens:    Tropical. Ost-Afrika: Uganda: Ruwenzori Exped. (Scott-Elliot); Naivasha (Johnston n. 249!). Kenya-Territ.: südlich von Mombassa (A. Whyte I). Deutsch-Ostafrika: Karagwe, bei Bukoba (Stuhlmann n. 1618!); Uvinsa 1060m (A.Peter n. 36 214!); Irangi, am Sseria-See 1350 m (A. Peter n. 44 537!); Kilimandjaro (A. Peter n. 2718!); West-Usambara, Heputu bei Bungu 900 m (A. Peter n. 15 743!); Matanana (v. Prittwitz n. 89!); Usaramo (Stuhlmann n. 8617!). Zanzibar (Hildebrandt n. 1193!). Mozambique: Borror, in Sümpfen (Peters!). Nordost-Rhodesia : Bangweolo, in Sümpfen bei Kamindas (R. E. Fries n. 919!). Nordwest-Rhodesia : Victoria-Fälle.

Description:   Anther connective not or scarcely extending beyond tips of anther locules, ovoid or subglobose, red.  Glume apices rounded.  (Translated after Kükenthal, 1935).  Anther connectives beyond globoid to sub-globose.  Racemes lax. Translated after Chiov. 1931). 

_____ ssp. zairensis (Chiov.) Kük., in Engler (ed.), Pflanzenreich, IV 20, booklet 101: p. 48, (1936).

C. zairensis Chiov., Lav. R. Ist. Bot. Modena, 1, p. 76, tab. 2, fig. 1-2 and tab. 4, fig. 1-2, (1931).

Lectotype:      Congo Belge, fleuve Congo, s. d., Curror (K).

Specimens:    Im Stromgebiet des Kongo: Französ. Kongo: Loango (Soyaux n. 106!, Naumann n. 138!). Belgisch- Kongo : Ponte da Lenha (Naumann n. 1391); Kisantu (Gillet n. 511); am Kongoufer (Smith n. 7, Gurror); im Süden des Kongostaates im Bezirk Lunda, amLomanifluß (Poggen. 1574!), am Luluafluß (Poggen. 1585!), am Mt. Lumbala bei Mutumbo (Quarré n. 25461). Angola: Loanda (Welwitsch n. 7085); Golungo Alto (Welwitsch n. 7104); Guanza Norte (Gossweiler n. 8779!).

Lectotype:      Nyasaland, fleuve Shire, 1887, L. Scott (K)

Description:   Anther connective 2-3 X anther width, subulate whitish. Spikelet wings acute.  Spikes conspicuously pedunculate.  Spikelets 3-7mm.  Glumes broadly elliptical.    (Translated after Kükenthal, 1935).  Anther connectives beyond locule apices 1-3 X anther breadth.  Glumes broadly elliptical, apex broadly rounded.  Spikelets 3.5mm.  (Translated after Chiov., 1931).  

Cyperus; Cyperus papyrus