Cyathula achyranthoides (Kunth) Moq.

Cyathula achyranthoides (Kunth) Moq. View in CoL in DC., Prodr. 13 (2): 326 (1849)

Fig. 1 View Figure 1

≡ Desmochaeta achyranthoides Kunth View in CoL , Nov. Gen. Sp. 2: 210 (1818).

≡ Cyathula prostrata var. achyranthoides (Kunth) Kuntze View in CoL , Revis. Gen. Pl. 2: 542 (1891).

= Desmochaeta densiflora Kunth View in CoL , Nov. Gen. Sp. 2: 211 (1818).

≡ Pupalia densiflora (Kunth) Mart. View in CoL , Nov. Gen. Sp. Pl. 2 (1): 61, t. 156, 158 (1826).

≡ Cyathula achyranthoides var. densiflora (Kunth) Moq. View in CoL in DC., Prodr. 13 (2): 327 (1849). Holotype: [ Colombia, Bolívar Department?] Rio Magdalena , [1801] Herb. Humboldt & Bonpland ( P 00670029 – image seen!; isotype P 00136028 – image seen!). Note. Kunth (1818) stated that the collections of both species, Desmochaeta achyranthoides View in CoL and D. densiflora View in CoL , originated from the same locality at Mompox. However, the original label of the type collection of the latter species does not mention Mompox, but only the Rio Magdalena River. This unnumbered collection could have originated elsewhere along the river, like at Bojorque [Bohorquez] as was previously stated on the type label. Kunth (1818) noted the differences between Desmochaeta achyranthoides View in CoL and D. densiflora View in CoL as follows: stems pubescent vs. glabrous, leaves longer vs. shorter, acuminate vs. acute, pubescent vs. less hairy, inflorescences slightly vs. densely pubescent, perianth [of the fertile flowers] throughout finely vs. basally and apically pubescent. These differences seem to be insignificant and likely fall within the natural variability of C. achyranthoides View in CoL . It should be noted that the perianth segments lose their pubescence during fruiting, which often makes them appear more glabrous than they do at flowering.

= Desmochaeta uncinata Willd. ex Roem. & Schult. View in CoL , Syst. Veg., ed. 15, 5: 554 (1819). Holotype: [ Colombia, Bolívar Department, Santa Cruz de Mompox, Rio Magdalena , April 1801] , A. Humboldt 1507 (B-W 05006-020 – image seen!; isotypes P 00670028, P 00136029). Note. As follows from the collection number and plant characters, this species was described on the basis of a duplicate specimen from the type collection of Desmochaeta achyranthoides Kunth View in CoL . Due to the brevity of the original description and the lack of the collection number in the protologue of Desmochaeta uncinata Willd. ex Roem. & Schult. View in CoL , Kunth (1820) was not able to recognise the synonymy when he aligned the South American taxa described from the collection of Willdenow with his own species.

= Cyathula achyranthoides var. glabrescens Moq. View in CoL in DC., Prodr. 13 (2): 327 (1849). Holotype: Guiane anglaise [ Guyana], 1839, Schomburgk 697 (G-DC [ G 00688954 !]; isotype P 00609947 – image seen!) . Note. Moquin-Tandon (1849) indicated that this variety was based on a single specimen at G-DC, which is the holotype. That specimen represents a stem collected in fruit. The hairs on the fertile flowers have partially fallen off; for this reason, the plant looks less hairy.

= Achyranthes hirtiflora A. Rich. View in CoL in Sagra , Hist. Fis. Cuba 11: 175 (1850). Lectotype (designated here). Cuba, Ramon de la Sagra s. n. ( P 04558206 – image seen!).

Holotype.

[ Colombia, Bolívar Department] Rio Magdalena, [Santa Cruz de] Mompox , [April 1801], Herb. Humboldt & Bonpland 1507 ( P 00670028 – image seen!; isotypes B-W 05006-020 – image seen!, P 00136029 – image seen!) . Note. The type specimens have been curatorially mislabelled as collected at Bojorque [Bohorquez], which is situated in the Atlántico Department of Colombia.

Note.

This species was cited as a synonym of Cyathula achyranthoides ( Standley 1917; Duke 1961; Acevedo-Rodríguez and Strong 2012; Greuter and Rodríguez 2017) or C. prostrata ( Grisebach 1866; Sauvalle 1870). In fact, one of the two specimens available, P 04558205, represents a true Achyranthes with glabrous flowers arranged in spikes. This specimen is A. fruticosa Lam. , which is native to Central America ( Sukhorukov et al. 2024). The other specimen, P 04558206, which is selected here as lectotype is Cyathula achyranthoides . This choice is in agreement with the original description of Achyranthes hirtiflora whose flowers were described as lanate ( Richard 1850).

Description.

Perennial herbs up to 1 m high, rooting at nodes (sometimes flowering in the first year and resembling an annual); leaf pairs 3–4 on each branch; leaves oblong, shortly petiolate (petioles up to 15 mm long), apically long-attenuate, 40–150 (170) × 20–50 (70) mm, sparsely pubescent below with appressed hairs mostly on the veins, sometimes glabrous on both sides; bract subtending each cyme (persistent on the inflorescence axis) 2.0 mm long; synflorescences rather dense or rarely interrupted in lower part, main florescence 70–150 (170–200) mm long, paracladia present; cymes (Fig. 2 A View Figure 2 ) asymmetric, with two unequal, narrowly oblong, mucronate first-ordered bracteoles (or br 1: Fig. 3 A, B View Figure 3 ) and consisting of fertile part of a cyme (one perfect flower) and sterile part mostly located on one side of the fertile flower (sometimes one very short and almost unnoticeable rudimentary flower is located on the other side of the fertile flower); sterile part with a pedicel 1.0– 1.5 mm long, bearing one sterile flower with perianth segments and two uncinate second-ordered bracteoles (br 2) 2.5–4.0 mm long, each br 2 has axillary highly rudimentary flowers accompanied by paired bracteoles beneath (br 3) shorter than br 2; sterile flower with a narrowly cylindrical perianth 1.5–2.5 mm long, acuminate or rarely two of five segments ± uncinate; thus, each cyme has at least 6 (8) unequal uncinae; fertile flower with five green perianth segments 3.0–4.0 mm long, pubescent with curved hairs and often glabrescent at fruiting, each segment mucronulate and three-nerved; pseudostaminodes 0.2–0.3 mm long, entire; anthers 0.3–0.4 mm long; style (with capitate stigma) 0.6–1.0 mm long; fruit (without style) 1.8–2.4 mm long; seed coat brown, thin; radicle pointing upwards.

Habitat.

Tropical forests up to 1150 (1500) m a. s. l. and rough ground; a common weed in many localities of Central America ( Standley and Steyermark 1946; Burger 1983). The presence at higher elevations in Colombia (1800 m a. s. l.: Schellenberg et al. (1914)) is either exceptional or erroneous. In the West Indies, the species is mostly restricted to the Greater Antilles ( Acevedo-Rodríguez and Strong 2012) and absent or very rare on the Lesser Antilles ( Kellogg 1988).

IUCN Category.

Besides the natural habitat of C. achyranthoides in forests, the species is also found on rough ground, being a noxious weed in many countries (see above). We evaluate C. achyranthoides as a Least Concern ( LC) species.

Distribution

(Fig. 4 View Figure 4 ). Belize: [Stann Creek Distr.] All Pines, 11 Mar 1932, W. A. Schipp 777 ( K 005773478).

Bolivia: [La Paz Dept.] Mapiri, 5000 ft [1524 m], Apr 1886, H. H. Rusby 1512 ( K 005773564); Yungas, 1890, M. Bang 505 ( K 005773563); Polo-Polo nr Coroico, 1912, O. Buchtien 686 ( G, K 005773562); Cochabamba Dept., San Rafaél, 24 Nov 1966, R. F. Steinbach 531 ( F 1645273, U 1049813); Pando Dept., Nicolás Suárez, 15 Jan 1983, J. F. Casas 257610 ( G); Beni Dept., José Ballivián Prov., 200 m a. s. l., 14 Dec 1994, J. Balderrama 373 ( B 100720368).

Brazil: nr Rio de Janeiro, Nov 1879, M. Glaziou ( K); Rio Jurua, Oct 1900, E. Ule 5230 ( G, K 00111421); [Ceará State] Serra de Baturité, Sep 1910, E. Ule 9027 ( G, K 001207192, L 1690980, NY 00837888); Amazonas State, Rio Branco, 1912, J. G. Kuhlmann 2851 ( RB 00038465); Espirito Santo State. Rio Doce, Sep 1950, J. N. Vieira 12 ( RB 00038438); Amazonas State, Benjamin Constant Mun., 7 Sep 1962, A. P. Duarte 6856 ( RB 00038470); Bahia State, Ilheus, 2 Apr 1965, R. P. Belém & M. Magalhaes 634 ( K 001207199, NY 00493802); [Acre State] Rio Jurua, nr Cruzeiro do Sul, 27 Oct 1966, G. T. Prance & al. 2907 ( F 1734964, K 001207198, NY 01212749, U 1049762); Amazonas State, Beruri, 7 Apr 1967, M. Silva 810 ( NY 01212751); Paraná do Autáz-Mirim, 26 Aug 1973, C. C. Berg & al. 19768 ( K 001207201, NY 01212750, U 1049754); Bahia State, Itabuna, 30 m a. s. l., 9 Mar 1977, R. M. Harley 19482 ( K 001207200, NY 00493799, P 05003751, U 1049753); Mun. Ilheus, area de CEPEC (Centro de Pesquisas do Cacau), 22 km from Rodovia, 4 Feb 1982, J. L. Hage 1627 ( K 001207202); Maranhão State, Caru Reserve, 17 Jul 1996, L. Cormier 164 ( NO 0052749).

Colombia (selected): [Magdalena Dept.] Santa Marta, 1898–1901, H. H. Smith 1203 ( G, K 005773537); [Chocó Dept.] nr La Esmeralda, 750 m a. s. l., 19 Mar 1927, E. P. Killip & A. C. Smith 20925 ( US 03543074); Boyacá Dept., Labranzagrande, 1150 m a. s. l., 1931, D. G. Amórtegui 339 ( US 03543072); Bogotá Region, Mt. Chapon, 3500 ft, 7 Jun 1932, A. E. Lawrance 171 ( G, K 005773538); Cauca Dept., between Gorgona & Puerto Cabuyo, 1000 m a. s. l., 4 Jun 1943, J. Cuatrecasas 14503 ( G); Antioquia Dept., Sonsón Mun., 23 Jan 1947, G. Gutiérrez 35544 ( F 1316964); [Antioquia Dept.] Heliconia, 28 Jun 1947, H. Daniel 3977 ( F 1283400); Rio Apaporis, between Rio Pacoa & Rio Kananari, 17 Jun 1951, R. E. Schultes & I. Cabrera 12605 ( K 005773539, US 03543061); Rio Cacuetá, 2 May 1952, R. E. Schultes & I. Cabrera 16350 ( K 005773540); [Chocó Region] Calima Valley, 2500 ft [762 m], 5 Sep 1962, D. Hugh-Jones 386 ( K 005773541); Santander Dept., Humid canyon, 700 m a. s. l., 18 Jul 1965, F. A. Barkley 35205 ( WAG 1290508); Chocó Dept., W of Unguía, 17 Jul 1975, A. Gentry & L. E. Aguirre 15226 ( F 1841364); between Rios Vaupes & Apaporis, 17 Sep 1976, J. L. Zarucchi 2066 ( K 005773530).

Costa Rica (selected): Guacimo, Aug 1901, anonymous 14669 ( K 00573477); San José Prov., nr El General, 950 m a. s. l., Nov 1936, A. F. Skutch 2904 ( K 005773482); Alajuela prov., Atenas, 28 Feb 1940, A. Smith 2467 ( K 005773483); Heredia Prov., Finca la Selva, 100 m a. s. l., 1 May 1981, J. Folsom 9978 ( F 2004755); Limón Prov., Talamanca, 3 Oct 2000, L. Acosta & al. 2820 ( INB 0003812629); Cartago Prov., Turrialba, 15 Jan 2001, E. Mora & E. Rojas 1756 ( G); Puntarenas, 7 Nov 2006, L. D. Vargas & K. Rosales 1838 ( INB 0004204213).

Cuba: [Ciego de Ávila Mun.] La Cunagua, 19 Feb 1916, N. L. Britton & al. 14585 ( NY 01374422); Guantánamo, 26 Dec 1918, B. Huram 2280 ( NY 01374418); Cienfuegos Prov., Buenos Aires, Las Lagunas, 500–550 m a. s. l., 2 Feb 2023, E. R. Bécquer & al. 91773 ( B 101259197).

Dominican Rep.: Santo Domingo, 7 Feb 1929, E. L. Ekman 11477 ( U 1049885); [Monte Plata Prov.] Yamasá, 8 Apr 1978, Alain & Liogier 27503 ( NY 00493804).

Ecuador (selected): [Canar Region] Southern Naranjapata, 550 m a. s. l., H. J. F. Schimpf 482 ( G); Pastaza Prov., Tena, 3 Oct 1939, E. Asplund 9020 ( K 005773535); [El Oro Prov.] Zaruma, 1150 m a. s. l., 12 Aug 1947, R. Espinosa 1713 ( F 1727575); Los Ríos Prov., 65 m a. s. l., 26 Aug 1949, S. S. White 5615 ( NO 0052752); Napa-Pastaza Prov., Mera, 1100 m a. s. l., 20 Feb 1956, E. Asplund 19486 ( G, K 005773536); Napo Prov., Rio Napo, Chiroisla, 24 Aug 1979, L. Holm-Nielsen & al. 19781 ( K 005773544); Napo Prov., Rio Aguaroci, San Pablo de los Secoyas, 13 Feb 1980, L. Holm-Nielsen & al. 21074 ( K 005773502, U 1049812); Morona-Santiago Prov., Taisha, 400 m a. s. l., 24 Jun 1980, J. Brandbyge & E. Asanza 32240 ( K 005773546); Pastaza Prov., Montalvo, 300 m a. s. l., 28 Jul 1980, B. Øllgaard & al. 35420 ( F 2019727, K 005773545).

El Salvador: Ahuachapán Dept., San Francisco Menéndez, 380 m a. s. l., 13 Jan 2000, J. M. Rosales 32 ( B 100019089); Santa Ana Dept., Candelaria de la Frontera Mun., La Criba, 809 m a. s. l., 17 Apr 2013, D. Rodríguez & al. 4061 ( B 100043231).

French Guiana: Montagne Cacao, 6 July 2012, G. Léotard (https://www.gbif.org/occurrence/4978245017).

Guatemala: Alta Verapaz Dept., Cubilhuitz, 1901, H. von Türckheim 7966 & 11423 ( G, US 03543016); Izabal Dept., Los Amates to Izabal, 31 May 1919, S. F. Blake 7801 ( US 03543014).

Guyana: [without locality] 1841, Schomburgk 93 ( G, K 005773507); Barima-Waini Region, 5 miles [8 km] W of Arakaka, 20–80 m a. s. l., 7 Aug 1986, J. Pipoly & H. Lall 8348 ( US 00474940); Kanuku Mts., head of Nappi Creek, 31 Oct 1987, M. J. Jansen-Jacobs & al. 588 ( K 005773498).

Haiti: Dept. du Nord, 19 Nov 1924, E. L. Ekman 2574 ( G, K 005773493); nr Plaisance, 400 m a. s. l., 26 Jan 1926, E. C. Leonard 9198 ( NY 01374416); nr St. Louis du Nord, 1929, E. C. Leonard & G. M. Leonard 14360 ( K 005773486, NY 01374413).

Honduras: Atlántida Dept., Tela, 18 Aug 1979, R. Andino 55 ( US 03542992); Atlántida Dept., Tela Mun., 10 Sep 2002, J. Pipoly & C. Valle 24307 ( USF 277322).

Jamaica: [without exact location] Mar 1871, ex herb. Hookerianum 1237 ( K 005773491); Port Antonio, 7 Jan 1906, A. E. Wight 36 ( NY 01374425); Bluefields, 500 m a. s. l., Mar 1908, N. L. Britton & A. Hollick 1999 ( NY 01374424).

Mexico (selected): [Veracruz State] Orizaba, 1865–1866, M. Bourgeau 3040 ( K 005773473); Chiapas State, Escuintla, 22 Nov 1947, E. Matuda 17177 ( K 005773474); Veracruz State, San Andres Tuxtla, 450 m a. s. l., 12 Aug 1972, R. Cedillo 270 ( BR 0000027811986); Veracruz State, San Andres Tuxtla, 12 Aug 1974, R. Cedillo 270 ( K 005773476); State of Chiapas, Yajalón, 31 Aug 1983, A. M. Ton 6563 ( MA 614037); State of Puebla, Taxipehuatl, 2 Nov 2014, P. Acevedo-Rodríguez & al. 15965 ( US 01343465).

Nicaragua: Chontales [Dept.], 1867, R. Tate 334 ( K 005773472); Jinotega Dept., Wiwilí Mun., 165 m a. s. l., 10 Jun 2007, I. Coronado & A. Fernández 4019 ( BM 001172464).

Panama (selected): [without exact locality] Nov 1861, S. Mayrs 612 ( K 005773533); nr Paya, 12 Jun 1959, W. L. Stern & al. 402 a ( G); Darién Prov., Rio Chico, 19 Dec 1966, D. Burch & al. 1083 ( K 005773552); Canal Zone, Farfan Beach, 27 Dec 1966, D. Burch & al. 1403 ( K 005773551); Bocas del Toro, Jun 1967, W. H. Lewis & al. 1997 ( K 005773549); Bocas del Toro, Chiriqui Prov., nr Puerto Armuelles airport, 17 Feb 1973, T. B. Croat 21885 ( USF 110618); Cocle & Herrera prov., border of Verauas, 11 Feb 1982, S. Knapp & al. 3338 ( K 005773553); nr Tiger Key, 21 Feb 1989, P. M. Person & C. R. Annable 6957 ( K 005773548).

Peru (selected): [without exact location] 1832, Poppig 146 ( G 00688958); Huánuco Prov., 10 km S of Tingo Maria, 700 m a. s. l., 26 Oct 1938, H. E. Stork & O. B. Horton 9508 ( F 1078734, G, K 005773560); Ganso Azul, 1500 ft [457 m], Oct 1942, Sandeman 3364 ( K 005773559); San Martín Region, nr Tarapoto, 890 m a. s. l., 11 Mar 1947, F. Woytkowski 35229 ( F 1316573, G); Bagua prov., E of Olmos, 22 Jan 1964, P. C. Hutchinson & J. K. Wright 3717 ( F 1640136, K 005773561, P 05002116); [Lambayeque Dept.] nr Olmos, 500 m a. s. l., 22 Jan 1964, P. C. Hutchinson & J. K. Wright 3717 ( G); [Cuzco Prov.] Villa Carmen [Manu], 3 Mar 1964, C. Vargas 15199 ( WAG 1290507); Huánuco Dept., Pachitea Prov., 300 m a. s. l., 30 Jan 1967, J. Schunke 1579 ( G); Ayacucho Region, La Mar prov., between Tambo San Miguel & Ayna, 18 Aug 1968, T. R. Dudley 11821 ( F 1692339); Amazonas Dept., Quebrada Huampami, 14 Nov 1972, R. Kayap 64 ( F 1821272); San Martín Dept., towards Santa Rosa, 350–370 m a. s. l., 9 Aug 1973, J. S. Vigo 6783 ( L 1690989); Loreto Dept., nr Iquitos, 1977, J. Revilla 3027 ( F 2202869); Loreto Dept., Maynas Prov., nr Iquitos, 120 m a. s. l., 22 May 1978, A. Gentry & al. 22206 ( F 2028534); Amazonas Dept., nr La Poza, 9 Aug 1979, J. A. Leveau 42 ( P 05003752); Loreto Dept., Caserio Sapa Playa, 15 Aug 1979, M. Rimachi 4560 ( K 005773557); Rio Itayu, 18 July 1981, H. Murphy 29 ( K 005773570); Huánuco Region, Tingo María, 700–800 m a. s. l., 9 Dec 1981, T. Plowman & al. 11242 ( F 1894977); Ucayali Dept., Panguana, 9 Jul 1983, F. Seidenschwarz 13 / 1 ( F 1976892); Jaén Prov., Huahuaya, 870 m a. s. l., 31 Jul 1994, S. Leiva & al. 1210 ( F 2154290); Bagua Prov., Imaza Distr., Yamayakat community, Jan 1995, V. Hodges & J. Gorham 130 ( F 2239684); Cuzco Prov., San Martin, 15 Feb 1997, P. Nuñez 19016 ( US 03543088).

Saint Martin: [without exact location] 1856, J. J. Triana 19701 ( BM).

Saint Vincent and the Grenadines: St. Vincent Island, [without date, probably 1820 s] L. Guilding s. n. ( K 005773490).

Suriname (selected): Paramaribo, Apr 1916, J. A. Samuels 98 ( K 005773497); Domburg, 13 December 1960, K. U. Kramer & W. H. A. Hekking 2348 ( NO 0052754); nr Paramaribo, 10 January 1961, K. U. Kramer & W. H. A. Hekking 2618 ( BR 0000027812082); Brokopondo Distr., Brownsweg, 37 m a. s. l., 19 Aug 2006, S. Ruysschaert & al. 858 ( BBS 0000438).

Trinidad and Tobago: Trinidad, [without date] B. de Schach s. n. ( K 005773485); Trinidad, 17 Jan 1889, ex herb. Krug & Urban 3426 ( G); Maracas Falls, 30 May 1956, Simmonds 15409 ( K 005773489); St. Joseph Stock Farm, 8 Mar 1958, J. W. Purseglove 6133 ( K 005773496, L 1690992).

Venezuela: Aragua State, Henry Pittier NP, 850 m a. s. l., 7 Nov 1973, T. Romero 302 ( F 1768541); Bolívar State, Salto Pará, 80 m a. s. l., 12 Jan 1977, J. A. Steyermark & al. 112887 ( F 1804290); Zulia [State], distr. Bolívar, 4 Feb 1980, S. Bunting 8673 ( K 005773556); Yaracuy State, Bruzual distr., 12 Mar 1981, A. Julián & al. 124909 ( K 005773554); Portuguesa State, 30 km W of Guanare, 13 Mar 1982, R. Liesner & al. 12599 ( K 005773555, MA 391992); The Capital Distr., Cordillera de la Costa, El Ávila NP, 200–230 m a. s. l., 14 Apr 2006, W. Meier 13874 ( B 100527007).

General distribution.

Tropical America.