Ceroxylon pityrophyllum (Mart.) Mart. ex Wendl., Kerchove de Denterghem, Les Palmiers (1878)

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Distribution

Map uses TDWG level 3 distributions (http://www.nhm.ac.uk/hosted_sites/tdwg/geogrphy.html)
Known from the Andes in Bolivia (La Paz, Cochabamba, Santa Cruz and Chuquisaca) and Southeastern Andes of Peru (regions Cuzco and Puno;), at 1285-2800 m, on slopes in patches of cloud, semi-deciduous, or deciduous forest (in the Yungas humid forest, the deciduous mountain Chaco forest, and the Tucumano-Boliviano forest). It is often observed along with Parajubea (Kessler & al. 5683, in scheda), with Podocarpus parlatorei Pilg., Ternstroemia asymmetrica Rusby, or in cloud forests disturbed by grazing, or in transitional forests between cloud forest and dry forests, mixed with Podocarpus sp., Anadenanthera colubrina (Vell.) Brenan and other Fabaceae and Myrtaceae; above limits of subtropical deciduous dry forests (Vargas & al. 2270, in scheda), Lauraceae, Myrtaceae and Dictyocaryum lamarckianum (Mart.) H. Wendl. (Paniagua & al. 5869, in scheda), but never in dry forest (Nee & Vargas 38268, in scheda). It can form populations of over several hundred trees along ridgetop (Nee & Vargas 38268, in scheda), but is mostly seen as scattered sub-canopy individuals. In Perú, it is conserved in cultivated plots and favored in shade coffee plantations, where it can be more abundant than in natural conditions (F. Kahn et al., in scheda). Nevertheless, most of the natural habitat is being rapidly converted to pastures and cultivation farms (Nee & Solomon 30324, in scheda). (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))A

Discussion

  • Ceroxylon pityrophyllum is very distinct in its large anthers (the largest in the genus, and almost twice the size that in other species;). It resembles C. parvum in its habit and hanging dry leaves; nevertheless, C. parvum has (6-) 9-11 slender staminodial filaments instead of six stamens and staminodes with swollen bases.
    The habit of C. pityrophyllum can also resemble that of C. vogelianum, particularly in the case of slender individuals, but the texture of the fruit exocarp (smooth to minutely verrucose in C.pityrophyllum, whereas always furrowed in C. vogelianum), and the length of the anthers (4.5-7 mm in C.pityrophyllum), while 2-2.5 mm in C. vogelianum), are key characters for distinguishing these species apart.
    All the specimens from Bolivia that have been previously identified as C. parvum (Galeano 1995) or C. vogelianum belong in fact to C. pityrophyllum. C. parvum appears to be restricted to south-western Ecuador while C. vogelianum extends from Venezuela to southern Peru (Cuzco).
    Ceroxylon pityrophyllum is morphologically very variable: it varies in size from 4 to 22 m tall; the wax on the stem and abaxial surface of the leaves can be thin or very thick; the leaf rachis arching and flexible or stiff and horizontal, and the orientation and disposition of the pinnae in 2 or in many planes, and almost regular (or even regular in juveniles) to conspicuously grouped. However, the reproductive characters such as the stamen number, the size of the anthers, the width of the staminodial filaments, and the texture of the fruit exocarp, are always constant characters throughout its geographic distribution.
    Regarding the confusion with the name C. pityrophyllum, causing its designation as Species dubia by Burret (1929) and its exclusion as "Uncertain names" in Henderson et al. (1995), the following statements can be put forward. The holotype (of the name Cocos pityrophylla Martius) lost in P does not belong to the genus Cocos, as Govaerts & Dransfield (2005), and Glassman (1972) suggest. The corrections made by Martius, who saw the type, "pinnis ... subtus squamilus parvis lanceolatis albidis", can only refer to Ceroxylon, not Cocos. Even though Burret (1929) explains that the observation "infrafoliar inflorescences, and rachis branched only to first order" could only be an error, he does not exclude the possibility of this being Cocos; therefore his designation of the species as species dubia. On the basis of recent observations in the area, the only genus growing in or near the type locality that combines the characteristics mentioned by Martius (1847; abaxial surface of pinnae pale, lepidote, inflorescences infrafoliar, rachis branched to first order) is Ceroxylon.
    Lastly, many specimens of C. pityrophyllum from Bolivia had been determined as C. vogelianum, based on several premises: the first, that C. vogelianum actually grew in Bolivia, the second, that the staminate flowers with six stamens and the grouped pinnae inserted in many planes were characters found together only in this species. Fieldwork in several departments has been carried out, and all of the collections of Ceroxylon from Bolivia have now been revised, not one belonging to C. vogelianum (see discussion above).
    Ceroxylon weberbaueri Burret still grows in its type locality in Peru (Pintaud et al. 2010), area which forms an ecological continuum with the type locality of C. pityrophyllum in La Paz. All of the recent collections from both localities are the same species, and correspond to the original descriptions for both names. Therefore, the decision of which name to keep was taken regarding the date of publication (1847 vs. 1929). (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))A

Conservation

Common Name

Uses

Description

  • Stem 4-20(-22) m tall, 9.9-30.0(-40.0) cm diam., green, covered with a very thin layer of wax. Leaves 12-20(-25), in a spheric crown, often grayish and with persistent, dried hanging leaves; sheath 40-124 cm long, 3.8 cm wide at apex, petiole 10-105 cm long, adaxially flat and slightly raised in the middle, green, glabrescent towards the middle, lepidote towards margins, abaxially covered with thick layer of persistent, yellowish stiff scales; rachis arched, 130-290 cm long, adaxially flattened in ½-3/5 of its length, ending in a well-defined 0.6 m hastula-like projection, that appears as if leaned to one side, glabrescent, abaxially convex, covered with scales like those on petiole; pinnae (47-)85-134 on each side, pinnae disposition and insertion variable, arranged in groups of 2-8(-13), and separated by 2.5-5.5 cm, inserted in many planes, or sometimes in very slightly divergent planes, sometimes pinnae inserted regularly on the basal and/or distal third of the rachis and in one plane, and sometimes almost regularly arranged in juveniles, the adaxial surface glabrous, the midrib bearing some persistent scale bases, the abaxial surface and midrib covered with yellowish linear; the most basal filiform pinnae 15-32 × 0.2-0.5 cm, basal pinnae (10th from base) 32-63 × 0.6-1.0 cm; middle pinnae 41-83 × 2.2-4.0 cm, apical pinnae 15-60 × 0.2-2.0 cm, the apical pinnae sometimes united along margins.
    Staminate inflorescences 3-4 at one time, with a peduncle 40-121 cm long, glabrescent; prophyll 40 cm long, peduncular bracts 5-7, (24-)37-188 cm long, 6-8 cm wide; rachis 52.5-76.0 cm long, with 56 branches, rachis and branches glabrescent, longest branches 27-42 cm long.
    Pistillate inflorescences 4-8 at one time; peduncle 69-185 cm long, 2 cm wide at apex; prophyll 17-45 cm long; peduncular bracts 4-6(-8), 37-180 cm long; rachis 47-93 cm long, with 33-50 branches, each subtended by a 0.2-0.6 cm long, membranaceous bract, longest branches 21-54 cm long; prophyll, peduncle and bracts covered with persistent, brown to ferrugineous scales, rachillae glabrous.
    Staminate flowers: sepals 3, elliptic, acuminate, 0.9-1.5 mm long, connate in 0.3 mm (1/3-1/5 of total length), not reaching the total length of corolla tube; petals 3, elliptic, long-acuminate, 4.5-6.0 mm long, including an acumen of 0.4-1.5 mm, connate in 2.0 mm, thin and membranaceous; stamens 6, conspicously exserted from the petals, alternating one antesepalous stamen, and one antepetalous stamen, filaments 1.0-1.5 mm long, very thick and swollen at base, inserted at basal ¼ to center of anther, anthers 4.7-7.0 mm long, anther connective not projected.
    Pistillate flowers: sepals 3, broadly triangular, acuminate, 1.0-1.5 mm long, connate in 0.5-0.8 mm (½-2/3 of total length), not reaching the corolla tube; petals 3, elliptic, acuminate, 3.5-5.0 mm long, connate up to 1.2-1.5 mm, acumen narrow, 1 mm long; staminodes 6, alternating 1 antisepalous, 1 antipetalous, filaments 1 mm long, with very broad bases like filaments of stamens, abortive anthers 2.0-2.2 mm long.
    Fruits subglobose, orange-red when ripe, 1.7-1.9 cm diam., exocarp smooth when fresh and minutely verrucose when dry, with conspicuous black lenticels in maturation, sometimes the other 2 carpels developed to some point; fruiting perianth with sepals 1.0-1.5 mm long, connate in 0.5-0.8, lobes not reaching corolla tube, petals very wide at base, 3.5-5.0 cm including a 1 mm acumen, connate in 1-2 mm, staminodes 6, filaments very broad-based, and anthers caducous.
    Seeds 1.2-1.4 cm diam. (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))A

Materials Examined

  • BOLIVIA: Chuquisaca. Provincia Luis Calvo, Municipio Villa Vaca Guzmán, Comunidad Entierrillos, Parque Nacional y Área Natural de Manejo Integrado Serranía de Iñao, sendero de subida a la Laguna de Iñao, 19°32'36.8" S 63°54'99.6" W, 1700-1800 m, 27 November 2009, J. Orías, F. Borchsenius, MJ. Sanín, R. Millanes, D. Andrade y L. Terrazas, s.n. (fr.) (HSB, COL). Cochabamba: José Carrasco Torrico, 3 Km de Siberia hacia Karahuasi, 17°48'S 64°41'W, 2400 m, 18 October 1996, M. Kessler, J. González & A. Acebey 9130 (LPB); Provincia Chapare, Cantón El Sillar, Km 87, 1830 m, 16 March 1991, L. R. Moreno & O. Moreno 84 (immat.fr) (USZ, JBSC, NY). La Paz: Province Franz Tamayo, Parque Nacional Madidi, Senda Apolo-San José de Uchupiamonas, 5 Km antes de llegar a Río 3 de Mayo, 14°34'74" S 68°20'23" W, 1735 m, 30 December 2002, T. Miranda, C. Maldonado, F. Canqui, R. Alvarez, J. Tito 302(LPB); Senda Apolo-San José de Uchupiamonas, Piñalito, 14°32'12" S 68°19'29" W, 2239 m, 25 April 2003, F.Canqui, E. Alvarez & J. Tito 351 (st.fl.) (LPB); Mojos, Carjata, sobre la senda a Queara, 14°34'14" S 68°54'4" W, 1656 m, 7 May 2007, L. Cayola, N. Chapi, G. Chive, F. Andia & T. Alvarez 2866 (sterile) (LPB); Carjata, 14°34'05"S 68°54'01" W, 1712 m, 30 June 2005, A. Fuentes, E. Cuevas, R. Cuevas & Darte 9173 (veg) (LPB); Piedra Blanca, 10,4 Km en línea recta al S de Pata, sobre el camino a Santa Cruz del Valle Ameno, 14°38'40" S 68°37'24" W, 1915 m, 11 November 2003, N. Paniagua, A. Fuentes, L. Cayola, S. Whiteheat, R. Cuevas, C. Cuevas, E. Cuevas, L. Cuevas 5869 (LPB); Province Nor Yungas, 10 km N of Caranavi on road from town, 15°40'S 67°39'W, 1500 m, 6 December 1985, A. Henderson & J. Solomon 523 (mat.fr.) (COL, K, NY); 10 km N (ca. 5 by air) of Caranavi on road from town, (aprox.) 15°47'S 67°32'W, 1400 m, 1 November 1984, M. Nee & J. Solomon 30324 (immat.fr.) (holotype, LPB; Isotypes, NY!, MO); 3.6 Km NE of Chuspipata on road to Yolosa, 16°17' S 67°48' W, 2800 m, 24 September 1986, J. Solomon 15641 (LPB); Province Sud Yungas, 39 Km de la Plazuela a La Paz, 16°38'S 67° 34'W, 2150 m, 29 September 1995, M. Kessler, J. González, K. Bach, S. Hohnwald 5683 (LPB). Santa Cruz: Province Vallegrande, Alto Seco, one hour walk south of town in cloud forest, 64° 05' W, 18° 50' S, 1900 m, 13 May 1988, A. Henderson, M. Moraes & M. Saldias 761 (st.fl.) (COL, LPB, NY); Loma Larga a 60 Km al sur de Vallegrande, NW de Loma Larga, 18°45'20"S 63°54'28" W, 1851 m, 8 April 2006, R. Hurtado 695 (LPB); 62 Km de Vallegrande a Masicuri, 2000 m, 9 July 1995, M. Kessler, J. González, K. Bach, S. Hohnwald 5208 (LPB); 4 Km de Loma Larga a Masicuri, 18°47'S 63°53'W, 1800 m, 4 April 1996, M. Kessler, Jiménez & T. Krömer 6298 (LPB); Loma Larga, camino a Masicuri 18°45'19" S 63°54'12" W, 2030 m, 15 April 2006, L. Moreno & O. Moreno 349 (JBSC); Los Sitanos, 75 Km S of Vallegrande, 18°52' S 63°57' W, 1400 m, 28 December 1989, I.G. Vargas 360(mat.fr.) (NY,); 1 Km by road S of Chujllas, 18°48'S 64°01?'W, 2125 m, 26 December 1989, M. Nee & I. Vargas C. 38446 (NY); Postrer Valle, 4 km S from town, on the way to the waterfall of the electric central, 18°32' S 63°40' W, 1750 m, 14 May 1988, M. Moraes, A. Henderson & M. Saldias 1052 (pist.fl., immat.fr.) (COL, LPB, NY); Localidad Aguaditas, entre Khazamonte y Los Sitanos, 18°43' S 64°01' W, 2580 m, 1 May 1989, I. Vargas 175 (LPB); Provincia La Florida, Samaipato, Yungas de Mairana, 2500-2600 m, 10 March 1990, L. R. Moreno 28 (NY); 5 Km SW of Yerba Buena, top of Ridge of upper reaches of Quebrada Agua Blanca, 18°01' S 64°03' W, 1900-1950 m, 23 December 1989, M. Nee & I.G. Vargas 38268 (mat.fr.); La Chapa, ca. 5 km E de la Yunga, 18°06' S 63°55' W, ca. 2200 m, 29 June 1996, M. Saldias & L. Orozco 4216, (NY); Parque Nacional Amboró, Santa Rosa de Lima 5 km W, between Santa Rosa and Rio La Pajcha, 17°52.5'S 64°12.5'W, 1670-1800 m, 3-5 May 1993, I.G. Vargas, J. Belalcazar & J. Erihsson 2270 (st.fl.) (NY, US); Parque Nacional Amboró, 8 Km by air NE of Mairana, along trails on and near ridgetop, 5.5 Km N of Campamento "La Yunga" park station, 18°03.5' S 63°55' W, 2300 m, 30 March 2002, M. Nee, M. Sundue, M. Mendoza & A. Arbeláez de Churchill 5206 (NY); Province La Florida, Las Yungas de la Negra, Km 85, Road Santa Cruz-Cochabamba, 1600 m, 9 August 1982, M.J. Balick, D.C Daly & S. López-Pérez 1376 (immat.fr.) (NY). PERU. Cuzco: Provincia Quispicanchi, en laderas cerca al río Araza, cerca de 48 km en la carretera de Quincemil a Urcos, 1680 m, 12 June 1960, H. Moore et al. 8602 (pist.fl.) (BH, USM); Región Puno, Prov. Sandía, San Juan del Oro, 14°14'19,1" S, 69°, 10', 40,5"W, 1758 m, 10 November 2009, F. Kahn, J. C. Pintaud, M. Machahua 4478 (USM). (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))A