Ceroxylon Bonpl. ex DC., Bull. Sci. Soc. Philom. Paris 3: 239 (1804)

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Distribution

Map uses TDWG level 3 distributions (http://www.nhm.ac.uk/hosted_sites/tdwg/geogrphy.html)

12 species distributed throughout the Andes, in Venezuela, Colombia, Ecuador, Peru and Bolivia.

Ceroxylon is distributed from 11° 10' N in Distrito Federal, Venezuela, to 18°52' S in Cochabamba, Bolivia;
and from 76° 30' W in Valle del Cauca, Colombia, to 63°55'W in Cochabamba, Bolivia. It occurs along the
slopes and high inter-Andean valleys of the Central and Northern Andes, from the Cordillera de la Costa
(Venezuela) to the three cordilleras of Colombia, and South to Ecuador (east and west slopes), Peru, and to the
Department of Santa Cruz in Bolivia, from 900 m to 3500 m (Fig. 18?20). The individual species have a
restricted elevation range; C. amazonicum, for example, is found only between 800-1200 m. Eight species
have elevation ranges of 2000-2500 m, corresponding to montane cloud forest. (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))B
Eleven species occurring at high elevations in the Andes from Venezuela through Colombia, Ecuador, and Peru to Bolivia. (J. Dransfield & N. Uhl & C. Asmussen & W.J. Baker & M. Harley & C. Lewis, Genera Palmarum. The evolution and classification of palms. 2008)A

Habitat

Discussion

Diagnosis

Biology And Ecology

  • Species of Ceroxylon are some of the tallest palms. They occur in premontane to low and high montane forest often among clouds for most of the time. Today trees are frequently left standing in fields where the forest has been cleared. Ceroxylon parvifrons occurs at elevations in excess of 3500 m above sea level, the palm species with the highest elevational occurrence (Borchsenius and Skov 1997). (J. Dransfield & N. Uhl & C. Asmussen & W.J. Baker & M. Harley & C. Lewis, Genera Palmarum. The evolution and classification of palms. 2008)A
  • A detailed study of the floral biology of Ceroxylon species has not been completed to date. Knudsen et al. (2001) found that the pistillate and staminate flowers of C. echinulatum (as C. alpinum subsp. ecuadorense) emitted high levels of unsaturated aliphatic hydrocarbons, with 80% similarity to those emitted by Mauritia flexuosa, and Wettinia maynensis, suggesting adaptations to pollinators with comparable sensory preferences. The dominance of these closely related volatile compounds in floral scent, regardless of the sex of the plant, is usually associated with beetle pollination (Knudsen et al. 2001). Kirejtshuk & Couturier (2009) presented an overview of species of the genus Mystrops Murray, 1864 (Nitidulidae beetles) collected on male inflorescence of C. quindiuense in Peru, including the description of several new species of Mystrops.

    Mejía-Londoño (1999) reported the following bird species as fruit dispersors of C. alpinum: sickle-winged guan (Chamaepetes goudotii), golden-headed quetzal (Pharomachrus auriceps), blue-crowned motmot (Momotus momota), toucanetes (Aulachorhynchus haematopygus, A. prasinus) as well as a single species of bat: Artibeus jamaicencis. The following were reported by the same author as potential dispersors: tawny-breasted tinamou (Nothocercus julius), chestnut wood quail (Odonthophorus hyperythrus), masked trogon (Trogon personatus), red-ruffed fruitcrow (Pyroderus scutatus), green jay (Cyanocorax yncas), thrushes (Turdus ignobilis and T. fuscater). In Venezuela, the fruits are consumed by the groove-billed toucanet (Aulacorhynchus sulcatus; Brown 1976).

    The fruits of C. quindiuense are eaten by birds like thrushes (Turdus spp.), green jays (Cyanocorax yncas), emerald toucanets (Aulacorhynchus prasinus) and by parrots (Hapalopsitaca fuertesii, H. amazonina), including the critically endangered yellow-eared parrot (Ognorrhynchus icterotis), which is closely associated with this species (Galeano & Bernal 2005) Ceroxylon ceriferum is important for wildlife, especially birds. According to C. Olaciregui (pers. comm.), in the Sierra Nevada de Santa Marta, Colombia, the dead stems of C. ceriferum are used as nesting sites by the Santa Marta parakeet (Pyrrhura viridicata), the scarlet-fronted parakeet (Aratinga wagleri), and the strongbilled woodcreeper (Xiphocolaptes promeropirhynchus). The latter species also forages on beetles on the stems and among the pinnae. Toucans (Aulacorhynchus prasinus lautus and A. sulcatus calorhynchus), the sickle-winged guan (Chamaepetes goudoti sanctaemartae), and the golden quetzal (Pharomachrus fulgidus festatus) feed on the fruits of this species.

    Karsten (1856) reported that the spectacled bear (Tremarctos ornatus) fed on the palm heart of a wax palm in Volcán de Chiles, Colombia, and Llamosas et al. (2003) reported this use for C. ceriferum in Venezuela. Henderson et al. (1995) describes that it climbs up the stems of the lower individuals to reach the crown. (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))B

Conservation

  • All species are endangered (Moore 1977). (J. Dransfield & N. Uhl & C. Asmussen & W.J. Baker & M. Harley & C. Lewis, Genera Palmarum. The evolution and classification of palms. 2008)A
  • The conservation status of species of Ceroxylon has not been globally addressed. The Red Lists of endangered species are issued by national agencies, and Ceroxylon occurs in five countries (Bolivia, Peru, Ecuador, Colombia, Venezuela). Ecuador and Colombia are the only countries to have published Red Lists for palms (Borchsenius & Skov 1999, Galeano & Bernal 2005). Of the seven species of Ceroxylon occurring in Colombia, four are under threat. Of these, one is endemic and Critically Endangered (CR; C. sasaimae), and three are non-endemic, and Endangered (EN; C. alpinum, C. quindiuense, C. ventricosum). In Ecuador, Borchsenius & Skov (1999) categorized C. echinulatum, C. ventricosum, and C. amazonicum as Vulnerable, but the last has later been re-categorized as Endangered (Valencia et al. 2000). The conservation status of the species in Bolivia (two species), and Peru (six species) is unknown.

    The most important threats to all Ceroxylon species are: a) habitat loss due to forest clearing processes; b) low regeneration rates due to cultivation and cattle farming; c) leaf extraction for religious festivities. Forest clearing for agriculture and cattle farming has severely reduced most populations of Ceroxylon, in some cases to a few individuals. These activities prevent seedlings from developing; thus, not only are population sizes reduced, but regeneration is severely hampered. Additionally, in many areas, the developing leaves of the remaining individuals were extracted for use during Palm Sunday, affecting growth and reproduction, and sometimes causing the death of the palms. Although this activity is banned in Colombia and discouraged in other countries, it still continues to threaten some Ceroxylon species, especially C. ceriferum, C. parvifrons, C. quindiuense, and C. alpinum.

    Ceroxylon species have life history traits that make the design of conservation strategies especially challenging. All of the species are dioecious, most are expected to be long-lived (as observed in C. alpinum, and C. echinulatum), and all are adapted to a specific elevation range, precipitation regime, and soil drainage. All of these factors must be considered in the development of conservation strategies for each of the twelve species.
    (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))B

Etymology

Uses

  • Stems provide wax for candles and matches; fruit are used for cattle food. Over-exploitation of young fronds for Christian religious ceremonies has seriously endangered some species. Several species have become prized but slow-growing ornamentals. (J. Dransfield & N. Uhl & C. Asmussen & W.J. Baker & M. Harley & C. Lewis, Genera Palmarum. The evolution and classification of palms. 2008)A
  • Ceroxylon palms have been mainly used for either ceremonial (religious) purposes, or for house and fence construction. Both activities are destructive and unsustainable. Other minor uses are fruit consumption by livestock (especially pigs), and usage of the cooked basal part of the peduncle of immature inflorescences for human consumption (Borchsenius et al. 1998); the fruits of C. echinulatum (Borchsenius & Moraes 2006) and C. vogelianum (Van der Eyden 2004) are reported to be eaten by humans; C. parvifrons is used for thatch in Bolivia (Moraes 2004). In the past, the stem wax of C. quindiuense was exploited and sold in local and international markets (Boussingault 1849). Locally it was used mainly for producing candles, a practice that stopped with the advent of electricity. Wax extraction persisted until World War II, when artificial wax replaced it (Brown 1976). Yet, as late as 1945?1946, Colombian statistics reported wax export to France as an important activity (Pérez-Arbeláez 1956). In Peru (Ocol, Amazonas), wax extraction persists today. Palms are felled every year during October to make torches and candles that are lit during the local festivities. (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))B

Description

  • Tall to very tall, solitary, unarmed, pleonanthic, dioecious palms. Stems smooth, usually waxy, with prominent leaf scars. Leaves moderate to large, reduplicately pinnate, neatly abscising; sheath splitting opposite the petiole at maturity, usually not forming a crownshaft, leathery; petiole channeled adaxially, rounded abaxially; rachis adaxially flat to angled, glabrous, abaxially rounded, silvery-grey tomentose; leaflets acute, single-fold, evenly spaced or clustered, usually glossy adaxially, often waxy or tomentose-scaly abaxially, midrib conspicuous, larger adaxially, no other veins evident. Inflorescences interfoliar, solitary in the leaf axil, branched to 3–4 orders; peduncle elongate; prophyll tubular, 2-keeled, flattened, open apically, incompletely encircling the peduncle abaxially; peduncular bracts several (ca. 5–7), inserted near the base of the peduncle, the lower early in development, borne singly along the rachillae, pedicellate. ones open apically, the upper 3–4 terete, beaked, completely enclosing Staminate flowers with 3 sepals connate in a low, acutely or acuminately the inflorescence in bud, splitting abaxially at anthesis, the uppermost lobed cupule; petals 3, fleshy, acute or acuminate, briefly connate basally sometimes reduced and inserted higher than the others, prophyll and with each other and with the bases of antesepalous stamen filaments, peduncular bracts with indumentum or scales; rachis bracts small, open; separate above at anthesis; stamens 6–15(–17), filaments awl-shaped, not rachillae usually flexuous or zigzag, often short, the pistillate usually inflexed at the apex in bud, anthers basifixed, bifid basally, bifid to acute shorter than the staminate, glabrous or with indumentum, bearing small, or pointed apically; pistillode minute, conic, usually minutely trifid. pointed bracts subtending the flowers. Flowers ebracteolate, open from Pollen ellipsoidal or oblate triangular, asymmetric; aperture a distal sulcus or trichotomosulcus; ectexine tectate or semi-tectate, finely rugulate-reticulate, coarsely reticulate or gemmate-reticulate (muri comprise rows of gemmae), aperture margin similar or slightly finer; infratectum columellate; longest axis ranging from 32–46 µm [5/11]. Pistillate flowers similar to the staminate but staminodes usually smaller with halberd-shaped or sagittate abortive anthers; gynoecium ovoid, trilocular, triovulate, but 2 ovules usually aborting, stigmas 3, recurved at anthesis, ovules pendulous, hemianatropous. Fruit red, orange-red, or orange to purplish-black at maturity, globose, normally 1-seeded, stigmatic remains lateral near the base; epicarp smooth or minutely roughened, mesocarp fleshy with few fibres, endocarp thin, not adherent to seed. Seed globose, hilum basal, round, raphe branches obscure, ascending from the hilum, endosperm homogenous; embryo lateral near the base. Germination adjacent-ligular; eophyll elliptic or narrowly lanceolate. Cytology: 2n = 36. (J. Dransfield & N. Uhl & C. Asmussen & W.J. Baker & M. Harley & C. Lewis, Genera Palmarum. The evolution and classification of palms. 2008)A
  • Tall to medium-sized, solitary, unarmed, dioecious, pleonanthic palms. Stem erect, smooth, covered by a layer of wax, with conspicuous leaf scars. Leaves pinnate, numerous in crown; sheath leathery, open, not arranged in a crownshaft; petiole channeled adaxially, rounded abaxially; rachis adaxially flat but turning narrowing towards apex ending as a ridge (triangular in cross-section), glabrescent, rounded and covered by grey tomentum abaxially; pinnae regularly arranged along the rachis in one plane, or clustered in groups and oriented in several planes, horizontal or pendulous, linear-lanceolate, acute, with a prominent midrib, adaxial surface usually smooth and glossy or shiny, abaxial surface covered by a white or yellowish to brown tomentum of waxy scales. Inflorescences interfoliar, 2-6 (-8), in different stages of development, solitary in the axil of each leaf, branched up to second, third or fourth order; peduncle elongate; prophyll tubular, bicarinate, flattened, incompletely encircling the peduncle abaxially and open apically; peduncular bracts 4-7, inserted near the base of the peduncle, the lower ones open apically, the upper one beaked, completely enclosing the inflorescence in bud and opening abaxially at anthesis, and sometimes with an additional smaller, thinner bract inserted distally on the peduncle; prophyll and peduncular bracts covered by thick indumentum of light brown scales; rachillae flexuous or zig-zag, glabrous or covered with indumentum, subtended by short, triangular bracts. Staminate and pistillate inflorescences can be distinguished from afar, the staminate being smaller and with adpressed rachillae close and somewhat drooping at anthesis. Pistillate inflorescences massive, with upright rachis and spreading, loosely inserted rachillae. Pistillate inflorescences green from the beginning of fruiting stage, and staminate inflorescences cream-coloured, turning yellowish after anthesis, and then brown and dry. Flowers open from early in development, solitary, pedicellate. Sepals 3, basally connate, acute to acuminate; petals 3, fleshy, briefly connate basally and shortly adnate to the bases of the antesepalous stamen/staminode filaments, acute to long-acuminate; stamens/staminodes 6-19, filaments awl-shaped, straight from early in development, anthers/abortive anthers basifixed, bifid basally, apically bifid or long-acuminate; pistil trifid, smooth to warty, pistillode minute. Fruits globose, 1.5-2.5 cm diam., red to orange-red; epicarp smooth, verrucose, reticulate or covered with prominent, irregular and acute bulges; mesocarp fleshy with few fibres; endocarp thin and papyraceous, loose from the seed; seed globose, light brown, with basal hilum, raphe branches slightly visible; endosperm homogeneous; embryo lateral near the base. Eophyll elliptic to narrowly lanceolate, covered with silver scales on abaxial surface. (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))B

Anatomy

Fossil record

Observation

  • Vergara-Chaparro (2002) evaluated various remnant populations of C. alpinum in Salento, Quindío Department (Colombia), with the use of Lefkovitch matrices to estimate the growth rate of the population. Individuals were found to produce 0.9 leaves per year in the seedling stage, 2.3 during the establishment phase, 7 during the mature-vegetative phase, and up to 11 during the mature-reproductive phase. The last phase was accompanied by an abrupt reduction in stem diameter and in internode length: 0-5 m tall palms grew 55.7 cm per year, while 35-40 m tall palms grew 20.5 cm per year. She estimated that it took a palm 57 years to start producing a stem, 83 years to reach the reproductive phase, and 140 years to grow up to 25 m; life expectancy was estimated at 213 years. The ratios among sexes were balanced in the populations studied, and the reproductive cycles were annual, with the main flowering of both sexes occurring around September and fruit maturation seven months thereafter. Each adult produced 4-12 inflorescences per year and each infructescence 2000-3000 fruits; not all inflorescences matured simultaneously, so fruits were available during much of the year.

    Rudas (1998) studied the life cycle and demography of a population of C. sasaimae, and found that the total number of adults summed to about 100, with an effective population size of less than 50 individuals, due to dioecy. For this reason, and the high level of degradation of the natural habitats this species was categorized as Critically Endangered, according to the 2001 IUCN Red List Categories and Criteria, version 3.1 (Galeano & Bernal 2005). Paredes-Ruiz (1995) studied the population structure and some aspects of growth and development of a population of C. echinulatum in the Province of Napo (Ecuador): leaf production per year in adults was estimated at 2.74 (SD = 0.86) in the forest and 2.85 (SD = 1.14) in pastures, and the production of pistillate and staminate inflorescences was 3 (sd = 2) per year, regardless of whether the individuals were growing in pastures or in the forest. Individuals growing in open pastures produced significantly more leaves per year, attained greater heights and had substantially lower regeneration rates, as can be expected in areas used for livestock grazing. The flowering cycles were annual, the main event occuring in September?January (staminate) and October?November (pistillate), suggesting an extended pollen offer throughout the whole flowering season.

    Girón-Vanderhuck et al. (2001) studied the population structure of C. quindiuense in the Central Andes of Colombia and estimated leaf production in seedlings at 4.5 per year and at 3.92 in adults; flowering was also annual with the main event occurring in March?April, and occasionally in November.

Occurrence

  • Another factor that determines the distribution of Ceroxylon is humidity. Although no quantitative studies have been carried out to correlate this factor with Ceroxylon distributions, palm stands are never found in xeric conditions. For example, C. ceriferum is restricted to the moist western slopes facing the Caribbean on the Sierra Nevada de Santa Marta but it does not occur in the drier forests of the eastern face; C. pityrophyllum is never present in the dry forest but some scarce individuals dwell in transitional to humid montane forests. Cultivated seedlings perform better at high humidity levels. Ceroxylon palms are always found in humid or moist, montane forests or surviving in pastures. The only exception to this distribution is C. parvifrons which can grow at up to 3500 m elevation.

    Reconstructing the biogeographic history of Ceroxylon is complicated for several resasons: 1) the ranges of all the species of Ceroxylon are probably relictual and hardly a fair representation of their past distributions; as 90% of the Andean forests have been eradicated (Henderson et al. 1991), most individuals of Ceroxylon are found in highly reduced populations surrounded by large pastures or eroded terrains where no palms persist, 2) collections of Ceroxylon are still incomplete, especially around the lake Titicaca area, the eastern slopes of the Peruvian Andes, and around the Táchira Deflection in southwestern Venezuelan. This is due in part to the difficulties of collecting these species, because of their size. Their dioecious nature, and the fact that an individual seldom carries fruits and flowers at the same time, means that a complete collection comprising staminate and pistillate flowers, as well as fruits, will require the collection from at least three individuals and usually several field campaigns. However, even if some sampling gaps remain, the present study encompasses field work done over more than 20 years, which has covered many previously unexplored localities. For this resason, no new species are expected in Venezuela, Colombia, Ecuador, and northern Peru.

    Trénel et al. (2007b) put forward a comprehensive theory on the biogeography of subfamily Ceroxyloideae. In that study, 5.5 Kb of DNA from three plastid and two nuclear genomic regions were used to reconstruct a phylogeny for which divergence times were estimated. Tribe Ceroxyleae was found to include two clades, one comprising Ravenea and the other Oraniopsis, Juania and Ceroxylon. Dating suggested that the Eocene-Oligocene was a geologic period of major radiation. Disjunctions in tribe Ceroxyleae most like arose during the mid-Tertiary which is too recent for the formerly proposed Cretaceous (Godwanan) vicariance to have occurred.

    Trénel et al. (2007b) suggest that repeated trans-oceanic dispersal events were involved. Ceroxyleae are, in general, adapted to cool hardy environments and therefore, the authors hypothesized that if Eocene ancestors exhibited similar ecological characteristics, austral intercontinental dispersal could have been possible. (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))B

Phenology

  • In C. alpinum, seven months pass from the time of pollination to the complete ripening of the fruit. Fruit maturation is more or less synchronized within the same area, but differs in timing between areas (Brown 1976). According to Brown & Delascio (1987), the seeds of C. ceriferum take 6-24 months to germinate. Local informants report that the fruits of C. echinulatum mature each year during November-December. In Ecuador, the locals report that C. ventricosum fruits only every 6-7 years (Doyle 233, in scheda). In some populations fruits may be available during most of the year. (Maria Jose Sanin & Gloria Geleano in Phytotaxa 34 (2011))B

Relationships

Taxonomic accounts

Use Record

Uses

  • Ceroxylon Bonpl. ex DC.: Además las palmas de cera dan su producto para antorchas y cirios. (…). Su producto se raspa con cuchillos, se funde y se cuela. Se caracteriza por su alto punto de fusión. En Colombia se aplica mucho al alumbrado doméstico y religioso. (…). En salento me informaron que los cerdos comen los frutos de estas palmas y se ceban presto. (Pérez-Arbeláez, E., Plantas útiles de Colombia. 1956)
    Use CategoryUse Sub CategoryPlant PartHuman GroupEthnic GroupCountry
    Animal FoodFodderSeedsNot identifiedN/AColombia
    FuelLightingStemNot identifiedN/AColombia
  • Ceroxylon Bonpl. ex DC.: Street trees (Livistona chinensis, Trachycarpus fortunei, Pritchardia pacifica, Washingtonia robusta, Phoenix canariensis, Roystonea regia, Parajubaea cocoides, Cocos nucifera, Ceroxylon spp., Archontophoenix alexandrae, Jubaea chilensis, Phoenix reclinata). (Balslev, H., and A. Barfod, Ecuadorean palms- an overview. 1987)
    Use CategoryUse Sub CategoryPlant PartHuman GroupEthnic GroupCountry
    EnvironmentalOrnamentalEntire plantNot identifiedN/AEcuador