Cucurbita ficifolia

Cucurbita ficifolia, popularment coneguda com chilacayote (en Mèxic, Guatemala i El Salvador), chiverre (en Costa Rica), alcayota (Chile i regió De la qual cuyana d'Argentina), cayote noroest d'Argentina, lacayote (al sur de Perú) i carabassa de cabell d'àngel, pantana o cidra (en Espanya) coneguda en el nom de Victoria o carabassa de victòria en Colòmbia és una espècie trepadora de frut comestible de la família de les cucurbitácees. És l'espècie més important de carabasses en les regions de grans altituts del Neotrópico, incloent els sèt països de Sudamérica en els que es despleguen els Andes. Ademés de la seua extensiva àrea de cultiu tradicional, esta espècie s'ha dispersat en les últimes poques centurias a lo llarc de totes les regions tropicals i subtropicals de grans altituts del món i en zones templades suaus. La seua popularitat sembla deure's a la facilitat del seu cultiu en regions fresques baix condicions extremes de sequia o d'humitat. A baixes altures li la pot cultivar com a peu d'esgall d'atres enredaderas cucurbitàcees com a melons i pepinos. Els fruts posseïxen similars qualitats nutricionals o sabor que les demés espècies cultivades de Cucurbita, pero posseïxen una longevitat molt llarga deguda a les propietats del seu clafoll.

Els usos varien en diferents regions, pero principalment és utilisada com a aliment en el mateix lloc a on li la cultiva, en sistemes agrícoles chicotets i sustentables. Si no, els fruts són venuts en mercats locals. Usos més comercials es troben en Mèxic, Costa Rica, Colòmbia, Equador i Argentina, a on és utilisada per a confites i dolces i també és cuit en plats tradicionals no dolços.

El registre arqueològic mostra que va ser extensivamente utilisada i comercialisada en temps pre-incaicos en el nort de Perú. La variació morfològica de l'espècie, si ben comparativament chicoteta per a una Cucurbita domesticada, és més àmplia en la regió de Perú a Colòmbia, lo que indica que la seua domesticació potencial va ser en esta àrea. El ancestro selvat és desconegut. (Introducció presa de TC. Andres 2006).

En el mateix gènero es troben quatre espècies en varietats cultivades (Cucurbita pepo, C. maxima, C. moschata, C. argyrosperma), pero C. ficifolia està alluntada evolutivamente d'estes, la seua polpa no posseïx el característic "sabor a zapallo" (sabor a carabassa) de les demés i diferix en l'hàbitat al que està adaptada i en atres caràcters morfològics i moleculars. No híbrida en cap atra espècie cultivada ni varietat selvada. Com en les demés cucurbitàcees, les parts herbàcees jóvens i flors també poden ser aprofitades com a verdura, els seus fruts inmaduros poden consumir-se en forma similar a una carabasseta o un zapallito, i les seues llavors també són comestibles.

Classificació i descripció

Ha segut descrita en TC. Andrés (1990), en la monografia de cucurbitàcees llatinoamericanes d'importància econòmica de R. Lira Saade (1995) i i TC. Andrés (2006).

Plantes rastreras o trepadoras, monoicas , anuals encara que persistents per un cert periodo donant l'impressió de ser perennes de vida curta, sense raïls engrossades de reserva; resistents a baixes temperatures, pero no a gelades severes; vellosas a suaument pubescentes, en alguns aguijones curts i punchants escampats en les parts vegetatives. Tallos vigorosos, llaugerament angulosos. Fulls en peciols de 5-25 cm; ovat cordados a suborbicular-cordados, en o sense taques blanques en el fes, de tres a cinc lobuladas, en lòbuls redonejats o obtusos, apiculats, el central més gran que els laterals, màrgens denticulats tres o quatre zarcillos ramificats. Flores pentámeras, solitàries, axilars. Flors masculines largamente pediceladas; càliç campanulado, de 5-10 mm de llarc i casi igual d'ample, sépals lineares, de 5-15 x 1-2 mm; corola tubular- campanulada alguna cosa eixamplada cap a la base, de 6-12 cm de llarc, groga a anaranjada pàlida, tres estams. Flors femenines en pedúnculs robusts, de 3-5 cm de llarc; ovari ovoide a elíptic, multilocular, sépals ocasionalment foliáceos i corola alguna cosa més gran que en les masculines; estil engrossat, tres estigmes lobados. Fruts de 25-50 cm llarc, globosos a ovoide-elíptics de color vert clar o obscur, en o sense ralles o franges llongitudinals blanques cap a l'àpiç, diminutamente tacats de blanc o vert i blancs o crema, polpa blanca, dolça, clafoll rígit, persistent, sense costelles. Llavors ovado-elíptiques comprimides de 15 a 12 mm de color pardo obscures o negres. Alguns fruts de tamany mijà poden contindre més de 500 llavors. Raïls primàries i adventicias fibrosas.^[1]^[2]^[3]

C. ficifolia és en el seu hàbit una típica Cucurbita anual: trepadora, enredadera per zarcillos, rastrera (arrela en els nucs en contacte en la terra), sensible a gelades, de duració anual en zones templades, zones a on al final de la primera temporada va formar fruts madurs preparats per a sobreviure l'hivern en forma de llavor, sense òrguens d'almagasenament en l'adult. En les grans altures montanyoses de clima fresc i sense gelades a on sol ser cultivada, la seua longevitat és de varis anys, longevitat que no apleguen a tindre les demés Cucurbita cultivades degut a que el temps fret, encara que no gele, resent els seus teixits i les envellix ràpidament.^{[cita 1]} A on la seua longevitat és de més d'una temporada, despuix de la seua primera temporada cobra característiques semillenyoses. No hi ha races ni cultivars nomenats,^{[cita 2]} pero sí hi ha formes "de dia curt" i "de dia llarc" (o millor dit insensibles a la duració del dia), és dir que la primera fallarà en florir al nort o al sur de la zona tórrida.^{[cita 3]}

Sent en la seua morfologia una típica Cucurbita cultivada,^{[cita 4]} a diferència de les atres quatre és relativament homogénea, en particular el frut del que s'esperaria variabilitat deguda a les pressions de selecció, és inesperadament homogéneu en forma, clafoll i polpa, i relativament fàcil de distinguir dels fruts de les demés espècies.^{[cita 5]} El color exterior pot tindre bàsicament tres patrons de color: blanc; vert obscur (a voltes en dèu bandes llongitudinals blanques que s'estenen de l'extrem proximal a l'extrem distal); o un variegado d'estos dos colors reticulats, que pugues també tindre bandes blanques. El patró és a voltes similar i confòs en el patró de colors del meló d'Alger (Citrullus lanatus).^{[cita 6]} Forma i color exterior del frut també pot ser confòs en algunes races de C. maxima (o a lo manco en la subespècie selvada, C. maxima subsp. andreana. El tamany del frut és de 15 a 50 cm de llarc, els venedors al costat de la ruta comenten que s'han vist fruts de 100 cm de llarc. No s'han associat estes diferències en el color del frut en diferències en la composició de la polpa. La polpa seca, fibrosa, de color clar, com algunes formes de C. argyrosperma.

Cucurbita ficifolia
Flor femenina.
Frut.
Full

Ademés de la longevitat i el frut, atres dos caràcters solen ser mencionats com a diferencials del restant de les Cucurbita cultivades: la forma del seu full "com la de la higuera" (Ficus carica) i les llavors negres; els dos necessiten alguns aclariments.^{[cita 7]}

Les fulles com en les demés Cucurbita: en pecíol, de nerviación pentapalmada (o pot vore's com hepta-palmada?), de gran tamany. En esta espècie són color vert obscur, d'esquena pubescente, similars al full de l'higuera, d'això deriva el seu nom científic (ficifolia, "de fulls de higuera" en llatí). Pero atres espècies de Cucurbita poden presentar fulls com de higuera: la selvada C. lundelliana, la feral C. ecuadorensis, i les cultivades C. moschata i C. pepo.^{[cita 8]} S'ha informat una forma de fulls "tipo C. moschata" que no ha tingut encara confirmació visual per part d'un taxónomo.^{[cita 9]}

La majoria de les formes són de llavors negres, pero no totes les llavors de C. ficifolia ho són, algunes són marró obscur (dark brown) i unes atres d'un color buff-colored més semblat al d'atres espècies de Cucurbita. De les demés Cucurbita, algunes races de C. moschata del nort de Sudamérica són de llavors marrons.^{[cita 10]} No hi ha diferències regionals ni races associades al color de llavor.

L'espècie també pot diferenciar-se per la forma de la llavor, per la presència de tricomas en els filaments dels seus estams , en la presència de tricomas setáceos en el brot similars als de C. maxima, i en el pedúncul dur, d'ànguls redonejats, i llaugerament estés sobre el frut en l'unió a ell, com en C. ecuadorensis i algunes races de C. moschata.

Número cromosòmic

Vint parells de cromosomes com en Cucurbita.

Orige, cultiu i distribució

Alguns autors han propost que l'orige de C. ficifolia és centroamericà o sur-mexicà-centroamericà, mentres que uns atres sugerixen que s'ubica en Amèrica del Sur, més específicament en la zona dels Andes. L'àrea de distribució d'esta espècie comprén les zones miges o altes de pràcticament totes les cordilleres o cadenes montanyoses de Llatinoamèrica, des del nort de Mèxic fins a Chile i el noroest d'Argentina, no obstant, com en el cas del seu orige, el centre de domesticació i diversificació, encara representa un enigma a resoldre, no obstant, els restants arqueològics trobats en Perú, pogueren inclinar la balança cap a esta zona. Esta espècie originària d'Amèrica Llatina es va difondre com a cultiu tant dins com fora del continent americà, en Europa, en Àfrica, en Àsia, Índia i finalment en Oceania,^[4]^{[cita 11]} els primers fruts de C. ficifolia en aplegar a Europa aparentment varen prendre una ruta des de Sudamérica a la costa de Malabar de l'Índia a lo llarc de la molt recorreguda ruta comercial portuguesa i holandesa en els sigles XVI i XVII, d'a on varen aplegar finalment a Europa.

S'ignora en exactitut la seua regió d'orige; distintes llínees d'evidència apunten a Mèxic o a la cordillera dels Andes, pero no s'ha pogut corroborar cap hipòtesis. L'evidència llingüística favoriria un orige mexicà, ya que el nom amprat casi universalment semblara ser d'orige náhuatl; no obstant, els restants arqueològics més antics conservats provenen del Perú. Aparentment va ser ben coneguda a lo llarc de la costa peruana i es hipotetizó que despuix del seu establiment pot haver segut abandonada en favor de les espècies de zapallos, conservant-se el seu cultiu sol en les zones altes montanyoses. Es desconeix la varietat selvada de la que s'haja originat, i les hipòtesis apunten a una espècie encara desconeguda, possiblement nativa de la regió oriental de la cordillera andina.^{[cita 12]}

Hui es cultiva des de Chile i el noroest d'Argentina fins al sur dels Estats Units; pero solament en les regions montanyoses fredes a on no es donen be atres espècies de cucurbitàcees. l'Índia, el Japó i les Filipines, que hui són importants productors.

És la menys intensament cultivada de les espècies comercials del gènero Cucurbita, pero potser la que mostra una distribució geogràfica més àmplia; en estat selvat no és difícil trobar-la en les zones altes (1000 a 3000 m s. n. m.) i templades del continent americà. Esta facilitat es deu en part a la seua provada resistència a varis virus que afecten a atres espècies afins, lo que fa més problemàtica l'impossibilitat d'obtindre híbrits sense mijos molt sofisticats.

C. ficifolia requerix sols humits i clima templat; preferix condicions de dia llarc, encara que en regions càlides li la cultiva tot l'any. No és resistent a les gelades en el primer any de vida.

En alguns llocs li l'ampra com a patró per a l'esgall de plantes de meló (Cucumis melo, una atra cucurbitácea).

Ambient

Els cultivares formen part de agrosistemas i es troben generalment en horts familiars, se li pot localisar en climes templats i càlit-humits.^[3]^[2]

Quan és escapada al cultiu, pot constituir-se en distints tipos de vegetació d'acort a la regió del cultivar, des de vegetació ruderal, acahuales derivats de selves tropicals, bosc mesófilo de montanya i boscs templats.^[4]

Es cultiva en varis tipos de sol, encara que preferix aquells que són capaços de retindre humitat i en bon drenage, encara que no soporta sols totalment arcillosos. Tolera sols pobres en nutrients, molt humits i poc drenados, s'adapta tant en sol en pH bàsic, neutre i àcit. És una espècie cultivada, principalment en tota Amèrica Llatina, ocupant zones d'altitut mija a alta, en climes tant càlits com templats. Per a esta espècie el ranc d'altitut és de 1000-3000 m).

Estat de conservació

És l'espècie menys diversa de totes les espècies cultivades de Cucurbita i no se sap que existixquen cultivares comercials d'ella. Des del punt de vista agronómico és possible pensar en l'existència de certa diversitat genètica per a les poblacions de C. ficifolia, degut a que es cultiva en regions uniformes en altitut pero que segurament són més diverses sobre atres factors ecològics locals, ademés de ser una espècie cultivada tant en sistemes agrícoles d'alta competència (milpas de temporal), com d'un maneig més intensiu (milpas cultivades en époques de sequia) i recentment a la detecció de coleccions resistents a malalties virals^[3]^[2].

Els bancs genètics "ex situ" han representat la forma de conservació més important, degut a que la conservació "in situ" és casi nula, encara que deu implementar-se este tipo de conservació, ya que solament d'esta forma es mantindran vius.^[3] No és una espècie registrada baix alguna categoria de risc.

Ocupació

Les flors i brots tendres de C. ficifolia s'ampren en Mèxic i atres països americans com a verdura, de manera similar a la fiore vaig donar zucca (la flor de Cucurbita pepo i Cucurbita maxima) utilisada en la cuina italiana. També s'ampra el frut inmaduro, pelat i hervir.

El frut madur en l'agregat de sucres es consumix com a dolç i s'utilisa per a elaborar begudes. En confit s'ampra per a la confecció de cabell d'àngel, un dolç elaborat acaramelando els #bri de la polpa en aixarop de sucre. En Mèxic, de la mateixa manera que la carabassa, sol preparar-se cristalizado (confitado), i es coneix com «chilacayote en dolça». En Chile, la conserva de la fruta cuita en sucre es coneix com «melada de alcayota».

En Mèxic és amprat àmpliament en la seua gastronomia quan està en la seua forma més tendra. Alguns d'estos platillo són els chilacayotes en pipián, chilacayotes en salsa verda, sopa de chilacayote, chilacayotes farcidures, etc; posant de manifest la seua versatilitat.

Les llavors, riques en lípits i proteïnes, són l'ingredient principal d'una típica darreria de la regió mexicana de Chiapas, les palanquetas. Les llavors també es mengen salades i torrades en diverses regions d'eixe país.

En Equador, el frut tendre és usat per a preparar sopa de sambo, locro de sambo i fanesca. En el madur es fa dolç de sambo. A la llavor li la coneix com pepa de sambo i és utilisada torrada i mòlta en diferents receptes.^[5]^[6]^[7]

Taxonomia

Va ser nomenada per primera volta concorde a les regles del Còdic com Cucurbita ficifolia per Bouché en 1837. La cita completa, segons (1) Bailey (1929:105) i (2) segons Lira Saade (1995:67):

(1) Cucurbita ficifolia, Bouché, in Verh. der Vore. dones Gartenb. Berlín, xii, 205 (1837).
(2) Cucurbita ficifolia, Bouché, Verh. Vereins Beförd. Gartenbaues Königl. Preuss. Staaten 12: 205. 1837. Tipo: no conegut.

Atres noms:

Cucurbita melanosperma Aparentment ya li l'utilisava en els catàlecs de llavors baix eixe nom, i segons Andrés (1990^{[cita 13]}) baix eixe nom va ser descripta per Gasparrini en 1848, nom que va ser utilisat per autors posteriors (Braun et al. 1853, Naudin 1856, 1857, 1865, 1866, uns atres), presumiblement sense coneiximent de la primera descripció. Bailey (1929:105) en la seua descripció de C. ficifolia la llista com a sinònim de Cucurbita melanosperma, Gaspar., Act. Acad. Napol. (1847), and Ann. Sci. Nat. ser. iii, ix, p.208, t. 9; A. Braun, Cat. Hort. Karls. 1824, nomen nudum. Lira Saade (1995) la llista com Cucurbita melanosperma Gasp., Rendiconti Reale Accad. Sci., Napoli 6:448. 1847.
Pepo ficifolia, Britton, Sci. Survey Porte Riu and Virgin Is (N. I. Acad. Sci.), vaig vore, 266 (1925) (o 1825?) (com llistat en Bailey 1929:105). Pepo ficifolius (Bouché) Britton, Bot. Porte Riu 6: 266. 1925. (com llistat en Lira Saade 1995)
Pepo malabaricus (Sageret 1826) no va ser un nom vàlit degut a que no estava acompanyat d'una descripció o una indicació d'on trobar la descripció, ni un espècimen. (comentari d'Andrés 1990) Lira Saade (1995) la llista com Pepo malabaricus Sageret, Mém. Acad. Agric. France 1: 102. 1827. Nomen nudum.
Cucurbita mexicana Hort. (1891) aparentment descriu una varietat de l'espècie per lo que és un sinònim del primer nom (Andrés 1990). Lira Saade (1995) la llista com Cucurbita mexicana Hort. ex Dammann Cat. 42. 1890-1891. Nomen nudum.

Noms de varietat sinonimizados.

Cucurbita ficifolia var. mexicana G. Nichols., III. Dict. Gard., Suppl. 1: 272. 1900. (pres de Lira Saade 1995)

Cucurbita ficifolia f. leucosperma Zhiteneva, Bull. Appl. Bot. Genet. & Pl. Breed., Leningrad, Suppl. 47: 301. 1930. (pres de Lira Saade 1995)
Cucurbita ficifolia f. melanosperma Zhiteneva, Bull. Appl. Bot. Genet. & Pl. Breed., Leningrad, Suppl. 47: 301. 1930. (pres de Lira Saade 1995)

Nom comú

Originària dels Andes americans, és coneguda en una varietat de noms locals entre els que estan els que semblen ser derivats del mateix terme natiu: alcayota, cayote, lacayote, chiclayo, chilacayote (en canvi no chayote), ademés d'uns atres com chiverre, i uns atres d'orige espanyol com a birla, cabell d'àngel, carabassa confitera, cidra, vitoria, victòria o zambo. El nom natiu de moltes regions sembla derivat de tzilacayotli, nom nahuátl. El nom pot haver-se propagat en el d'una varietat exitosa. El vocable Chilacayota procedix del náhuatl “tzilac” = pla i ”ayotli” = carabassa, per lo tant es pot traduir com a carabassa plana. No obstant, l'orige de la planta no és de Mèxic, segons pareix l'evidències del seu orige es troba en la zona andina de Perú.

Per país

Argentina: alcayota (en Que el seu) o cayote (en el noroest argentí)
Bolívia: lacayote
Brasil: gila-caiota
Chile: alcayota
Colòmbia: carabassa, vitoria o birla
Costa Rica: chiverre
Equador: sambo
El Salvador: chilacayote
Espanya: cidra, sidra cayote, sidra, carabassa de sidra, pantana, cabell d'àngel.
Guatemala: chilacayote, cidracayote
Hondures: chiberro
Mèxic: chilacayote, chilaca, kan
Nicaragua: toncua o toncuá
Perú: chiuche (Cajamarca), carabassa blanca (Lima), lacayote (Arequipa), chiclayo (Amazones), zambumba o sambumba (Piura)^[8]
Portugal: chila, chila-caiota
Veneçuela: cabell d'àngel

Per nom

Alcayota (en Chile i en províncies de Que el seu: Mendoza, Sant Luis i Sant Joan, Argentina)
Alcayote
Ayote (Guatemala)
Blanca (Bolívia)
Birla (Colòmbia)
Cabell d'àngel (Veneçuela i algunes zones del sur d'Espanya com les comarques del nort almeriense: Els Vélez, Almanzora...)
Carabassa
Carabassa blanca (Perú)
Carabassa chinenca (Cuba)
Cayote (en Bota, Tucumán i Jujuy, Argentina)
Chiclayo (Perú)
Chila (Portugal)
Chilaca (Mèxic)
Chila-caiota (Portugal)
Chilacayote (Mèxic, Guatemala i El Salvador)
Chiverre (Costa Rica)
Cidra (Espanya)
Cidra cayote (Espanya)
Cidracayote (Guatemala)
Gila-caiota (Brasil)
Kan (Mèxic)
Lacayote (Perú i Bolívia)
Lacayute
Mexicà
Pantana (Canàries)
Sambo (Equador)
Sambumba (Perú)
Tzilicayotli (/chili-kaiotli/) en idioma náhuatl
Toncua (Occident de Nicaragua)
Vitòria (Colòmbia)
Zambumba (Perú)
Zapallo (Veneçuela)

Notes

↑ Andres (1990): "Several characters have been incorrectly used to separate C. ficifolia from the other four domesticated species of squash, the most notable being perennial versus annual habit. Taxonomic keys to the squashes invariably delineate C. ficifolia as the only perennial cultigen in the genus (Whitaker and Bemis 1964, Whitaker and Davis 1962); however, no morphological features llaure ever given by which the duration of the plant ca be recognized. To further confuse matters, it is sometimes stated that C. ficifolia is perennial but is often cultivated as an annual (Bailey 1948, Dieterle 1976). - It is now evident that C. ficifolia does not differ in longevity from the other squash species. All domesticated species of Cucurbita have extensive fibrous root systems and indeterminate growth. Under suitable growing conditions, each of these species, with the exception of bush varieties, will continue to grow indefinitely when the stems llaure permitted to root at the nodes. Cucurbita ficifolia is tolerant of low temperatures, although it and all species of Cucurbita llaure frost-tendir (Tachibana 1987). Cucurbita ficifolia, however, is grown in the high-altitude tropics, where it will often maintain vigor through the cool winters while the other, less cold-tolerant species perish, thereby appearing to differ from the other cultivated cucurbits grown in the same region. In warmer climates, landraces of C. moschata have been reported to last for several years (Morton 1975), although this species is generally reported to be an annual. All of the domesticated squash species llaure usually grown as annuals, but in their native habitats, i.i., the presumed habitat of their wild progenitor, they may grow spontaneously as short-lived herbaceous perennials."
↑ Andres (1990): "What little variation in C. ficifolia appears to occur throughout its range. The buff-seeded form, occurs througout the range, usually in the same field with black-seeded plants. The fruits have basically three different color patterns that occur throughout Mexico, Central America, and South America (Figure 2): all white; a distinct reticulated pattern of green on white, sometimes with tingues radial stripes spreading from the apex to the base; and dark green, without reticulations but sometimes also with white llongitudinal stripes. Slight variations of the reticulations and stripes occur, but nothing like the wide assortment of color variants, warts, corky outgrowths, and shapes and sizes that occur in the fruits of the other domesticated species. There is no association between seed color and fruit color. Uses of these various forms of C. ficifolia llaure generally the same and no named cultivars have been recognized."
↑ Andres (1990): "Whitaker and Bohn (1950) report that C. ficifolia seems to require a short photoperiod for flowering, whereas the other domesticated species llaure insensitive to day lenght. While this is generally true, not all accessions of C. ficifolia require short-days for flowering, and many landraces of other Cucurbita species llaure short-day plants. There llaure accessions of C. ficifolia that produïx fruits at the extreme latitude of Norway (Janson 1923). On the other hand, there llaure accesions of squash, primarily in the species C. ficifolia and C. moschata, that do not flower north or south of the torrid zone."
↑ Andres (1990): "Biosystematic evidence indicates that the species is not as highly distinguished morphologically and genetically from the other domesticated squashes as was onze believed; it does not differ from these other species in having perennial rather than annual growth habit as has been widely reported, and it may be hybridized to them by the use of embryo culture."
↑ Andres (1990): "In contrast (with C. pepo, C. maxima, C. moschata, C. argyrosperma), C. ficifolia is relatively homogeneous in morphology and genetic composition. This facilitates recognition of the species but has given a misleading impression of its uniqueness. The fruits, which llaure the part of the plant most likely to have undergone diversification as a result of human selection, llaure relatively uniform in shape, rind and flesh characteristics. This is in marked contrast to the other species of squash, in which numerous cultivars of striking variation exist."
↑ Andres (1990): "Other morphological features of C. ficifolia have been adequately described elsewhere (Bailey 1937, Bailey 1948, Bukasov 1930, Naudin 1856, Whitaker and Bohn 1950). The stem trichomes of C. ficifolia llaure setaceous, like those of C. maxima. The peduncle is hard, smoothly angled, and slightly flaring at the attachment to the fruit, like that of C. ecuadorensis and some landraces of C. moschata. The exterior color and shape of the fruit llaure very similar to some landraces of C. maxima; and the fruit flesh is white, somewhat dry, and coarsely fibrous, like some forms of C. argyrosperma. The fruits of C. ficifolia, which range from 15 to 50 cm long, with reports by roadside vendors of up to 100 cm long, llaure sometimes even confused with watermelon, Citrullus lanatus (Thunb.) Matsum. & Nakai, a case of convergent rather than parallel evolution."
↑ Andres (1990): "...In summary, of the three characters most often used in taxonomic keys to identify C. ficifolia, one is misinterpreted (perennial growth habit), one is not unique to C. ficifolia (leaf shape), and the third is not always true for C. ficifolia (black seeds)."
↑ Andres (1990): "A final character that is often mistakenly used to distinguish C. ficifolia from the other species of Cucurbita llaure the leaves shaped like those of Ficus carica L., hence the specific epithet. Other species, such as C. lundelliana L. H. Bailey, C. ecuadorensis Cutler & Whitaker, and some landraces of C. moschata and C. pepo may also have figlike leaf shapes. Furthermore, most species of Cucurbita and many other related genera llaure heterophyllous, with clapix-developing leaves generally more deeply lobed than those produced early in the growth cycle. Unfortunately, most herbarium specimens of cucurbits include only one stage of leaf development. This variation in leaf shape has perhaps contributed to an excesive splitting of the number of described species of Cucurbita and led Bailey (Bailey 1948) to believe there was another undescribed pecies of black-seeded gourd.Nota de l'autor: "Bailey grew the unidentified accession with black seeds in Ithaca, New York, but the plants were evidently sensitive to a short-day photoperiod and never produced fruits. The voucher specimens indicate that they llaure C. ficifolia.""
↑ Andres (1990): "Several investigators have attempted to cross C. ficifolia with the other cultivated species. Successful hybrids have been produced in a very low percentage of the pollinations between C. ficifolia and C. maxima, C. moschata, and C. pepo (Drude 1917, Grebenscikov 1965, Naudin 1865, Van Eseltine 1936, Weiling 1955, Whitaker 1956, Whitaker and Davis 1962, T.C. Andres, unpublished data). Only partially developed embryos form, and embryo culture must be employed to obtain subsequent generations. Unfortunately, sterility barriers have thus far prevented obtaining progeny beyond the F1 and first backcross generations. Variable results, however, have been obtained when different cultivars of a species llaure used. It is hoped that more compatible cultivar combinations may exist. Whitaker and Bemis (1964) state that a fig-leafed form of C. ficifolia is probably allied with C. pepo or C. maxima, whereas a form with moschata-like leaves is probably closely related to C. moschata. Evidence for these two forms was not documented nor have they been observed by the author."
↑ Andres (1990): "One character used to delineate C. ficifolia is the color of its seeds. It is the only species of Cucurbita with black seeds, but not all C. ficifolia seeds llaure black; some may be dark brown, while others llaure more or less pale buff-colored like those of most other species of Cucurbita (Figure 1). Bukasov (Bukasov 1930) listed two forms of C. ficifolia based on the seed colors, C. ficifolia f. melanosperma and C. ficifolia f. leucosperma. There llaure landraces of C. moschata grown in northern South America that also have dark brown seeds."
↑ Andres (1990) p.109-110: "Origin and Distribution. Like all other species of Cucurbita, C. ficifolia originated in the Americas. It is presently grown in many tropical highland regions of the world, including south-central Àsia and the Philippines (Herklots 1972). However, it is primarily used under sustainable agriculture systems in Latin America, where personal observations and herbarium records show that it is generally grown between 1000 and 2800 m above siga level. The Latin American distribution of C. ficifolia exceeds that of all the other cultivated Cucurbita species, extending from the highlands of northern Mexico, through Central America, and throughout Andean America from Colòmbia and Veneçuela to central Chile and northwestern Argentina. The climate in these areas is generally very moist and too cool for any other species of squash to grow, except for some short-season landraces of C. pepo and C. maxima. - The first fruits of C. ficifolia to reach Europe apparently took a circuitous route from South America to the Malabar coast of Índia along the much traveled Portuguese and Dutch trade routes in the 16th and 17th centuries, before finally reaching Europe (Merrill 1954). Bouché (1837) did not know where his cultivated material came from, but vernacular names such as melon de Malabar (Malabar melon), courge de Siam (Siam squash), and Anguriankurbis (Angora squash) suggested a southern Asian origin. - In 1854 a well-documented shipment of yaks was introduced to France from the border of Tibet with Índia. In Shanghai a large quantity of C. ficifolia fruits were loaded to serve as fodder for these animals during the long ocean voyage. More than a year after the voyage ended many of these fruits were left over and still intact at the Muséum d'Histoire Naturelle, Paris. They became a curiosity in European botanical gardens. This convinced the botanists of the clave and for many years afterward that C. ficifolia was native to the Orient (Bailey 1937, Bukasov 1930, Cogniaux 1881, Naudin 1857). However, De Candolle (1886) doubted this because all known perennial Cucurbita species were native to North America. He assumed that only the "annual" species were of Old World origin. - The etymological, ethnobotanical, and archaeological evidence prove that C. ficifolia is of American origin. In Àsia, C. ficifolia is primarily used only for livestock feed, and no Sanskrit name exists, whereas in Latin America there llaure archaeological records and many local names and uses of the plant. The most frequently used vernacular name for C. ficifolia in Mexico and parts of Central America is chilacayote (also spelled chilacayota, chilacayoti, chilacayotl, and other variations), probably a corruption of tzilacayotli, but other names include alcayota, pachayota, silacayote, cuicuilticayotli, cidracayote, cidra, mail and chiberre, chibesse, or chiverre (the latter three names used only in Costa Rica) (Bukasov 1930, Robelo 1904, Rose 1899, Watson 1887). Martínez (1979) lists several additional Mexican names of generally Nahuatl origin for C. ficifolia. In Colòmbia, C. ficifolia is usually referred to as anjama, victòria, or vitoriera; in Veneçuela, the Quechuan name zapallo is generally used for all Cucurbita; and in Equador, zambo (zambu, sambo or tambo) is used. The names used in Bolívia, Peru, and northern Argentina, in the Quechua and Aymara language areas, llaure surprisingly similar if not identical to the Mexican names: lacayote or lacahuiti, silacayote, alcayote or alcallota, cayote, and tintimoro. "
↑ Andres (1990) p.110-114: "While C. ficifolia was clearly cultivated in Mexico prior to the Spanish conquest, it is uncertain where it was first domesticated in its extensive Latin American range. The etymological evidence is inconclusive at best in this regard The Nahuatl names could be derived from or precedix the similar sounding Quechuan and Aymara names. For example, Francisco Hernández in the 16th century was the first to clearly descriu the chilacayote in Mexico (see Urbina 1902-1903). He speculated that the Nahuatl name, tzilacayotli, is derived from either the words tzilac-ayotli, meaning "smooth carabassa or squash", or tzilictic-ayotli, meaning "squash that resonates or sounds" when struck. The name cuicuilticayotli literally means the "painted squash". Even if Hernández is correct, these names may still be derivations from South America that have taken on slightly new meanings in Nahuatl by folk etymology.
Sahagún (1956), a Franciscan friar, recorded in the 16th century that the hollowed-out fruit of tzilacayotli was used by the Aztecs as a bowl to contain offerings, such as pulque, for their deities. The bowl was plagau in front of the image or images, and with their marbled appearance, were said to be made of precious stones which the Aztecs called chalchihuitl. This religious practice suggests an ancient use, if not origin, in Mexico and subsequent spread to the south.
The archaeological record tells a different story. Numerous seed remnants and pedicels of C. ficifolia have been recovered from archaeological sites along the central to northern coastal desert of Peru. These have been recovered from several horizons spanning several thousand years, beginning from a pre-ceramic, pre-maize horizon dated at 3000 B.C. (Bonavia 1982, Towle 1961). In contrast, no definitive archaeological specimen of C. ficifolia has been found north of South America. A single seed, recovered from the Valley of Oaxaca, dated at 700 A.D., and tentatively identified as C. ficifolia (Whitaker 1981a, Whitaker and Cutler 1971), appears instead to be C. pepo.
Despite the archaeological record, C. ficifolia is generally considered to have originated in Mesoamerica, specifically southern Mexico and Guatemala (Bukasov 1930, Hurd et al. 1971, Sauer 1969, Vavilov 1949-1950, Whitaker 1980, Whitaker 1981b). This conclusion is based on several lines of circumstancial evidence in addition to that already mentioned and suggests a pattern of dispersal evident in the often closely associated crops maize, beans, and the squash species, C. pepo. On the basis of the estimated clave taken for C. ficifolia to have diffused from Mesoamerica to Peru by 3000 B.C., Cutler and Whitaker (1961) concluded that it must be the oldest cultivated plant in the Americas.
Sauer (1969) noted that C. ficifolia is absent today from the hot, dry lowland region where the archaeological specimens were found. He concluded from this that C. ficifolia must be an ancient yet still primitive cultigen that was dropped from cultivation except in the cool high country when better squashes were developed. He did not consider the possibility that these archaeological squashes were not autochthonous but were instead brought down from farms at higher altituts. Cohen (1977) presented evidence that the early Peruvian coastal cultures #dependre primarily on resources from the ocean and trading with the Andean cultures. Agriculture was initially of lesser importance in the lowlands than at the higher elevations. The earliest archaeological seeds of C. ficifolia llaure the same size as those today, which suggests that it was an imported domesticate.
Hurd et al. (1971) used two genera of solitary bees (Peponapis and Xenoglossa), commonly known as squash and gourd bees, to help discern the species relationships in the genus Cucurbita. These bees depend almost exclusively on squash flowers for pollen and nectar for themselves and their larvae, and therefore appear to have coevolved with the cucurbits. The center of diversity of these bees occurs in southern Mexico, where the greatest concentration of Cucurbita species is located.
There is one species of squash bee, P. atrata (Smith), which is reported to be restricted to the pollen of C. ficifolia (Hurd et al. 1971). It is found only in the highlands of Mexico and adjacent Central America and not in South America. There llaure other more generalized species of squash and gourd bees, which occur throughout the Americas and use the pollen of all the cultivated species of squash, including C. ficifolia. The absence of P. atrata in South America is used as evidence to support the hypothesis that C. ficifolia was introduced by humans into South America from Mexico (Hurd et al. 1971). This is based on the assumption that P. atrata did not concordantly extend its range into South America with C. ficifolia because it was unable to cross the intervening lowland areas. Before reaching a definitive conclusion about human dispersal of C. ficifolia based on the distribution of its pollinators, a thorough sampling needs to be conducted on the species of pollinators visiting C. ficifolia flowers throughout its range, including South America. Peponapis atrata may not be as monolectic as reported in the literature; a female P. atrata was collected from a flower of C. argyrosperma in Chiapas, Mexico in December, 1985 (specime collected by T.C. Andres and species determined by W.I. LaBerge; deposited in the Cornell University Insect Collection, lot number 1166).
The hypothesis resulting from the squash and gourd bee study of Hurd et al. (1971) llaure not conclusive. While they llaure derived from extensive field observations, they have not been tested experimentally. The study did not demonstrate whether the various species of bees coevolved with specific species of Cucurbita or with the genus in general. The latter is the rule among oligolectic bees (G.C. Eickwort, pers. comm., 1988). The fact that these bees and Cucurbita species share centers of diversity may not be due to their coevolution but rather to other causes such as their independent adaptation to the extreme habitat diversity of the region. Experimental tests need to be conducted to determine if species such as P. atrata, given the choice between C. ficifolia and other species of Cucurbita, llaure able to differentiate between them.
Hurd et al. (1971) stated that the pollen collecting devices of the bees llaure species-specific, being adapted to carry the pollen grains of different Cucurbita species, which vary in size and structure. However, Andres (1981) showed by light and scanning electron microscopy that the pollen exine characteristics of Cucurbita do not show significant variation between species and therefore probably do not play an important role in the evolution of the pollinators.
There llaure also few consistent morphological differences in the flowers of Cucurbita species. For example, the ultraviolet patterns, i.i. "nectar guides", llaure indistinguishable among Cucurbita species (T.C. Andres, unpublished observation). Whether the bees llaure able to cue in on other interspecific differences in the cucurbit flowers has yet to be determined.
Whitaker (1980, 1981b) and Whitaker and Davis (1962) also assume that C. ficifolia was domesticated in Mesoamerica rather than South America. This conclusion is based on compatibility data and the distribution of the possible wild progenitors. Cucurbita ficifolia and C. moschata llaure the only domesticated species of squash with no known logical wild ancestor. Whitaker (1980) suggests that C. ficifolia may be derived from the wild mesophytic species, C. lundelliana or C. martinezii L. H. Bailey. Evidence from comparative morphology, geographic distribution, ecology, and genetic relationships do not, however, indicate a very close relationship between these wild species and C. ficifolia. Cucurbita lundelliana is the only wild mesophytic cucurbit that has been reported to cross with C. ficifolia, but attempts to obtain subsequent generations have failed because of sterility factors (Whitaker 1980). Furthermore, pollen fertility was reported to be higher in the F₁ hybrids between C. lundelliana and the other domesticates than between C. lundelliana and C. ficifolia (Whitaker 1956). Neither wild species occurs in the cool highland areas where C. ficifolia is grown; C. lundelliana is confined to the Yucatan peninsula, which includes a part of the countries of Mexico, Belize, and northern Guatemala, whereas C. martinezii extends from southern Tamaulipas, through Veracruz to northern Chiapas, Mexico. These two species llaure closely related and both have small gray colored (green when wet) seeds that bear little resemblance to those of C. ficifolia.
A recent extensive crossing program showed that when C. ficifolia is used as the pistillate parent it may be crossed with the interfertile xerophytic species, C. foetidissima and C. pedatifolia, without the need for embryo culture (Andres 1987). Sterility barriers prevented obtaining subsequent generations. These two xerophytic species and C. radicans share with C. ficifolia pubescent filaments but otherwise bear little resemblance to C. ficifolia in morphology and ecology. In the study on the relationships of Cucurbita species with the squash and gourd bees (Hurd et al. 1971), however, C. pedatifolia was grouped with C. ficifolia.
In South America there llaure less likely candidates for the wild progenitor of C. ficifolia. Only two wild species of Cucurbita llaure endemic to South America. One, C. andreana Naud., is fairly certain to be the wild progenitor of C. maxima (Millán 1945). The other, C. ecuadorensis, has some phenetic similarity to C. ficifolia, such as the similar leaf shape already mentioned and an unusually large ruit for a wild cucurbit, but crossing studies indicate it is more closely aligned with C. maxima and the C. lundelliana group (Cutler i Whitaker 1969).
There is perhaps more persuasive evidence suggesting that C. ficifolia originated in northern South America rather than Mesoamerica. Intensive searches for a compatible wild progenitor of C. ficifolia in Mexico have produced no evidence of any such populations. Although less intensive searches have been conducted south of Mexico, there llaure sketchy reports of possible wild or at least spontaneous populations. Dieterle (1976) reports populations of C. ficifolia in some highland localities of Guatemala to "have become thoroughly naturalized and look like native plants." The difference between wild and naturalized populations of cucurbits is often ambiguous or at least difficult to determine. In Colombian markets Cardenas (1969) reported seeing "victòria" squashes with "necks like a bottle". This shape has not been found elsewhere and, if this report is correct, may represent a primitive character or a derived morph from an ancestral population. In Bolívia, Cardenas (1969) reported that C. ficifolia plants with viviparous seeds appear wild because they llaure spontaneous and hanging off the mountain sides. No description or documentation is provided on the morphology of the plants. Granat (1931) stated that C. ficifolia originated in Bolívia, but he did not provide any evidence. Agronomists at Inquisivi, Departament La Pau, reported a wild form of C. ficifolia, but this has not yet been confirmed (M. Nee, pers. comm., 1988).
The hypothesis of an Andean origin and domestication of C. ficifolia rather than a Mexican origin explains the archaeological record and does not neccesitate a change in habitat as Sauer proposed (Sauer 1969). Cucurbita ficifolia is most likely uniquely adapted to cool, moist conditions because it evolved in a habitat characterized by such a climate. The preponderance of short-day flowering accesions points to an area of origin within the torrid zone. If domestication occurred in the northern Andes around or before 3000 B.C., there was ample clave for a gradual spread of the crop to the north and use by the Aztecs in Mexico beginning in the 12th century. A similar situation occurred in C. pepo, which spread northward during pre-Columbian claves from central and northern Mexico to southwestern and eastern Unites States to as far north as adjacent Canada (Whitaker and Bohn 1950, Whitaker and Davis 1962). However, in the case of C. pepo, this dispersal may have occurred several millennia before C. ficifolia was even domesticated, thus allowing sufficient clave for secondary diversification of cultivars to occur within the new range (see Decker, this volume). Other examples of New World crops involved in very early long distance cultural exchanges include peanuts and guava, which spread from Peru to Mexico by 200 B.C., and maize, which spread from Mexico to Peru by around 3000 B.C. (Flannery 1973).
It is conceivable that there llaure still extant wild conspecific populations of C. ficifolia in some isolated cool forests on the eastern flank of the Andes in Veneçuela, Colòmbia, Equador, Peru, or Bolívia. Furthermore, in such an area this species may contain greater genetic diversity than has been found elsewhere. Germplasm collecting and evaluations need to be conducted in this region. There llaure few dry caves in this region where archaeological preservation could have occurred, but those fruits that were traded with cultures along the dry Peruvian coast may have been preserved. This type of trading of the highland grown C. ficifolia for lowland crops, such as C. moschata, still takes plau today in much of Latin America."
↑ Andres (1990): "Nomenclature. Cucurbita ficifolia was described by Bouché in 1837 (Bouché 1837), however, the species was first reported in Europe and named C. melanosperma by Braun in 1824 in a seed catalog from Carlsruhe Garden, Germany, but without a description or indication of plau of origin (Bois i Gerome 1920, Walpers 1857). Gasparrini (1848), apparently unaware of either earlier publication, also used the name C. melanosperma in his 1848 description of the species. Braun et al. (1853) later described C. melanosperma in detail, and this name was subsequently used by Naudin (1856,1857,1865,1866) and others. However, the name C. ficifolia has priority. In addition to C. melanosperma, another name precediu Bouché's. Pepo malabaricus was proposed by Sageret in 1826 (Sageret 1826, Sageret 1827) for the "melon du Malabar", a vernacular name used in France for the fig-leaf gourd. Because there was no accompanying description or reference to a publication or a specimen, the name was not validly published (Bailey 1929). In the clapix 1800's C. mexicana Hort., originally described in the Dammann Catalogue (1890-1891), was cited (Anonymous 1891) as a unique species from Mazatlan, Mexico, "very similar to C. melanosperma, but with the leaf of a different shape, and flower of a different hue, the seeds llaure large and black". There is nothing in the description to indicate this name to be anything but a synonym of C. ficifolia, the only known species of Cucurbita with black seeds. Two species have been incorrectly ranked as varieties or cultivars of C. ficifolia in the past, including C. argyrosperma Huber (Bailey 1937, Hutchins i Sando 1941) and C. moorei L. H. Bailey (Filov 1966), which is probably a synonym for C. pedatifolia L. H. Bailey (T. C. Andres, unpublished observation).

Referències

Bibliografia

Hernández Bermejo, J. I.; #León, J. (eds.) (1994). Neglected crops: 1492 from a different perspective, Roma: FAO.
USDA, ARS, National Genetic Resources Program. GRIN. National Germplasm Resources Laboratory, Beltsville, Maryland. https://web.archive.org/web/20141110134226/http://www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?12589 (8 feb 2008)

Enllaços externs

http://www.conabio.gob.mx/conocimiento/bioseguridad/pdf/20833_especie.pdf
Enciclovida té un artícul sobre [1]Cucurbita ficifolia.
[2]Naturaliste.

UC.cl («alcayota» en el Proyecte Hortalices, de l'Universitat Catòlica de Chile).
Web.Chasque.net (fotos i preparació del dolç de cidra).
Este artícul conté una traducció derivada de «Cucurbita ficifolia» de Wikipedia en castellà publicada baix la Llicència de documentació lliure de GNU i la Llicència Creative Commons Reconeiximent-CompartirIgual 4.0 Internacional.

↑ Hernández Bermejo J. I. i J. #León. 1992. Cultius Marginats, una atra perspectiva de 1492. Organisació de les Nacions Unides per a l'Agricultura i l'Alimentació. Roma. 345 p.
↑ ^2,0 ^2,1 ^2,2 National Research Council (1989). Squashes and Their RelativesLost Crops of the Incas: Little-Known Plants of the Andes with Promise for Worldwide Cultivation
↑ ^3,0 ^3,1 ^3,2 ^3,3 R. Lira S. (1995). Estudis Taxonòmics i Ecogeográficos de les Cucurbitaceae Llatinoamericanes deImportancia Econòmica. Systematic and Ecogeographic Studies on Crop Genepools. 9.. International Plant Genetic Resources Institute, Roma, Itàlia
↑ ^4,0 ^4,1 R. Lira S., I. Rodríguez-Arévalo (1999). Cucurbitaceae A.L. Juss.. En: Flora del Valle de Tehuacán-Cuicatlán. Fascículo 22. Universitat Nacional Autònoma de Mèxic. Mèxic, D.F
↑ «a-les-pepas.html Del sambo són útils fins a les pepas». El Comerç. Consultat el 14 de juliol de 2018.
↑ Hora, Diari La. «Un emprendimiento lojano es basa en la pepa de sambo - L'Hora», L'Hora Notícies d'Equador, les seues províncies i el món. Consultat el 14 de juliol de 2018 (en es).
↑ «Sambo - Patrimoni Alimentari» (en es). patrimonioalimentario.culturaypatrimonio.gob.ec. Archivat des de l'original, el 14 de juliol de 2018. Consultat el 14 de juliol de 2018.
↑ Hocquenghem, Anne-Marie; Monzon, Susana (4 d'abril de 2013). La Cuina Piurana: Ensaig d'antropologia de l'alimentació (en es), Institut français d’études andines. ISBN 9788489302235. Consultat el 21 de giner de 2019.

[Andres_1990_annual-4] Andres (1990): "Several characters have been incorrectly used to separate C. ficifolia from the other four domesticated species of squash, the most notable being perennial versus annual habit. Taxonomic keys to the squashes invariably delineate C. ficifolia as the only perennial cultigen in the genus (Whitaker and Bemis 1964, Whitaker and Davis 1962); however, no morphological features llaure ever given by which the duration of the plant ca be recognized. To further confuse matters, it is sometimes stated that C. ficifolia is perennial but is often cultivated as an annual (Bailey 1948, Dieterle 1976). - It is now evident that C. ficifolia does not differ in longevity from the other squash species. All domesticated species of Cucurbita have extensive fibrous root systems and indeterminate growth. Under suitable growing conditions, each of these species, with the exception of bush varieties, will continue to grow indefinitely when the stems llaure permitted to root at the nodes. Cucurbita ficifolia is tolerant of low temperatures, although it and all species of Cucurbita llaure frost-tendir (Tachibana 1987). Cucurbita ficifolia, however, is grown in the high-altitude tropics, where it will often maintain vigor through the cool winters while the other, less cold-tolerant species perish, thereby appearing to differ from the other cultivated cucurbits grown in the same region. In warmer climates, landraces of C. moschata have been reported to last for several years (Morton 1975), although this species is generally reported to be an annual. All of the domesticated squash species llaure usually grown as annuals, but in their native habitats, i.i., the presumed habitat of their wild progenitor, they may grow spontaneously as short-lived herbaceous perennials."

[Andres_1990_fruit_color-5] Andres (1990): "What little variation in C. ficifolia appears to occur throughout its range. The buff-seeded form, occurs througout the range, usually in the same field with black-seeded plants. The fruits have basically three different color patterns that occur throughout Mexico, Central America, and South America (Figure 2): all white; a distinct reticulated pattern of green on white, sometimes with tingues radial stripes spreading from the apex to the base; and dark green, without reticulations but sometimes also with white llongitudinal stripes. Slight variations of the reticulations and stripes occur, but nothing like the wide assortment of color variants, warts, corky outgrowths, and shapes and sizes that occur in the fruits of the other domesticated species. There is no association between seed color and fruit color. Uses of these various forms of C. ficifolia llaure generally the same and no named cultivars have been recognized."

[Andres_1990_dia_curt_dia_llarc-6] Andres (1990): "Whitaker and Bohn (1950) report that C. ficifolia seems to require a short photoperiod for flowering, whereas the other domesticated species llaure insensitive to day lenght. While this is generally true, not all accessions of C. ficifolia require short-days for flowering, and many landraces of other Cucurbita species llaure short-day plants. There llaure accessions of C. ficifolia that produïx fruits at the extreme latitude of Norway (Janson 1923). On the other hand, there llaure accesions of squash, primarily in the species C. ficifolia and C. moschata, that do not flower north or south of the torrid zone."

[Andres_1990_típica_Cucurbita-7] Andres (1990): "Biosystematic evidence indicates that the species is not as highly distinguished morphologically and genetically from the other domesticated squashes as was onze believed; it does not differ from these other species in having perennial rather than annual growth habit as has been widely reported, and it may be hybridized to them by the use of embryo culture."

[Andres_1990_homogeneous-8] Andres (1990): "In contrast (with C. pepo, C. maxima, C. moschata, C. argyrosperma), C. ficifolia is relatively homogeneous in morphology and genetic composition. This facilitates recognition of the species but has given a misleading impression of its uniqueness. The fruits, which llaure the part of the plant most likely to have undergone diversification as a result of human selection, llaure relatively uniform in shape, rind and flesh characteristics. This is in marked contrast to the other species of squash, in which numerous cultivars of striking variation exist."

[Andres_1990_other_morpho-9] Andres (1990): "Other morphological features of C. ficifolia have been adequately described elsewhere (Bailey 1937, Bailey 1948, Bukasov 1930, Naudin 1856, Whitaker and Bohn 1950). The stem trichomes of C. ficifolia llaure setaceous, like those of C. maxima. The peduncle is hard, smoothly angled, and slightly flaring at the attachment to the fruit, like that of C. ecuadorensis and some landraces of C. moschata. The exterior color and shape of the fruit llaure very similar to some landraces of C. maxima; and the fruit flesh is white, somewhat dry, and coarsely fibrous, like some forms of C. argyrosperma. The fruits of C. ficifolia, which range from 15 to 50 cm long, with reports by roadside vendors of up to 100 cm long, llaure sometimes even confused with watermelon, Citrullus lanatus (Thunb.) Matsum. & Nakai, a case of convergent rather than parallel evolution."

[Andres_1990_caract_dif-10] Andres (1990): "...In summary, of the three characters most often used in taxonomic keys to identify C. ficifolia, one is misinterpreted (perennial growth habit), one is not unique to C. ficifolia (leaf shape), and the third is not always true for C. ficifolia (black seeds)."

[Andres_1990_fig_leaf-11] Andres (1990): "A final character that is often mistakenly used to distinguish C. ficifolia from the other species of Cucurbita llaure the leaves shaped like those of Ficus carica L., hence the specific epithet. Other species, such as C. lundelliana L. H. Bailey, C. ecuadorensis Cutler & Whitaker, and some landraces of C. moschata and C. pepo may also have figlike leaf shapes. Furthermore, most species of Cucurbita and many other related genera llaure heterophyllous, with clapix-developing leaves generally more deeply lobed than those produced early in the growth cycle. Unfortunately, most herbarium specimens of cucurbits include only one stage of leaf development. This variation in leaf shape has perhaps contributed to an excesive splitting of the number of described species of Cucurbita and led Bailey (Bailey 1948) to believe there was another undescribed pecies of black-seeded gourd.Nota de l'autor: "Bailey grew the unidentified accession with black seeds in Ithaca, New York, but the plants were evidently sensitive to a short-day photoperiod and never produced fruits. The voucher specimens indicate that they llaure C. ficifolia.""

[Andres_1990_cross-12] Andres (1990): "Several investigators have attempted to cross C. ficifolia with the other cultivated species. Successful hybrids have been produced in a very low percentage of the pollinations between C. ficifolia and C. maxima, C. moschata, and C. pepo (Drude 1917, Grebenscikov 1965, Naudin 1865, Van Eseltine 1936, Weiling 1955, Whitaker 1956, Whitaker and Davis 1962, T.C. Andres, unpublished data). Only partially developed embryos form, and embryo culture must be employed to obtain subsequent generations. Unfortunately, sterility barriers have thus far prevented obtaining progeny beyond the F1 and first backcross generations. Variable results, however, have been obtained when different cultivars of a species llaure used. It is hoped that more compatible cultivar combinations may exist. Whitaker and Bemis (1964) state that a fig-leafed form of C. ficifolia is probably allied with C. pepo or C. maxima, whereas a form with moschata-like leaves is probably closely related to C. moschata. Evidence for these two forms was not documented nor have they been observed by the author."

[Andres_1990_seed_color-13] Andres (1990): "One character used to delineate C. ficifolia is the color of its seeds. It is the only species of Cucurbita with black seeds, but not all C. ficifolia seeds llaure black; some may be dark brown, while others llaure more or less pale buff-colored like those of most other species of Cucurbita (Figure 1). Bukasov (Bukasov 1930) listed two forms of C. ficifolia based on the seed colors, C. ficifolia f. melanosperma and C. ficifolia f. leucosperma. There llaure landraces of C. moschata grown in northern South America that also have dark brown seeds."

[Andres_1990_americà-15] Andres (1990) p.109-110: "Origin and Distribution. Like all other species of Cucurbita, C. ficifolia originated in the Americas. It is presently grown in many tropical highland regions of the world, including south-central Àsia and the Philippines (Herklots 1972). However, it is primarily used under sustainable agriculture systems in Latin America, where personal observations and herbarium records show that it is generally grown between 1000 and 2800 m above siga level. The Latin American distribution of C. ficifolia exceeds that of all the other cultivated Cucurbita species, extending from the highlands of northern Mexico, through Central America, and throughout Andean America from Colòmbia and Veneçuela to central Chile and northwestern Argentina. The climate in these areas is generally very moist and too cool for any other species of squash to grow, except for some short-season landraces of C. pepo and C. maxima. - The first fruits of C. ficifolia to reach Europe apparently took a circuitous route from South America to the Malabar coast of Índia along the much traveled Portuguese and Dutch trade routes in the 16th and 17th centuries, before finally reaching Europe (Merrill 1954). Bouché (1837) did not know where his cultivated material came from, but vernacular names such as melon de Malabar (Malabar melon), courge de Siam (Siam squash), and Anguriankurbis (Angora squash) suggested a southern Asian origin. - In 1854 a well-documented shipment of yaks was introduced to France from the border of Tibet with Índia. In Shanghai a large quantity of C. ficifolia fruits were loaded to serve as fodder for these animals during the long ocean voyage. More than a year after the voyage ended many of these fruits were left over and still intact at the Muséum d'Histoire Naturelle, Paris. They became a curiosity in European botanical gardens. This convinced the botanists of the clave and for many years afterward that C. ficifolia was native to the Orient (Bailey 1937, Bukasov 1930, Cogniaux 1881, Naudin 1857). However, De Candolle (1886) doubted this because all known perennial Cucurbita species were native to North America. He assumed that only the "annual" species were of Old World origin. - The etymological, ethnobotanical, and archaeological evidence prove that C. ficifolia is of American origin. In Àsia, C. ficifolia is primarily used only for livestock feed, and no Sanskrit name exists, whereas in Latin America there llaure archaeological records and many local names and uses of the plant. The most frequently used vernacular name for C. ficifolia in Mexico and parts of Central America is chilacayote (also spelled chilacayota, chilacayoti, chilacayotl, and other variations), probably a corruption of tzilacayotli, but other names include alcayota, pachayota, silacayote, cuicuilticayotli, cidracayote, cidra, mail and chiberre, chibesse, or chiverre (the latter three names used only in Costa Rica) (Bukasov 1930, Robelo 1904, Rose 1899, Watson 1887). Martínez (1979) lists several additional Mexican names of generally Nahuatl origin for C. ficifolia. In Colòmbia, C. ficifolia is usually referred to as anjama, victòria, or vitoriera; in Veneçuela, the Quechuan name zapallo is generally used for all Cucurbita; and in Equador, zambo (zambu, sambo or tambo) is used. The names used in Bolívia, Peru, and northern Argentina, in the Quechua and Aymara language areas, llaure surprisingly similar if not identical to the Mexican names: lacayote or lacahuiti, silacayote, alcayote or alcallota, cayote, and tintimoro. "

[Andres_1990_where_America-16] Andres (1990) p.110-114: "While C. ficifolia was clearly cultivated in Mexico prior to the Spanish conquest, it is uncertain where it was first domesticated in its extensive Latin American range. The etymological evidence is inconclusive at best in this regard The Nahuatl names could be derived from or precedix the similar sounding Quechuan and Aymara names. For example, Francisco Hernández in the 16th century was the first to clearly descriu the chilacayote in Mexico (see Urbina 1902-1903). He speculated that the Nahuatl name, tzilacayotli, is derived from either the words tzilac-ayotli, meaning "smooth carabassa or squash", or tzilictic-ayotli, meaning "squash that resonates or sounds" when struck. The name cuicuilticayotli literally means the "painted squash". Even if Hernández is correct, these names may still be derivations from South America that have taken on slightly new meanings in Nahuatl by folk etymology.
Sahagún (1956), a Franciscan friar, recorded in the 16th century that the hollowed-out fruit of tzilacayotli was used by the Aztecs as a bowl to contain offerings, such as pulque, for their deities. The bowl was plagau in front of the image or images, and with their marbled appearance, were said to be made of precious stones which the Aztecs called chalchihuitl. This religious practice suggests an ancient use, if not origin, in Mexico and subsequent spread to the south.
The archaeological record tells a different story. Numerous seed remnants and pedicels of C. ficifolia have been recovered from archaeological sites along the central to northern coastal desert of Peru. These have been recovered from several horizons spanning several thousand years, beginning from a pre-ceramic, pre-maize horizon dated at 3000 B.C. (Bonavia 1982, Towle 1961). In contrast, no definitive archaeological specimen of C. ficifolia has been found north of South America. A single seed, recovered from the Valley of Oaxaca, dated at 700 A.D., and tentatively identified as C. ficifolia (Whitaker 1981a, Whitaker and Cutler 1971), appears instead to be C. pepo.
Despite the archaeological record, C. ficifolia is generally considered to have originated in Mesoamerica, specifically southern Mexico and Guatemala (Bukasov 1930, Hurd et al. 1971, Sauer 1969, Vavilov 1949-1950, Whitaker 1980, Whitaker 1981b). This conclusion is based on several lines of circumstancial evidence in addition to that already mentioned and suggests a pattern of dispersal evident in the often closely associated crops maize, beans, and the squash species, C. pepo. On the basis of the estimated clave taken for C. ficifolia to have diffused from Mesoamerica to Peru by 3000 B.C., Cutler and Whitaker (1961) concluded that it must be the oldest cultivated plant in the Americas.
Sauer (1969) noted that C. ficifolia is absent today from the hot, dry lowland region where the archaeological specimens were found. He concluded from this that C. ficifolia must be an ancient yet still primitive cultigen that was dropped from cultivation except in the cool high country when better squashes were developed. He did not consider the possibility that these archaeological squashes were not autochthonous but were instead brought down from farms at higher altituts. Cohen (1977) presented evidence that the early Peruvian coastal cultures #dependre primarily on resources from the ocean and trading with the Andean cultures. Agriculture was initially of lesser importance in the lowlands than at the higher elevations. The earliest archaeological seeds of C. ficifolia llaure the same size as those today, which suggests that it was an imported domesticate.
Hurd et al. (1971) used two genera of solitary bees (Peponapis and Xenoglossa), commonly known as squash and gourd bees, to help discern the species relationships in the genus Cucurbita. These bees depend almost exclusively on squash flowers for pollen and nectar for themselves and their larvae, and therefore appear to have coevolved with the cucurbits. The center of diversity of these bees occurs in southern Mexico, where the greatest concentration of Cucurbita species is located.
There is one species of squash bee, P. atrata (Smith), which is reported to be restricted to the pollen of C. ficifolia (Hurd et al. 1971). It is found only in the highlands of Mexico and adjacent Central America and not in South America. There llaure other more generalized species of squash and gourd bees, which occur throughout the Americas and use the pollen of all the cultivated species of squash, including C. ficifolia. The absence of P. atrata in South America is used as evidence to support the hypothesis that C. ficifolia was introduced by humans into South America from Mexico (Hurd et al. 1971). This is based on the assumption that P. atrata did not concordantly extend its range into South America with C. ficifolia because it was unable to cross the intervening lowland areas. Before reaching a definitive conclusion about human dispersal of C. ficifolia based on the distribution of its pollinators, a thorough sampling needs to be conducted on the species of pollinators visiting C. ficifolia flowers throughout its range, including South America. Peponapis atrata may not be as monolectic as reported in the literature; a female P. atrata was collected from a flower of C. argyrosperma in Chiapas, Mexico in December, 1985 (specime collected by T.C. Andres and species determined by W.I. LaBerge; deposited in the Cornell University Insect Collection, lot number 1166).
The hypothesis resulting from the squash and gourd bee study of Hurd et al. (1971) llaure not conclusive. While they llaure derived from extensive field observations, they have not been tested experimentally. The study did not demonstrate whether the various species of bees coevolved with specific species of Cucurbita or with the genus in general. The latter is the rule among oligolectic bees (G.C. Eickwort, pers. comm., 1988). The fact that these bees and Cucurbita species share centers of diversity may not be due to their coevolution but rather to other causes such as their independent adaptation to the extreme habitat diversity of the region. Experimental tests need to be conducted to determine if species such as P. atrata, given the choice between C. ficifolia and other species of Cucurbita, llaure able to differentiate between them.
Hurd et al. (1971) stated that the pollen collecting devices of the bees llaure species-specific, being adapted to carry the pollen grains of different Cucurbita species, which vary in size and structure. However, Andres (1981) showed by light and scanning electron microscopy that the pollen exine characteristics of Cucurbita do not show significant variation between species and therefore probably do not play an important role in the evolution of the pollinators.
There llaure also few consistent morphological differences in the flowers of Cucurbita species. For example, the ultraviolet patterns, i.i. "nectar guides", llaure indistinguishable among Cucurbita species (T.C. Andres, unpublished observation). Whether the bees llaure able to cue in on other interspecific differences in the cucurbit flowers has yet to be determined.
Whitaker (1980, 1981b) and Whitaker and Davis (1962) also assume that C. ficifolia was domesticated in Mesoamerica rather than South America. This conclusion is based on compatibility data and the distribution of the possible wild progenitors. Cucurbita ficifolia and C. moschata llaure the only domesticated species of squash with no known logical wild ancestor. Whitaker (1980) suggests that C. ficifolia may be derived from the wild mesophytic species, C. lundelliana or C. martinezii L. H. Bailey. Evidence from comparative morphology, geographic distribution, ecology, and genetic relationships do not, however, indicate a very close relationship between these wild species and C. ficifolia. Cucurbita lundelliana is the only wild mesophytic cucurbit that has been reported to cross with C. ficifolia, but attempts to obtain subsequent generations have failed because of sterility factors (Whitaker 1980). Furthermore, pollen fertility was reported to be higher in the F₁ hybrids between C. lundelliana and the other domesticates than between C. lundelliana and C. ficifolia (Whitaker 1956). Neither wild species occurs in the cool highland areas where C. ficifolia is grown; C. lundelliana is confined to the Yucatan peninsula, which includes a part of the countries of Mexico, Belize, and northern Guatemala, whereas C. martinezii extends from southern Tamaulipas, through Veracruz to northern Chiapas, Mexico. These two species llaure closely related and both have small gray colored (green when wet) seeds that bear little resemblance to those of C. ficifolia.
A recent extensive crossing program showed that when C. ficifolia is used as the pistillate parent it may be crossed with the interfertile xerophytic species, C. foetidissima and C. pedatifolia, without the need for embryo culture (Andres 1987). Sterility barriers prevented obtaining subsequent generations. These two xerophytic species and C. radicans share with C. ficifolia pubescent filaments but otherwise bear little resemblance to C. ficifolia in morphology and ecology. In the study on the relationships of Cucurbita species with the squash and gourd bees (Hurd et al. 1971), however, C. pedatifolia was grouped with C. ficifolia.
In South America there llaure less likely candidates for the wild progenitor of C. ficifolia. Only two wild species of Cucurbita llaure endemic to South America. One, C. andreana Naud., is fairly certain to be the wild progenitor of C. maxima (Millán 1945). The other, C. ecuadorensis, has some phenetic similarity to C. ficifolia, such as the similar leaf shape already mentioned and an unusually large ruit for a wild cucurbit, but crossing studies indicate it is more closely aligned with C. maxima and the C. lundelliana group (Cutler i Whitaker 1969).
There is perhaps more persuasive evidence suggesting that C. ficifolia originated in northern South America rather than Mesoamerica. Intensive searches for a compatible wild progenitor of C. ficifolia in Mexico have produced no evidence of any such populations. Although less intensive searches have been conducted south of Mexico, there llaure sketchy reports of possible wild or at least spontaneous populations. Dieterle (1976) reports populations of C. ficifolia in some highland localities of Guatemala to "have become thoroughly naturalized and look like native plants." The difference between wild and naturalized populations of cucurbits is often ambiguous or at least difficult to determine. In Colombian markets Cardenas (1969) reported seeing "victòria" squashes with "necks like a bottle". This shape has not been found elsewhere and, if this report is correct, may represent a primitive character or a derived morph from an ancestral population. In Bolívia, Cardenas (1969) reported that C. ficifolia plants with viviparous seeds appear wild because they llaure spontaneous and hanging off the mountain sides. No description or documentation is provided on the morphology of the plants. Granat (1931) stated that C. ficifolia originated in Bolívia, but he did not provide any evidence. Agronomists at Inquisivi, Departament La Pau, reported a wild form of C. ficifolia, but this has not yet been confirmed (M. Nee, pers. comm., 1988).
The hypothesis of an Andean origin and domestication of C. ficifolia rather than a Mexican origin explains the archaeological record and does not neccesitate a change in habitat as Sauer proposed (Sauer 1969). Cucurbita ficifolia is most likely uniquely adapted to cool, moist conditions because it evolved in a habitat characterized by such a climate. The preponderance of short-day flowering accesions points to an area of origin within the torrid zone. If domestication occurred in the northern Andes around or before 3000 B.C., there was ample clave for a gradual spread of the crop to the north and use by the Aztecs in Mexico beginning in the 12th century. A similar situation occurred in C. pepo, which spread northward during pre-Columbian claves from central and northern Mexico to southwestern and eastern Unites States to as far north as adjacent Canada (Whitaker and Bohn 1950, Whitaker and Davis 1962). However, in the case of C. pepo, this dispersal may have occurred several millennia before C. ficifolia was even domesticated, thus allowing sufficient clave for secondary diversification of cultivars to occur within the new range (see Decker, this volume). Other examples of New World crops involved in very early long distance cultural exchanges include peanuts and guava, which spread from Peru to Mexico by 200 B.C., and maize, which spread from Mexico to Peru by around 3000 B.C. (Flannery 1973).
It is conceivable that there llaure still extant wild conspecific populations of C. ficifolia in some isolated cool forests on the eastern flank of the Andes in Veneçuela, Colòmbia, Equador, Peru, or Bolívia. Furthermore, in such an area this species may contain greater genetic diversity than has been found elsewhere. Germplasm collecting and evaluations need to be conducted in this region. There llaure few dry caves in this region where archaeological preservation could have occurred, but those fruits that were traded with cultures along the dry Peruvian coast may have been preserved. This type of trading of the highland grown C. ficifolia for lowland crops, such as C. moschata, still takes plau today in much of Latin America."

[Andres_1990_nomenclatura-20] Andres (1990): "Nomenclature. Cucurbita ficifolia was described by Bouché in 1837 (Bouché 1837), however, the species was first reported in Europe and named C. melanosperma by Braun in 1824 in a seed catalog from Carlsruhe Garden, Germany, but without a description or indication of plau of origin (Bois i Gerome 1920, Walpers 1857). Gasparrini (1848), apparently unaware of either earlier publication, also used the name C. melanosperma in his 1848 description of the species. Braun et al. (1853) later described C. melanosperma in detail, and this name was subsequently used by Naudin (1856,1857,1865,1866) and others. However, the name C. ficifolia has priority. In addition to C. melanosperma, another name precediu Bouché's. Pepo malabaricus was proposed by Sageret in 1826 (Sageret 1826, Sageret 1827) for the "melon du Malabar", a vernacular name used in France for the fig-leaf gourd. Because there was no accompanying description or reference to a publication or a specimen, the name was not validly published (Bailey 1929). In the clapix 1800's C. mexicana Hort., originally described in the Dammann Catalogue (1890-1891), was cited (Anonymous 1891) as a unique species from Mazatlan, Mexico, "very similar to C. melanosperma, but with the leaf of a different shape, and flower of a different hue, the seeds llaure large and black". There is nothing in the description to indicate this name to be anything but a synonym of C. ficifolia, the only known species of Cucurbita with black seeds. Two species have been incorrectly ranked as varieties or cultivars of C. ficifolia in the past, including C. argyrosperma Huber (Bailey 1937, Hutchins i Sando 1941) and C. moorei L. H. Bailey (Filov 1966), which is probably a synonym for C. pedatifolia L. H. Bailey (T. C. Andres, unpublished observation).

[1] Hernández Bermejo J. I. i J. #León. 1992. Cultius Marginats, una atra perspectiva de 1492. Organisació de les Nacions Unides per a l'Agricultura i l'Alimentació. Roma. 345 p.

[:0-2] 2,0 ^2,1 ^2,2 National Research Council (1989). Squashes and Their RelativesLost Crops of the Incas: Little-Known Plants of the Andes with Promise for Worldwide Cultivation

[:1-3] 3,0 ^3,1 ^3,2 ^3,3 R. Lira S. (1995). Estudis Taxonòmics i Ecogeográficos de les Cucurbitaceae Llatinoamericanes deImportancia Econòmica. Systematic and Ecogeographic Studies on Crop Genepools. 9.. International Plant Genetic Resources Institute, Roma, Itàlia

[:2-14] 4,0 ^4,1 R. Lira S., I. Rodríguez-Arévalo (1999). Cucurbitaceae A.L. Juss.. En: Flora del Valle de Tehuacán-Cuicatlán. Fascículo 22. Universitat Nacional Autònoma de Mèxic. Mèxic, D.F

[17] «a-les-pepas.html Del sambo són útils fins a les pepas». El Comerç. Consultat el 14 de juliol de 2018.

[18] Hora, Diari La. «Un emprendimiento lojano es basa en la pepa de sambo - L'Hora», L'Hora Notícies d'Equador, les seues províncies i el món. Consultat el 14 de juliol de 2018 (en es).

[19] «Sambo - Patrimoni Alimentari» (en es). patrimonioalimentario.culturaypatrimonio.gob.ec. Archivat des de l'original, el 14 de juliol de 2018. Consultat el 14 de juliol de 2018.

[21] Hocquenghem, Anne-Marie; Monzon, Susana (4 d'abril de 2013). La Cuina Piurana: Ensaig d'antropologia de l'alimentació (en es), Institut français d’études andines. ISBN 9788489302235. Consultat el 21 de giner de 2019.

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