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Diversity of the Calabash Tree (Crescentia Cujete L.) in Colombia

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Diversity of the Calabash Tree (Crescentia Cujete L.) in Colombia Agroforest Syst (2009) 76:543–553 DOI 10.1007/s10457-009-9207-0 Diversity of the calabash tree (Crescentia cujete L.) in Colombia Johanna Arango-Ulloa Æ Adriana Bohorquez Æ Myriam C. Duque Æ Brigitte L. Maass Received: 26 October 2007 / Accepted: 4 January 2009 / Published online: 28 January 2009 Ó The Author(s) 2009. This article is published with open access at Springerlink.com Abstract Germplasm of the calabash tree (Crescen- Overall, genetic diversity was high (mean Nei and Li’s tia cujete L.) was collected in five major regions of coefficient of 0.43). No relations could be established Colombia, i.e. the Andes, Caribbean, Amazon, Ori- between either geographical provenance or fruit mor- noco, and Pacific regions. Collecting this multipurpose phology and patterns of genetic diversity. Concerning tree was guided by the indigenous knowledge of the outgroups, the C. amazonica accession appeared to farmers and artisans in each region. Large variation in be a distinct species. The C. alata accession, however, fruit shapes and sizes was found, of which some forms did not seem to be sufficiently distinct from C. cujete to were typical for certain regions. Overall 56 accessions merit species status. The latter material may in fact be a were collected and roughly classified into 22 types by hybrid or serve to challenge the validity of interspecific eight fruit shapes and eight sizes. Molecular markers organization of the genus Crescentia. (Amplified fragment length polymorphisms) were applied to leaf tip tissue originating from vegetatively Keywords AFLP Á Bignoniaceae Á propagated plants in order to assess the diversity Calabash tree Á Genetic diversity Á available in the germplasm collected as well as to Homegarden Á Molecular marker Á detect patterns of geographical or morphological Multipurpose tree Á Non-timber forest product Á similarity. One accession each of C. alata H.B.& Plant genetic resources Á Underutilized species K. and C. amazonica Ducke were used as outgroups. Abbreviations AFLP Amplified fragment length polymorphism J. Arango-Ulloa Á B. L. Maass (&) GIS Geographic information systems Department of Crop Sciences: Agronomy in the Tropics, PCR Polymerase chain reaction Georg-August-University Go¨ttingen, Grisebachstr. 6, 37077 Go¨ttingen, Germany e-mail: [email protected] Introduction Present Address: The calabash (Crescentia cujete L.: Bignoniaceae) is J. Arango-Ulloa Resid. Vista Hermosa Casa 8/Barrio San Martin, a small tree with multiple uses, originating from Santa Rosa de Copan, Honduras tropical America and now widely distributed in the e-mail: [email protected] tropics (Burger and Gentry 2000; Widodo 2001). It is a typical component of homegardens not only in A. Bohorquez Á M. C. Duque Centro Internacional de Agricultura Tropical (CIAT), Mexico (e.g., Vogl et al. 2002), Central (e.g., Bass A.A. 6713, Cali, Colombia 2004) and South America (e.g., Lamont et al. 1999; 123 544 Agroforest Syst (2009) 76:543–553 Gari 2001), but also in Africa and Asia (Widodo research has been undertaken to underpin the further 2001). It is also grown as a living fence (Avendan˜o- development of this multipurpose tree. This study Reyes and Acosta-Rosado 2000), for fuelwood, and was performed within a larger initiative of the as an ornamental and shade tree alongside urban non-governmental organization Centro para la Inves- streets (Pe´rez-Arbelaez 1990). tigacio´n en Sistemas Sostenibles de Produccio´n Pe´rez-Arbelaez (1990) described the ‘totumo’ or Agropecuaria (CIPAV) that aims to further develop ‘jı´caro’ as a ‘‘popular panacea’’ with many diverse underutilized multipurpose tree species for their use uses. The bottle-like dry and empty calabash fruits by smallholders in silvo-pastoral production systems. serve as containers, for home-made utensils, and to The aims of this project were to collect germplasm, prepare handicrafts, and the pulp and foliage are used which represented the morphological and geographic as livestock feed (Cajas-Giron and Sinclair 2001; Bass diversity of the species based on indigenous knowl- 2004; Ibrahim et al. 2006). The tree has also known edge from farmers and other users; and to assess that medicinal properties (Gentry 1980;Pe´rez-Arbelaez diversity with a view to its inclusion in future 1990; Widodo 2001). research. Considerable morphological variation has been observed in C. cujete, particularly in fruit shape and size (Pe´rez-Arbelaez 1990). Gentry (1980) suggested Materials and methods that two variants of this polymorphic species may in fact deserve taxonomic recognition. Both variants Germplasm collection differ from typical C. cujete in possessing smaller, more coriaceous leaves and fruits. Gentry (1973, 1980) Germplasm and indigenous knowledge of the cala- also reported that apparent hybrids in Costa Rica had bash tree (C. cujete) were collected from five major the small fruit characteristic of C. alata H.B.&K. but regions in Colombia (van Wyngaarden and Fandin˜o- the simple leaves of C. cujete and only occasional Lozano 2005; Table 1). The collecting strategy was branches bore 3-foliolate leaves typical of C. alata. guided by Guarino et al. (1995) and based on The cultivated C. cujete, a native of Mexico and information provided by artisans, livestock produc- Central America, is often confused with C. alata ers, and/or people applying natural medicines. (Gentry 1973), a wild relative with a more restricted Specific morphological fruit forms were collected distribution, but often dominant in the dry forest only once from any one region. As the morphological savannas of the Pacific coast from Mexico to Costa forms were mostly planted as individual trees along- Rica (Gentry 1973, 1980; Bridgewater et al. 2002). side houses, in backyards, as living fences or Whether or not the natural distribution of C. cujete scattered in pasture paddocks, a vegetatively propa- extends to South America has not been established gated sample from such an individual tree represents (Gentry 1973; Burger and Gentry 2000). one accession. One accession each of C. alata and Although the calabash tree is widely distributed C. amazonica Ducke were collected from a roadside and used in Colombia (Pe´rez-Arbelaez 1990), little in the town of Cali (Valle del Cauca department) and Table 1 Geographic distribution and summary of collection and length of dry season extracted from Worldclim and data of Crescentia cujete germplasm collected in five regions Bioclim databases; the number of dry months per year were of Colombia; mean annual temperature, annual precipitation defined by precipitation with less than 60 mm Colombian region Collected Range of Range of Elevation Mean annual Annual Dry accessions latitude N longitude W (m asl.) temperature precipitation months (no.) (°C) (mm) (no.) Caribbean 9 8° 510–11° 150 73° 180–76° 160 20–94 27.0–28.3 1,148–1,497 4 Mompox island 6 9° 040 74° 410 51–56 27.9 2,257 3 Orinoco 13 3° 320–5° 230 72° 130–76° 460 267–548 24.8–26.9 2,485–4,597 0–3 Amazon 10 1° 270–1° 360 75° 350–75° 410 205–286 25.7–26.0 3,541–3,674 0 Andes 10 1° 600–6° 320 75° 020–77° 060 500–1,365 21.6–26.3 1,407–2,981 0–3 Pacific 8 1° 390–5° 450 76° 320–78° 100 41–160 25.7–26.8 5,861–7,498 0 123 Agroforest Syst (2009) 76:543–553 545 Table 2 Primer combinations and polymorphic bands produced when applying AFLPs on germplasm accessions of Crescentia cujete collected in five regions of Colombia Primer Total bands Polymorphic Polymorphic Accessions with combination (no.) bands (no.) bands (%) unique bands (no.) EAAG-MCAC 150 145 96.6 23 EACA-MCTG 107 79 73.8 5 Total (mean) 257 224 (86.8) 28 a gallery forest in the Orinoquian region, respec- polymorphism and resolution (Table 2). This selec- tively, to serve as outgroups. tion was based on previous experiences by Roa et al. (1997) and Caicedo (1996, cited by Segura et al. Morphological and molecular germplasm 2002) that two or three primer combinations were characterization sufficient to analyze genetic differences among populations or species by applying AFLPs as long A preliminary visual assessment of forms and sizes of as these provided a high level of polymorphic bands. mature fruits was performed during collecting. Trees Electrophoresis and detection of PCR products were and fruits were also photographically documented. carried out on 6% polyacrylamide gel by silver nitrate Mature fruits were visually classed into 8 sizes (1/ staining following Bassam et al. (1991) with modi- 2 = miniature; 3/4 = small; 5/6 = medium; 7/8 = fications. Only strong bands were scored visually as large) and 8 shapes (flattened, oblong, cuneate, elon- present or absent. Eleven accessions had to be gated, globular, rounded-drop-shaped, oblong-drop- excluded from further analysis because they did not shaped, kidney-shaped). From rooted stakes established amplify. in the CIPAV field at Jamundı´, Valle del Cauca department, 1–2 fully expanded leaves per accession Data analysis were obtained from 3 to 4 month old plants to determine leaf shape (lanceolate, oblong, spatulate, oblanceolate, Passport data (longitude, latitude and altitude) from the obovate, or elliptic), size (length and width in cm) and collecting points served to generate a map of potential form of the apex (acute, acuminate, or obtuse), which areas of distribution by applying FloraMapÒ (Jones were then documented photographically. and Gladkov 2005), a GIS-based program that helps After harvesting young leaf tips (300 mg) from identify areas with similar climates (Jones et al. 1997, one well-established plant per accession in the field at 2002). The map was produced on the basis of four Jamundı´, the tissue was immediately stored on ice in principal components that included 97.1% of the data a cooling box before freezing the sample at -80°Cat variance.
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