To Check About Presence of Populations of Wild Phaseolus Species Found in May 1987 and May 1988

FIELD TRIP REPORT – CUZCO, PERU, 2-3 JULY 2016

Purpose: to check about presence of populations of wild Phaseolus species found in May 1987 and May 1988.

Travelers: D.G. Debouck, N. Palmer.

Report and photographs by: D.G. Debouck

Routing: July 2 (Urubamba province): Cusco- Sacsayhuaman- Pucara- Corao- Pisaq- Coya- Lamay- Calca- Huarán- Yucay- Urubamba- Ollantaytambo- Huaynapuquio- return; total 258 km. July 3 (Anta province): Cusco- Poroy- Izcuchaca- cross Zurite- Ancahuasi- Pampaconga- Limatambo- Puente Achaco- return: total 156 km.

Background

In the area of study – upper Urubamba valley and valley of river Limatambo – the only Phaseolus species that can be found as wild plants are Phaseolus augusti Harms and a wild form of Phaseolus vulgaris L. (also previously reported as P. aborigineus Burkart). The distribution of the former extends from Azuay, Ecuador (Debouck et al. 1989) down to Tucuman, Argentina (Palacios & Vilela 1993), while the latter extends from the western Venezuelan Andes (Berglund-Brücher & Brücher 1976) down to Córdoba, Argentina (Drewes 2008; Toro-Chica et al. 1990). The presence of these two species in the department of Cuzco has been documented in several works (Table 1, and: Berglund-Brücher & Brücher 1976, Brako & Zarucchi 1993, Debouck 1987, Debouck & Tohme 1988, Macbride 1943, Weberbauer 1945). Two germplasm explorations were carried out in 1987 and in 1988 in order to increase the representativeness of these two species in the germplasm banks of INIA and CIAT.

Table 1 – Some old records about the presence of Phaseolus augusti and wild P. vulgaris in the area of study.

Herbarium specimen Species Place Date found Source OF Cook 239 augusti between Pisac and Calca 20/04/1915 Debouck 2015a OF Cook 511 augusti Ollantaytambo 01/05/1915 Debouck 2015a FL Herrera 1376 augusti near Yucay ?/01/1927 Debouck 2015a AW Hill 138 augusti Urubamba ?/01-03/1903 Debouck 2015a E Petersen 1535 augusti Calca 24/04/1953 Debouck 2015a Marín 63 augusti Pisac ?/01/1942 Debouck 2015a J Soukup 726 augusti near Yucay 28/12/1937 Debouck 2015a C Vargas 7110 vulgaris wild Limatambo, río Blanco 26/03/1948 Debouck 2015b - - - vulgaris wild 30 km NW Machupicchu ?/?/ 1967 Berglund-Brücher & Brücher 1976

Results The two populations of P. augusti were found at the sites where they were found in May 1987. The population # 2312 (introduced in CIAT genebank in late May 1987 and kept as G40725) was found on July 2, 2016, 3 km W from Lamay at 2,941 masl (GPS: S13o 21’ 08.6”, W071o 56’ 12.8”). At this altitude and at this time of the year, as expected, it had been affected by freezing temperatures. It was found at the latest time of seed dispersal (Fig. 1). This population seems to be 2 stable because the topography does not allow farming, and no housing has occurred yet. No grazing by goats was seen at the site.

Figure 1 – Population # 2312 of P. augusti (left) (dry pods can be seen on the vine climbing on Opuntia); site where found (right).

Population # 2313 of P. augusti (introduced in CIAT genebank in May 1987 and kept as G40771) was found on July 2, 2016, 2 km NW of Ollantaytambo at 2,833 masl (GPS: S13o 15’ 08.5”, W072o 16’ 25.1”). As expected, rains would have stopped 4-6 weeks ago; consequently the vegetation of the understory was dried and already affected by low temperatures. This P. augusti was found at late seed dispersal stage (Fig. 2). As compared to site of #2312, the slope is gentle and less protected. Although already late for assessment, the population seems smaller in 2016 as compared to 1987 (effect of grazing?). No housing would be allowed there because of the proximity of the ruins of Ollantaytambo declared ‘Patrimonio histórico de la Nación’ (historic heritage of Peru).

Figure 2 – Population # 2313 of P. augusti (left) (four trifoliolate leaves can be seen in center of picture); site where found (right).

Population # 2580 of wild Phaseolus vulgaris (introduced in CIAT genebank in early June 1988 and kept as G23454) was found on July 3, 2016, at 2,433 masl (GPS: S13o 29’ 57.0”, W072o 28’ 06.6”) 5 km W from Limatambo and 0.5 km from Puente Achaco. This is about 2-3 km further west as compared to the location of May 1988 where housing and walls to protect the road against flooding have eliminated much of the original vegetation. As it can be seen in Fig. 3, the plant material is certainly wild: small seeds with black speckled brown background and twisting pod valves. This clarifies the doubt left in the work by Brako & Zarucchi (1993): apart from escapes from cultivation, P. vulgaris is certainly wild in Peru. Other works (e.g. Bitocchi et al. 2013, Kwak & Gepts 2009, Tohme et al. 1996) have also demonstrated this point. The pods (Fig. 3 left) could sometimes be cut off as compared to normal ones with recurved pod beak (Fig. 3 right) due to damages by birds that feed on the young developing seeds. The population was found after seed dispersal with twisted pods and dry vines climbing on Opuntia, Lantana, Caesalpinia, Celtis, Acacia and Schinus bushes. The habitat is that of a subhumid montane forest with many open spots where wild P. vulgaris can climb on tall Composites and grasses. The different variants found in this population in 1988 and studied later (Beebe et al. 1997) were not seen on this visit, because it was too late for such kinds of observations.

Figure 3 – Population # 2580 of wild P. vulgaris (left); open dry pods (right) twice as long as the one to the left (compare pedicels).

Figure 4 – Population # 2580 of wild P. vulgaris climbing on Opuntia (left); site where found (right). 5

Figure 5 – Site of population # 2580 of wild P. vulgaris; the wild bean thrives on bushes surrounding small maize fields (forefront) but not in the dry scrub on the back slope.

Discussion The above facts elicit the following points for discussion. First, the coordinates reported by the collector and the genebank (http://www.ciat.cgiar.org/urg) allow to find the populations twenty- nine (Debouck 1987) or twenty-eight (Debouck & Tohme 1988) years after the original finding, even if global positioning system was not available at that time. The capacity of going back to the original site can be critically important if there is a doubt about taxonomical identification (see the case of the lost teosintes: Iltis et al. 1986), or the need to re-sample the original population (the strategy followed by the Millennium Seed Bank for many wild endangered species), or the interest in checking extinction (our current purpose). Second, the populations of these two wild species are still there after twenty-nine/ twenty-eight years, as an indication that if habitats are left with no disturbance the wild species will survive. While this seems to be true for both populations of P. augusti, that was not exactly the case of wild P. vulgaris #2580 found 2-3 km further west from the original site of 1988. Wild P. vulgaris sets a challenge for in situ conservation: in case of a drastic change in the habitat, the population will go extinct, while limited periodical disturbance would favor survival. The possibility of these populations being there for long is also shown by the collection dates tracing back to the early/ mid 1900s (Table 1). Third, the late visit to the 6 populations, even though it was too late for an appropriate sampling for germplasm conservation (it was not the purpose of the visit, starting from the fact that no collecting permit was asked for nor granted), allowed a couple of interesting observations. On the one hand, some populations of P. augusti could be tolerant to low temperatures, because of the combination of high altitude (almost 3,000 masl!) and early frosts. This trait if confirmed could be of interest to Lima bean breeding, as P. augusti seems to belong to the secondary genepool of P. lunatus (Caicedo et al. 1999, Serrano-Serrano et al. 2010). On the other hand, in contrast to P. augusti, wild P. vulgaris seems more mesophytic (no extreme temperatures, adequate water supply: not present in flooding areas of Limatambo river, but neither on the dry slopes). In both species pods unopened at this time had all grains damaged by insects. But pods damaged by birds were seen only in wild P. vulgaris. Similar damages by birds have been seen on wild P. vulgaris in southern Ecuador (Debouck et al. 1993). Birds feeding on young developing seeds could have indicated a potential food source to would-be domesticators (Debouck 2016). Finally, the early including of samples of these three populations into genebanks has made possible many studies to advance our understanding of bean genetic diversity and evolution (e.g. Beebe et al. 1997, Bitocchi et al. 2013, Caicedo et al. 1999, Chacón-Sánchez et al. 2005), and from there to prepare the next steps of plant breeding.

Acknowledgments Two persons have expressed interest into this field verification: M. Gamarra-Flores, J. Tohme; this interest is deeply acknowledged. The help of Josefina Martínez to check the manuscript throughout has been much appreciated.

Literature cited Beebe, S.E., O. Toro-Chica, A.V. González, M.I. Chacón-Sánchez & D.G. Debouck. 1997. Wild-weed- crop complexes of common bean (Phaseolus vulgaris L., Fabaceae) in the Andes of Peru and Colombia, and their implications for conservation and breeding. Genet. Resources & Crop Evol. 44 (1): 73-91.

Berglund-Brücher, O. & H. Brücher. 1976. The South American wild bean (Phaseolus aborigineus Burk.) as ancestor of the common bean. Econ. Bot. 30 (3): 257-272.

Bitocchi, E., E. Bellucci, A. Giardini, D. Rau. M. Rodríguez, E. Biagetti, R. Santilocchi, P. Spagnoletti- Zeuli, T. Gioia, G. Logozzo, G. Attene, L. Nanni & R. Papa. 2013. Molecular analysis of the parallel domestication of the common bean (Phaseolus vulgaris) in Mesoamerica and the Andes. New Phytol. 197 (1): 300-313.

Brako, L. & J.L. Zarucchi. 1993. Catalogue of the flowering plants and gymnosperms of Peru. Monographs in Systematic Botany, vol. 45, Missouri Botanical Garden, St. Louis, Missouri, USA. 1286p.

Caicedo A.L., Gaitán E., Duque M.C., Toro Ch. O., Debouck D.G. & J. Tohme. 1999. AFLP fingerprinting of Phaseolus lunatus L. and related wild species from South America. Crop Sci. 39 (5): 1497-1507.

Chacón-Sánchez, M.I., B. Pickersgill & D.G. Debouck. 2005. Domestication patterns in common bean (Phaseolus vulgaris L.) and the origin of the Mesoamerican and Andean cultivated races. Theor. Appl. Genet. 110 (3): 432-444.

Debouck, D.G. 1987. Recolección de germoplasma de Phaseolus en el centro y centro-sur del Perú. International Board for Plant Genetic Resources, Rome, Italy. Mimeographed, 36p.

Debouck, D.G. 2015a. Cahiers de phaséologie: section Paniculati. International Center for Tropical Agriculture (CIAT), Cali, Colombia. Available at http://www.ciat.cgiar.org/urg. Accessed on 9 July 2016. 392p.

Debouck, D.G. 2015b. Cahiers de phaséologie: section Phaseoli. International Center for Tropical Agriculture (CIAT), Cali, Colombia. Available at http://www.ciat.cgiar.org/urg. Accessed on 9 July 2016. 190p.

Debouck, D.G. 2016. Your beans of the last harvest and the possible adoption of bright ideas. In: “Ethnobotany of Mexico: interactions of people and plants in Mesoamerica”, R. Lira, A. Casas & J. Blancas (eds.). Springer, New York, New York, USA. Pp. 367-387.

Debouck, D.G., R. Castillo T. & J.M. Tohme. 1989. Observations on little-known Phaseolus germplasm of Ecuador. FAO/IBPGR Plant Genet. Resources Newsl. 80: 15-21.

Debouck, D.G. & J. Tohme. 1988. Recolección de germoplasma de Phaseolus en el centro-sur del Perú. International Board for Plant Genetic Resources, Rome, Italy. Mimeographed, 31p.

Debouck, D.G., O. Toro, O.M. Paredes, W.C. Johnson & P. Gepts. 1993. Genetic diversity and ecological distribution of Phaseolus vulgaris (Fabaceae) in northwestern South America. Econ. Bot. 47 (4): 408-423.

Drewes, S.I. 2008. Prospección y colecta de germoplasma silvestre de Phaseolus vulgaris en la zona central de Argentina. Plant Genetic Resources Newsletter 155: 9-14.

Iltis, H.C., D.A. Kolterman & B.F. Benz. 1986. Accurate documentation of germplasm: the lost Guatemalan teosintes (Zea, Gramineae). Econ. Bot. 40 (1): 69-77.

Kwak, M. & P. Gepts. 2009. Structure of genetic diversity in the two major gene pools of common bean (Phaseolus vulgaris L., Fabaceae). Theor. Appl. Genet. 118 (5): 979-992.

Macbride, J.F. 1943. Flora of Peru – Family Leguminosae. Field Mus. Nat. Hist. vol. 13, part 3, num, 1, 507p.

Palacios, R.A. & A.E. Vilela. 1993. Las especies argentinas de Phaseolus (un recurso genético que necesita protección). In: Actas del II Simposio Latinoamericano sobre Recursos Genéticos de Especies Hortícolas, A.M. Clausen, E.L. Camadro, A.F. López Camelo & M.A. Huarte (eds.). Instituto Nacional de Tecnología Agropecuaria, Balcarce, Argentina. Pp. 123-139.

Serrano-Serrano, M.L., J. Hernández-Torres, G. Castillo-Villamizar, D.G. Debouck & M.I. Chacón. 2010. Gene pools in wild Lima bean (Phaseolus lunatus L.) from the Americas: evidences for an Andean origin and past migrations. Molecular Phylogenetics and Evolution 54 (1): 76-87.

Toro-Chica, O., J. Tohme & D.G. Debouck. 1990. Wild bean (Phaseolus vulgaris L.): description and distribution. Centro Internacional de Agricultura Tropical, Cali, Colombia, 106p.

Tohme, J., D.O. González, S. Beebe & M.C. Duque. 1996. AFLP analysis of gene pools of a wild bean core collection. Crop Sci. 36 (4): 1375-1384.

Weberbauer, A. 1945. El mundo vegetal de los Andes peruanos. Editorial Lumen, S.A., Lima, Peru. 776p.