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Atlas of Wild Potatoes SYSTEMATIC AND ECOGEOGRAPHIC STUDIES ON CROP GENEPOOLS 10 Atlas of Wild Potatoes Robert J. Hijmans, David M. Spooner, Alberto R. Salas, Luigi Guarino and Jorge de la Cruz SYSTEMATIC AND ECOGEOGRAPHIC STUDIES ON CROP GENEPOOLS Atlas of Wild Potatoes Robert J. Hijmans1, David M. Spooner2, Alberto R. Salas1, Luigi Guarino3 and Jorge de la Cruz1 1 International Potato Center, Apartado 1558, Lima 12, Peru 2 USDA, Agricultural Research Service, Department of Horticulture, University of Wisconsin, 1575 Linden Drive, Madison, Wisconsin 53706-1590 USA 3 International Plant Genetic Resources Institute, Office for the Americas, c/o CIAT, AA 6713, Cali, Colombia ii SYSTEMATIC AND ECOGEOGRAPHIC STUDIES ON CROP GENEPOOLS ATLAS OF WILD POTATOES iii TABLE OF CONTENTS Acknowledgments ix Abstract x 1. Introduction 1 2. Wild potatoes 2 Habitats 2 Shapes and colours 3 3. Potato taxonomy 6 A dynamic taxonomy 6 Ingroup and outgroup relationships 12 Cultivated species 14 Complicating factors for a stable potato taxonomy 14 Breeding barriers 16 4. Data and software 17 Collecting expeditions 17 Data sources 17 Data quality 18 Information quality 19 Software 23 5. Distribution maps 24 Methods 24 Guatemala, Mexico, and USA 26 Central America 37 Northern Andes 40 Peru 45 Bolivia 61 Southern Andes 67 Southern cone 79 Distribution of series 84 6. Ecogeographic statistics 92 Distribution by country 92 Distribution by species 93 Range size statistics 94 Altitude and climate 103 Species distribution by latitude 109 7. Diversity 111 Species richness 111 Scale effect 116 Series 118 8. Conclusion 122 9. References 123 iv SYSTEMATIC AND ECOGEOGRAPHIC STUDIES ON CROP GENEPOOLS List of Tables Table 1. Wild potato species (Solanum sect. Petota) and three outgroup relatives in sect. Etuberosum, their standard three- letter codes, the number of the map in chapter 5 showing their distribution, series membership (or section membership for Etuberosum), and ploidy and Endosperm Balance Number (EBN) where known. 7 Table 2. Names of series in Solanum sect. Petota, and number of species in each series 13 Table 3. Wild potato distribution by country. 92 Table 4 Ecogeographic distribution statistics for wild potato species. Order and three-letter species codes follow Table 1. Number of observations (Obs); Range size: MaxD and CA50; Mean and standard deviation (SD) of altitude (Alt), annual average temperature (Temp) and annual precipitation (Rain). 96 Table 5. Grid-based species richness statistics by country. 115 Table 6. Number of species and number of observations (Obs) in our database, and average number of observations over species for each series. 118 List of Figures Figure 1. Wild potato habitats. A: dry deciduous forest in Jalisco, Mexico (2080 m); B: paramo in Mérida State, Venezuela (3050 m); C: seashore in the Chonos Archipelago, Región Aisén, Chile; D: humid highlands in La Paz Department, Bolivia (3900 m). 2 Figure 2. Wild potato plant forms. A: Solanum hougasii; B: S. agrimonifolium; C: S. morelliforme; D: S. infundibuliforme. 4 Figure 3. Wild and cultivated potato flowers. A: Solanum bulbocastanum; B: S. paucijugum; C: S. tuberosum (cultivated species); D: S. colombianum. 4 Figure 4. Wild potato fruits. A: Solanum verrucosum; B: S. andreanum; C: S. schenckii; D: S. moscopanum. 5 Figure 5. Wild potato tubers. A: Solanum sparsipilum; B: S. polyadenium; C: S. acaule; D: S. chiquidenum; E: S. commersonii; F: S. piurae. 5 Figure 6. Number of accepted potato species over time. 6 Figure 7. Expected and observed distance from the road of locations where wild potatoes were observed in central Bolivia. 22 Figure 8. Number of observations by species in our database. 94 Figure 9. Illustration of the calculation procedure of MaxD (A) and CA50 (B) for S. jamesii in the USA and Mexico. 95 Figure 10. Two measures of range size of wild potato species. Maximum distance between two observations of one species (MaxD) and circular area (CA50). 101 ATLAS OF WILD POTATOES v Figure 11. Maximum distance (MaxD) versus circular area (CA50) for all wild potato species. 101 Figure 12. Circular area (CA50) versus number of observations of wild potato species. 102 Figure 13. Altitude distribution of wild potatoes, for observations and species. 103 Figure 14. Altitude distribution by latitude of wild potato observations. 107 Figure 15. Temperature distribution of wild potatoes, for observations and species. 108 Figure 16. Temperature distribution of wild potato (over observations) by latitude. 108 Figure 17. Precipitation distribution of wild potatoes, for observations and species. 109 Figure 18. Precipitation distribution of wild potato (over observations) by latitude. 110 Figure 19. Number of wild potato species by latitude. 110 Figure 20. Frequency distribution of the number of wild potato species per 50 x 50 km grid cell. 114 Figure 21. Ratio of the number of wild potato species to number of observations for each grid cell (obs. = 1247). 116 Figure 22. Number of series versus number of species by grid cell. 121 vi SYSTEMATIC AND ECOGEOGRAPHIC STUDIES ON CROP GENEPOOLS List of Maps Map 1. Collecting sites and roads in central Bolivia. 21 Map 2. Area of known distribution of wild potatoes (gray shade). 25 Map 3. First-level administrative subdivisions in Guatemala, Mexico, and the USA. 26 Map 4. Distribution of S. guerreroense and S. stoloniferum.27 Map 5. Distribution of S. brachistotrichum, S. bulbocastanum, and S. hjertingii.28 Map 6. Distribution of S. leptosepalum, S. polyadenium, S. stenophyllidium, and S. ×semidemissum.29 Map 7. Distribution of S. demissum, S. hougasii, S. macropilosum, and S. matehualae.30 Map 8. Distribution of S. fendleri, S. hintonii, S. ×michoacanum, S. ×sambucinum, S. tarnii, and S. trifidum.31 Map 9. Distribution of S. cardiophyllum, S. jamesii, and S. ×vallis-mexici.32 Map 10. Distribution of S. lesteri, S. pinnatisectum, S. schenckii, and S. verrucosum.33 Map 11. Distribution of S. ×edinense, S. oxycarpum, and S. wightianum.34 Map 12. Distribution of S. iopetalum, S. morelliforme, and S. nayaritense.35 Map 13. Distribution of S. brachycarpum and S. papita.36 Map 14. First-level administrative subdivisions in Central America. 37 Map 15. Distribution of S. agrimonifolium and S. woodsonii.38 Map 16. Distribution of S. clarum and S. longiconicum.39 Map 17. First-level administrative subdivisions in the northern Andes. 40 Map 18. Distribution of S. albornozii, S. colombianum, and S. paucijugum. 41 Map 19. Distribution of S. calacalinum, S. flahaultii, S. pamplonense, S. regularifolium, S. subpanduratum, and S. tundalomense.42 Map 20. Distribution of S. andreanum, S. burtonii, S. chilliasense, S. donachui, S. garcia-barrigae, S. lobbianum, and S. moscopanum.43 Map 21. Distribution of S. minutifoliolum, S. neovalenzuelae, S. orocense, S. otites, S. solisii, S. sucubunense, and S. tuquerrense.44 Map 22. First-level administrative subdivisions in Peru. 45 Map 23. Distribution of S. acroglossum, S. anamatophilum, S. bill-hookeri, S. chancayense, S. chomatophilum, S. guzmanguense, and S. velardei.46 Map 24. Distribution of S. acroscopicum, S. ambosinum, S. ayacuchense, S. aymaraesense, S. chiquidenum, and S. wittmackii.47 Map 25. Distribution of S. albicans, S. buesii, S. burkartii, S. sandemanii, S. tapojense, S. trinitense, and S. yamobambense.48 Map 26. Distribution of S. amayanum, S. ancoripae, S. ×arahuayum, S. sarasarae, S. tacnaense, S. taulisense, and S. pillahuatense.49 Map 27. Distribution of S. cajamarquense, S. coelestipetalum, S. dolichocremastrum, S. gracilifrons, and S. hypacrarthrum.50 Map 28. Distribution of S. chillonanum, S. hastiforme, S. huarochiriense, S. incahuasinum, S. ingifolium, and S. saxatilis.51 ATLAS OF WILD POTATOES vii Map 29. Distribution of S. ancophilum, S. ariduphilum, S. huancabambense, S. humectophilum, S. incasinum, and S. ortegae.52 Map 30. Distribution of S. irosinum, S. jaenense, S. jalcae, S. laxissimum, S. longiusculus, and S. multiinterruptum.53 Map 31. Distribution of S. augustii, S. limbaniense, S. lopez-camarenae, S. marinasense, S. medians, and S. ×blanco-galdosii.54 Map 32. Distribution of S. huancavelicae, S. mochiquense, S. olmosense, S. simplicissimum, S. sogarandinum, and S. ×neoweberbaueri.55 Map 33. Distribution of S. immite, S. nemorosum, S. neovargasii, S. nubicola, and S. raphanifolium.56 Map 34. Distribution of S. cantense, S. contumazaense, S. lignicaule, S. paucissectum, S. peloquinianum, and S. santolallae.57 Map 35. Distribution of S. orophilum, S. pampasense, S. piurae, S. puchupuchense, S. sawyeri, and S. taratapanum.58 Map 36. Distribution of S. raquialatum, S. rhomboideilanceolatum, S. salasianum, S. scabrifolium, and S. urubambae.59 Map 37. Distribution of S. bukasovii.60 Map 38. First-level administrative subdivisions in Bolivia. 61 Map 39. Distribution of S. avilesii, S. gandarillasii, S. neocardenasii, S. soestii, and S. ugentii.62 Map 40. Distribution of S. arnezii, S. boliviense, S. circaeifolium, S. neovavilovii, and S. ×litusinum.63 Map 41. Distribution of S. alandiae, S. flavoviridens, S. hoopesii, S. virgultorum, and S. ×sucrense.64 Map 42. Distribution of S. achacachense, S. ×doddsii, and S. violaceimarmoratum.65 Map 43. Distribution of S. berthaultii and S. bombycinum.66 Map 44. First-level administrative subdivisions in the southern Andes. 67 Map 45. Distribution of S. acaule.68 Map 46. Distribution of S. ×bruecheri and S. okadae.69 Map 47. Distribution of S. ×setulosistylum and S. vidaurrei.70 Map 48. Distribution of S. tarijense.71 Map 49. Distribution of S. candolleanum, S. infundibuliforme, and S. yungasense.72 Map 50. Distribution of S. megistacrolonum.73 Map 51. Distribution of S. microdontum.74 Map 52. Distribution of S. brevicaule and S. vernei.75 Map 53. Distribution of S. oplocense.76 Map 54. Distribution of S. neorossii, S. sparsipilum, and S. spegazzinii.77 Map 55. Distribution of S. leptophyes.78 Map 56. First-level administrative subdivisions in the Southern cone. 79 Map 57. Distribution of S. calvescens, S. chacoense, S. maglia, and S. palustre.80 Map 58. Distribution of S. commersonii, S. fernandezianum, S. sanctae-rosae, S. ×rechei, and S. ×viirsoii.81 viii SYSTEMATIC AND ECOGEOGRAPHIC STUDIES ON CROP GENEPOOLS Map 59. Distribution of S.
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