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Taxonomy of the Genus Arachis (Leguminosae)

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Taxonomy of the Genus Arachis (Leguminosae) BONPLANDIA16 (Supi): 1-205.2007 BONPLANDIA 16 (SUPL.): 1-205. 2007 TAXONOMY OF THE GENUS ARACHIS (LEGUMINOSAE) by AntonioKrapovickas1 and Walton C. Gregory2 Translatedby David E. Williams3and Charles E. Simpson4 director,Instituto de Botánicadel Nordeste, Casilla de Correo209, 3400 Corrientes, Argentina, deceased.Formerly WNR Professor ofCrop Science, Emeritus, North Carolina State University, USA. 'InternationalAffairs Specialist, USDA Foreign Agricultural Service, Washington, DC 20250,USA. 4ProfessorEmeritus, Texas Agrie. Exp. Stn., Texas A&M Univ.,Stephenville, TX 76401,USA. 7 This content downloaded from 195.221.60.18 on Tue, 24 Jun 2014 00:12:00 AM All use subject to JSTOR Terms and Conditions BONPLANDIA16 (Supi), 2007 Table of Contents Abstract 9 Resumen 10 Introduction 12 History of the Collections 15 Summary of Germplasm Explorations 18 The Fruit of Arachis and its Capabilities 20 "Sócias" or Twin Species 24 IntraspecificVariability 24 Reproductive Strategies and Speciation 25 Dispersion 27 The Sections of Arachis ; 27 Arachis L 28 Key for Identifyingthe Sections 33 I. Sect. Trierectoides Krapov. & W.C. Gregorynov. sect. 34 Key for distinguishingthe species 34 II. Sect. Erectoides Krapov. & W.C. Gregory nov. sect. 40 Key for distinguishingthe species 41 III. Sect. Extranervosae Krapov. & W.C. Gregory nov. sect. 67 Key for distinguishingthe species 67 IV. Sect. Triseminatae Krapov. & W.C. Gregory nov. sect. 83 V. Sect. Heteranthae Krapov. & W.C. Gregory nov. sect. 85 Key for distinguishingthe species 85 VI. Sect. Caulorrhizae Krapov. & W.C. Gregory nov. sect. 94 Key for distinguishingthe species 95 VII. Sect. Procumbentes Krapov. & W.C. Gregory nov. sect. 99 Key for distinguishingthe species 99 VIII. Sect. Rhizomatosae Krapov. & W.C. Gregorynov. sect. 113 Key for distinguishingthe species 114 Ser. Prorhizomatosae Krapov. & W.C. Gregory nov. ser. 114 Ser. Rhizomatosae 117 IX. Sect. Arachis 123 Key for distinguishingthe species 124 Arachis hypogaea L 167 Key for distinguishingthe taxa of A. hypogaea 171 Taxa of dubious position 175 Acknowledgements 175 Literature Cited 176 Numerical List of the Taxa 181 List of Collectors 182 List of Parent Numbers used in Crosses 192 Abbreviations of Collectors' Names 194 InterspecificCrosses Conducted 195 List of ScientificNames Cited in Text 203 8 This content downloaded from 195.221.60.18 on Tue, 24 Jun 2014 00:12:00 AM All use subject to JSTOR Terms and Conditions A. Krapovickasand W.C. Gregory, Arachis (Leguminosae) TAXONOMY OF THE GENUS ARACHIS (LEGUMINOSAE) byAntonio Krapovickas and Walton C. Gregory Abstract: Krapovickas,A. & W.C. Gregory.2007. Taxonomyof the genus Arachis (Leguminosae).Translated by D.E. Williams& C.E. Simpson.Bonplandia 16 (Supl.): 1-205. ISSN: 0524-0476. Almost100 years elapsed betweenLinnaeus' naming the thenlone species of Arachis(A. hypogaeaL.) knownto Europeans,and the first taxonomie treatment ofthe genus by Bentham in 1841. Duringthe next 100 yearsfive to tenadditional species descriptionsappeared, assigning differentspecies to the same names,and differentnames to the same species. By mid-20th Century,itwas impossibleto examineany herbarium collection of Arachis and assignany epithet withany assurance to anyspecimen (which was nota typecollection) except to A. hypogaea, A. guaranitica, A. tuberosaand A. villosulicarpa. In ourtreatment, the literatureof this botanical chaos in Arachisis reviewedin detailand an assessmentis made ofthe foundationsfor its occurrence.It is shownthat the bases forthe confusionlay in the combinationof the esotericnature of the differentiatingmorphological featuresof Arachis, the fragmentaryearly collections, and the representationof species by seedlingspecimens. Also,it is relatedhow, in 1959,we decidedto re-explorethe type locality of each speciesthen known,collect therein complete plant specimens and therebyresolve the problem.Thirty-five years,two generations of plant collectors and around2000 collectionslater, we presenthere 69 speciesdescriptions of Arachis, species distributed inSouth America east ofthe Andes, south of theAmazon, north of La Plataand from NW Argentina to NE Brazil. We soon discoveredthat the mostsignificant characters of Arachislay in theirunderground structures,including their fruits, rhizomatous stems, root systems and hypocotyls. We showedthat these definingcharacters tended to clusterthe collectionsinto groups which wereassociated with generally different geographic areas and ecologicalfeatures. We drewa sample of 100 collectionsrepresenting these clusters,areas and features,and arrangedthem in a hybridizationdiallel and showed,in crosses between collections representing differentclusters of characters, areas and features,a remarkablenumber of complete failures to cross-fertilizeand, in those hybridsthat were recovered, a highdegree of F., hybridinfertility. When these cross-incompatibilitiesand polleninfertilities were combinedwith the data on characterclustering, the ninedistinct sections of the genus presentedhere then crystallized. Figuresimposed upon maps of SouthAmerica illustrate the geographicdistributions ofthese sections. The collectionswere then assigned to thedifferent sections on thebases ofcross-incompatibility and exo-morphologiccharacter clustering. When these groupswere made, the esoteric characteristicsreferred to above,so confoundingwhen applied across sectionallines, became highlypertinent when applied to theproblem of species differentiation between collections within sections. These characteristics,applied in conjunctionwith chromosome cytology, chromatographicand antigenicreactions, variations in intra-sectionalhybrid fertility and adaptationsof plant form, and annualand perennialhabit, allowed us to assemblethe following taxa ofthe genus Arachis : Section I. TRIERECTOIDESnov:. 1. A. guaranitica, 2. A. tuberosa. SectionII. ERECTOIDES nov.: 3. A. Martii, 4. A. brevipetiolatanov., 5. A. Oteroinov., 6. A. Hatschbachiinov., 7. A. cryptopotamicanov., 8. A. majornov., 9. A. Benthamii,10. A. douradiana nov.,11. A gracilisnov., 12. A. Hermanniinov., 13. A. Archerinov., 14. A. stenophyllanov., 15a. 9 This content downloaded from 195.221.60.18 on Tue, 24 Jun 2014 00:12:00 AM All use subject to JSTOR Terms and Conditions BONPLANDIA16 (Supl.),2007 A paraguariensissubsp. paraguariensis, 15b. A paraguariensissubsp. capibarensisnov. SectionIII. EXTRANERVOSAEnov:. 16. A. setinervosanov., 17. A. Macedoinov.t 18. A. marginata, 19. A. prostrata, 20. A lutescens, 21. A retosanov., 22. A Burchelliinov., 23. A Pietrarelliinov., 24. A villosulicarpa.Section IV. TRISEMINATAEnov.: 25. A inseminatanov. SectionV. HETERANTHAEnov:. 26. A Giacomettiinov., 27. A. sylvestris,28. A pusilla,29. A Dardaninov. SectionVI. CAULORRHIZAEnov: 30. A repens,31. A Pintoinov. Section VII. PROCUMBENTESnov: 32. A lignosanov. comb., 33. A. Kretschmerinov., 34. A Rigonii,35. A chiquitananov., 36. A. matiensisnov., 37. A. appressipilanov., 38. A. Vallsiinov., 39. A. subcoriaceanov. Section VIII. RHIZOMATOSAE nov., Series PRORHIZOMATOSAE nov: 40. A. Burkartii.Series RHIZOMATOSAEnov: 41. A. pseudovillosanov. comb., 42a. A. glabratavar. glabrata,42b. A. glabratavar. Hagenbeckii.Section IX. ARACHIS : 43. A. glandulifera,44. A. cruziananov., 45. A. montico/a,46. A magnanov., Al. A. ipaënsisnov., 48. A. validanov., 49. A. Williamsiinov., 50. A. Batizocoi,51. A. duranensisnov., 52. A. Hoehneinov., 53. A. stenosperma nov.,54. A. praecoxnov., 55. A. palustrisnov., 56. A. benensisnov., 57. A. trinitensisnov., 58. A. decoranov., 59. A. Herzogiinov., 60. A. microspermanov., 61. A. villosa,62. A. helodes,63. A. correntinanov. comb.,64. A. Simpsoniinov., 65. A. Cardenasiinov., 66. A. Kempff-Mercadoi nov.,67. A. Diogoi,68. A. Kuhlmaniinov., 69a. A. hypogaeasubsp. hypogaeav ar. 1. hypogaea, var.2. hirsuta,69b. A. hypogaeasubsp. fastigiatav ar. 1. fastigiata,var. 2. peruviananov., var. 3. aequatoriananov., var. 4. vulgaris. The autogamousreproductive systems, agametic reproduction, underground fruiting habit and the limitedmeans of seed dispersalare shownto be logicallytied to thedrift in chromosomal organizationwhich gives riseto noticeableincreases in infertilityin crosses betweendifferent collectionsof the same species,to a variablyhigher infertility incrosses between species within sections,to a neartotal infertility incrosses between species from different sections. Theevolutionary and phylogeneticrelationships between the different sections are discussedand are furthershown in a sequenceof diagrams illustrating the ideas presented.It is evidentthat the geneticdistances separating the sectionsare farfrom being of the same magnitude.The presumablyolder sections (Triseminatae,Trierectoides, Erectoides, Extranervosae and Heteranthae),except for section Erectoides, are muchmore isolated from the remaining sections and fromeach otherthan those taken to be ofmore recent origin (Procumbentes , Caulorrhizae, Rhizomatosaeand Arachis). SectionArachis is byfar the largest, containing about 40% ofthe species described. Species of thissection appear to be spreadinginto new territory and to be invadingareas occupiedby species ofother sections. They grow intermixed with populations of Extranervosae in the upper Paraguaybasin and occupycommon ground with section Procumbentes in the Gran Pantanal. Theyhave reachedthe shores of La Plata and thesoutheastern coast ofBrazil and growfrom Yala in NW Argentinato the Tocantinsin NE Brazil.They include the worldwidecultivar, A. hypogaea. Essentiallyevery published work on thebotanical history and taxonomy
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