Taxonomy of the Genus Arachis (Leguminosae)

BONPLANDIA16 (Supi): 1-205.2007

BONPLANDIA 16 (SUPL.): 1-205. 2007

TAXONOMY OF THE GENUS ARACHIS (LEGUMINOSAE)

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.

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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

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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.

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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 of Arachis is presented herein the individualspecimen references and in the generalbibliography. The historyof A. hypogaeafrom the early 16th Century to morerecent times, along with the common names in severalnative American languages, provide a perspectiveon theantiquity ofthis cultivar and the levelof civilization required for its creation. Six appendicesprovide supporting data and mattersof record. Diagnostic keys to thesections andto the species within each sectionselect the more sharply distinguishing characteristics ofthe sectionsand species. Nineteenline drawings capture the sectionaland species structuresof wholeplants, root systems, fruit orientations, agametic reproductions from fruiting structures, carpelshapes and surfacefeatures of leaves and stems. Key words: Arachis,peanut, groundnut, Leguminosae, history, taxonomy, genetic resources, phytogeography

Resumen: 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.

Pasaron casi 100 años entrela designaciónpor Linneode la entoncesúnica especie de

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Arachis(A. hypogaeaL.) conocidapor los europeos,y el primertratamiento taxonómico del géneropor Benthamen 1841. Durantelos siguientes100 años, aparecieroncinco a diez descripcionesde especies adicionales,que asignaban diferentesespecies a los mismos nombres,y diferentesnombres a las mismasespecies. A mediadosdel SigloXX, era imposible examinarun ejemplar de herbariode Arachisy asignarcon algunacerteza algún epíteto a algún espécimen(que no fueraun ejemplar tipo) excepto a A. hypogaea, A. guaranitica,A. tuberosay A. villosulic arpa. En nuestrotratamiento, la literatura de este caos botánicoen Arachisesta revisadaen detalley se hace un análisisde los fundamentosde su ocurrencia.Se demuestraque las bases de la confusiónmoran en la combinaciónde la naturalezaesotérica de los caracteresmorfológicos diferenciadosde Arachis, de los especímenesfragmentarios de antaño, y de la representación de especiespor plántulas. Además,se relatacómo, en 1959,decidimos reexplorar la localidadtipo de cada especiehasta entoncesconocida, y recolectarallí especímenesde las plantasenteras y así resolverel problema.Después de treintay cincoaños, dos generacionesde coleccionistasde plantas,y alrededorde 2000 colecciones,presentamos aquí las descripcionesde 69 especiesde Arachis, especiesdistribuidas en Sudaméricaal este de los Andes,al sud delAmazonas, al nortede La Platay desde el noroesteargentino hasta el nordestede Brasil. Descubrimosmuy pronto que los caracteresmás significativosde Arachisresiden en sus estructurassubterráneas, incluyendo sus frutos,tallos rizomatosos,sistemas radicales e hipocótilos. Demostramosque estos caracteresdeterminantes tienden a aglomerarlas coleccionesen gruposque se asociancon áreas geográficasyformaciones ecológicas generalmente diferentes. Hicimosun muestreode 100 materialesrepresentativos de aquellos grupos, áreas, y formacio- nes y los arreglamosen unexperimento dialélico de cruzamientosy mostramos, en cruzamientos entremateriales de los diferentesgrupos, un númeronotable de fracasoscompletos en la fertilizacióncruzada y, en aquellos híbridosque se lograron,se observóuna alta tasa de infertilidaden la Fr Cuandose combinaronestas incompatibilidadese infertilidades de polen híbridocon los datosde agrupamientode caracteresmorfológicos, se cristalizaron entonces las nuevedistintas secciones del géneroaquí presentadas.Las figurasimpuestas sobre mapas de Sudaméricailustran las distribucionesgeográficas de estas secciones. Las colecciones,entonces, fueron asignadas a las diferentessecciones sobre la base de las incompatibilidadesde cruzamiento y de los agrupamientosde caracteresexo-morfológicos. Al hacerestos grupos,las caracteristicasesotéricas a las cuales se hace referenciaarriba, tan confusascuando se aplicana travésde los límitesseccionales, se volvieronaltamente pertinen- tes al ser aplicadasal problemade la diferenciaciónespecífica entre materiales dentro de las secciones.Estas características,aplicadas en conjuntocon la citologíacromosomica, las reac- cionescromatográficas y antigénicas, las variacionesen la fertilidadhíbrida intra-seccional y las adaptacionesde formade planta,y de hábitoanual o perenne,nos permitiódefinir los siguientes taxadel géneroArachis: Sección I. TRIERECTOIDESnov.: 1. A. guaranitica, 2. A. tuberosa.Sección II. 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. douradiananov., 11. A gracilisnov., 12. A. Hermanniinov., 13. A. Archerinov., 14. A. stenophyllanov., 15a. A. paraguarìensissubsp. paraguariensis,15b. A. paraguarìensissubsp. capibarensis nov. Sección III. EXTRANERVOSAE nov.: 16. A. setinervosanov., 17. A. Macedoinov., 18. A. marginata,19. A. prostrata, 20. A. lutescens,21. A retusanov., 22. A. Burchelliinov., 23. A. Pietrarelliinov., 24. A. villosulicarpa. SecciónIV. TRISEMINATAEnov.: 25. A. inseminatanov. Sección V. HETERANTHAEnov.: 26. A. Giacomettiinov., 27. A. sylvestris,28. A. pusilla,29. A. Dardani nov. Sección VI. CAULORRHIZAEnov.: 30. A. repens,31. A. Pintoinov. Sección VII. PROCUMBENTES nov.: 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. Sección VIII. RHIZOMATOSAE nov., Serie PRORHIZOMATOSAE nov.: 40. A. Burkartii.Serie RHIZOMATOSAEnov.: 41. A. pseudovillosanov. comb., 42a. A. glabratavar. glabrata, 42b. A. glabratavar. Hagenbeckii. Sección IX. ARACHIS: 43. A. glandulifera,44. A. cruziananov., 45. A. montícola,46. A. magnanov., 47. A. ipaënsisnov., 48. A. validanov., 49. A. Williamsiinov., 50. A. Batizocoi,51. A. duranensisnov., 52. A. Hoehneinov., 53. A. stenospermanov., 54. A. praecox

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nov.,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-Mercadoinov., 67. A. Diogoi, 68. A. Kuhlmaniinov., 69a. A. hypogaeasubsp. hypogaea var. 1. hypogaea,var. 2. hirsuta,69b. A. hypogaeasubsp. fastigiata var. 1. fastigiata,var. 2. peruviananov., var. 3. aequatoriananov., var.4. vulgaris. Se demuestracómo los sistemasreproductivos autógamos, la reproducciónagamética, el hábito de fructificaciónsubterránea y el mecanismolimitado de dispersiónde semillasestán lógicamen- te ligadoscon la derivade organizacióncromosomica que da origena incrementosnotables de infertilidaden cruzamientos entre diferentes accesiones de la mismaespecie, a una infertilidad variablementemás altaen cruzamientosentre especies dentro de las secciones,hasta una casi totalinfertilidad entre especies de diferentessecciones. Las relacionesevolutivas y filogenéticasentre las diferentessecciones están discutidasy tambiéndemostradas en una secuenciade diagramas¡lustrando las ideas presentadas.Es evidenteque las diítandas genéticasque separanlas seccionesestán lejos de ser todas de la mismamagnitud. Las seccionespresumiblemente más antiguas(Triseminatae, Trierectoides, Erectoides,Extranervosae y Heteranthae), excepto por la secciónErectoides, están mucho más aisladas de las seccionesrestantes y entresí que aquellas seccionesque se considerande origenmás reciente(Procumbentes, Caulorrhizae, Rhizomatosae y Arachis). La secciónArachis es pormucho la más grande,pues contienecerca de 40% de las especies descritas.Parece que las especiesde esta secciónse estánexpandiendo hacia nuevosterrito- riose invadiendoáreas ocupadas porespecies de otrassecciones. Crecen entremezcladas con poblacionesde Extranervosaeen la cuenca alta del ríoParaguay y ocupanterrenos comunes con la secciónProcumbentes en el GranPantanal. Han llegado hasta las orillasde La Platay a la costasureste de Brasil,y crecendesde Yala en el noroestede la Argentinahasta el ríoTocantins en el nordestede Brasil.Incluyen el cultígenode importanciamundial, A. hypogaea. Esencialmentecada trabajopublicado sobre la historiabotánica y la taxonomíade Arachisestá presentadoaquí en las referenciasa especímenes individuales y en la bibliografíageneral. La historiade A. hypogaeadesde principiosdel Siglo XVI hasta tiempos más recientes,junto con los nombrescomunes en variosidiomas autóctonos americanos, nos dan una perspectivasobre la antigüedadde este cultivoy el nivelde civilizaciónrequerido para su creación. Seis apéndicesproporcionan datos de apoyoe informaciónde archivo. Claves diagnósticas a las seccionesy a las especiesdentro de cada secciónseleccionan los rasgosmás distintivosde las seccionesy especies.Diecinueve dibujos de líneacapturan las estructurasclaves para distinguir las secciones y especies, incluyendoplantas enteras, sistemas radicales, orientaciones de frutos,reproducción agamética a partirde estructurasfructíferas, formas de carpelos,y la fisionomíasuperficial de hojas y tallos.

Palabras clave: Arachis, maní, cacahuete, Leguminosae,historia, taxonomía, recursos genéticos,fitogeografia.

Introduction In 1939, Burkartseparated Zornia , placing it in the subtribePoiretiinae . This point of view was continuedby Schulze-Menz (1964) The genus Arachis was placed in the tribe and Rudd (1981), who placed the genus Hedysareae subtribe Stylosanthinae by Arachis in the tribeAeschynomeneae subtribe Bentham (1865) togetherwith Chapmannia, Stylosanthinae(Benth.) Rudd, togetherwith Stylosanthes and Zornia. Bentham's criteria the genera Arthrocarpum (2 spp. from were continuedby Mohlenbrock (1962) who Somalia and Socotra), Pachecoa (1 sp. from also included Pachecoa , and Hutchinson Mexico and Guatemala), Chapmannia (1 sp. (1964), who elevated the group to the fromFlorida, USA) and Stylosanthes(ca. 25 categoryof tribe. spp. fromthe Old and New World tropicsand

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subtropics). Of all these genera, that with Burkart(1939), afterstudying the Argentine greatest affinityto Arachis is Stylosanthes species, followed Chevalier's 1933 criteria with which it shares fused stipules and the and recognized five species: A. marginata same basic chromosomenumber (x=10). (now A. Burkartii),A. prostrata(=A. glabrata), By studyingthe pollen morphologyof the A. villosa, A. pusilla (= A. duranensis and A. Hedysareae, Pire (1974) foundthat the genus montícola) and A. hypogaea. Burkart was Zornia shows greatest affinity with the correctin the delimitationof the species, but Stylosanthinae, which would support the withthe material at his disposal he was unable criteriaof Hutchinson(1964). Two types of to correctlyinterpret the names. He was the pollen grains are found in the genus Arachis, first author to describe and illustrate a one tricolpate,in A. hypogaea and similarto biarticulatedsubterranean fruit, corresponding those of Zornia, and the othersyncolpate, si- to A. montícola, and shortly afterwards milar to those of Stylosanthes. (Burkart 1942) added those of A. villosa, The firsttreatment of the genus Arachis is observed fromcultivated specimens. owed to Bentham (1841), who described the At about the same time, Hoehne (1940) firstfive wild species. Later, in 1859, he lists published a revisionof the genus Arachis. He seven species, includingA hypogaea. presenteda key based almost exclusively on Chevalier(1929a), in an essay on systematic leaflet characters.He relied upon collections classification,tried to definethe relationships made by Otero and Archer,and made use of among the species, which he grouped into (1) live materialof various species. The work is A. prostrata with its races villosa, glabrata, well illustratedand the figures demonstrate marginata and pusilla, (2) A. tuberosa with interestingvegetative characters, especially the related forms A. guaranitica and A. undergroundones, that he did not take into paraguariensis and (3) A. Diogoi withnarrow account in his classification scheme. Hoehne leaflets. He found that A. pusilla, being the recognized 11 species, some with numerous only annual wild species, was the most forms, which are described without Latin analogous withthe cultivatedpeanut. diagnoses and consequently are invalid, Shortly thereafter, Chevalier (1929b, having been published after 1935 (Art. 36 of 1929c) describedthe new species A. sylvestris theInternational Code ofBotanical Nomenclature). and became the firstauthor to describe and The interpretationof the majority of the illustrate the subterranean fruit of a wild species is confusingbecause Hoehne did not species. In thiscase, the fruitis constitutedby attach importanceto the biological forms. only a single article. An importantcontribution was made by Later, Chevalier (1933) made the first Mendes (1947) who,utilizing the nomenclature attemptat classificationby producinga key. of Hoehne, established the diploid character He separated perennialswith slightlymarked of various wild species. His work is perfectly leafletmargins (A. glabrata), erectperennials documented, enabling us to correlate his (A. paraguariensis , A. tuberosa and A. results with the following species: A. Oteroi guaranitica), perennials with subterranean (V.82), A. Benthamii (Y.83), A. Archeri stolons (A. marginata and A. Hagenbeckii), (V.84) and A. major (V.85), for which he perennials with stolons growing upon the determined2n=20 chromosomes. ground (A. villosa and A. Diogoi), and In 1949, Krapovickas and Rigoni obtained annuals (A. pusilla, A. sylvestris and A. the firsttriploid interspecific hybrid (2n=30) hypogaea). He consideredA. sylvestristo be aftercrossing A. correntina (2n=20) with A. the ancestor of some varieties of cultivated hypogaea (2n=40). peanut. AlthoughChevalier was unfortunatein In 1951, Krapovickasand Rigoniencountered his interpretationof some species such as A. the first wild species with the same glabrata and A. marginata,his systemwas an chromosomenumber as the cultivatedpeanut importantcontribution because it is thefirst to (2n=40), thisbeing A. montícola,which at the utilize vegetative characters, both aerial timethey identified as A. pusilla. and subterranean,to distinguishthe species. In 1951, Gregory, Smith and Yarbrough,

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 followinga thoroughmorphological analysis, Dardani (Hoehne 1940), A. hypogaea laid the foundationsof the presenttaxonomy (Hoehne 1940) and A. glabrata var. glabrata of cultivated peanut by distinguishingthe (Burkart1939, Hermann1954). branchingpatterns that Bunting(1955) would The name A. glabrata was used for: A. later denominate"alternate" and "sequential" glabrata (Bentham 1859), A. helodes and Krapovickas and Rigoni (1960) would (Bentham 1859, Chevalier 1933, Hoehne utilize to establish the nomenclaturefor the 1940), A. prostrata (Chevalier 1933, intraspecifictaxa of A. hypogaea. Hermann 1954), A. repens (Chevalier 1933, In 1954, Hermannpublished a synopsis of Hermann 1954), A. stenosperma (Chevalier the genus Arachis, using the new collections 1933, Hermann 1954), A. pseudovillosa of Stephens and Hartley (1949), in which he (Hoehne 1940), A. glabrata var. glabrata attemptsto reconcile the criteriaof Chevalier (Hoehne 1940, Hermann 1954), A. glabrata (1933), Burkart (1939) and Hoehne (1940). var. Hagenbeckii (Hoehne 1940, Hermann Hermann's treatmentrecognizes nine species 1954), A. Archeri (Hermann 1954), A. and suffersfrom the same shortcomingsas the appressipila (Hermann 1954) and A Dardani previousschemes because of the unavailability (Hermann 1954). of livingmaterial. The name A. pusilla was used for: A. In 1957, Krapovickas and Rigoni describe pusilla (Bentham 1859, Chevalier 1933, the wild tetraploidspecies A. montícola and Krapovickas & Rigoni 1957), A. Dardani the obtentionof fertilehybrids when crossing (Bentham 1859, Chevalier 1933, Hermann it withA. hypogaea. 1954), A. sylvestris (Chevalier 1933), A. With few exceptions,such as A. guaranitica montícola (Burkart 1939, Krapovickas & and A. tuberosa, the species were very Rigoni 1951) and A. duranensis (Burkart difficultto interpretdue to the great apparent 1939, Krapovickas & Rigoni 1957). similaritythey have withone another,to such In response to this chaotic situation, we an extentthat in some cases it is practically began in 1959 a methodicalexploration to try impossible to separate them if informationis to collect herbarium specimens and living unavailable on their biological form,corolla plants in the type localities of all the species - color, peg position and fruitmorphology describedup to thatpoint and whereverelse a informationcommonly absent fromherbarium herbarium specimen of Arachis had been specimens. made. To give some idea of these difficulties,we This new material was studied from offer a few interpretations of names differentperspectives. To thefield observations frequentlyfound in the literature. were added data from interspecific crosses For example, the name A. marginata has (Gregory, W.C. & Gregory 1967, Gregory, been used to identifythe followingspecies: A. M.P. & Gregory 1979), chromosomestudies marginata (Bentham 1859, Chevalier 1933), (Smartt1964), chromatography(Krapovickas A. Oteroi (Bentham 1859, Chevalier 1933, & al. 1973, Krapovickas & al. 1974) and Hoehne 1940, Hermann 1954), A. Burkartii palynologicalstudies (Pire 1964). (Bentham 1859, Chevalier 1933, Burkart In 1973, Gregory& al. enumeratedthe ma- 1939, Hoehne 1940, Hermann 1954), A. terial of the wild species and cultivated pseudovillosa (Chevalier 1933, Hoehne 1940, peanut obtained during the aforementioned Hermann 1954), A. glabrata var. glabrata explorations and presented an outline in (Chevalier 1933, Burkart1939, Hoehne 1940, which names for sections within the genus Hermann 1954) and A. lignosa (Hermann were proposed. 1954). Another approximation was offered The nameA prostrata has been used for:A. (Krapovickas 1973) in a seminar given in prostrata (Bentham 1859), A. lutescens 1969 in which, besides presentinga scheme (Bentham 1859), A. stenosperma (Bentham forordering the genus, various "genocenters" 1859, Hoehne 1940), A. sylvestris(Hoehne of cultivatedpeanut in South America were 1940), A. Cardenasii (Hoehne 1940), A. proposed.

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A very importantcontribution towards the When Bentham (1859) published the definitionof the sections and establishingthe Arachis treatmentin the Flora Brasiliensis of phylogeneticrelationships among them was Martius, very few collections were added. based on the resultsof diallelic crosses made Among these were the collections of using material from 100 sources (Gregory, Gaudichaud in Rio de Janeiro (A. M.P. & Gregory 1979). In light of this stenosperma),Arsène Isabelle in Rio Grande information'sgreat value in the delimitation do Sul (A. Burkartii),Manso in Cuiabá, Mato of the taxa, we reprint all of the crosses Grosso (A. helodes), Martius in northeast obtained, with the updated species names, in Brazil, SaintHilaire in southernBrazil and Uru- the chapter"Interspecific Crosses Conducted" guay {A. Burkartii, A. glabrata and A. (p. 195). villosa) and Weddell in Mato Grosso (A. Resslar (1980) compares the attemptsat lutescens). All of these herbariumspecimens orderingthe genus Arachis of Gregory& al. lack thesubterranean portions of theplants, for (1973) and Krapovickas (1973), and introdu- which, in general, they have been ces one morename: sect.Goniorhiza, appearing misinterpreted. in a mimeographed note (Krapovickas A heftyincrease in knowledge about the 1969b), and therefore not effectively genus Arachis was produced with the published (Art. 29 of the International Code collections made by E. Hassler in Paraguay, of Botanical Nomenclature). publishedby Chodat & Hassler (1904). Nota- Recently, Smartt (1990) did an excellent ble among the 20 collectionsmade by Hassler job of bringingthings up to date by compiling are the new species A. guaranitica, A. and analyzing the existinginformation about paraguariensis, A. lignosa and A. A. hypogaea and its relationshipwith the wild pseudovillosa, as well as demonstratingthe species. ample distributionof A. glabrata in eastern Paraguay. In September of 1908, Cesar Diogo Historyof the Collections collected a wild Arachis around Lagoa Gahiba, on the Paraguay River north of Corumbá, that Hoehne (1919) would later Up until 1841, the year in which Bentham name A. Diogoi. describedthe firstwild species ofArachis, the Between 1908 and 1914, F. C. Hoehne and only known species was A. hypogaea L., the J. C. Kuhlmannparticipated in theexploration cultivated peanut, described by Linnaeus in of Mato Grosso by the Comissão Rondon. The 1753. The five species describedby Bentham scantiness of theircollections is conspicuous were the product of collections made by -only eightspecimens- and was surelydue to travelersin Brazil at the beginningof the 19th the factthat the routeof the telegraphline, the Century.Johann E. Pohl was in Trahiras(Oct. principal objective of the Comissão Rondon, 18-24, 1819), Goiás, where he collected A. took them over the higherparts of the Serra prostrata. Ludwig Riedel passed by Rio Par- dos Parecis, an obstacle that constitutesthe do, in Mato Grosso do Sul (Oct. 1826), where northernlimit of the genus Arachis in Mato he collectedthe typespecimens of A. glabrata Grosso. Kuhlmann collected A. glabrata in and A. tuberosa. Jacques S. Blanchet, who Diamantina and A. nambyquarae in Pimenta began collecting in Bahia in 1828, collected Bueno. Hoehne collectedA. Hoehnei in Amo- A. pusilla at "Serra Jacobina."In 1832, James lar and A. appressipila in Corumbá, on the Tweedie traveledthe Uruguay River wherehe Paraguay River. encounteredA. villosa. Two singular events occurred during the George Gardner, in October of 1839, War of the Chaco, betweenBolivia and Para- discovered A. marginata at "Mission of guay. In October 1934, M. Cárdenas, a Duro" (today Dianópolis in the state of memberof the health services of the Bolivian Tocantins), a species that Gardner himself army, discovered A. Cardenasii in Roboré, describedin 1842. notfar from the front. In March 1935, T. Rojas,

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 who accompanied the vanguard of the Between 14 November 1947 and 11 April Paraguayan army, collected A. duranensis in 1948, a tripwas conductedby J. L. Stephens Carandayti,shortly before the end of the war (Division of Plant Exploration and (14 June 1935). Introduction, USDA) and W. Hartley In December 1932 and January1933, Jorge (CSIRO, Australia) with the principal Ramos de Otero, who was interested in objective of collectingseeds and livingplants studying the grazing potential of Mato of cultivatedpeanut varieties as well as wild Grosso, made his firsttrip to Campo Grande species of Arachis (Hartley 1949). They where he collected A. Archeri,A. Martii and collected wild species in northernand central A. Oteroi forthe firsttime. Corrientes, southern Misiones and F. C. Hoehne and A. Gehrt collected A. northeasternEntre Rios (Argentina);in Rive- Oteroi and A. Archeri in Campo Grande (5 ra and Colonia (Uruguay); in Asunción, August 1936), which they cultivated at the Paraguarí,Puerto Casado and Villa Rica (Pa- Institutode Botánica de São Paulo and at the raguay); and in Cuiabá (Mato Grosso, Brazil). InstitutoAgronómico de Campinas. The herbariummaterial was used by Hermann An importanttrip was made by W. Andrew (1954) forhis synopsisof the genus Arachis. Archer of the USDA who visited Campo In Argentina,explorations dedicated to the Grande (MS) with A. Gehrt(1-11 September collection of Arachis germplasm began in 1936) where he collected A. glabrata, A. 1945 with the initiation of plant breeding Archeri,A. Oteroi and A. Benthamii.He later programs at the Manfredi Agricultural traveledto Rio Grande do Sul and to Uruguay ExperimentStation (Cordoba) and with the (Rivera) (1 1-15 November 1936) to collect.4. organization of the Department of Plant Burkartii.He thenwent into Argentina where, Exploration and Introduction(DEIP) of the aftervisiting the Botanical Garden of the Fa- Ministeriode Agriculturade la Nación, under cultad de Agronomía de Buenos Aires (27 the directionof E. C. Clos. November 1936), he collected samples of A. The firstintroductions of wild peanuts are villosa and then traveled to Misiones where, owed to J. R. Báez and V. A. Rigoni (director in Posadas and the surrounding area (10 and subdirector,respectively, of the Manfredi December 1936), he collected A. glabrata. ExperimentStation), who in 1947 obtained From there,Archer crossed into Paraguay to samples ofA. correntinain northernCorrien- collect in Encarnación (14 December 1936), tes, and to J. R. Báez who collectedA. villosa Villa Rica (16 December 1936) and the area in Colonia, Uruguay in 1949. Plants of A. - around Asunción (24 December 1936 9 glabrata, provided by the DEIP, were also January 1937) where he made herbarium available. This was the material with which specimens of the two varietiesof A. glabrata. V. A. Rigoni attemptedthe firstinterspecific He endedhis collectionwith A. villosa whichhe crosses, obtainingthe firsthybrids between A. foundin Colonia, Uruguay,on 17 January1937. hypogaea and A. correntina(Krapovickas & In 1939, J. Ramos de Otero returnedto Rigoni 1949, 1951). Mato Grosso. Between April 14 and May 28 In May of 1950, Báez, Rigoni and he collected along the Amambay River and at Krapovickas traveledto theprovinces of Salta Campanario, Ponta Porā, Bela Vista, and Jujuy (Argentina) where, after visiting Aquidauana and Campo Grande (Otero 1941, several localities where herbariumspecimens 1952). had been made of wild species, they were The material collected by Otero, Hoehne only able to finda few plants at Yala (Jujuy) and Archerwas utilized by F. C. Hoehne for which served as the basis for the description his monographof the genus Arachis (Hoehne of A. montícola (Krapovickas & Rigoni 1940) and by A. J. T. Mendes (1947) for the 1957). first chromosome counts of wild peanut In March of 1953, Rigoni, Pietrarelliand species, determiningthat 2n=20 in A. Oteroi, Krapovickas made a trip to Corrientes and A. Benthamii, A. Archeri, A. major and A. Misiones (Argentina) to collect living villosulicarpa . specimens of wild species and samples of

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) cultivatedpeanuts. Grosso); and São Paulo. Then, between May In May of 1953, Krapovickas collected 11-19, 1961, W. C. Gregory and A. some fruitsat Campo Durán (Salta, Argenti- Krapovickas traveled in Mato Grosso to Rio na) that later produced plants of A. Brilhante,Dourados, Ponta Porā; and in Para- duranensis. guay to P. J. Caballero, Ñu Porá, Concepción In 1954-1955, Krapovickas, thanks to a and PuertoCasado. grant fromthe GuggenheimFoundation, had In 1967, Gregoryand Krapovickas traveled the opportunityto studyherbarium specimens to northeastBrazil, where theyvisited Bahia, at Brazilian and NorthAmerican institutions, the eastern part of Pernambuco and the thatenabled him to plan futureexplorations. northwestof Ceará. In January and February of 1958, with The tripsfrom 1959 to 1967 are discussed funds from the Instituto Miguel Lillo of by Gregory(1973). The numbersassigned to Tucumán (Argentina), Krapovickas traveled the collections are from the herbarium of to Bolivia where he searched between Krapovickas, with the collectors listed on the Cochabamba and Santa Cruz, without specimens in alphabetical order: Gregory, success. He began encounteringwild species Krapovickas and Pietrarelli. of Arachis in Santa Cruz de la Sierra (A. Between April and June of 1968, R. 0. Rigonii), in Roboré (A. Cardenasii) and along Hammons and W. R. Langford (CRD, ARS, the Parapeti River (A. Batizocoi). USDA) traveledto Argentina,Brazil and Uru- With financialsupport from their respecti- guay. Together with J. R. Pietrarelli (EEA ve institutions,W. C. Gregory(USDA), J. R. Manfredi, Cordoba), they explored the Pietrarelli (INTA) and A. Krapovickas departmentsof Colonia, Salto, Rivera and (CONICET), undertook an extensive Tacuarembó in Uruguay. With the exploration, between January and April of collaborationof Krapovickas, theytraveled to 1959, with the objective of obtaining living the followinglocalities: Corrientes(Argenti- materialfrom wherever herbarium specimens na); Uruguaiana, Porto Alegre, Itajai, of Arachis had been collected,trying as much Curitiba, Antonina, São Paulo, Porto 15 de as possible to visit the type localities. The Novembro,Campo Grande,Aquidauana, Bela itineraryof Corrientes,Posadas (Argentina); Vista and Ponta Porā (Brazil). One of themost Encarnación,Asunción, Caacupé, PuertoEm- importantacquisitions made was that of A. balse, Asunción, P. J. Caballero (Paraguay); stenosperma, collected in Antonina and Ponta Porā, Bela Vista, Amambai, Campo Paranaguá, thanksto the collaboration of G. Grande, Aquidauana, Porto 15 de Novembro, Hatschbach,of Curitiba. Campo Grande,Rondonópolis, Cuiabá, Rosa- From May 1 to June9, 1971, with funding rio Oeste, Chapada dos Parecís, Cuiabá, Jataí, fromFAO, Rome, Italy, A. Krapovickas, L. Ituiutaba, São Paulo, Corumbá (Brazil); A. Mroginski and A. Fernández traveled to Roboré, Santa Cruz (Bolivia); and Salta (Ar- Orán (Salta, Argentina) and from there gentina) was traveledusing various means of crossed into Bolivia to make the journey transportation. When an automobile was between Bermejo, Tarija, Villa Montes and available, stops were made every ten km, as Yacuiba. They encountered wild Arachis was the case betweenCorrientes and Posadas, species only in the Bolivian "chaco," near Vi- and in Mato Grosso and Minas Gerais lla Montes, where they collected A. ipaënsis (Pietrarelli1961). and A. duranensis. Between April 3 and May 8, 1961, W. C. A proper analysis of the germplasm is Gregory,J. R. Pietrarelliand A. Krapovickas presentedby Banks (1976), which shows the visited São Paulo, Ituiutaba, Capinópolis need to make additional collections of the (Mato Grosso); Itumbiara, Caldas Novas, cultivatedpeanut and the wild species before Piracanjuba, Anápolis, Niquelãndia, Ceres, the destruction of their present habitats Brasilia, Formosa, Dianópolis (Goiás); occurs. Barreiras (Bahía); Brasilia, Patos de Minas, With the supportof the InternationalBoard Tres Marias, Diamantina,Sete Lagoas (Mato for Plant Genetic Resources (IBPGR); FAO,

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Rome; and the InternationalCrops Research trips were conducted with the added Institute for the Semi-Arid Tropics participationof C.E. Simpson (Texas A&M (ICRISAT), Patancheru,A.P., India, another University,Stephenville, Texas, U.S.A.), J. F. 17 expeditionswere undertakenbetween 1976 M. Vails, A. C. Allem and L. Coradin of the and 1983 to the centersof originand diversity Centro Nacional de Recursos Genéticos of Arachis in South America, surveyingthe (CENARGEN-EMBRAPA), Brasilia, Brazil, entire area of distribution of the genus which contributedits infrastructureto ensure (Simpson 1984b, Vails & al. 1985). These the success of these expeditions.

Summaryof Germ plasm Explorations

Year Date Team* RegionExplored

1932-33 24/Dec-30/Jan O BRASIL,MS. 1936-37 18/JuH7/Jan Ar ARGENTINA,BRASIL, PARAGUAY, URUGUAY. 1939 14/Apr-28/MayO BRASIL,MS (Otero1941, 1952). 1947-48 14/Nov-ll/Apr StHt ARGENTINA,BRASIL, PARAGUAY, URUGUAY (Hartley 1949). 1947 BaRi ARGENTINA,Corrientes. 1950 20-26/May BaRiK ARGENTINA,Salta, Jujuy. 1953 1-13/Mar RiKP ARGENTINA,Corrientes, Misiones, E.Ríos. 1953 27-3l/May K ARGENTINA,Salta, Campo Durán. 1958 3/Jan-28/Feb K BOLIVIA,Cochabamba, S.Cruz, Roboré, Parapetí. 1959 16/Jan-8/Apr GKP ARGENTINA,BOLIVIA, BRASIL, PARAGUAY.(Pietrarelli 1961). 1961 20/Mar-20/MayGKP BRASIL,MG GO, BA, MS, PARAGUAY. 1967 28/Mar-21/AprGK BRASIL,BA, CE, PE. 1968 16/May-14/JunHLPK ARGENTINA,BRASIL, URUGUAY. (Pietrarelli 1968, Krapoviekas1969a). 1971 l/May-9/Jim KMoF BOLIVIA, Bermejo,Tarija, Villa Montes. 1976 29/Nov-31/Dec GKAOk BRASIL,MS, MT. (Gregory1977, Gregory & Simpson1978). 1977 15/Mar-19/MayGKBSPSc ARGENTINA,Salta, Jujuy. BOLIVIA,Yacuiba, Camiri, Sta.Cruz BPZ BOLIVIA,S.Cruz, Cochabamba, La Paz. GKSSc BOLIVIA,S.Cruz 1977 9/Jun-7/Jul GKPSc PARAGUAYFlorida, Tobati, Bella Vista. BRASIL,PPorá, Pto.Murtinho, Corumbá. (Gregory 1977). 1978 31/May-13/JunS ARGENTINA,Mdi, Ctes, BRASIL, IAC. (Gregory& Simpson 1978). 1979 28/Mar-30/AprGKSPScGb BOLIVIA, Sta.Cruz,Trinidad. (Gregoiy 1979).

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Year Date Team* RegionExplored

1980 26/Mar-10/MayKSBScCo NW ARGENINA, BOLIVIA. (Simpson1980). BZC BOLIVIA, Sta.Cruz,La Paz. BZCJk BOLIVIA, La Paz. KSSc BOLIVIA, Sta.Cruz,S.Matías. S.José. 1980 10-24/May SPAi PERU,Lima, Cuzco, Quillabamba. (Simpson 1981). 1981 10-7/Jun WeSv BRASIL,GO, BA, PE. 1981 28/Apr-3/Jun SPZ PERU,Lima, T.Maria, Tarapoto, Iquitos, Ayacucho. 1981 14/Aug-9/Sep VSGr BRASIL,GO, MT. (Simpson1981). 1981 27/Sep-lO/Oct PZi BOLIVIA, La Paz, Beni,Chuquisaca. (Pietrarelli 1982). 1982 9-22/Mar ScVn ARGENTINA,Salta, Jujuy. 1982 2-21/Mar VKRSv BRASIL,GO, BA, PI, PE. 1982 17/Apr-16/MayVSW BRASIL,GO, MQ BA. (Simpson1982). 1983 5-20/May VKSvVe BRASIL,BA, PI. (Vails1983). 1983 18/Apr- 11 /May KSScCr BOLIVIA, Tanja.ARGENTINA, Salta. 1983 10-30/May VSMSvGe BRASIL,MQ RS, PA, SP, RJ.(Simpson 1984a, Vails 1984). 1983 10-24/May BPZ ECUADOR (Banks1984). 1984 10-28/Apr VRGeSv BRASIL,MQ MT. 1984 31/Jul-2/Sep VSGdSaW BRASIL,GO, MT. (Simpson1 986a). 1985 6-17/Mar B PERU,Lima, Casma, Chibóte, Trujillo. (Banks 1985). 1985 14-28/Mar WeSv BRASIL,GO, TO, MA, PI. 1985 15-26/Apr VPoPeJAj BRASIL,MS. 1985 ll/May-10/JunVKSSv BRASIL,MT, RO. (Simpson1986b). 1985 9-29/Oct VPoBi BRASIL,MS, MT. 1986 15/Mar-15/AprVSW BRASIL,MS, MT, GO. (Simpson1987). 1986 22/Oct-4/Nov VPoJSv BRASIL,MT, MS. 1987 16-30/Apr VRSv BRASIL, BA, PE, CE, PI. 1988 Jun VQFdSv BRASIL,MS. 1988 8/Sep-13/Dec Wi BOLIVIA, S.Cruz,Beni, La Paz, Pando.(Williams 1989). 1989 24-25/Jan VK BRASIL,MT. 1989 lO/Jun-5/Oct Wi BOLIVIA, S.Cruz,Beni 1990 26/JuHO/Sep Wi BOLIVIA, Beni (Williams1991). 1990 Jun VGaRoSv BRASIL,MT. 1991 May VFaPzSv BRASIL,MG 1992 May VPzVaW BRASIL, MG 1992 May VSPmWiSv BRASIL,SP. VSPmPzRs BRASIL,GO. 1992 Wi La Sud 11-29/Sep BOLIVIA, Paz, Yungas. * For key to participants'names see page 194.

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The Fruitof Arachis and its Capabilities factthat all of its species are geocarpie,that is, theyonly produce undergroundfruit. The first authorto correctlyexplain fruitdevelopment The genus Arachis is characterizedby the in A. hypogaea was Poiteau (1802, 1806). He

Fig. 1. Fruitarticles: 1, A. guaranitica(V.7704). 2, A. tuberosa(K.34497). 4, A. brevipetiolata(G.10138). 5, A. Oteroi(G. 10541). 6, A. Hatschbachii(G.9863). 7, A. cryptopotamica(K.30026). 8, A. major(K.30022). 9, A. Benthamii(G.9761). 10, A. douradiana(G.10554). 11, A. gracilis(G.9772). 12, A. Hermannii(G.9841). 13, A. Archeri(G.9835). 14, A. stenophylla(K.30013). 15a, A. paraguariensisssp. paraguariensis (K.30013). 15b, A. paraguariensisssp. capibarensis (K.30134). 17, A. Macedoi(G. 10127). 18, A. marginata(V.6652). 19, A. prostrata (G.10240). 20, A. lutescens(V.7741). 21, A. refusa (V.9950). 22, A. Burchellii (V.6556). 23, A. Pietrarellii (V.9000). 24,A. villosulicarpa(V.8818, without the villous epicarp). Scale = 1 cm.

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Fig.2. Fruitarticles: 25, A. inseminata(V.6772). 26, A. Giacomettii(W.201). 27, A. sylvestris(V.6767). 28, A. pusilla(V.6110). 29, A. Dardani(V. 10963). 30, A. repens(Conagin 1). 31, A. Pintoi(V.6728). 32, A. lignosa (K.14248). 33, A. Kretschmeri(V.7631). 34, A. Rigonii(G. 10034). 35, A. chiquitana(K.36027). 36, A. matiensis (V.6324).37, A. appressipila(K.30003). 38, A. Vallsii(V.8678). 39, A. subcoriacea(V.8922). 40, A. Burkartii (K.38473).41, A. pseudovillosa (G. 10566). 42a, A. glabrata var. glabrata (K.30135). Scale = 1 cm.

studiedthe plant in 1797 in Santo Domingo and Poiteau himself (1853) and enumerated by described the flower with its calyx tube at Smith(1950), who made an in-depthstudy on whose base is found the sessile ovary which, the flowering and fruitingof A. hypogaea, afterflowering, is borne by a stipe or pedicel confirmingPoiteau's observations. that elongates considerably towards the soil In the flower of A. hypogaea, the ovary is where the fruit will mature. Nevertheless, sessile and has a basal meristemthat, after various other interpretationswere refutedby fertilization, grows and gives rise to a

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Fig.3. Fruitarticles: 43, A. glandulifera(K.30091). 44, A. cruziana(K.36024). 45, A. montícola(K.30062). 46, A. magna(K.30093). 47, A. ipaënsis(K.30076). 48, A. valida(V.9153). 49, A. Williamsii(Wi.1118). 50, A. Batizocoi (K.30083).51, A. duranensis(K.30067). 52, A. Hoehnei(K.30006). 53, A. stenosperma(V.7382-Atlantic coast). 53a,A. stenosperma(V.7762-Mato Grosso). 54, A. praecox(V.6416). 55, A. palustris(V.6536). 56, A. benensis (Wi.860).57, A. trinitensis(Wi.1117). 58, A. decora(V.9955). 59, A. Herzogii(K.36029). 60, A. microsperma (V.7681).61 , A. villosa(HLP.8). 62, A. helodes(G.9926). 63, A. correntina(K.7830). 64, A. Simpsonii(K.36009). 65, A. Cardenasii(K.36016). 66, A. Kempff-Mercadoi(K.30088). 67, A. Diogoi(G. 10602). 68, A. Kuhlmannii (V.8935).Scale = 1 cm. postfloralaxis or peduncle that carries at its one. Smith (1950: 806) gives morphological apex the ovary with the fertilized ovules. reasons for discarding the term gynophore Once the ovary is buried, it develops, that is used to describe the peduncle, producing a unilocular legume that encloses commonly known in English as the "peg" from2 to 5 seeds or, throughabortion, only (paxillus in Latin) and in Spanish "clavo," a

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) termused by farmers. zontal position or downward,as did the peg, In the wild species, the fruitis articulatedor according to the sections of the genus. This lomentiform,with two articles, as in the isthmushas a similar appearance to the peg, majority of species, or with three as in A. withthe differencethat it carriesonly a single triseminata (Krapovickas & Rigoni 1957: ovule at its apex in the two-seeded fruits,or 436, fig. 1; Gregory & al. 1973: 68). The two in A. triseminata. The unequal articlesare separatedby an intercalarypeg or developmentof the ovules withthe formation isthmusof various centimetersin length.The of an intercalaiytissue of a structuresimilar to articles of the fruitmature successively. The the peg, was observed by Conagin (1959: 58, proximal article develops firstand, when it fig. 3). acquires a certain size, an intercalary In A. hypogaea the isthmusis missing but meristemthat will formthe isthmusdevelops the seeds grow differentially,as in the wild in the apex. The isthmuswill grow in a hori- species, althoughmuch closer in time to one

Fig.4. A-D,Development ofbuds on the tips of the peg in A. paraguariensis ssp. paraguariensis : a, destroyedovaiy; b,hypertrophy oftissue next to the ovaiy; c, adventitious floriferous buds; d, pegs of the new flowers; e, subterranean flower;f, leaf of the adventitious bud; g, soil level (KC 11488, cult. Alva, 1993).

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another,giving the impressionof simultaneous so similar to one another that they are maturation.Smith (1956: 236) demonstrated practically indistinguishablewithout all of thatin both "Virginia"and "Spanish" peanuts, the plant parts, many times missing on theembryo and theendosperm of thedistal seed herbarium specimens. Some cases are so do not grow as quicklyas thoseof theproximal strikingthat they are worthpointing out. seed of thesame fruit. Arachis pusilla vs. A. duranensis. These The peg not onlyhas the role of interringthe two species are very much alike, both in the fertilizedovules and of distancingthe seeds from wild and as herbariumspecimens. The first themother plant, but it also has some verypecu- lives along the length of the São Francisco liar abilities.In thesection Extranervosae, many River in the Brazilian state of Bahia and the species frequentlyproduce adventitious roots on second grows in northwesternArgentina and the pegs, as was observedin A. villosulicarpa southernBolivia. The differencesare found (Gregory 1946: 42, fig. 26). These roots can primarilyin the flowers: dimorphicwith the formtubers similar to thoseof the motherplant standardhaving reddishlines on both faces in and can also form shoots. The formationof A. pusilla, and normal flowers with reddish shootson rootswas observedonly in thesections lines only on the ventralface of the standard Extranervosae and Erectoides, and the in A. duranensis. formationof shoots on pegs only in the section Arachis montícolavs. A. magna. These two Extranervosae(Gregory & al. 1973: 60). species are almost indistinguishable.The li- In the section Erectoides, adventitious ving plants look alike, the distributionof inflorescences were observed on the pegs tomentumis similar in both, and both have (Gregory& al. 1973: 60). fruitswhose articles are the same size and In A. paraguariensis ssp. paraguariensis, with a strongly reticulate pericarps. Both materialfrom specimen KC 1 1488, cultivated species belong to the sectionArachis, but A. in Alva, FL (USA), one of us (W.C.G.) montícolais tetraploidand has the small pair observed that when the ovary is destroyedat of chromosomescharacteristic of the species the tip of the peg, possibly by the action of related to A. hypogaea. Arachis magna, on nematodes, insects or fungi, or fertilization the otherhand, is a diploid that is lacking the failure,the tissue becomes hypertrophiednext small pair of chromosomes. to theovary at thepeg tip (fig.4, A, b). On this Arachis helodes vs. A. lignosa. Both swollen apex appear adventitiousfloral spikes species have the same habit,with procumbent (fig. 4, B, c), on which subterraneanflowers branches,leaves of the same shape and size, are formedthat manage to fertilizethemselves almost completelyglabrous. The firstlives in and produce new pegs (fig. 4, C). In the event the vicinityof Cuiabá, in Mato Grosso, and that the adventitious inflorescence emerges the second along the Paraguay River northof fromthe soil, it may produceleaves at itsapex. Concepción, in northernParaguay. The most The production of flowers on pegs with important morphological difference is the damaged ovaries was also observed in species growth of the peg: nearly vertical in A. of the sections Extranervosae and Arachis helodes, and horizontal,very long and super- (Simpson, pers. com.). ficial in A. lignosa. Still, they are very differentcytologically. Although both have 20 chromosomes,A. helodes has the large "Sosias5" or Twin Species satelliteand the small pair of chromosomes, while A. lignosa has a dot-likesatellite and is lackingthe small pair. One of the greatestdifficulties presented by the genus Arachis is the existence of species IntraspecificVariability 5 "Sosia"is a termused by Monteiro Filho (1949: 509); fromthe dramatist Pļautus, Sosia resembledanother characterwith whom he couldbe easilyconfused. Another problem presented by the

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 is its similarityof bands. An interestingcase is that intraspecificvariability. There is some proof of samples 30065, 30067 and 30068, collected of its existence. between Senda Hachada and General In A. inseminata, a veryuniform and easily Ballivián, in the northeastern part of the distinguishable species, upon crossing two province of Salta (Argentina) along a 20 km accessions fromlocations 20 km distantfrom stretchof the Seco River, in which, of the 22 one another,in the neighborhoodof Juazeiro, bands involved only 8 (36%) coincided. The in Bahia (parents 97 x 98), hybrids were samples 30065 and 30067 were collected only obtained with 50.7 and 68.2% stained pollen. 3 km apart and in these, of the 18 bands These values indicate the presence of some involved, 13 (72%) coincided. On the other geneticbarrier unrelated to exomorphological hand, the accessions thatdemonstrate greatest characters. similarityin the bands are 7988 (Campo Du- Upon crossing two accessions of A. ran,500 m) and 36036 (cityof Salta, 1250 m), glabrata var. glabrata separated from one localities separatedby some 400 km. The two anotherby 20 km between Encarnación and localities belong to differentwatersheds: the Trinidadin southeasternParaguay (parents 10 ItiyuroRiver that passes by Campo Duran and x 84), a hybrid was obtained with a high pertains to the watershed of the Bermejo degree of sterility,with 12.9% pollen stained. River; and the Arias River thatpasses through In seven other crosses, involvingaccessions the cityof Salta and pertainsto the watershed much more distant from each other, the of the Salado or JuramentoRiver. hybridsproduced pollen that stained between The karyotypesof nine accessions of A. 45.2 and 92.9%. We are dealing witha taxon duranensis also show variabilitythat is not that has plenty of variability, both associated witheither geographic distribution exomorphologicaland genetic. or withelectrophoretic bands of seed proteins In accessions of A. valida coming fromthe (Fernández & Krapovickas 1994). same locality,collected at the same place but on differentdates, it was foundthat one (KG 30011) had a "B" chromosome with a Reproductive Strategies and Speciation satellite,while the other(KGPSc 30147) had two morphologically different "B" chromosomes.These two accessions grownin Arachis is a genus in which all of its species Manfredi are exomorphologically identical produce only subterraneanfruit. One could (Fernández & Krapovickas 1994). say thatthese species are "pegged" to the soil, In A. montícola, morphologicallydistinct and beneath the soil they have produced a "B" (SAT) chromosomes were observed in great differentiation.There is variability in two accessions collected in Yala (BaKRi the roots, and thereare rhizomes,stolons and 7264) and in Lozano (KG 30063), localities in branchesproduced fromadventitious buds on the province of Jujuy (Argentina) separated the roots. There is variation in the position by only 5 or 6 km (Fernández & Krapovickas and growthof the pegs. On the pegs, which 1994). are a part of the fruit, there can form Arachis duranensis is a taxon witha fairly adventitiousroots, shoots and also floralbuds. uniformappearance thatshows a greatdeal of Due to these characteristics,the species of variability unrelated to the geographic Arachis can disperse their seeds no farther distributionof the accessions. In a recent than 1 m, and often less. Occasionally the work(Bianchi-Hall & al. 1993), 14 accessions seeds are transportedby water during floods of thisspecies wereanalyzed electrophoretically and can be carriedby animals, and possibly by and these showed great variability,to such a humans, to become "pegged" again in a new degree that these accessions are distributed place. Self pollination is the norm, although randomly in the illustration presented the flowersare visited sporadicallyby insects (Bianchi-Hall & al. 1993: 9, fig. 2) and there of limited radius of action, and there is are no two accessions that show a total evidence of parthenogenesis. Under these

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 conditions the rate of speciation depends isolated species such as A. Rigonii, A. directlyupon the rate of mutationand upon duranensis and A. paraguariensis . It is the accumulationof old mutations.Gene flow possible that, in these cases, the selection is very limitedand restrictedto withinsmall pressure was not sufficientto raise genetic populations. We rarely observed extensive barrierswhose presence would not have been populations; what is usually seen is the necessary as an isolation mechanismin these formationof small populations due to their allopatìe species. adaptation to special, primarilysandy, soil The percentage of fertilityin the hybrids types. In this manner,the species of Arachis allows evaluation of the genetic barriersthat behave as apomictic species despite their can be very important in speciation in sexuality,due to a gene flow thatis restricted sympatricpopulations. Nevertheless, there are within small populations and almost non- some evidentexamples of low percentagesof existentbetween populations separated from pollen staining in hybrids obtained with one another. parents that we suppose belong to the same It can be expected that random changes species due to theirvery similar appearance. occur within the same species at different The barrier is not always associated with isolated localities and that mutations are exomorphologicaldifferentiation, it could be accumulated with visible effects,as well as genetic as in A. duranensis, or geographic as intra-chromosomal transpositions of in A stenosperma. importantDNA sequences that make meiotic The chromosomesare good indicators.For pairing difficult. example, the ploidy level is indispensablefor A good measure of thissituation is givenby separating the two series of the section the resultsof crossing experiments(Gregory, Rhizomatosae or for definingA. montícola. M.P. & Gregory 1979) that show low The morphology of the "B" (SAT) percentagesof pollen stainingin some crosses chromosome gives very good information. between accessions of the same species from For example, the presence of a dot-shaped differentlocalities. satelliteis highlyimportant for delimitingthe It is very difficultto establishthe extentto section Procumbentes, but dot-shaped whichthe lack of fertilityof the hybridsis due satellitesare also foundin A sylvestrisand A. to a specific or intraspecificlevel of parental benensis, species that, based on differences.Nevertheless, it is highlyvaluable exomorphology, have other affinities. The informationfor determiningthe degree of presence of the "A" chromosome, or small genetic isolation. Above 50% pollen pair, is an unsurpassable indicatorof genetic stainability,the dehiscence of the anthersis affinitywith A hypogaea, but it does not seem normal. Below 50% the anthersbegin to open to be associated with any exomorphological withdifficulty. Around 25% the anthersare no characterso as to be useful taxonomically. longer dehiscent but their pollen may be All of this informationhas permitted,in capturedby bees. Below 15% the anthershave manycases, the appraisal of exomorphological to be dissectedto extractthe pollen for staining. charactersthat would have otherwise passed Below 10% it is questionable if the fewgrains unnoticed. thatdo stain could functionin pollination. There are cases in which the combination To define or delimitthe species one must of the available informationwas not sufficient reconcile or harmonize the exomorphology and it was preferredto adopt a conservative withall the otherinformation, such as genetic, position,thereby leaving some taxa withpro- chromosomalor geographic,because none by visional solutions. Such is the case with A. itselfwould sufficein absolute terms. sylvestrisand A. benensis, whose placement The ability to produce experimental in theirrespective sections is not definitive. hybrids is of great importancefor detecting Some species have a broader distribution affinitiesand was used in large measure to when relatedmaterial is included in the lists. establish or confirmthe sections. A greater In these cases we have divided the studied ability to cross was observed in peripheral, material.

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Dispersion ourselves by the small excavations left by these animals in our search for wild species, as in the case of A ipaënsis. We considerthat the genus Arachis originated The action of humans must have been in the Sierra de Amambay, on the border important.The domesticationprocesses of A. betweenMato Grosso do Sul (Brazil) and Pa- hypogaeaand A. villosulicarpaalready implies a raguay,where A. guaranitica grows, possibly great understandingand managementof wild the oldest species of the genus (Gregory& al . peanuts by Amerindians. In northeastern 1980). It is difficultto understandhow this Brazil, wild peanuts are called by names like genus could have extended its range some "amendoim" of Portuguese origin, and 4000 km, both towards the NE, up to the "mendubi" or "mundubí" of Guarani origin, Amazon, as well as to theW, up to theAndes. while, on the otherhand, cultivatedpeanut is By its own means, no species of Arachis known only by the name "amendoim," which can propagate itself more that one meter's suggestsan earlierknowledge of the wild ma- distance per year due to its peculiar type of terial. Childrenare in the habit of seekingand fruiting.To cover 4000 km, a theoretical eating the fruitsof wild peanuts such that, minimum of 4,000,000 years would be thanks to their help, we made our first required.And so, theremust have been other collections of A. montícola. means of dispersal that explain the abilityto There is a fact that proves the action of occupy an area as extensiveas the one thatthe humans in the transportof wild peanut fruits. genus inhabitsin South America. On the Pacific coast of Peru, in Bermejo (Dep. Fluvial dispersionmust be very important, Ancash, 10°33'S, 77°53IW), in a prehistoric such that many of the species have a excavation, archeologistsdiscovered remains distributionassociated with the watershedof of one-seededpeanut articles (12 mm long x 7 the great Paraguay, Uruguay and Paraná or mm wide), with a slightlyreticulate pericarp. São Francisco Rivers. The species generally Evidently we are dealing with some wild live near watercourses, in places where the peanut that so far has not been collected in water evidentlyreaches only duringthe higher Peru. They are quite different from the floods. In A. hypogaea the maturefruits can archeological peanuts frequentlyencountered float,such thatin India a harvestingtechnique on the coast of that countrywhich by their was developed in which the peanut field is notablepod reticulationcan be identifiedas A. flooded, the soil is disturbed and the fruits hypogaea ssp. hypogaea var. hirsuta. The float to the surface where the wind carries closest wild peanuts live in the Beni regionof them to a corner where they are collected Bolivia, on the otherside of the Andes. Of the (Rusby 1901). It is conceivable that in this threespecies thatwe know fromthat area, the same manner fruitsof wild species could be most similar is A. Williamsii,from Trinidad, transportedby water duringfloods. at 256 m elevation and 1500 km distantfrom Dispersalby animalsshould not be discarded. Bermejo as the crow flies. The transportation Birds, hogs, rodentsand armadillos seek out by humansis evidentin thiscase, to have been the wild fruits. Although upon eating the able to cross thebarrier of theAndean Cordille- seeds theydestroy the embryo,the possibility ra, whosepasses are above 4500 m in elevation. exists of whole fruitsbeing carried to their Transportationby human action is also a burrows.Names like "amendoimde carcará," plausible explanation for interpretingthe "mundubíde carcará" (caracará peanut),or its disjunctrange of A. stenosperma. equivalent "hire mať ni" of the Carajás Indians of the Ilha do Bananal, reveal the predilectionof the "carcará" bird ( Anhinga The Sections of Arachis anhinga) for the fruitsof wild Arachis. The name "mendoimde porco" (hog peanut) also shows the appetite of this animal for wild In Diagram I we presentthe relationships peanut fruits.In some cases we have guided betweenthe sections of the genus Arachis, trying

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 to combine previous schemes (Krapovickas higherelevations in the Andes, of the section 1973; and Gregory,M.P. & Gregory1979). The Arachis. sectionsare orderedvertically according to their The sections Trierectoides,Erectoides and affinitywith Stylosanthes and accordingto their Triseminataehave species of erector decumbent ploidy level. Above the upper line are the habit. In the restof the genus the species have tetraploids(2n=40) and below it are thediploids prostratebranches. The exceptionwould be A. (2n=20). The sectionsare orderedaccording to appressipila,the only species witha decumbent the number of characters in common with habitin thesection Procumbentes. Stylosanthes; on the lower part of the diagram The sectionsExtranervosae and Heteranthae are foundthe sectionswith greateraffinity to have flowerswhose standardhas redlines on the that genus. On the horizontalaxis we try to interiorface, as is frequentin Stylosanthes. representthe geographic distribution, placing to It is possible that these five sections, as the leftthose sectionsthat reach the foothillsof theyare located near the bottomof Diagram I, the Andes, and to the rightthe more eastern are the most primitive of the genus. This sections(figs. 5-9). This geographicscheme is suppositionis supportedby the highdegree of valid for almost all the sections,except forthe genetic isolation that is found among them section Arachis, whose species cover a large and in the comparativemorphology of the "B" part of the range of the genus, extendingfrom (SAT) chromosome and the absence of the the Atlanticcoast (A. stenosperma)over to the "A" pair (Fernández & Krapovickas 1994). Andes (A. duranensisand A montícola). In the seven sectionsthat are foundtowards The lines that unite the sections represent the bottomof Diagram I, the peg of the fruitis degrees of genetic affinity,according to the verylong and superficial,in contrastwith the resultsof crossingexperiments carried out by peg in the sectionsRhizomatosae and Arachis W.C. Gregory (1967) and M.P. Gregory & that is shorter and growing in a vertical Gregory(1979) (see page 195). The drawing positionor close to it. of the lines was done while tryingto avoid, as Towards the top of the diagram are found much as possible, having themcut across one the sections Caulorrhizae and Rhizomatosae another. withnew methodsof vegetativemultiplication: Diagram I shows the greatest number of stolons in the formerand rhizomes in the lines towards the left,while downwards and latter. towards the rightthere is a markedreduction It is possible that the annual character in them. No successful intersectionalcross representsan adaptive advantage thatpermits could be obtained with section Triseminatae; avoidingadverse seasons such as the droughts withA. Burkartii(R,), the only diploid in the in northeast Brazil ( Heteranthae) and the section Rhizomatosae',nor with the perennial droughtsin the foothillsof the Andes, as well species (A,) and tetraploids(A3) of the section as the flooding of the Paraguay River Arachis. watershed( Arachis Aj). Diagrams II, III and IV illustratethree of It is noteworthythat fruitswith reticulate the species with best success in the crossing pericarpsonly appear in two perennialspecies experiments,representing different sections. (A,), various annual species (A,) and in all of With A. these, paraguariensis ssp. the polyploid species (A3) of the section paraguariensis, A. Rigonii and A. duranensis, Arachis (fig. 3). almost all of the intersectionalaffinities that appear in Diagram I can be established. It is strikingthat all threeof these species are from Arachis L. marginal areas. Arachis paraguariensis ssp. paraguariensis is the southernmosttaxon in the section Erectoides, A. Rigonii is the Linnaeus,C., Sp. pl. 741, 1753. Linnaeus,C., westernmostspecies in the section Procumbentes, Gen.pl., ed. 5, 329, 1754. and A. duranensis is one of the most western Arachidna[Plum.] Boehmer in C. G. Ludwig, species and one of the ones that grow at Def. gen.pl. ed. 3, 255, 1760 nom.illeg. Moench,

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DiagramL Relationshipsbetween the sections of the genus Arachis.

DiagramDL Crosses of A paraguariensisssp. paraguariensis with species of other sections. The numbers indicatethe parent materials used in the crosses.

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Diagramm. Crossesof A Rigoniiwith species of other sections. The numbers indicate the parent materials usedin the crosses.

DiagramIV. Crossesof A duranensiswith species of other sections. The numbers indicate the parent materialsused in the crosses.

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Fig.5. Rangeof sections Trierectoides, Extranervosae and Triseminatae.

Meth.pl., 121, 1794. branches thickened or not. Tetrafoliolate Mandubí [Marcgr.]Adans., Fam. pl. 2: 323, leaves, sometimestrifoliolate, generally those 579, 1763.nom. illeg. of the mainstem (n) somewhat larger and differentlyformed than those of the secondary Annual, biennial or perennialplants; erect, (n+1) and tertiary(n+2) branches. Stipules decumbent or procumbent; sometimes partially fused to the petiole, sheathing, rhizomatous or stoloniferous.Taproot, with forming a vagination, with free portions

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Fig.6. Rangeof sections Erectoides and Heteranthae . sharply pointed. Petiole and rachis long or shortbristle; outstanding venation on canaliculate, channel generally interruptedat the upper leaf surface,especially on the mar- the firstpair of leafletsby a transverseline of ginal vein. Pauciflorous axillary spikes, hairs. Leaflets from suborbicular to arrangedalong the lengthof the branches or lanceolate. Indumentumconstituted by long grouped around the collar of the plant. silky hairs, short adpressed hairs, cilia and Flowers sessile, hypanthiumwell developed. emergences with conical bases ending in a Bilabiate calyx, upper lobe wider and 4-

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) toothed,lower lobe falcate. Corolla orange or withthree uniseminate articles or witha single yellow. Standard with red lines on the upper, 1- to 5 -seminatearticle; pericarp smooth or lower, or both faces. Subterraneanfruit; peg reticulate,covered with a layer of finehairs of short and vertical or horizontal up to more variable density. Seed smooth, seed coat than 1 m long, sometimes with adventitious whitishor yellow-brownin the wild species or roots that may tuberize; two uniseminate various colors in the cultivatedpeanut. articles a filiform separatedby isthmus,rarely Type species: Arachis hypogaea L.

Key for Identifyingthe Sections

A. Trifoliolateleaves. Tuberiform hypocotyl. Erect plants. Flowers and fruits grouped around the collar of theplant. Pegs horizontal,superficial, very long. I. Trierectoidespage 34 A1.Tetrafoliolate leaves. Cylindrical hypocotyl. B. Plantswithout rhizomes. C. Fruitswith 2-3 articles. Branches decumbent. Flowers and fruits along length of branches. Standard withreddish lines on bothsurfaces. Cotyledons with veins on uppersurface very sunken. IV. Triseminataepage 83 C. Fruitswith 2 articles.Cotyledons with smooth upper surface. D. Standardwith red lines on lowersurface or on bothsurfaces. Procumbent branches. E. Perennialplants, roots with thickenings. Standard with red lines only on the lower surface. All flowersnormal, with expanded corolla. III. Extranervosaepage 67 E' Annualplants, roots not thickened. Standard with red lines on thelower surface or on both surfaces.Flowers dimorphic, normal, open or verysmall withthe corollanot extending beyondthe calyx. V. Heteranthaepage 85 D' Standardwith red lines on upperface. F. Plantserect or decumbent.Flowers densely grouped around the collar of the plant, those thatnormally produce fruit. Towards the base ofthe branches, only those flowers which are buriedproduce fruit. Roots with swollen branches (except in A. stenophyllaand A. paraguariensis). II. Erectoidespage 40 P. Branchesprocumbent. Collar of the plant without flowers; inflorescences and fruits along lengthof branches. In A appressipila(sect. Procumbentes) the branches are decumbent, butthe flowers are notgrouped around the collar of the plant. G. Stemswith roots at thenodes. VI. Caulorrhizaepage 94 G' Stemswithout roots at thenodes, sometimes at theunderground basal internodes. H. Pegshorizontal, very long and superficial. VII. Procumbentespage 99 H'. Pegsalmost vertical.

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ĪX.Arachispage 123 B'. Plantsrhizomatous. VIE. Rhizomatosaepage 113 I. Leafletscoriaceous, with the margin outstanding on bothsurfaces. Standard orange with red lines on bothsurfaces. Diploid. SeriesProrhizomatosae page 114 r. Leafletsmore or less soft, with the margin slightly marked. Standard orange or yellow, with red lines onlyon thefront. Tetraploid. SeriesRhizomatosae page 117

I. Sect. TrierectoidesKrapov. & W.C. this section live in the highest places of the Gregory nov. sect divide between the watersheds of the Para- and Paraná between400 and 700 Fig. 5 guay rivers, m above sea level. The northernlimit is found in in 700 m Sect. TrierectoidesKrapovickas, Agricultural Jataí, Goiás, at some above sea level near the divide between the Genetics. Selected Topics: 137, 1973, nomen Araguaia nudum. and Paranaiba rivers. Sect. Erectoidesser. TrifoliolataeKrapov. & Obs. The two W.C. Gregory,in W.C Gregory,M.P. Gregory, species of this group have thatare in Krapovickas,Smith & Yarbrough,in Peanuts- various characters unique the genus Cultureand Uses: 93, 1973,nomen nudum. Arachis, such as trifoliolate leaves, the tuberiformhypocotyl, and the well-developed Perennes. Hypocotyluscrassus, fusiformis. basal tube of the stipules.Because of the erect habit and the thickened root this Radieis ramificationesincrassatae. Stipulae branches, is related the section marginibusbasi connatistubum internodia 1-2 section closely to includentemefficientes. Folia trifoliolata.Vexillum Erectoides, in which are found some species aurantiacum,supra lineis rubescentibus ornatum. with basal tubes on the stipules, but much Fructus subterraneus biarticulatus paxillo reducedand not always constant. Trierectoides horizontālipaulo profundo,pericarpio laevi. The section is easily differentiatedfrom all of the othersections; it Typussectionis : Arachis guaranitica Chodat & is genetically very isolated and it is quite Hassl. probablythe most primitiveof the genus. For all of these reasons, it is much more convenient to treat this as an Perennial plants. Hypocotyl thickened, group fusiform.Root withthickened branches. Erect independentsection. stems. Stipules with the margins fused, forming a tube that encloses one or two internodes. Trifoliolate leaves. Flowers Key fordistinguishing the species clusteredat the base of the plant. Hypanthium well-developed. Standard orange withreddish A. Leafletslinear-lanceolate, rigid, up to 11.5 cm lines on the front. Fruit subterranean, longX 0.6 cmwide. biarticulate;peg veryextended, up to 1 m long, 1. A. guaranitica not isthmus horizontal, deep; well-developed; A'. Leafletsobtuse, elliptical-lanceolate, ca. 2.5 cm articles smooth.2n=20 single-seeded;pericarp long X 1 cm wide,obovate and smallertoward chromosomes. thebase ofthe branches. 2. A. tuberosa Geographic distribution.The two species of

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Fig.7. Rangeof sections Procumbentes and Caulorrhizae.

1. Arachis guaranitica Chodat & Hassl. 13(146-147):764, 1933. Hoehne, Flora Brasílica 25(2) 122: táb. 1940. Figs. 1,1; 10,F part. 10, 1, fig.2, Hermann,Agrie. Monogr.USDA 19: 7, fig.2, 1954.

Chodat& Hassler,Pl. Hassl. 2: 449-450, 1904. Erect perennial, 30-50 cm tall. Hypocotyl Chevalier, Rev. Int. Bot. Appl. Agrie. Trop. enlargedto forma tubersome 3 cm long x 1.5

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Fig.8. Rangeof section Rhizomatosae : RI, 2n=20ser. Prorhizimatosae , and R2, 2n=40 ser. Rhizomatosae. cm wide, with some adventitiousroots toward coriaceous. Fused portionof the stipuleswith the base; the main root deep, without fused marginsforming a long tube or sheath thickened branch roots. The collar bears which encloses up to two internodes;sheath several unbranched stems. Internodes long, closed along almost its entire length,30-60 completelyenveloped by the stipular sheath. mm long but open at thetop for5- 10 mm; free Leaves trifoliolate, distichous, glabrous, part of the stipules abruptlytriangular, up to

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Fig.9. Rangeof the wild species of section Arachis.

12 mm long x 2 mm wide; sheath and free sharp-pointed; upper surface smooth with portion with numerous, well-marked prominentmargin; lower surface with very longitudinalveins. Petiole short,10 mm long, prominent mid- and marginal veins and canaliculate. Leaflets, more or less equal in markedsecondary veins. Flowers clusteredat size withina leaf, linear-lanceolate,rigid, up the base of the plant, surroundedby filiform to 11.5 cm long x 0.6 cm wide, apex acute, bristly bracts. Hypanthium 4-8 cm long,

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Fig.10. Arachis tuberosa : A, schematic ofthe plant; B, leaf,upper surface; C, basalleaf, upper surface; D, leaf,lower surface;E, basalleaf, lower surface (Otero 192). A. guaranitica : F, leaf,lower surface (G.9701). villous. Calyx 5-6 mm long, bilabiate, with hairs which retain a thin coat of soil. 2n=20 silky hairs and bristles ca. 1 mm long. Stan- chromosomes (Fernández & Krapovickas dard orange, 10-18 mm long x 10-18 mm 1994). wide. Fruit Particulate; peg horizontal, up to 50 cm long; isthmus well developed; Holotype: PARAGUAY. "In campo Ipe hu, articles 15-19 mm long x 7 mm wide, Sierra de Maracayu,oct [1898] Hassler 4975''' pericarp smooth, covered with very short (G!) (photoF. 27924!). Isotypes:NY!, UC!.

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Additionalmaterial: BRAZIL. Mato Grosso do Sul. 1841,"Ad Rio Pardo.Brasiliae (v.s . comm.a Mus. RioAmabaí, entre Ponta PorãyAmambaí, 17-11-1959, Acad. Petrop. cum.fl.)." Bentham, Fl. bras.15(1): 88, " Gregory& al. 9665(GH, LIL, MO, NY, SI,SP, US); 14- 1859, Habitatin campisCamapuensibus ad Rio V-1961,Gregory & al. 10568(LIL, US); 13-VI-1968, Pardo, Brasiliae meridionalis,Riedel." Hoehne, Hammons&al.574(CTES); 13kmNdorioAmambai,Flora Brasílica 25(2) part.122: 9-10,táb. 1, fig.1, 24-IV-1984,Vails & al.7694 (CEN, CTES); 8,5 km Sdo 1940.Hermann, Agrie. Monogr. USDA 19: 7, fig.1, rioAmambaí, 24-IV-1984, Valls & al. 7700 (CEN, 1954.Otero, 52, 1941. CTES); Dourados,km 28,5 da BR-463,25-IV-1984, Valls & al. 7704 (CEN, CTES); entrePonta Porā e Erect perennialup to 40 cm tall. Hypocotyl Dourados,25-IV-1985, Valls & al. 8728(CEN, CTES); a fusiformtuber, ca. 3 cm long x 1 cm thick, AntonioJoão (Capitan Bado), 18-11-1959,Gregory & with adventitiousroots toward the base, the al. 9680(LIL, NY, US); 5 kmde Amambaí,camino a taproot usually forming additional PontaPorā, 19-11-1959, Gregory &al. 9701(CTES, GH, concatenated enlargements. Stems simple, LIL,MO,NY,SI, SP,US);9kmWdeCaiapó, 54°55W, unbranched,several occurringat the collar of 22°35'S,19-1-1979, Krapovickas & al. 34335(CEN, the plant, internodesvillous, usually covered CIES). by the stipules. Leaves trifoliolate.Stipules PARAGUAY.Amambay. Pedro Juan Caballero, sheathingthe lengthof the internodeabove or 19-X-1986,Pedersen 14681 (CTES, Heib. Pedersen). somewhat shorter,with marked longitudinal veins; the portion fused with the petiole Geographie distribution. This species (sheath) 18-20 mm long, completelyclosed, grows along the Cordillera de Amambay, on forminga tube at thebasal 5 mm; freeportion the border between Paraguay and Mato acute, some 10 mm long x 2 mm wide. Petiole Grosso do Sul (Brazil), fromIpe Jhuto Pedro canaliculate, short, 3 mm long. Leaflets Juan Caballero. The eastern part of its area obtuse, elliptical-lanceolate, some 25 mm extendsto Dourados (MS), located some 120 long x 10 mm wide, in well-developed leaves km fromP.J. Caballero. It prefersopen areas themiddle leaflet is somewhatlarger (up to 40 in "campo cerrado," near rivers, where it mm long x 1 1 mm wide) than the laterals (up grows in grasses and is very difficult to to 35 mm long x 9 mm wide), smaller leaves distinguishfrom the surroundingvegetation with obovate leafletstowards the base of the except forits characteristicflowers at thebase branches. Sheath and free portion of the of the plant. stipules withglabrous surfaces; marginswith long silkyhairs. Petiole glabrous on the dorsal Obs. Arachis guaranitica is the most surface and the marginsof the canal villous. differentiated species of the genus and Leaflets glabrous on both surfaces; upper cannot be confused with any other because surface smooth, shiny green; lower surface its lanceolate leaflets make it easy to with midvein very prominent and marked mistake for a grass. We have crossed it with secondary veins; margin prominenton the A. tuberosa, the closest genetically, and lower surface, villous and with some short, with A. gracilis, A. major and A. rigid bristles. Flowers clusteredat the collar paraguariensis ssp. paraguariensis of of theplant but also presentat the basal nodes section Erectoides, and withA. appressipila, of the branches. Hypanthiumfiliform, pilose, the only decumbent species of section 4-7 cm long. Calyx bilabiate, villous, wide Procumbentes.In all cases the pollen fertility lobe 7 mm long, narrow lobe falcate, 8 mm was low, demonstratingthe genetic isolation of long. Standard 10-16 mm long, orange. Fruit the species. biarticulate, peg horizontal, sinuous, elongated; isthmusca. 8 cm long; articles 10- 2. Arachis tuberosa Bong, ex Benth. 15 mm long x 7 mm wide; pericarp smooth, Figs. 1,2; 10, A-E fragile.

Holotype:BRAZIL. Mato Grosso.Rio Pardo, Bentham,Trans. Linn. Soc. London18(2): 159, X-1826,Riedel 605 (K!). Isotype:P!.

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Selected additional material: BRAZIL. II. Sect. Erectoides Krapov. & W.C. Goiás. Jataí,18-XII-1948, Macedo 1535 (LIL, Gregory nov. sect Macedo 2693 NY); Balsamo, l-XI-1950, (US); Fig. 6 10 km W de Jataí,16-III-1959, Gregory& al. 9942 (GH, LIL, MO, NY, SI, US); id., Valls & al. 63 10 (CEN); Serra do Caiapó, 30 kmN de Sect. TetraerectoidesKrapovickas, Agricultural Jataí,24-X-1964, Irwin & al. 7281 (K, NY, RB, Genetics.Selected Topics: 137, 1973, nomen nudum. US); 10 kmN ofJataí, Eiten & al. 9340-B (UB); Sect.Erectoides ser. TetrafoliolataeKrapov. & 10 km N de Jataí, 14-X-1968, Fonseca & al. W.C. Gregory,in W.C. Gregory,M.P. Gregory, 1551 (UB); Un kmW de Jataí,Valls & al. 6305 Krapovickas,Smith & Yarbrough,in Peanuts- (CEN); km 193,2da BR-364, 10-IV-1984,Valls Cultureand Uses: 93, 1973,nomen nudum. & al. 7545 (CEN, CTES); 10,2kmNW de Jataí, BR-364, ll-IV-1984, Valls & al. 7547 (CEN, Perennes. Radieis ramifìcationes saepe CTES). Mato Grosso do Sul. Campo Grande, incrassatae.Rami erectivel decumbentes.Folia Fazenda Imbirussú,31-1-1933, Otero 192 (RIZ, quadrifoliolata. Vexillumaurantiacum, supra SP); id., Otero 452 (K, SP); id., 25-11-1959, lineisrubescentibus ornatum. Fructus subterraneus Gregory& al. 9818 (GH, LIL, MO, NY, SI, US); biarticulatuspaxillo horizontāli,valde elongato, alrededores de Campo Grande, camino a paulo profundo,pericarpio laevi. Sidrolandia, 10-11-1979,Krapovickas & al. 34497 (CEN, CTES, F, G, MO, SP, UC, US); 54 Typussectionis: Arachis Benthamii Handro kmN de Campo Grande,5-III-1959, Gregory & al. 9837 (GH, LIL, MO, NY, SI, SP, US); Campo Perennial plants. Root frequently has Grande, Imburussú,16-IV-1984, Valls & al. thickened branches. Branches erect or 7607 (CEN, CTES); Campo Grande, bairro decumbent. Leaves tetrafoliolate. Flowers Batistão, 4-IV-1986, Valls & al. 9878 (CEN, clusteredat the base of the plant. Hypanthium CTES). well-developed.Standard orange with reddish lines on the front. Fruit subterranean, Common name, "amendoim de Particulate;peg verylong, horizontal, not deep; tuberas" (Otero 1952: 178). isthmuswell-developed; articles single-seeded; pericarpsmooth. 2n=20 chromosomes. Geographic distribution. Grows in Mato Grosso do Sul (Brazil), in the hills Geographic distribution.Two groups can of Maracaju along the divide between the be established according to the presence of watersheds of the Paraguay and Paraná roots with thickened branches or without rivers, from Campo Grande to the them.The first,constituted by the majorityof headwaters of the Pardo River. It species fromthis section, is characteristicof appears in the area of Jataí in the south the "cerrado" with red soil which surrounds of Goiás. In the "cerrado," it prefers the Mato Grosso Pantanal and is almost shady places under trees. exclusive to Mato Grosso do Sul, except for A. brevipetiolatafrom the extreme south of Obs. With some frequency, the plants Goiás on the Paranaiba River, A. have small basal leaves with four cryptopotamicawhich barely extendsbeyond leaflets. thenorthern boundary of Mato Grosso do Sul, The only crosses obtained with this and A. major which extends as far as the species have been with A. guaranitica departmentof Amambay,in Paraguay. It lives of the same section and with A. gracilis on both sides of the divide between the and A. major of section Erectoides . In watershedsof the Paraguay and Paraná rivers all cases the pollen fertilityof the hybrids and barelypenetrates into the Pantanal. is very low, except in the case of A. The second group, constituted by A. gracilis in which the pollen fertility stenophylla and A. paraguariensis, lives in fluctuates between 11.8 and 18.7%. the SW extremeof the section's range. The

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taxon of widest distribution is A. Obs. The second group, that includes A. paraguariensis ssp. paraguariensis , whose paraguariensis, perhaps deserves to be range extendsfrom the northof the Sierra de considered as an independentsection, given Bodoquena, in Mato Grosso do Sul, to close to the low genetic affinityexhibited in crosses Paraguari, in Paraguay. The other taxa live with the rest of the species of the section withinthis range, along the border between Erectoides. Moreover, it shows some Mato Grosso do Sul and Paraguay, between suggestive differences,such as the lack of Bela Vista and Porto Murtinho.They preferto thickened branches on the roots, and the live outside the "cerrado," in soils of light- flowers,although more frequentat the base of colored sand and, while they may be found the plant,do not formsuch a dense clusteras near watercourses,they preferhigher places, in the otherspecies, and the fruitshave larger away fromperiodic flooding. articles.

Key fordistinguishing the species

A. Rootswith thickened branches. Branches straight, not undulate. B. Leafletssuborbicular, oval or oval-lanceolate, ratio length/width less than 4: 1 (mayreach a maximumof 4.2:1, in A douradiana). C. Distalleaflets obovate to suborbicular and the proximal leaflets elliptical, small, 7-12 mm long x 4-9 mmwide, upper surface with very short hairs, lower surface villous, with bristles. Stipules villous, withoutbristles. 3. A. Martii C' Leafletsgreater than 20 mmlong. D. Petioleshort, 2-4 mmlong. Stipulesand petiolevillous, without bristles. Leaflets with both surfacesvillous. Seed ellipsoid,1 1 mmlong x 3.5 mmwide. A.A. brevipetiolata D' Petiolegreater than 5 mmlong. Seed thicker, 5-7 mm wide. E. Leafletswith upper surface glabrous and lowersurface with adpressed hairs. F. Leafletswith the margin very marked on bothsurfaces (when dry), noticeably citiate. Leafletsgenerally obovate and frequently with bristles on thelower surface. Stipules and petiolewithout bristles. Petiole 8-15 (20) mmlong. 5. A. Oteroi F'. Leafletswith the margin somewhat marked only on thelower surface, commonly elliptical. G. Stipulesand petiole with bristles. Petiole 15-28 mm long. 6. A. Hatschbachii G'. Stipulesand petiole without bristles. Petiole 30-38 mm long. I.A. cryptopotamica E'. Leafletswith short hairs on theentire upper surface, and sometimes only toward the base and on themidvein. H. Largeplants, highly branched, n+3 branchespresent. Leaflets commonly without bristles on thelower surface. 8./1.major

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H1.Smaller plants, little branched, secondary branches erect, very slightly arched at thebase, thebranching ends with short n+2 branches,not longer than 10 cm. J.Leaflets with length/width ratio 1.5-2.5:1 (rarely 3), frequentlywith bristles on thelower surface,ciliate margin, very rarely with an occasionalbristle. Petiole 10-30 mm long. 9. A. Benthamii J'.Leaflets with length/width ratio 2.3-4.2: 1, without bristles. Petiole commonly 6 mm long, rarelyup to9 mm. 10.A. douradiana B'. Leafletselongatedly elliptical, ovate-lanceolate, length/width ratio 3-8: 1 (thereis usuallya lesserratio towardthe base ofthe branches, but always, on thesame plant, the leaflets towards the ends of the branchespresent a ratiogreater than 5:1), generallywith bristles on thelower surface. K. Petioleand stipules with bristles, stipules with the margins fused forming a very short tube. Length/ widthratio of the leaflets 4-8: 1. Petiole 10-25 mm long. H.A. gracilis K'. Petioleand stipuleswithout bristles. L. Tubeat the base of the stipules up to 9 mmlong. Leaves spaced out, internodes up to 50 mmlong, petiole15-35 mm long. Length/width 3-7: 1. 12.A Hermannii V. Tube at thebase of the stipulesup to 4 mmlong. Leaves groupedtowards the apex of the branches,internodes 10-15 mm long. Petiole commonly 10-15 mm long (rarely up to 30 mm). 13.A Archen A'. Taproot,with slender ramifications, without thickenings. Stems undulate or somewhat twisted. Margin ofthe leaflets marked. M. Leafletslinear-lanceolate, less than 10 mmwide, length/width ratio 7-12:1. 14.A. stenophylla M'. Leafletsmore than 10 mm wide, length/width ratio less than 5:1. N. Length/widthratio of the distal pair of leaflets 2.6-3.4: 1 andof the proximal pair 3.3-4.3: 1. Margin and veinsvery marked on thelower surface. 15a.A. paraguariensisssp. paraguariensis N'. Leafletswider, with veins and margin not so outstanding;length/width ratio of the distal pair 2-3:1 andof the proximal pair 2.5-3.5: 1. 15b.A. paraguariensisssp. capibarensis

3. Arachis Martii Handro Perennial plant, sub-erect,20-30 cm tall. Taproot,with enlarged branches. Crown thick, Handro,Arq. Bot. Estado São Paulo 3: 179-180, with remains of inflorescences. Stems tab.46, 1958. decumbent,10-30 cm long, internodeshirsute, A. helodesMart, ex Hoehne,Flora Brasílica 25(2) covered by the stipules. Leaves tetrafoliolate. part.122: 13,táb. 7, 1940,nomen nudum pro parte Stipuleswith the sheath4-5 mmlong, striated, (Otero174), excl.specimina ex Cuiabá. marginscontiguous; the freeportion 8-10 mm A. pusilla auct.nonBenth., Otero, Serv. Inform. long,linear-lanceolate. Petiole canaliculate, 8- Agrie. Min. Agrie.: 37, 1941, photographof 10 mm long. Rachis canaliculate, 2.5-3.5 mm specimenOtero 174. long. Leaflets subsessile, 7-12 mm long x 4-9

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mm wide; the two apical leaflets obovate to 4. Arachis brevipetiolataKrapov. & W.C. suborbicular,the two basal elliptical. Stipules Gregory nov. sp. those of the villous, hairs caducous, margins Figs. 1,4; 11,D-E more persistent.Petiole, rachis and pulvinus densely villous. Upper surface of the leaflets with short, sparse, more or less erect hairs; Herbaperennis, erecta. Radieis ramißcationes the lower surface villous, some bristles may tenerrimae,partes incrassatas5-10 mm longas, be present mainly on the secondary veins, subglobosasvel cylindricasconcatenatas geren- midvein hirsute; margin ciliate. Spikes few- tes. Caulis angulosus, villosus. Stipulae flowered, short, clustered principally at the marginibusbasi connatistubum usque ad 9 mm base of the plant but also occurringalong the longumx 1.5-2 mm latumefficientes, villosae. branches. Hypanthiumfiliform, villous, up to Petiolus 2-4 mm longus. Folíola elliptica vel 5 cm long. Calyx villous, 5-6 mm long. Stan- obovatiora epiphyllo villoso usque lanato, dard ca. 12 mm long x 14 mm wide, orange hypophyllo dense adpresso-villoso. Fructus with a yellow base. Fruits biarticulate, biarticulatuspaxillo horizontālica. 20 cm longo, clustered at the collar of the plant and along articulis13 mmlongis x 6 mmlatis, pericarpio those stems in contact with the soil; peg laevi. horizontal, some 25 cm long, isthmus 2 cm long; articles 8-10 mm long x 4-5 mm wide, Holotype:BRAZIL. Goiás. Itumbiara,borde pericarp smooth. del camino a Goiania, 5-IV-1961, Gregory, Krapovickas& Pietrarelli10138 (LIL). Isotypes: Holotype:BRAZIL. Mato Grossodo Sul. Cam- CTES, GH, MO, NY, US. po Grande,Fazenda Imbirussú,30-1-1933, Otero 174 (SP!). Isotype:RIZ!. Perennial plant, erect, without rhizomes, villous, reddish-brown.Taproot, with slender Additionalmaterial: BRAZIL. Mato Grosso branches and with subsphericalor cylindrical do Sul. Campo Grande,Imbirussú, 8-VI-1968, concatenate enlargements 5-10 mm long. Hammons& al. 525 (CTES) and 526 (CTES, Stems erect,up to 40 cm long, wavy toward SI). the apex; internodesusually shortbut toward the base of the plant may reach 55 mm in Geographic distribution.Known only from length,somewhat angular, veryhairy. Leaves the type locality. We collected it in places tetrafoliolate.Stipules withthe fused base 7-9 with deep red soil where the arboreal mmlong x 1.5-2 mmwide and the freeportion vegetation had been recently removed. 11-14 mm long x 1 mm wide, rigid, with Currently, with its demographic growth, prominentveins; the stipularsheath somewhat Imbirussúhas been transformedinto a suburb closed, forminga short tube at the base, or of Campo Grande, and consequently it is open; densely villous on the fused portion presumablethat we are witnessinga species in withlong, brownhairs; on the freeportion the the process of extinction. externalsurface is villous but the epidermisis visible. Petiole 2-4 mm long, villous. Rachis Obs. Artificialhybrids were obtained with 4-5 mm long, villous. Leaflets elliptical or A. gracilis and A. stenophylla,of the same somewhat obovate, obtuse to moderately section, and with A. Rigonii and A. acute, upper leaf surface villous to woolly, appressipila of the section Procumbentes.In with hairs more or less curly but epidermis almost all cases the pollen fertilitywas very visible; lower surface densely villous, hairs low, less than 1%, except withA. gracilis in adpressed. Apical pair of leaflets 24-25 mm which the hybridshowed 6% pollen fertility. long x 9-10 mmwide, basal pair of leaflets21- It is worthnoting that while A. Mardi has sub- 25 mm long x 9-10 mm wide; midvein orbiculateleaflets, it crosses with the species prominent beneath, margin and secondary with the narrowest leaflets of the section veins slightlymarked. Flowers clusteredat the Erectoides . base of the plant. Fruit subterranean;pegs

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Fig. 11. ArachisOteroi: A, schematicof theplant; B, leaf,lower surface; C, leaf,upper surface (G.9790). A. brevipetiolata: D, leaf,upper surface; E, leaf,lower surface (G.10138). horizontal,ca. 20 cm long, growingca. 2 cm Geographic distribution.Known only from deep; articles 13 mm long x 6 mm wide, the type locality.This is the species in section epicarp smooth; seed 11 mm long x 3.5 mm Erectoides thatgrows in the northeasternmost wide. edge of its range. It was collected near the

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Paranaiba River on an open trail in a forested fusiformenlargements. Basal partof the stems environment.Today the place has been totally beneath the soil with numerous subterranean changed by the extensivecultivation of sugar flowersand pegs. Stems erect,20-30 cm, up to cane and by the increasingurbanization of the 55 cm long,little branched, frequently arching area. toward the soil; internodesfurrowed, villous, withlong silkyhairs, covered by the stipules. Obs. This species is close to A. Benthamii Leaves tetrafoliolate.In well-developedleaves fromwhich it may be separatedby its narrow, thefused portion of the stipulesreaches 20 mm ellipsoid seed, its veryshort petiole, its woolly in lengthwith margins 4 mm wide, the free upper leaflet surface, and by the concatenate portionup to 30 mmin length.Petiole usually 8- thickeningsof the roots. 15 mmlong, up to 20 mm.Rachis some 10 mm long. Leaflets oblong to obovate, rigid, 5. Arachis Oteroi Krapov. & W.C. Gregory coriaceous; apex obtuse, mucronulate; thick nov. sp. margin, very prominent on both surfaces, Figs. 1,5; 11, A-C especially when dry; the apical pair some 55 mm long x 21 mm wide, the basal pair A. marginataauct. non Gardner, Hoehne, Flora somewhatsmaller, 50 mmlong x 16 mmwide. Brasílica 25(2): 16, p.p. táb. 10, 1940. Mendes, The fused base of the stipulesvillous, the free Bragantia7: 262, 1947,2n=20. Handro,Arq. Bot. portion acute, rigid, subglabrous, margin Estado São Paulo 3(4): 177,táb. 44, 1958. denselyciliate, hairs long. Petiolevillous on the A. OteroiKrapov. & W.C. Gregory,in W.C. back, the margin of the canal strongly Gregory,M.P. Gregory,Krapovickas, Smith & thickened,ciliate. Pulvinus densely villous. Yarbrough,in Peanuts- Cultureand Uses: 101, Upper surface of the leaflets glabrous, the 1973,nomen nudum (Hammons & al. 521,522, 523 lower surface with weak adpressed hairs and and524). frequentlywith long scatteredbristles, midvein hirsute,margin with long, dense, spreading cilia. Spikes few-flowered, very short, Herbaperennis. Radix fitsiformis ramificationibus occurring along the branches or densely incrassatis.Caule erecti,villosi. Stipulae basi grouped around the collar of the plant where villosaemarginibus dense ciliatae. Folíola oblon- theiraxis may be up to 10-20 mm long and 5- ga usque obovata, rigida, coriacea, supra flowered.Flowers in the axil of two bracts, 10 subtusqueinsigniter marginata, epiphyllo glabro, mm long, one entire and single-veined,and hypophyllopilis adpressissetulis sparsis saepe the other bifid and two-veined. Hypanthium immixtisvestito, nervo medio hirsuto,margine 30-85 mm long, villous, hairs copper-colored. ciliis longis, densis, patentibus ornate. Calyx 6 mmlong, villous. Standard 11-13 mm Hypanthium3-8.5 cm longum,villosum. Calyx 6 long x 14-16 mm wide, orange toward the mmlongus, villosus. Vexillum 11-13 mm longum x marginand yellow at thebase; wings 7-10 mm 14-16 mm latum, aurantiacum. Fructus long. Ovary sessile, biovular. Fruit biarticulatuspaxillo horizontāliusque ad 60 cm subterranean,Particulate; peg up to 60 cm longo, isthmousque ad 10 cm longo, articulo long somewhatsinuous, isthmusup to 10 cm proximali10-13 mmlongo x 5-6 mmlato, distali long; basal article 10-13 mm long x 5-6 mm usque ad 21 mm longo x 7.5 mm lato apice wide, apical articlesomewhat larger, up to 21 recurvo,pericario laevi usqueleviter reticulato. mm long x 7.5 mm wide, beak developed; pericarp smoothor lightlyreticulate, covered Holotype:BRAZIL. Mato Grossodo Sul. Cam- with small hairs that retain particles of soil. po Grande,Fazenda Imbirussú,30-1-1933, Otero Seeds cylindrical, filling the cavity of the 194 (SP 30163), (Handro,1958, táb. 44, sub. A. articles. 2n=20 chromosomes(Mendes 1947, marginata).Isotypes: RIZ, CTES. sub A. marginata,no. 82).

Erect perennial.Main root fusiform,deep, Additionalmaterial: BRAZIL. Mato Grosso do often with constrictions, branches with Sul. In locissubhumidis campi Camapuan, Riedel

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604, Herb. Acad. Petrop.418 (K); Ribas do Rio (GH, LIL, MO, NY, SI, US); 9 km E de Pardo,3-III-1959, Gregory & al. 9828 (GH, LIL, Entroncamento,ll-V-1961, Gregory& al. 10549 MO, NY, US); 38 kmW de Ribasdo Rio Pardo,25- (LIL, US); TresLagoas, 3-XII-1943,Baldwin Jr. 1-1979,Krapovickas & al. 34409 (CTES); Ribasdo 3139 (US); TresLagoas, Valls & al. 11746 (CEN). Rio Pardo,Valls & al. 11768 (CEN); CampoGran- Mun. Campo Grande;terrenos baldios junto ao de aeropuerto,22-11-1959, Gregory & al. 9735 antigocolegio Batista Matogrossensedo Bairro (GH, LIL, MO, NY, SI, US); id., 9-V-1968, Batistão,4-IV-1986, Valls & al. 9875 (CEN, CTES); Hammons& al. 548 (CTES, SI); Campo Grande, antigo campo de esportes do Colegio Batista Faz. Imbirussú,27-V-1939, Otero & al. 453 (SP); Matogrossenseno bairroImburussu, 17-IV-1984, 25-11-1959,Gregory & al. 9819 (GH,LIL, MO, NY, Valls & al. 7618 (CEN, CTES); cercade 12 kmao US); id., 8-VI-1968,Hammons & al. 524 (CTES, longoda BR-163 a partirdo perimetrourbano de US); Campo Grandel-IX-1936, Archer & al. (SP CampoGrande, 9-IV-1986, Valls & al. 9937 (CEN, 36470); Campo Grande,Faz. das Moças, 9-IX- CTES). Mun. Terenos: ca. 1 km ao longo da 1936, Archer& al. (Hoehne, 1940, tab. 10) (SP estrada de acesso a Fazenda Modelo da 36473); CampoGrande, Estaca- Jaraguary, 10-IX- EMBRAPAa partirda BR-262,5-IV-1986, Valls & 1936,Archer & al. 151 (SP 36474); CampoGrande, al. 9882 (CEN, CTES); ao longo da rodoviade 5-VIII-1936(Mendes, 1947, 2n=20), Hoehne & al. acesso a antigafazenda modelo do Ministerioda (SP 35773); PedroCelestino, 57 kmW de Campo Agriculturaa partir da BR-262,18-IV-1984, Valls Grande,9-VI-1968, Hammons & al. 549 (CTES); 37 & al. 7621 (CEN, CTES). Mun. Rio Verdede M. kmN de CampoGrande, 5-III-1959, Gregory & al. Grosso: 20 km ao sul de Rio Verde e 1,2 km ao 9832 (CTES, GH, LIL, MO, NY, SI, SP, US); nortedo corregoMatadeira a W da BR-163, 15- Congonha, 102 km N de Campo Grande,6-III- IV-1984,Valls & al. 7598 (CEN, CTES). Mun.São 1959,Gregory & al. 9853(LIL, SI, SP); 135km N de Gabrieldo Oeste:km 55 1 da BR-163e 62 kmao sul CampoGrande, ruta a Cuiabá,6-III-1959, Gregory de Rio Verde,15-IV-1984, Valls & al. 7599 (CEN, & al. 9855 (GH, LIL, MO, NY, US); 163 kmN de CTES); id., Valls & al. 7600 (CEN, CTES). Mun. Campo Grande,6-III-1959, Gregory& al. 9858 Bandeirantes:93 kmS de Rio Verdee 105km N de (CTES, GH,LIL, MO, NY, SI, SP, US); 3 kmN de CampoGrande a W de BR-163,15-IV-1984, Valls Bataguaçú, camino a Xavantina, 26-11-1959, & al. 7602 (CEN, CTES). Mun.Jaraguari: km 463 Gregory& al. 9747 (GH, LIL, MO, NY, SI, US); da BR-163e 48 kmao N de CampoGrande, a E da Faz. 3 Barras,entre Xavantina y PortoUeré (río rodovia, 15-IV-1984, Valls & al. 7603 (CEN, Pardo),26-11-1959, Gregory & al. 9763 (GH, LIL, CTES). Mun. Anaurilandia:km 90,6 da BR-267 a MO, NY, US); Faz. Pouso Alto, 30 km SE de contardo rioParana e a W de Bataguassu,26-IV- Xavantina,26-11-1959, Gregory & al. 9765 (GH, 1984, Valls & al. 7717 (CEN, CTES): id., 7718 LIL, MO, NY, SI, US); id.,Gregory & al. 9766 (GH, (CEN). LIL, MO, NY, SI, SP, US); entrePouso Alto y Xavantina,27-11-1959, Gregory & al. 9770 (LIL, Geographie distribution. This species US); 42 kmW de Porto15 Novembro,7-VI-1968, inhabits the central part of Mato Grosso Hammons& al. 521 (CTES, US); 50 km W de do Sul (Brazil), in high places, covering a Bataguaçú,28-11-1959, Gregory & al. 9786 (LIL, range that extends fromNova Alvorada to US); 10-15 km W de Douradinho,ll-V-1961, Rio Verde and from Pedro Celestino (57 Gregory& al. 10541 (CTES, LIL, MO, NY, US); km W of Campo Grande) to Bataguaçu 135km W de Bataguaçú,28-11-1959, Gregory & al. and Tres Lagoas, near the Paraná River. 9790(CEN, CTES, G, GH,K, LIL, MO, NY, SI, SP, It prefers the deep red soils of the "cerra- US); 45 kmW de Douradinho,1l-V-1961, Gregory do." & al. 10543(LIL); 150km W de Bataguaçú,28-11- 1959,Gregory & al. 9792 (GH, LIL, MO, NY, US); Obs. Arachis Oteroi produces hybrids 40 kmE de Entroncamento(mun. Rio Brilhante), with A. Hermannii and with A. 1l-V-1961, Gregoiy & al. 10545(LIL, NY, US); 25 paraguariensis ssp. paraguariensis of its kmE de Entroncamento,1-III- 1959, Gregory & al. own section, and with A. Rigonii of the 9793 (LIL, MO, NY, US); 16 km E de section Procumbentes. The hybrids Entroncamento,1-III-1959, Gregory & al. 9810 obtained with the firstspecies had a pollen

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stain of 17.6%, while those obtained withthe villous toward the base of the fused portion, last two are very sterile. margin ciliate and with bristles on the back of the fused portion. Petiole and rachis 6. Arachis Hatschbachii Krapov. & W.C. canaliculate, canal margin ciliate, glabrous Gregory nov. sp. on the back or with a few scatteredhairs and Fig. 1,6 always with sparse bristles, 1-2 mm long. The bristles on the back of the fused portion of the stipules and on the back of the petiole Herba perennis erecta. Radix polaris and rachis are always present on the apical ramificationibuspartes incrassatas elongatas leaves, but may be lacking on basal leaves gerentibus.Stipulae subglabrae, interdumbasi of the same branch. Leaflets elliptical to villosiusculae,dorso setulosae,margine ciliatae. ovate, apex sub-acute, mucronulate; upper Folíola elliptica usque ovata apice subacuta, surface smooth, glabrous, with the margin mucronulata,epiphyllo laevi, glabro, hypophyllo little- or unmarked; lower surface glabrous glabrovel paucis pilis diminutisadpressis vestito, or with scattered, diminutive adpressed nervo medio prominente, villoso, margine hairs, midvein prominent, villous, margin manifestiore,ciliato. Hypanthiumca. 5 cm somewhat marked, ciliate, with hairs exten- longum. Calyx 6 mm longus, villosus setulis ded in young leaves and curved in mature nonnullislongis immixtis. Vexillum aurantiacum leaves, rarely some short bristles occur on supra lineis rubescentibusornatum. Fructus the underside. Flowers clustered on the subterraneus, biarticulatus, paxillo glabro basal nodes buried in the soil; the axes of horizontāli15-60 cm longo, isthmoca. 2 cm the inflorescences covered by the bases of longo,articulis 9-12 mmlongis x 5.5-6 mmlatis, the stipules. Hypanthium ca. 5 cm long, pericarpiolaevi. villous. Calyx 6 mm long, villous and with some long bristles. Standard orange, yellow Holotype:BRAZIL. Mato Grossodo Sul. 25 km at the base and with red lines on the front; N de Bandeirantes,100 kmN de Campo Grande, wings orange with the apex yellow. Fruit erecta,clavos largos,horizontales, mato cerrado, subterranean; pegs horizontal, glabrous, suelo arena roja, 6-III-1959, Gregory,Krapo- somewhatsinuous, ca. 15-20 cm long and up vickas& Pietrarelli9848 (CEN). Isotypes:CTES, to 60 cm; isthmusca. 2 cm long; articles9-12 G, LIL, P, RB, SI, SP. mmlong x 5.5-6 mmwide, pericarpsmooth to lightlyreticulate, covered by a dense coat of Erect perennial.Deep-rooted, branch roots small hairs. Seeds 9-10 mm long x 5 mmwide. with elongated enlargements. One to four 2n=20 chromosomes(Smartt 1964, Smartt& erectbranches, some 20-40 cm long and up to Gregory1967, GKP 9848). 1.2 m tall, grow up fromthe collar withlittle or no furtherbranching. Internodes2-5 cm Additional material: BRAZIL. Mato long, quadrangular, with corners very Grosso do Sul. Capão Redondo (mun. Rio prominent, glabrous or somewhat villous, Verde), 20-V-1973, Hatschbach 32105 with scattered hairs on the young parts; (CTES, MBM); 190 Ian N de Campo Grande, bristles present toward the apex of the camino a Cuiabá, 6-III-1959, Gregory & al. branches. Leaves tetrafoliolate. In well- 9863 (GH, LIL, MO, NY, SI, US); 195 km N developed leaves, the fused portion of the de Campo Grande, camino a Cuiabá, 6-HI- stipules 11-19 mm long x 5-7 mm wide, the 1959, Gregory& al. 9865 (LIL, MO, NY, SI, freetips acute, 13-25 mm long. Petiole ca. 20 US); 220 kmN de Campo Grande,6-III-1959, mm long (15-28 mm). Rachis 9-12 mm long. Gregory& al. 9869 (GH, LIL, MO, NY, SI, Apical pair of leaflets30-5 1 mm long x 16-20 SP, US); 11 km N de Rio Verde, 7-III-1959, mm wide, the basal pair somewhat smaller, Gregory& al. 9875 (GH, LIL, MO, NY, SI, 27-45 mm long x 12-16 mm wide. Stipules US); 32 km N de Rio Verde, 7-III-1959, with prominent veins, both surfaces Gregory& al. 9880 (GH, LIL, MO, NY, SI, subglabrous or occasionally somewhat US).

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Geographicdistribution. This species grows and cylindrical toward the base of the in a small area centeredaround Rio Verde de branches,usually villous on the young parts Mato Grosso (MS), between 270 and 500 m and glabrousfrom the middle to thebase of the elevation. branches. Leaves tetrafoliolate. In well- developed leaves, the fused portion of the Obs. Of all the crosses attempted,success stipules 14-18 mm long x 6 mmwide, the free was only obtained with A. Archeri, of the portionacute, 18-26 mm long; petiole 30-38 same section, but the resultinghybrid was mmlong; rachis 14-18 mmlong; apical pair of totallysterile. leaflets42-55 mm long x 20-24 mmwide, and the basal pair 39-50 mm long x 17-19 mm We dedicatethis species to thebotanist Gert wide. Stipules with surfaces glabrous to Hatschbach who collected one of the subglabrous;back of the fusedportion villous specimensstudied of this species. to subglabrous;margin densely ciliate. Petiole and rachiscanaliculate, margin of both canals 7. Arachis cryptopotamicaKrapov. & W.C. villous, the remainder subglabrous.Leaflets Gregory nov. sp. elliptical to ovate, apex obtuse to sub-acute, surface Fig. 1,7 mucronulate;upper smooth,glabrous, withthe margininconspicuous; lower surface glabrous or with some diminutive, very Herba perenniserecta. Radix ramiflcationibusadpressed hairs, rarely with some short tenuibus,partes incrassataselongatas fusiformes bristles, midvein prominentwith some long efficientibus.Caules quadrangulares, novelli hairs toward the apex, margin littlemarked, villosi. Stipulae glabrae subglabraeve dorso ciliate. Flowers denselyclustered at the collar villosae usque subglabrae, margine dense of the plant and also present along the ciliatae. Folíola elliptica usque ovata, apice branches; axis of the inflorescences very obtusa subacutave, epiphyllo laevi, glabro, short. Hypanthium5.5-8.5 cm long, weakly hypophylloglabro aut pilis nonnullisdiminutis villous. Calyx 6 mm long, villous and with a adpressis, raro setulis immixtisvestito, nervo few scatteredbristles. Standard 15 mm long x medio prominentepilos longos nonnullosge- 18 mmwide, orange;wings 8-10 mmlong, half rente,obscure marginata,ciliata. Hypanthium yellow and half orange. Fruit subterranean, 5.5-8.5 cm longum,laxe villosum.Calyx 6 mm pegs growinghorizontally, articles 15-17 mm longus, setulis nonnullis sparsim immixtis long x 6-8 mmwide, recurvedapex, blackish, villosus. Vexillum15 mmlongum x 18 mmlatum, pericarp smooth with lightly marked aurantiacum. Fructus biarticulatus articulis longitudinalveins. 15-17 mmlongis x 6-8 mmlatis, apice recurvo, pericarpio laevi. Selectedadditional material: BRAZIL. Mato Grosso do Sul. Mun. SonoraGaucho (Benjamín Holotype:BRAZIL. Mato Grosso.Itiquira, rio Constant):108 km N de Coxim,54°48'W, 17°40'S, Correntes,12 kmW del km352 de la rutaCuiabá- 8-III-1959,Gregory & al. 9889 (LIL, US). Mun. Campo Grande,54°57'W, 17°37'S, 14-XII-1976, Coxim:km 379, caminoCuiabá a CampoGrande, Krapovickas& Gregory30026 (UB). Isotypes: 54°57'W,18°24'S, 14-XII-1976, Krapovickas & al. CEN, CTES, G, GH, K, LIL, MO, NY, P, RB, SI, 30024 (CEN, CTES, GH, MO, NY, US); Fazenda SP, US. Buriti,Pantanal do Paiaguas, 65-70 km W del caminoCuiabá a CampoGrande, km 379, 55°15'W, Erect perennial. Main root deep with 18°S,14-XII-1976, Krapovickas & al. 30025(CEN, elongated enlargements,slender branch roots CTES, G, GH, K, MO, NY, RB, US); 28 kmW del with elongated fusiformenlargements. A few caminoCuiabá a Campo Grande,20 kmN de Rio sparse distichous, erect branches, 30-40 cm Verde, 55°10'W, 18°40'S, 13-XII-1976, long (up to 60 cm) are bornefrom the collar of Krapovickas& al. 30023 (CEN, CTES, GH, MO, the plant. Internodes25-35 mm long, up to 60 NY, US); 32 kmW del caminoCuiabá a Campo mm long, quadrangular with marked comers Grande,20 kmN de Rio Verde,55° W, 18°S, 13-

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XII-1976,Krapovickas & al. 29949 (CEN, CTES, 352,figs. 3,4 & 11, est. 4, 1962.Conagin, Bragantia US). Mun. PedroGomes: 17 kmN de Coxim,BR- 22(11):126, 1963. 163,faz. Palmeira,12-IV-1984, Valls & al. 7563 (CEN, CTES); id.,Valls & al. 7565 (CEN); id.,Valls Herba perennis. Radix ramificationibus & al. 7566 (CEN, CTES); Sumidourodo rio incrassatis. Caulis principalis erectus, rami Corrente,15 km NW da BR-163 (98 km N de decumbentesramosissimi. Stipulae marginibus Coxim),13-IV-1984, Valls & al. 7568(CEN, CTES); basi connatis tubumbrevem 1.5 mm longum 200 m ao N do corregoGaucho, BR-163, km 748,5, efficientes,subglabrae, dorso villosae, margine 13-IV- 1984, Valls & al. 7572 (CEN, CTES). Mun. ciliatae. Foliola elliptica,acuta, epiphyllolaevi, Coxim:l,5kmSdorioPiquiri, BR-163, 13-IV-1984, pilis brevissimis erectis vestito, hypophyllo Valls & al. 7574 (CEN, CTES). Mun. Rio Verde: villoso, nervo medio prominente, obscure Entradada rodoviaque liga a BR-163 ao corrego marginata, ciliata. Hypanthium3.5-11.5 cm Feiosona serrada Alegria,14-IV-1984, Valls & al. longum,villosum. Calyx 5-7 mm longus pilis 7588 (CEN, CTES); id., Valls & al. 7590 (CEN, longissericeis setulis nonnullis immixtis vestitus. CTES); Sope da serrada Alegriana estradapara Vexillum9-19 mm longumx 14-22 mm latum, oeste que parte da BR-163, a 18 km N de Rio aurantiacum. Fructus biarticulatus paxillo Verde,14-IV-1984, Valls & al. 7593 (CEN, CTES); horizontāli5-30 cmlongo, isthmo 2-20 mmlongo, 12 km S de Rio Verde e 4 km ao nortede corr. articulis 8-20 mm longis x 5-8 mm latis, Pereirinhaa lesteda BR-163,15-IV-1984, Valls & pericarpiolaevi. al. 7596 (CEN, CTES). Mun. Coxim: faz. Santa Rosa, Paiaguás, 22-V-1986, Pott2191 (CTES). Holotype: BRAZIL. Mato Grosso do Sul. Mun. Rio Verde: 1 kmW da BR 463, ao longoda Aquidauana, 19-V-1939, Otero 423-A (SP estradapara Rio Negro,29-X-1985, Valls & al. 41208!). 9455(CEN, CIES). Perennial plant. Main taproot deep, with Geographie distribution. This species enlarged branch roots. Collar thick, with grows in the northernextreme of the range numerousclustered flowers which in fruiting of section Erectoides, from the Correntes produce a "mare's tail" of pegs. Mainstem River, the border between the states of erect, up to 80 cm tall, much branched, Mato Grosso and Mato Grosso do Sul, to the forminga bush of ca. 80 cm in diameter. vicinityof Rio Verde do Mato Grosso, in the Secondary branches decumbent, 30-75 cm "cerrado." long, tertiarybranches 12-65 cm long and quaternarybranches up to 45 cm long. Stem Obs. The name of this species derives softly villous, internodes to 51 mm long, fromthe character of the place fromwhich covered by the stipulestoward the apex of the the type was collected. There the Correntes branches. Leaves tetrafoliolate.Fused part of River plunges into a great hole and runs fora the stipules 10-31 mm long, the freepart 12- distance below ground. 37 mm long; the margins usually united forminga shorttube up to 1.5 mm long toward 8. Arachis major Krapov. & W.C. Gregory the base of the plant. Petiole 20-5 1 mm long; nov. sp. rachis up to 13 mm long. Leaflets elliptical, Figs. 1,8; 12,A-B apical pair 16-58 mm long x 6-19.5 mm wide, thebasal pair 12-51 mmlong x 5-17 mmwide; on the mainstem the leaves are somewhat ArachisDiogoi Hoehnesubsp. major Hoehne, larger, the apical pair reaching 70 mm x 19 Flora Brasílica 25(2) part. 122: 12, fig. pág. 4, mm; length/widthratio 2-3.7:1. Stipules with 1940,nomen nudum. Otero, Serv. Inform. Agrie. prominent longitudinal veins, fused part Min. Agrie. 25-26, 1941. Mendes,Bragantia 7: villous, the free apices with subglabrous 262 (2n=20), 1947. Otero,Serv. Inform.Agrie. surfaces and marginswith long cilia. Petiole Sér. Did. 11: 172-175, 1952. Conagin,Bragantia and rachis villous; very infrequentlybristles 18(5): 67, 1959. Conagin,Bragantia 21(21): 349- occur on the base of the stipules and on the

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 petiole. Leaflets smooth above, with al. 551 (CEN, CTES, US); id., 9-VI-1968, abundant,very small erecthairs, especially on Krapovickas& al. 14447(CTES); RodoviaCampo the young leaves, sometimesthese small hairs Grande-Aquidauanaao longo dos kms52 a 110, are localized toward the base of the leaflet, 25-1-1979,Leitão Filho & al. 9301 (UEC); Estrada along the midvein; lower surface with CampoGrande-Aquidauana, km 110, 14-XII-1976, prominentmidvein, villous, rarelywith hairs Shepherd & al. 4084 (UEC); 29 km O de more or less adpressed or witha few bristles; Aquidauana,BR-262, 13-11-1993,Hatschbach & marginbarely marked,ciliate, and also rarely al. 59010 (CTES, MBM); 12 km E de Miranda, with a few bristles. Spikes short, few- caminoa Aquidauana,12-XII-1976, Krapovickas flowered,along the lengthof thebranches and & al. 30016(CEN, CTES, G, GH,IAC, K, LIL, MO, more frequentlyclustered on the half-buried NY, SI, SP, US); id., 12-XII-1967,Allem 699 collar of the plant. Hypanthium3.5-11.5 cm (CEN); Miranda,XII-1944, Octacillo (IAC-7891, long,softly villous. Calyx villouswith long, silky SP); Miranda,km 2-3 da rod.para Agachi, 13-11- hairs and some scatteredbristles; lower lobe 1993,Hatschbach & al. 59045 (CTES, MBM); id., subfalcate,6-7 mmlong, upper lobe broad,5-6 Hatschbach& al. 59049 (CTES, MBM); 40 kmS mm long. Standard9-19 mm long x 14-22 mm de Aquidauana,camino a Nioaque, 10-VI-1968, wide, orange with a central yellow spot and Hammons& al. 559 (CEN, CTES, NY, SI, SP, US); with rosy lines on the upper surface; wings 41 kmS de Aquidauana,Fazenda Varzea Alegre, yellow. Fruitingclustered at the base of the caminoa Nioaque,30-VI-1977, Krapovickas & al. plant and on the branches in contactwith the 30144 (CEN, CTES, NY, SP, US); 5 km S de soil. Fruit Particulate; peg 5-30 cm long, Nioaque, 10-VI-1968,Krapovickas & al. 14424 somewhatvillous towardthe base, of horizon- (CEN, CTES); 8 kmW de Jardim,camino a Porto tal growth,articles 2-10 cm deep; isthmus2- Murtinho,10-VI-1968, Hammons & al. 562 (CEN, 20 mm long; articles 8-20 mm long x 5-8 mm CTES, SP, US); BR-262, 8 km W do acesso a wide, pericarp smooth,covered with a dense Palmeirase 10 kmE do rio Dois Irmãos,18-IV- coat of diminutivehairs. Seeds 9-12.8 mm 1984,Valls & al. 7628 (CEN, CTES); id.,Valls & al. long x 3.2-5.3 mm wide, testa ochraceous or 7630 (CEN, CTES); 32 km W do acesso a white. 2n=20 chromosomes (Mendes 1947 Aquidauana,BR-262, 19-IV-1984, Valls & al. 7632 sub A. Diogoi ssp. major no. 85). (CEN, CTES); 15 kmda intersecçãocom a rodovia Guia Lopes-Jardimna estradaBonito-Jardim, 21- Selectedadditional material: BRAZIL. Mato IV-1984,Valls & al. 7644 (CEN, CTES); 8 kmS da Grosso do Sul. Aquidauana, 18-V-1939,Otero BR-262 no trechoAquidauana-Nioaque, 16-IV- 423 (SP-41207); Aquidauana,campo de aviação, 1985,Valls & al. 8530 (CEN, CTES). Mun. Rio 22-V-1939,Otero 439 (SP 41210); Aquidauana,27- Negro: 11,6 km SE da pontesobre o rioNegro na H-1959,Gregoiy & al. 9825 (GH,LIL, MO, NY, SI, estradapara Aquidauana,29-X-1985, Valls & al. US); Aquidauana, aeròdromo, 27-11-1959, 9468 (CEN, CTES); id.,9469 (CEN); BR-262,km Gregoiy& al. 9826 (GH,LIL, MO, NY, SI, SP, US); 462, 17 kmE do Anastacio,5-IV-1986, Valls & al. Aquidauana,24-IV-1977, Gibbs & al. 5467 (UEC); 9887 (CEN, CTES); 8 kmS da BR-262 na estrada 4 kmW de Aquidauana,30-VI-1977, Krapovickas Aquidauana-Nioaque,7-IV-1986, Valls & al. 9918 & al. 30145 (CTES); Fazenda Santa Virginia,7 (CEN, CTES); Fazenda Dallas, logo a NW de leguas de Aquidauana, 20-V-1939, Otero 438 Aquidauana,29-X-1986, Valls & al. 10384(CEN, (RBR,SP 41209,SP 41210); 2 kmE de Piraputanga CTES); 52 kmNW de Aquidauanana estradaque 28 kmE de Aquidauana,9-IV-1968, Hammons & penetrano Pantanal,29-X-1986, Valls & al. 10397 al. 555a (CEN, CTES, MO, NY, SI, SP,US); 23 kmE (CEN, CTES); Saida N de Aquidauana, 30-X- de Aquidauana, 3-VII- 1977, Krapovickas& al. 1986,Valls & al. 10407(CEN, CTES); Bela Vista, 30151(CEN, CTES, G,MO, NY, SP, US); id.,3-VII- Fazenda Formosa,a 4,5 leguas da cidade de Bela 1977,Krapovickas & al. 30153 (CEN, CTES, US); Vista,1 l-V-1939, Otero 402 (RBR, SP); 20-25km 33 kmE de Aquidauana,camino a CampoGrande, de Bela Vista,camino a PontaPorā, 14-11-1959, 3-VII-1977,Krapovickas & al. 30154(CEN, CTES, Gregory& al. 9651 (LIL, US); 37 km E de Bela G, NY, SP, US); 4 kmE de JoaquimMurtinho, 70 Vista,26-VI-1977, Krapovickas & al. 30128 (CEN, kmW de CampoGrande, 9-VI-1968, Hammons & CIES, NY, US); 50 kmE de Bela Vista,26-VI-1977,

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Fig.12. Arachis major : A, branch (G.9825); B, leaf(G. 10580). A. Benthamii : C, leaf(G.9753). A. douradiana : D, leaf (G.9799).

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Krapovickas& al. 30129 (CEN, CTES); Estancia Hammons & al. 551 and 555a consist of Dr. CastroPinto, 11-16 km W de Colonia Penso, individuals with and without bristles on the actualmenteAntonio João, camino Ponta Porã a petiole and on the base of the stipules. It is Bela Vista, 13-11-1959,Gregory & al. 9641 (LIL, interestingto note that the closest species, US); 10 kmW de Colonia Penso (AntonioJoão), A. Benthamii,very frequentlyshows bristles 13-11-1959,Gregory & al. 9638(LIL, MO, NY, US); on the back of the leaflets, but totally lacks id., 14-11-1959,Gregory & al. 9652 (LIL, NY, US); bristles on the petioles and stipules. 15 kmW de Bela Vistana estradapara São Carlos, Arachis major is very similar to A. 22-IV-1984,Valls & al. 7672 (CEN, CTES); 16 km Benthamii,with which it differsprimarily by W de Bela Vista, 22-IV-1984, Valls & al. 7676 its greater size. In various specimens from (CEN, CTES). west of the Sierra de Maracajú, which is the PARAGUAY. Amambay. 30 km W de Pedro boundary between the two species, the JuanCaballero, camino a Concepción,12-11-1959, chromatographicprofiles obtained had very Gregory& al. 9632 (LIL, NY, US); 48 kmSW de few blots. In contrast,the material fromthe PedroJuan Caballero, 16-V-1961, Gregory & al. east of the same mountains, which 10573(GH, LIL, MO, NY, US); 54 kmSW de Pedro corresponds to A. Benthamii, has more JuanCaballero, 1 6- V- 1961, Gregory& al. 10575 complex profiles and always with the (CTES, GH, LIL, MO, NY, US); 62 km SW de presence of one very notable blot (no. 8) PedroJuan Caballero, 16-V-1961, Gregory & al. (Krapovickas 1973: 139 and Seeligmann, 10576 (LIL, MO, NY, US); 70 km SW de Pedro pers. comm.). JuanCaballero, 16-V-1961, Gregory & al. 10580 In crosses between two populations of A. (CTES, LIL, MO, NY, US); CerroCorá (ca. 40 km major, one fromthe northof its range (HLK W de Pedro Juan Caballero), 14-VII-1968, 559) and the other from the extreme south Krapovickas& al. 14444(CEN, CTES, G, K, NY, (GK 10582), we obtained hybridswith very SI, SP, US); CerroCorá (13 kmW de PedroJuan little pollen fertility (11.7%). These are Caballero), erecta,borde de selva, 24-11-1968, possibly two differententities, but more Krapovickas& al. 14168(BAA, CEN, CTES, G, K, study would be necessary to define them. MO, US); Colonia Yvypyté,56°W, 23°S, aero- Arachis major shows very good affinity puerto,20-VIII-1989, Schinini & al. 20467 (CEN, withA gracilis, with 30% pollen stain in the CTES, G, MO, SI); id.,20-VIII-1980, Schinini & al. hybrids, and with A. Archeri (17.6 and 20469 (CTES); 45 kmS de Bella Vista,camino a 40%). It is easy to distinguish.4. major from Ruta 5, 26-11-1994,Krapovickas & al. 45037 these two species because their leaflets are (CTES). Concepción.2 kmE de Ñu Porá(antiguo lanceolate, not elliptical like ^4. major. camino Pedro Juan Caballero-Concepción), With A. paraguariensis ssp. 56°21'W,23°8'S, 16-V-1961,Gregory & al. 10582 paraguariensis, of the same section, it (LIL, US); Ñu Porá, 16-V-1961,Gregory & al. produces hybrids that are highly sterile 10588(LEL, US). (0.5% pollen stain). We also obtained hybrids by crossing A. major with A. Common name. "amendoim de guaranitica and A. tuberosa of section Aquidauana" (Otero 1952: 172). Trierectoides, with A. repens of section Caulorrhizae, with A. appressipila and A. Geographic distribution.Species grows to Rigonii of section Procumbentes, and with the south of the Gran Pantanal in a zone that A. glabrata var. glabrata, the tetraploid extends west of the Serra de Maracajú taxon of section Rhizomatosae; in all cases (Mato Grosso do Sul) and the Sierra with a very high degree of sterility. Amambay in Paraguay, usually below 300 m elevation. It prefers deep red soils in open 9. Arachis Benthamii Handro woodlands. Figs. 1,9; 12,C

Obs. Usually, Arachis major shows no bristles on the leaves, but the specimens Handro,Arq. Bot. Estado São Paulo 3: 179,táb.

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45, 1958. flowered, occurring all along the branches, "Brasil,Estado de MatoGrosso, Campo Grande, and more frequently clustered densely on cultivadaen São Paulo,fl. 12/1/1957,fr. VIII/1957, the half-buried collar of the plant. leg.Oswaldo Handro 682 (SP 55525TYPUS)." Hypanthium3-8 cm long, more or less villous Arachis marginata Gardner forma with silky hairs. Calyx densely villous with submarginataHoehne, Flora Brasílica25(2) part. long, silky hairs and some scattered, long 122: 16-17,táb. 11, fig. 1, 1940,nomen nudum, pro bristles; lower lobe subfalcate, 7-8 mm long, parte, speciminaMato Grossi tantuminclusa. upper lobe broad, 6-7 mm long. Standard 7- Mendes,Bragantia 7: 262,fig. 1 & 2 (2n=20),1947, 14 mm long x 12-23 mm wide, orange with a as A. marginatavar. submarginata. centralyellow spot and with rosy lines on the upper surface; wings yellow with orange Perennial plant. Taproot deep, with apex. Fruiting clustered at the base of the thickened branch roots, which commonly plant. Fruit biarticulate, peg 6-42 cm long, also arise on the hypocotyl.Collar enlarged, growinghorizontally, isthmus 0.5-5 cm long; with numerous clustered flowers, which articles 10-16 mm long x 5-7 mm wide, upon fruitingproduce a "mare's tail" of pericarp smooth, covered by a dense coat of pegs. Stems erect, up to 75 cm tall, and also diminutive hairs. Seed coat ochraceous or somewhat decumbent, angular, villous, white. 2n=20 chromosomes (Mendes 1947, glabrescent on the basal nodes. Toward the no. 83). apex of the branches the upper third is totally covered by the stipules. Secondary Holotype:BRAZIL. Mato Grossodo Sul. Cam- branches short,the largest up to 30 cm long, po Grande,cultivada em São Paulo, fl. 12-1-1957, the branching ordinarily terminates with fr.VIII-1957, Handro682 (SP 55525!). Isotypes: short tertiarybranches up to 10 cm long. K!, US!. Leaves tetrafoliolate. Fused part of the stipules 8-22 mm long, the free part 12-23 Selectedadditional material: BRAZIL. Mato mm long, toward the base the margins unite Grosso do Sul. Mun. Campo Grande: Campo forming a tube up to 4 mm long. Petiole Grande,Capão Bonito,1 -IX- 1936, Archer3982 usually 10 mm long but up to 30 mm long. (SP); CampoGrande, 10-11-1979, Krapovickas & Rachis some 5 mm long but up to 10 mm al. 34549 (CEN, CTES, G, K, MO); 17 km S de long. Leaflets elliptical, the lower pair Campo Grande,3-VII-1977, Krapovickas& al. always narrower and somewhat shorter, 30157 (CEN, CTES, US); id.,30158 (CEN, CTES, apex obtuse and sometimes sub-acute; NY, US); RodoviaCampo Grande-SãoPaulo, km length/widthratio commonly2: 1, in extreme 25, 10-VI-1976,Leitão Filho 21 19 (UEC); Rodovia cases it can reach 1.5:1 and 3:1; in typical Coxim-Cpo.Grande, a 471 kmde Cuiabá, 23-IV- well-developed leaves, the apical pair 1978,Shepherd & al. 7589 (UEC). Mun. Terenos: measures 37 mm long x 18 mm wide, the 59 km W de Campo Grande, camino a basal pair 35 mm long x 16 mm wide. The Aquidauana,3-VII-1977, Krapovickas & al. 30156 basal part of the stipules striated, villous (CTES); 4 kmE de Murtinho(70 kmW de Campo with abundant silky hairs; the free part Grande),9-VI-1968, Hammons & al. 550 (CEN, lanceolate, acuminate, with marked CTES). Mun. Anastácio:2 kmW de Palmeira(50 longitudinal veins, villous, and with ciliate kmE de Aquidauana),9-VI-1968, Hammons & al. margins. Petiole and rachis canaliculate, 554 (CEN, CTES, SP, US); 80 km W de Campo villous. Leaflets with upper surface smooth, Grande(BR-262), 28-1-1979,Ferreira M.B. s/n hirsute, covered with very short hairs that (CEN, CTES); 85 kmW de Campo Grande,28-1- permit seeing the epidermis; lower surface 1979,Krapovickas & al. 34434(CEN, CTES); 4 km villous, entirely covered with loose, silky a lestede Palmeiras,BR-262, 18-IV-1984, Valls & hairs, frequentlywith some bristles on the al. 7622 (CEN, CTES); id.,Valls & al. 7627 (CEN, secondary veins, midvein very prominent, CTES). Mun. Sidrolandia:70 km S de Campo margin somewhat marked, generally wavy Grande,BR-163, 1-III-1959,Gregory & al. 9805 when dry, ciliate. Spikes short, few- (GH, LIL, MO, NY, SI, US). Mun. Rio Brilhante:

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Rio Anhandui, 23-X-1970, Hatschbach 25104 Pedersen);Tres Lagoas, 23-11-1981,Oliveira 21 (CTES, MBM); EntroncamentoBR- 163 y BR-267 (UEQ. (Nova Alvorada),1-III-1959, Gregory & al. 9803 (LIL); 5 km N de Entroncamento,7-VI-1968, Geographic distribution. This species Hammons& al. 523 (BAA, CTES, SI, UC); 10km E grows in east central Mato Grosso do Sul, to de Entroncamento,1-III-1959, Gregory & al. 9795 the east of the Serra de Maracajá, in a broad (GH, LIL, MO, NY, SI, US); 9 km E de area whose rivers flow toward the Rio Entroncamento,ll-V-1961, Gregory& al. 10547 Paraná. It is found preferablyin the "cerra- (LIL, US); id.,10548 (LIL, NY, US); BR-267,5 kma do," in places more or less shaded. leste do Entroncamentocom a BR-163, 26-IV- 1984, Vails & al. 7712 (CEN, CTES). Mun. Obs. This species is very similar in Brasilandia:7 kmN de portoUeré, río Pardo, 26-11- appearance to A. major, from which it 1959,Gregory & al. 9761 (GH,LIL, MO, NY, SI, SP, differsprimarily by having fewer branches. US); 30 kmSE de Xavantina,26-11-1959, Gregory However, the differencesare more notable & al. 9764 (GH, LIL, MO, NY, SI, SP, US); entre in the chromatographicprofiles. The mate- Xavantina y fazenda Pouso Alto, 27-11-1959, rial fromthe west of Serra de Maracajú, that Gregory& al. 9769 (GH, LIL, MO, NY, SI, US); we identifyas A. major, has very simple Xavantina,27-11-1959, Gregory & al. 9774 (LIL, profileswith few blots, in contrast,the mate- US). Mun. Bataguaçú: CórregoFeio, 15-11-1970, rial fromeast of those mountains,which we Hatschbach23570 (CTES, MBM); 42 km W de consider to be A. Benthamii, has more Porto 15 de Novembro,Hammons & al. 520 complex profiles and with the presence of (CTES, SI. US); Bataguaçú,26-11-1959, Gregory & blot "8" (Krapovickas & al. 1974: 139 and al. 9753 (CTES, GH,LIL, MO, NY, SI, SP, US); 25 Seeligmann, pers. comm.). km W de Bataguaçú,28-11-1959, Gregory & al. Artificial hybrids were obtained with A. 9776 (GH. LIL, MO, NY, SI, SP, US). Mun. Hermannii and A. paraguariensis ssp. Anaurilandia:50 kmW de Bataguaçú,BR-267, 28- paraguariensis, of the same section; with 11-1959,Gregory & al. 9784 (GH, LIL, MO, NY, A. appressipila and A. Rigonii of section US). Mun. Rio Verde: Rio Verde, 13-XII-1976, Procumbentes and with A. glabrata var. Krapovickas& al. 30018 (CEN, CTES, GH, MO, glabrata of section Rhizomatosae. All the NY, SP, US); Base de la escarpade SerraAlegre, hybrids are highly sterile, except that ca. 20 km N de Rio Verde, 13-XII-1976, obtained with A. Hermannii, which had a Krapovickas& al. 30022 (CTES, US); 52 kmN de 19. 1% pollen stain. Rio Verde,7-III-1959, Gregory& al. 9883 (LIL, US); BR-163 a 13,2km sul do rioTaquarí, 14-IV- 10. Arachis douradiana Krapov. & W.C. 1984,Valls & al. 7585 (CEN, CTES); BR-163,18,7 Gregory nov. sp. kmS do rio Valls & al. 7586 Taquarí. 14-IV-1984, Figs. 1,10; 12,D (CEN, CTES); 2,6 km N da vila de Perdigão,na estrada de Rio Verde a Rio Negro, 29-X-1985, Valls & al. 9456 (CEN, CTES). Mun. Coxim:Rio Herba perenniserecta. Radix ramificationibus Taquarí,13-XI-1973, Hatschbach & al. 33 179;late uniformitercrassis. Caulis quadrangularis, ClubeRio Verdena margemsul do rioTaquarí, 12- villosus.Stipulae basi marginibusconnatis tubum IV-1984,Valls & al. 7557(CEN, CTES); BR-163,1,5 usque ad 3 mm longumejjicientes, pilis plus kmN de Coxim,14-IV-1984, Valls & al. 7578 (CEN, minusveadpressis vestitae, dorso hirsutae, margi- CTES); BR-163, 5 kmN de Coxim, 14-IV-1984, ne ciliatae. Folíola ellipticaapice acuto, leviter Valls& al. 7581 (CEN, CTES); 1 kmNde Coxim,9- plicato, epiphyllolaevi pilis brevissimisplus 11-1975,Anderson 11298 (MBM); Coxim,BR-163, minusve erectis copiose vestito, hypophyllo 1900 m N del rioTaquarí, 3 1-1-1989,Krapovickas villoso,nervo medio prominente, hirsuto, margine & al. 43 165 (CEN, CTES). Mun.Pedro Gomes: rod. manifestiore,laxe ciliata. Hypanthium0.5-3 cm BR-163, 12-XI-1975,Hatschbach 37425 (CTES). longum,villosum. Calyx 4 mm longus,villosus. Mun. Guia Lopes da Laguna: 12 kmW de Ervania, Vexillum6 mm longum x 7 mmlatum, aurantiacum. BR-267,24-X-1986, Pedersen 1471 1 (CTES, Herb. Fructusbiarticulatus, paxillo horizontaleusque

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) ad 30 cm longo, isthmousque ad 5 cm longo, plant,in veryshort spikes. Hypanthium0.5-3 articulis10-12 mmlongis x 5-6 mmlatis, apice cm long, villous. Calyx villous, lower lobe paulo recurvo,pericarpio laevi. falcate, 4 mm long; upper lobe broad, 3.5 mm long. Standard orange, 6 mm long x 7 Holotype:BRAZIL. Mato Grossodo Sul. Mun. mm wide (dry). Fruiting clustered at the Dourados,5 kmNE de Dourados,lugar abierto en base of the plant or on the branches in bosque,erecta, 12-V-1961, Gregory & Krapovickas contact with the soil. Fruit biarticulate; peg 10556(LIL). Isotypes:GH, MO, NY, US. up to 30 cm long, growinghorizontally from 1-3 cm deep in the soil; isthmus up to 5 cm Perennial plant, erect, small. Taproot long; articles 10-12 mm long x 5-6 mm wide, small, 5 cm in diameternear the collar, with with beak little-marked; pericarp smooth, branch roots uniformlyenlarged. Flowers covered by a dense coat of diminutivehairs. densely clustered at the collar of the plant, covered by the soil, that upon fruitingform a Additionalmaterial: BRAZIL. Mato Grosso do "mare's tail" of pegs. Mainstem erect, some Sul. Mun. Rio Brilhante: 10 km E de 30 cm tall, little-branched, with some Entroncamento,rutas Pto. 15 Nov.-Campo Gran- reproductive branches, and secondary de-Dourados,1-III- 1959, Gregory & al. 9799 (GH, branches (n+1) somewhat arched at the LIL, MO, NY, US); id., Gregory& al. 9800 (LIL, base and later uprightbut not surpassing the GH, MO, NY, US); 5 kmSW de Rio Brilhante,12- mainstem,little or no branchingof the n+1. V-1961,Gregory & al. 10554 (LIL, US); 30 kmW Tertiarybranches (n+2), if present,are very de Rio Brilhante,estrada a Maracajá,20-VII-1977, short,not more than 5 cm long. Frequently, Gibbs& al. 5338(UEC); Rio Brilhante,26-IV-1984, the plants have only a mainstem. Stem Valls & al. 7709 (CEN, CIES); BR-267,5 kmE do quadrangular,villous, glabrescenttoward the entroncamentocom a BR-163,26-IV-1984, Valls & base of the plant; internodes 1-3 cm long. al. 7711 (CEN, CTES). Mun.Domados: 28 kmSW Leaves tetrafoliolate. Fused part of the de Dourados, 12-V-1961,Gregory & al. 10557 stipules 8-11 mm long, with the margins (LIL, US); Dourados,25-IV-1984, Valls & al. 7707 fused at the base forminga short tube up to (CEN, CTES). Mun. Maracajú: Maracajá, 3-II- 3 mm long; the free part 10-15 mm long. 1952, Kuhlmann (SP 69929); 4 km S do rio Petiole 6 mm long (rarely up to 9 mm). Brilhante,estrada Maracajá-Sidrolandia, 20-IV- Rachis 6-7 mm long. Leaflets oblong, apex 1985,Valls & al. 8644 (CEN, CTES). Mun. Anto- acute, lightly plicate, base somewhat nio João:5 kmNW de ColoniaPenso (actualmen- asymmetric;apical pair 18-42 mm long x 6- te Antonio João), 22°10'S, 56°W, 13-11-1959, 11 mm wide (length/widthratio 2.3-4.2:1), Gregory& al. 9636(GH LIL,MO, NY, US); 10km W basal pair 17-38 mm long x 5-12 mm wide; de ColoniaPenso, 13-11-1959, Gregory & al. 9639 toward the base of the plant the leaflets may (LIL). Mun.Caarapó: 9 kmW de Caarapó,19-1-1979, be elliptic. Stipule surfaces with hairs more Krapovickas& al. 34336(CEN, CTES, G,K, SP,US). or less adpressed, smaller toward the apex of the free tips; the back of the fused part Geographiedistribution. This species grows hirsute with long hairs, 1-1.5 mm long; in the southof Mato Grosso do Sul in the zone marginlongly ciliate. Petiole and rachis with of "campos limpos" (Andrade-Lima 1966), in the back villous to hirsute,canal of both very the county of Dourados and neighboring narrow, subglabrous. Pulvinus villous. counties.Grows in open places, in grasslands. Leaflets with smooth upper surface, with abundant, very short, more or less erect 11. Arachis gracilis Krapov. & W.C. hairs; lower surface villous with long soft Gregory nov. sp. midvein with hairs, prominent,hirsute, long, Figs. 1,11; 13 more or less straighthairs 1-1.5 mm long; margin somewhat marked on the underside of the leaflet, longly ciliate. Flowers Herbaperennis. Radix polaris ramificationibus clustered principally at the collar of the incrassatis. Caulis principalis erectus ramis

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 procumbentìbus,arcuatis, gracilibus. Caules scattered adpressed hairs, rarely somewhat quadrangulares, subvillosi. Stipulae basi villous, and always with rigid bristles, marginibusconnatis tubumbrevem efficientes, midvein with hairs somewhat longer and dorsovillosae usque subglabrae,setulosae, mar- marginwith scattered cilia. Flowers densely gine ciliatae. Foliota longe elliptica, acuta, clustered at the base of the plant and also aliquanto discolora: epiphyllo glabro presentalong the lengthof the stems; axis of subglabrove basi pilos nonnullis gerente, the inflorescences very short. Hypanthium hypophyllopallidiore pilis adpressis et setulis 3-8 cm long, shorterin basal flowers,villous. rigidis sparsis, in nervo medio aliquanto Calyx villous and withscattered bristles, 4-5 majoribusvestito, margine ciliata. Hypanthium 3- mm long, narrow lobe subfalcate, 6-7 mm 8 cm longum,villosum. Calyx 4-5 mm longus, long. Standard orange, 10-12 mm long x 1 1- villosus,setulis sparsim immixtis. Vexillum 10-12 13 mm wide, wings 6-6.5 mm long, yellow. mm longumx 11-13 mm latum,aurantiacum. Fruitsubterranean, Particulate; peg growing Fructus biarticulatuspaxillo horizontāliusque horizontally,up to 32 cm long,thin, glabrous; ad 32 cmlongo, isthmo 5 cmlongo, articulis 10.5- isthmus some 5 cm long; articles 10.5-12.5 12.5 mmlongisx 5-6 mmlatis, pericarpio laevi. mm long x 5-6 mm wide, pericarp smooth; seed 8.5 mm long x 4 mm wide. 2n=20 Holotype:BRAZIL. Mato Grossodo Sul. Mun. chromosomes (Fernández & Krapovickas Nova Andradina,Fazenda São Jorge,80 kmW de 1994). Bataguaçú, 275 m s.m., 28-11-1959,Gregory, Krapovickas& Pietrarelli9788 (LIL). Isotypes:GH, Selectedadditional material: BRAZIL. Mato MO, NY, US. Grosso do Sul. Mun.Brasilandia: Xavantina, 27- 11-1959,Gregory & al. 9772 (GH, LIL, MO, NY, Taprooted perennial withenlarged branch US). Mun. Bataguaçú:42 kmal W de Porto15 de roots. Mainstem more or less erect, little Novembro, 7-VI-1968, Krapovickas 14389 branched, lateral branches 30-50 cm long, (CTES); 25 km W de Bataguaçú, 28-11-1959, procumbent, arched, wavy, gracile. Gregory& al. 9775 (GH, LIL, MO, NY, US). Mun. Internodes 2-3 cm long, exceeded by the Rio Brilhante: río Anhandui, 23-X-1970, stipules, longer toward the base of stems,up Hatschbach25149 (CTES, MBM); 5 km E de to 6.5 cm, quadrangular, subvillous in the Entroncamentoestrada 15 Nov.-Campo Grande- young parts, later subglabrous. Leaves Dourados,ll-V-1961, Gregory& al. 10551 (LIL, tetrafoliolate. Fused portion of the stipules US). Mun.Ribas de Rio Pardo:5 kmE de Ribasde 9-15 mm long, briefly tubular toward the Rio Pardo, rio Pardo, 3-III-1959, Gregory& al. base, the free portion very acute, 11-21 mm 9829 (LIL, US); Ribasdo Rio Pardo,río Botas, 25- long x 1-1.5 mm wide. Petiole 10-18 mm 1-1979,Krapovickas & al. 34361(CEN, CTES, US). long, up to 25 mm. Rachis 4-10 mm long. Mun. Anaurilandia:km 90,6 da BR-267,a contar Leaflets long elliptic, somewhat asymmetric do rio Paraná e a W de Bataguassú,26-V-1984, at the base, acute, mucronulate,length/width Valls& al. 7715 (CEN, CTES); id.,Valls & al. 7716 ratio 4-8:1, apical pair 23-47 mmlong x 3-10 (CEN,CTES). mm wide, the basal pair somewhat smaller, 21-41 mm long x 3-8 mm wide. Stipules Geographiedistribution. This species grows rigid, with marked veins, villous to in areas adjoiningthe Pardo River,in the SE of subglabrous, with bristles on the back; free Mato Grosso do Sul, fromRibas de Rio Pardo portion with citiate margin. Petiole and in the north,to Bataguaçú in the southeast.It rachis more or less villous and with rigid preferssandy, ashen soils near watercourses. bristles, more abundant on the back of the petiole. Leaflets somewhat two-toned, Obs. Arachis gracilis is a species that has margin somewhat thickened on the back; a special abilityto cross between sections. It upper surface subglabrous to glabrous, with crosses withA. tuberosa (18.7% pollen stain) some short hairs clustered at the base of the and with A. guaranitica (10.3%) of section midvein; lower surface lighterin color, with Trierectoides, with A. Rigonii (12.2%) of

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)

Fig.13. Arachis gracilis: plant (K.34361).

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section Procumbentes and with A. predominantlyreproductive. Internodes up to duranensis (0%) of section Arachis. Within 50 mm long, quadrangular, subglabrous to its own section it produced hybridswith A. slightlypubescent. Leaves tetrafoliolate,well paraguariensis ssp. paraguariensis (1.7%), separated.Basal partof the stipules 14-25 mm with A. Martii (6.2%), with A. Hermannii long, withthe fused marginsforming a basal (20.1%) and withA. major (30%). As in the tube some 9 mm long which,with the growth case with other Arachis species with good of the stem, is broken, leaving the margins crossing ability, A. gracilis grows on the necrotic;the freeparts very acute, 20-35 mm eastern edge of the range of section long x 1-1.5 mmwide. Petiole 15-35 mmlong, Erectoides and does not grow togetherwith rachis 7-11 mm long. Leaflets ovate- any of the taxa with which it would produce lanceolate, mucronulateat .the apex, length/ hybrids. width ratio 3-7:1, the apical pair 29-62 mm long x 8-10 mmwide, thebasal pair 27-59 mm 12. Arachis Hermannii Krapov. & W.C. long x 6-8 mm wide. Stipules with very Gregory nov. sp. prominentveins, subglabrous surfaces, with Fig. 1,12 diminutive adpressed hairs and with long hairs on theback of the fusedportion; the free tips are usually longerthan the petiole,linear, Herba perennis. Radix polaris profunda, rigid,with citiate margin. Petiole and rachis crassa, fusiformis,ramificationibus incrassatis. withadpressed hairs on the back, the margins Caulis principaliserectus ramis decumbentibus. of both canals ciliate. Upper surface of the Caulis subglaber. Stipulae marginibus basi leaflets subglabrous, with some hairs at the connatistubum usque ad 9 mmlongum efficientes, base of the midvein,very young leaves with pilis diminutisadpressis dorso longis vestitae, very small scattered hairs, smooth, with margineciliatae. Folíola ovato-lanceolataepiphyllo inconspicuous margin; lower surface with sub-glabro, basi pilos nonnullis gerente, inconspicuous adpressed hairs, scattered hypophyllopilis adpressis paulo conspicuis bristlesand long hairs,more or less adpressed, setulisque sparsis, in nervo medio prominenti along themidvein, veins and marginprominent; majoribusvestito, conspicue marginata, margine margin with more or less dense, recurved pilis recurvis plus minusve densis instructo. hairs. Flowers and fruitsclustered at the collar Hypanthium5-10 cm longum,villosum. Calyx 6-7 of the plant, flowersalso occurringalong the mmlongus, setulis nonnullis immixtis subvillosus. lengthof the branches, but these are usually Vexillum14-16 mmlongum x 15-18 mmlatum, unfruitfulifthey are notin contactwith the soil. aurantiacum. Fructus biarticulatus, paxillo Hypanthium 5-10 cm long, villous. Calyx horizontāliusque ad 43 cmlongo, isthmo 0.5-11.5 subvillouswith a fewbristles, the wider lobe 6- mmlongo, articulis 9-12 mm longis x 5-7mm latis, 6.5 mm long, the narrowerlobe subfalcate, 7 pericarpiolaevi. mm long. Standard orange, 14-16 mm long x 15-18 mm wide; wings 7-9 mm long. Fruit Holotype:BRAZIL. Mato Grossodo Sul. 54 km subterranean, biarticulate; peg growing N de CampoGrande, camino a Cuiabá,5-III-1959, horizontally,commonly ca. 17 cm long and up Gregory,Krapovickas & Pietrarelli9841 (LIL). to 43 cm long, glabrous; isthmus fromvery Isotypes:GH, MO, NY, US. short (5-10 mm long) up to 11.5 cm long; articles9-12 mmlong x 5-7 mmwide, the distal Erectperennial. Taproot deep, withfusiform one always larger than the basal, pericarp thickenings and with enlarged branches. smooth. Seed 10-13 mm long, 5-6 mm wide. Mainstem erect with a predominance of 2n=20 chromosomes (Smartt 1964; GKP vegetativebranches on the first14 nodes, on 9843; Smartt & Gregory 1967, sub A. the remainder(up to node 18) reproductive Diogoi). branches predominate. Lateral branches decumbent, 50-80 cm long, n+1 branches Additionalmaterial: BRAZIL. Mato Grosso do predominantlyvegetative, and n+2 branches Sul. 54 km N de Campo Grande, 5-III-1959,

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Gregory& al. 9839 (GH,LIL, MO, NY, SI, SP, US); There are also differences in the id.,9843 (LIL, SI). Mun.Coxim: proximo a BR-163, ao chromatographic profiles in that A. longoda Estradaparalela ao rioTaquari, para oeste, paraguariensis has two blots, 7 and 8, which margemsul, 12-IV-1984,Valls & al. 7555 (CEN, are missing in A. Hermannii (Krapovickas CTES); 1,5km oeste da BR-163,ao longoda estrada 1973: 139 and Seeligmannpers. comm.). paralelaao rioTaquari, 12-IV-1984, Valls & al. 7560 (CEN, CTES); Coxim,3 l-X-1986,Valls & al. 10426 Obs. 3. The specimen Vails 7594 is (CEN, CTES); id.,Valls & al. 10427(CEN, CTES). differentiatedfrom the other material by the Mun. Rio Verdede M. Grosso:Corrego Fundo na presence of bristles on the stipules and the BR-163,a 8 kmN doRio Verde, 14-IV-1984, Valls & petiole, but is the same in the shape of the al. 7594 (CEN, CTES). Mun.Aquidauana: entre as leaflets and in the nature and distributionof fazendasBuritizinho e Boa Vista, ao longo da the tomentum. estradaque sai para noroestede Aquidauana,29- X-1986,Valls & al. 10386(CEN, CTES); id.,Valls & We dedicatethis species to the botanistF.J. al. 10387(CEN, CTES); entreas fazendasBoa Vista Hermann (Horticultural Crops Research e Alegreteao longoda estradaque sai noroes- Branch, authorof "A of the para USDA)," Synopsis tede Aquidauana,29-X-1986, Valls & al. 10390;52 Genus Arachis (1954). km NW de Aquidauanaao longo da estradaque penetrano Pantanal,29-X-1986, Valls & al. 10396 13. Arachis Archeri Krapov. & W.C. (CEN,CIES). Gregory nov. sp. Fig. 1,13 Geographie distribution. This species grows in the centerof Mato Grosso do Sul, to the northof Aquidauana and Campo Grande, A. Diogoi auct non Hoehne, Hoehne, Flora extendingto Coxim, between 160 and 330 m Brasílica25(2) part. 122: 10, 1940, tab. 2, 1919.Otero, elevation. Serv.Inform. Agrie. Min. Agrie., figs. pags. 30, 34, 35, 51. (n. 454), 1941. Mendes,Bragantia 7: 262, Obs. 1. Hybridshave been obtainedbetween 2n=20,1947. Otero, Serv. Inform. Agrie. Sér. Did. 11: A. Hermannii and A. appressipila and A. 170-172,1952. Conagin, Bragantia 21: 352, est.5, Rigonii of section Procumbentes', with A. figs.5 A, 5B,e 12,1962. duranensis, an annual species of section A. Diogoi Hoehne f. sericeo-villosaHoehne, Arachis' and with A. glabrata var. glabrata Flora Brasílica25(2) part. 122: 11, táb. 4. 1940, and A. pseudovillosa of sectionRhizomatosae. nomennudum. The hybridswith sectionRhizomatosae never A. Diogoi Hoehnef. subglabrata Hoehne, Flora floweredand the other intersectionalhybrids Brasílica25(2) part.122: 11, táb. 3, 1940,nomen have a verylow percentageof fertilepollen. nudum. Within its own section,A. Hermannii pro- A. Diogoi Hoehne f. minor Hoehne, Flora duces hybridswith A. Benthamii,A. gracilis Brasílica25(2) part.122: 12, táb. 5, 1940,nomen and A. Oteroi, with pollen fertilityof 17.6 to nudum. 20.1%. On the other hand, with A. paraguariensis ssp. paraguariensis the Herba perennis. Radieis ramificationes hybrids produced pollen with a very low incrassatae, fusiformes, elongatae. Caulis pollen stain (0.9 to 4.3%). principaliserectus ramis decumbentibus.Caulis quadrangularis, subglaber usque villosus. Obs. 2. In the shape of its leaflets, A. Stipulae marginibusbasi connatistubum 4 mm Hermannii has a great resemblance to A. longumefficientes, subglabrae, margine modice paraguariensis ssp. paraguariensis , ciliatae. Folíola oblongo-lanceolata, rigida, notwithstanding these two taxa are acuta,epiphyllo laevi, praeter aliquot pilos longos differentiatedby the erect stems and enlarged in base nervi medii glabro, hypophyllopilis roots of the first,and the sinuous stems and brevibusadpressis setulisque sparsis immixtis rootswithout enlarged branches of the second. vestito, margine pilis recurvis, caducis.

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Hypanthium2-10 cm longum,villosum. Calyx 6-8 with some long hairs at the base of the mm longus, villosus,setulae nonnullaeadsunt. midvein; lower surface with short, very Vexillum8-16 mm longumx 15-22 mm latum, adpressed hairs, hairs somewhat longer on aurantiacum. Fructus biarticulatus paxillo the midvein and usually with scattered horizontaleusque ad 50 cm longo,isthmo usque bristles; midvein very marked, margin and ad 12 cmlongo, articulis 8-13 mm longisx 5-7 mm secondary veins marked; margin with latis,pericarpio laevi. recurved, caducous hairs. Inflorescences densely clustered at the undergroundbase of Holotype:BRAZIL. Mato Grossodo Sul. Cam- the branches and also occur along the length po Grande,Cidade Universitaria,área verdecerca of the stems; floral axis up to 2 cm long de la piscina,24-1-1979, Krapovickas & Cristóbal below the soil and very short on the aerial 34340 (CEN). Isotypes:CTES, G, F, K, MBM, parts. Hypanthium of the subterranean MO, NY, P, RB, SI, SP, US. flowers 2-4 cm long, that of the aerial flowersup to 10 cm long, villous. Calyx 6-7 Erect perennial. Taproot deep with mm long, villous and with some bristles, thickened, elongated branches. Mainstem lower lobe falcate, acute, 7-8 mm long. Stan- erect, up to 65 cm tall; secondary branches dard 8-16 mm long x 15-22 mm wide, decumbent,up to 35 cm long, arched at the orange, withyellow basal spot and pink lines base such that the firstnodes are in contact on the front.Fruit subterranean,Particulate; with the soil; tertiarybranches up to 29 cm pegs growing horizontally,2-5 cm deep, up long. Stems with persistentstipules toward to 50 cm long; isthmus up to 12 cm long; the base and denselyfoliated toward the apex; articles 8-13 mm long x 5-7 mm wide, distal internodesusually 10-15 mmlong but varying article somewhat larger than the basal, up to between 5 and 27 mm, quadrangular, from 18 mm long x 8 mm wide; pericarp smooth; subglabrous to villous, especially along the seed with a white seed coat with pink tinge. corners.Leaves tetrafoliolate.Stipular sheath 2n=20 chromosomes (Mendes 1947 sub A. that completely envelops the stem, with the Diogoi f. typica no. 84). margins fused some 4 mm at the base, the remainder with the margins more or less Additionalmaterial: BRAZIL. Mato Grosso do contiguous; with the growth in thicknessof Sul. Campo Grande,5-VIII-1936, Hoehne & al. the stem, the part with fused margins is (SP 35775, SP 36478); id., 28-V-1939,Otero 457 broken,leaving the marginsnecrotic, and the (RIZ, SP); id., 23-XI-1844,Octacilio 7890 (IAC, whole sheath separates fromthe stemforming SP, UC); id., 23-III- 1948, Stephens& al. SH 259 a veryacute angle. Stipular sheath 10-25 mm (LIL); id.,22-11-1969, Gregory & al. 9736 (GH,LIL, long, the free part 10-30 mm long, linear- MO, NY, SI, SP, US); id.,9-VI-1968, Hammons & lanceolate, rigid,with prominentlongitudinal al. 547 (CTES); id.,Valls & al. 11778 (CEN); 7 kmS veins. Petiole usually 10-15 mm long, rarely de CampoGrande, 23-11-1959, Gregory & al. 9811 up to 30 mm, rachis 5-10 mm long. Leaflets (LIL, NY, SI, US); id., Gregory& al. 9812 (LIL, oblong-lanceolate, rigid, acute, somewhat MO, NY, SI, US); 17 kmS de Campo Grande,3- asymmetricat the base, mucronulateat the VII-1977, Krapovickas & al. 30159 (CTES); apex; in well-developedleaves the apical pair Imbirussú,15 km SW de Campo Grande,30-1- reaches 60 mm long x 10 mm wide and the 1933,Otero 172 (RIZ, SP); 27-V-1939,Otero 455 basal pair 55 mm long x 8 mm wide; with a (SP); 25-11-1959,Gregory & al. 9820 (LEL,NY, SI, length/widthratio of 4.5:1 to 7:1, the first US); Gregory& al. 9821 (GH, LIL, MO, NY, SI, leaves, however,usually have a smallerratio. US); InduBrasil, 18 km S de CampoGrande, 24-11- Stipules withsurfaces subglabrous, with some 1959,Gregory & al. 9817 (LIL, NY, US); Fazenda hairs towardthe margin of the sheath;margins Exper.Criação, 30 kmS de CampoGrande, 25-V- somewhat ciliate. Petiole and rachis with the 1939,Otero 442 (RIZ, SP); Otero444 (RIZ); Otero backs subglabrous or with scattered hairs, 445 (SP); 24-11-1959,Gregory & al. 9814 (LDL,NY, both canaliculate with villous canal margins. US); Lagoinha,30 kmSE de CampoGrande, 6-IX- Upper surfaceof theleaflets smooth, glabrous, 1936, Archer & al. 3968 (K, NA, NY, SP);

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Anhandui,60 kmS de CampoGrande, 1-III-1959, Among the diverse forms described by Gregory& al. s/n(LIL 462183); 59 kmW de Cam- Hoehne, it is worthpointing out A. Diogoi f. po Grande,camino a Aquidauana, 3-VII- 1977, sericeo-villosa Hoehne, 1940: 11, tab. 4 Krapovickas& al. 30155 (CEN, CTES, NY, US); ( nomen nudum), which, although it is very Rodovia Campo Grande-Aquidauanaao longo similar to A. Archeri, is differentiatedby the dos kms 52 a 110, 25-1-1979,Leitão Filho & al. more abundant and silky tomentumon the 9306 (UEC); 2 1 kmN de CampoGrande, camino a stems and stipules and by its leaflets with Cuiabá,5-III-1950, Gregory & al. 983 1 (GH, LIL, very short,more or less uprighthairs on the MO, NY, US); CampoGrande Fazenda Xarqueada upper surface and a villous lower surface. Velha,3-IX-1936, Archer 3967 (NA, SP); Campo The material we studied of this variant Grande,"amendoim erecto", 9-IX-1936, Archer & also came from Campo Grande and its al. 138 (K, NA, US); CampoGrande, 3-XII-1943, environs:Campo Grande, 24-XII-1932, Ote- Baldwin Jr. 3140 (US). Mun. Campo Grande: ro 1 (SP); Faz. Experimentalde Criação, 25- AntigaFazenda Modelo do Ministerioda Agri- V-1939, Otero 444 (SP), Otero 444 bis (RIZ, cultura,17-IV-1984, Valls & al. 7614 (CEN, CTES); SP), 24-11-1959, Gregory & al. 9816 (LIL, km 374, da rodovia de Campo Grande a US); 48 km N de Campo Grande, camino a Sidrolandia,16-IV-1984, Valls & al. 7606 (CEN, Cuiabá, 5-III-1 959, Gregory& al. 9835 (GH, CTES). Mun. Terenos:BR-262, acesso a Antiga LIL, MO, NY, SI, SP, US). Fazenda Modelo, 18-IV-1984,Valls & al. 7619 In the experimental crosses, A. Archeri (CEN, CTES); id., 7620 (CEN, CTES). Mun. produced hybrids with fertility that Jaraguari:Jaraguari, 9-IV-1986, Valls & al. 9941 fluctuatedbetween 17.6 and 40.4% with A. (CEN, CTES). major and almost completely sterile hybrids with A. Hatschbachii, of its same section, Cultivated material: U.S.A. Florida. and with A. Rigonii of section Gainesville,cult., originally from Campo Grande, Procumbentes. Because of its oblong- MS (Archer138), 7-V-1939, Carver s/n (SP 42275). lanceolate leaves, A. Archeri is easily BRAZIL. São Paulo. Campinas,Instituto Agro- distinguishedfrom all of these species and is nómico, V. 128, originally from the Sec. not sympatricwith any of them. Agrostologia,km 47, RJ,Conagin 4 (IAC 18131); id.,V.128, Conagin 12 (IAC 18669). We dedicate this species to W.A. Archer, a scientist with the U.S. Department of Common name, "amendoim do campo Agriculture,who collected it in 1936. limpo" (Otero 1952: 170). 14. Arachis stenophyllaKrapov. & W.C. Geographic distribution.This species has Gregory nov. sp. been collected in the area around Campo Fig. 1,14 Grande (MS) in an area thatextends some 60 km to the west, 60 km to the south and 20 km to the north.It grows in "cerrado," in deep red A. angustifoliaauct. non (Chodat & Hassl.) soils. Killip ex Hoehne,Hoehne, Flora Brasílica 25 (2) part. 122: 12-13,táb. 6, 1940. Hermann,Agrie. Obs. Arachis Archeri has been confused Monogr.USDA 19: 8, fig.3, 1954. Both authors with A. Diogoi Hoehne, a species likewise referto specimenFiebrig 4277. The typeof the with oblong-lanceolate leaflets, but which taxonof Chodat & Hassler shows rhizomesand belongs to the sectionArachis forits prostrate forthat reason it is treatedas a synonymof A. to semi-prostratehabit, for fruiting only along glabrata. the lengthof its branches,and forits failureto A. Diogoi auct. non Hoehne, Burkart, cluster its flowers at the collar, as we have Darwiniana3(2): 282, 1939. It likewiserefers to been able to verifyby collectingit (A. Diogoi) specimenFiebrig 4277. in its type locality (Mato Grosso, Lagoa A. Diogoi Hoehne forma submarginata Gaiba). Hoehne,Flora Brasílica 25 (2) part.122: 12, 1940

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 nomenpro parte (Otero 407). scattered on the back and abundant on the marginsof the canal, interiorof the canal with Herbaperennis. Radix profunda, fusiformis, ad scattered short, silky hairs; pulvinus light medium incrassata, ramificationibus non green, densely pilose; upper surface of the incrassatis.Rami suberectiusque decumbentes. leaflets smooth, glabrous, the lower surface Caulis villosus. Stipulae pubescentes, dorso withvery prominent midvein and marginsand villosae et marginepilis longisserieeis ornatae. markedsecondary veins, the surface scattered Foliola lineari-lanceolata,rigida, epiphyllo laevi, withadpressed hairs and withabundant long, glabro,hypophyllo pilis adpressissparsim vestito, silky hairs on the midvein and margin. nervomedio margineque prominenti copiose pilis Reproductivespikes subterranean,very short, longis serieeis obtectis, nervis secundariis 3-flowered,along the lengthof the stems and manifestis.Hypanthium 4-10 cm longumpilis also denselyclustered at the collar. Flowers in serieeissparsim vestitum. Calyx 8-9 mmlongus, theaxils of thebracts, the lower bractpointed, pilis longis serieeis ootectus. Vexillum16 mm 9 mm long x 3 mm wide, the upper bract 10 longum X 19 mm latum, luteo-aurantiacum. mm long x 3 mm wide, 2-veined and split at Fructusbiarticulatus paxillo horizontāliusque ad the apex. Hypanthium 4-10 cm long with 50 cm longo, isthmo2.5-5.5 cm longo,articulis scattered,silky hairs. Calyx with long, silky 10-16 mmlongis x 7-8 mmlatis, apice recurvo, hairs, the larger lobe 8 mm long x 5.5 mm pericarpionervis modice manifestis. wide, the smaller lobe falcate, 9 mm long x 1 mm wide. Standard 16 mm long x 19 mm Holotype:ARGENTINA. Corrientes.Facultad wide, up to 24 mmx 30 mm,orange-yellow all de CienciasAgrarias, Corrientes, cult., individuos over and with soft red lines on the upper transplantadosprocedentes de Brasil,Mato Grosso surface. Fruit Particulate. Pegs of various do Sul, 87 km E de PortoMurtinho, BR-34 (leg. lengths according to the position of the Hammons,Langford, Krapovickas & Hemsy572), flower,those originatingat the collar of the 25-11-1969,Krapovickas 15412 (CEN). Isotypes: plant up to 50 cm long, and those originating CTES, G, K, SP, US. along thebranches 7-20 cm long. Basal article 3-15 cm deep, 10-13 mmlong x 7-8 mmwide, Erect or decumbentperennial. Root deep, isthmus2.5-5.5 cm long,and the apical article fusiform,thickened toward the middle, with 13-16 mm long x 7-8 mm wide, beaked; peg unthickened branch roots. Collar with and isthmus covered with small, caducous numerousburied compact inflorescencesthat hairs, epicarp not prominently veined, upon maturityform a dense "mare's tail" of covered witha dense coat of diminutivehairs. fruits. Branches up to 1 m long, erect or Seed cylindrical, completely tilling the decumbent;stems green, furrowed, somewhat article's cavity.2n=20 chromosomes(Fernán- twisted; internodeslong, up to 60 mm long, dez & Krapovickas 1994). with long, silky hairs. Leaves distichous, tetrafoliolate,on the lower nodes trifoliolate Additionalmaterial: BRAZIL. Mato Grosso do leaves are often present. Stipules with the Sul. S/l. Campo,G.T. Bertonis/n (LPS 23126). fusedpart 20-23 mmlong, free part 30-35 mm Mun.Porto Murtinho: 87 kmE de PortoMurtinho, long x 2 mm wide. Petiole 20-40 mm long, camino a Jardim,en campo con palma enana canaliculate; rachis 6-10 mm long. Leaflets {Butta sp.), suelo rojo arcilloso-arenoso,12-VI- linear-lanceolate, stiff, the basal pair 1968,Hammons & al. 572 (CEN, CTES, US); 90 somewhat smaller than the apical pair, in km E de Porto Murtinho, 12-VI-1968, well-developedleaves the basal leaflets45-5 1 Krapovickas& al. 14442 (CTES); 67 km E de mm long x 4-7 mm wide, and the apical PortoMurtinho, 29-VI-1977, Krapovickas & al. leaflets 55-62 mm long x 7-9 mm wide. 30136 (CEN, CTES, US); 70 km E de Porto Stipules with long, silky hairs on the margin Murtinho29-VI-1977, Krapovickas & al. 30137 of the freeparts and on the back of the fused (CTES). Mun. Bela Vista: 7 kmE de Bela Vista, part,both surfaces of the freeparts pubescent; 14-11-1959,Gregory & al. 9648 (GH, LIL, MO, petiole and rachis with long, silky hairs NY, US); 7 km E de Bela Vista, 26-IV-1977,

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Krapovickas& al. 30126 (CEN, CTES); Bela Vista, long,with margins somewhat imbricated at the 10-V-1939,Otero 407 p.p. (RBR). base; the freepart up to 30 mm long, 2-3 mm PARAGUAY: Concepción.Zwischen rio Apa wide, very acute. Petioles up to 30 mm long, und rio Aquidaban,Centurión, 57°35'W, 22°18'S, canaliculate,angular. Rachis ca. 10 mm long. 25-XI-1908,Fiebrig 4277 (G, K, SI, US). Leaflets oblong-lanceolate,acute, apical pair up to 62 mmlong x 2 1 mmwide, basal pair up Geographie distribution. This species to 60 mm long x 14 mm wide. Stipules with inhabitsthe northernpart of easternParaguay surfaces pubescentto subglabrous, long silky and southwesternMato Grosso do Sul, in a hairs on the margins,sometimes with bristles band whose axis is constitutedby the Rio on the fused part; petiole and rachis villous, Apa. It grows in red soils with rocky canaliculate, canal softly pubescent, outcroppingsin grasslands withabundant low sometimeswith bristles on the back; leaflets palms (Butia ). with smooth upper surface, glabrous to subglabrous and with the margin somewhat Obs. The only hybridsthat were obtained marked, the lower surface with midvein, with A. stenophylla were with A. Martii secondary veins and margins very marked, (0.07% pollen fertility) and with A. surfacewith loose silkyhairs, frequentlywith paraguariensissubsp. paraguariensis (16.8%). some bristles, midvein and margins with With its sinuous stemsand its rootswithout abundant long, silky hairs. Inflorescences enlargements,A. stenophylla is very much clusteredunderground on the collar and on the like A. paraguariensis, from which it is lower nodes, flowers with epigeous corolla distinguishedby its linear-lanceolateleaflets. that ordinarilyproduce fruit.There are also aerial flowers along the length of the 15a. Arachis paraguariensis Chodat & branches, which only fruit if the branch is Hassl. subsp. paraguariensis decumbentand the flowersoccur near the soil. Figs. 1,15a; 4; 14,A-B; Plate II Hypanthium4.5 to 12 cm long, villous. Calyx bilabiate, 6-11 mm long with silky hairs and some bristles on the outer surface. Standard Chodat& Hassler,Pl. Hassl. 2: 449, 1904,"/« 14-21 mm long x 16-23 mm wide, entirely arenosispr. Tobaty,Sept., n. 6358." Chevalier, orange or orange with a small basal yellow Rev.Int. Bot. Appl. Agrie. Trop. 13: 763, 1933. spot and with lightlymarked red lines on the A. prostrataBenth. var. intermediaChodat & upper surface;wings yellow. FruitParticulate; Hassl.,Pl. Hassl. 2: 449, 1904,"in campishumidis pegs 10-75 cm long, growing horizontally; pr. Concepción,Sept., n. 7542." isthmus2-10 cm long; articlesusually 15 mm A. Diogoi Hoehneforma submarginata Hoehne, long x 7 mmwide (up to 17x8 mm), pericarp Flora Brasílica25(2) part. 122: 12, 1940,nomen smoothwith a dense coat of diminutivehairs p.p. (Otero393, 407 & 409). and curved apical beak. 2n=20 chromosomes (Smartt 1964, Smartt& Gregory 1967, GKP Perennial plant. Taproot somewhat 9646). fusiform,up to 10 mm in diameter in its thickest part, deep, with branch roots that Holotype: PARAGUAY. Dep. Cordillera. taper uniformly,without enlargements. Collar Tobaty, IX-1900, Hassler 6358 (G!). Isotype: with numerous subterranean inflorescences BM!. with fertileflowers that forma "mare's tail" of pegs. Stems sinuous. Mainstem erect,late- Selectedadditional material: BRAZIL. Mato ral branches decumbent, 30-85 cm tall; Grosso do Sul. N del Aquidabán,G.T. Bertonis/n internodesangular, up to 7 cm long,very short (LPS 23127). Mun. Miranda: 16 km S of toward the apex of the branches, villous, or Guaicurus, 56°45'W, 20°10'S, ll-XII-1976, with long silky hairs on the angles, or Krapovickas& al. 30013 (CEN, CTES, GH, LIL, glabrous. Leaves distichous, tetrafoliolate. MBM, MO, NY, RB, SP, US); 30 km S de The fused part of the stipules up to 20 mm Guaicurus,1 l-XII-1976, Krapovickas & al. 30014

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(CEN, CTES. EAC, GH, IAN,K, MO, NY, SI, SP, KC 11488, XI-1993, Gregoiy s/n (CTES 195522). US); EstanciaEsmeralda, 45 kmS de Guaicurus, 11-XII- 1976, Krapovickas& al. 30015 (CEN, Geographic distribution. This species CTES, G, GH, MO, NY, P, RB, SI, SP, US); 30 km grows in SW Mato Grosso do Sul, in Brazil, na SerraBodoquina, 35 km depois da curvaP 2 and in eastern Paraguay, in the basin of the (RADAM), 20-X-1980,Pires & al. 17200(CTES). Paraguay River. It is the southernmosttaxon Mun. Porto Murtinho: 93 km E de Porto of section Erectoides. Its range extends Murtinho,29-VI-1977, Krapovickas & al. 30142 from the Serra da Bodoquena, near (CEN, CTES, US). Mun. Jardim:71 km W de Guaicurus (MS) to the department of Jardim,56°48'W, 21°15*S,480 m, 27-VI-1977, Paraguarí, in Paraguay. Krapovickas& al. 30133 (CTES, US). Mun. Bela Vista:Bela Vista,8-V-1939, Otero & al. 393 (SP); Obs. 1. The specimen Krapovickas & al. Bela Vista,faz. Formosa,14-V-1939, Otero 407 1 1488 (Paraguay, dep. Cordillera, 2 km de p.p. (SP); y Otero409 (SP); Bela Vista, 14-III- Ruta 2, camino a San Bernardino, 26-IV- 1959, Gregory& al. 9646 (CTES, GH, LIL, MO, 1964; CTES, G, MO, NY, US) has broad NY, SI, SP, US). leaflets and does not correspond to eitherof PARAGUAY. S/l.6-1-1919, G.T. Bertoni684 the subspecies we recognize. We visited the (LPS 23123). Concepción.Concepción, XI-1901, same locality later and could not find this Hassler 7542 (G, holotypeof A. prostratavar. variantagain. intermediaChod. & Hassler, BM, K, NY); Ñu Porá, 16-V-1961,Gregory & al. 10585 (LIL, MO, Obs. 2. The subspecies paraguariensis NY, US); 3 1 kmE de la uniónde ratas3 y 5, 18-11- and capibarensis are difficult to separate 1968,Krapovickas & al. 14006(BAA, CTES, IPA, using exomorphologicalcharacters. We treat LL, MBM). Amambay.Estancia San Luis,20 km themas two distincttaxa because, in addition NW de PedroJuan Caballero, 13-V-1961, Gregory to havingdifferent geographic distributions, the & al. 10562 (LIL, US); 1 kmN del rioAquidabán, firsthas a SAT chromosomewith differential camino de ruta 5 a Bella Vista, 23-VI-1977, stainingdetected in 1 1 accessions, and in the Krapovickas& al. 30115 (CTES); 9 kmSE de Bella chromosomes of the second, observed in 5 Vista, 24-VI-1977, Krapovickas & al. 30118 accessions, thisdifferentiation was notpresent (CTES, US); 20 kmSE de Bella Vista,24-VI-1977, (Fernández & Krapovickas 1994). Krapovickas& al. 30124 (CTES, US); Colonia Yvypyté,56°W, 23°S, 20-VIII-1980, Schinini & al. Obs. 3. This taxon occupies a key position 20470 (CTES); Bellavista, 19-X-1981,Schinini forits ability to producehybrids, both within its 21353a(CTES, G, MO); 9 kmSE de BellaVista, 20- section as well as with other sections (Plate X-1981,Schinini 21387 (CTES, G, K, LIL, MO); II). id., Schinini21388 (CTES); 12 km SE de Bella In crosses made betweenplants of the same Vista,Ea. Primavera,22-X-1981, Schinini 21516 subspeciesfrom different localities, the fertility (CEN, CTES, G, K, LIL, MO, P). Cordillera. of the pollen fluctuatedbetween 24 and 70%. Tobatí,24-V-1964, Krapovickas & al. 11462(AS, The hybrid with the most closely related C, CTES, F, G, GH,MO,NY, LIL, MBM,P, SI, US); species, A. stenophylla, gave 16.8% pollen 2 kmN de Tobatí,17-VI-1977, Krapovickas & al. fertility.The hybridsobtained with the other 30110 (CTES); 17 km de Paraguarí,camino a species of section Erectoides, such as A. Piribebuy,6-II-1966, Krapovickas& al. 12456 Benthamii, A. gracilis, A. major and A. (CTES, NY, US); 1 km N de ruta2, camino de Oteroi, gave very low values of fertility,less Ipacarai a San Bernardino, 17-VI-1977, than 1.8%, and withA. Hermannii less than Krapovickas& al. 30109 (CTES, US). Paraguarí. 4.3%. Saltos del Chololó,57°2'W, 25°33'S, 14-XI-1978, This taxon could be crossed with A. Arbo& al. 1762(CTES, G, LIL, MO). guaranitica of section Trierectoides (1.1%), with A. Dardani of section Heteranthae Cultivatedmaterial: U.S.A. Florida. Alva. (2%), withA. appressipila (1.3 to 4.5%) and Adventitiousshoots on the end of thepegs, cult. with A. Rigonii (1 to 4.9%), of section

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Fig.14. Arachis paraguariensis ssp. paraguariensis : A, schematicof the plant; B, leaf(K.l 1462).A. paraguariensis ssp.capibarensis : C, leaf(K.30134).

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Procumbentes, with A. repens and A. Pintoi somewhat villous to subglabrous. Leaves (3.5%) of section Caulorrhizae and with A. tetrafoliolate,distichous. In wild material,the glabrata var. glabrata and A. pseudovillosa, fused part of the stipules 9-17 mm long, the tetraploidspecies of section Rhizomatosae. freeparts 10-16 mm long x 1.5 mm wide (in Together with A. stenophylla, it forms a cultivatedmaterial, the apices up to 28 mmx 3 group withinthe section Erectoides, being the mm); petiole 22-42 mm long (cult, up to 60 only ones thatgrow in the southeastextreme of mm), rachis 7 mm long (cult, up to 20 mm); the range, and which have roots withoutthe leafletselliptical, apical pair 25-35 mm long x thickeningsthat are so characteristicof the rest 10-12 mmwide (cult, up to 75 mm x 24 mm), of the section. basal pair 24-31 mm long x 9-10 mm wide The two subspecies of A. paraguariensis, (cult, up to 65 mm x 18 mm). Base of the as well as A. stenophylla,all show a blot at no. stipules somewhat imbricated,with marked 7 in thechromatographic profiles (Krapovickas longitudinal veins, base subglabrous or 1973: 139), which is very infrequentand was somewhat pubescent, with hairs somewhat previouslydetected only in A. Archeri and A. longer on the back, apices subfalcate, acute, pseudovillosa (Seeligmann pers. comm.). glabrous with the margin somewhat ciliate. Petiole subglabrous or with scattered silky 15b. A. paraguariensis subsp. hairs. Leaflets with upper surface smooth, capibarensis Krapov. & W.C. Gregory glabrous, with the margin slightly or not nov. subsp. marked, lower surface villous, midvein and veins markedwith Figs. 1,15b; 14,C secondary marked,margin shorthairs directedtoward the apex. Flowers are clusteredtoward the base of the plant but Herba perennis. Radix profunda, leviter also occur along the branches,in veryshort 4- fusiformis, ramificationibus tenuibus non floweredspikes. Hypanthium4.5-16 cm long, incrassates. Caulis principalis erectus ramis with silky hairs. Calyx with silky hairs decumbentibus. Caules flexuosi, subvillosi. somewhat shorter than those of the Stipulaedorso pilis longisvestitae, quod superest hypanthiumand withsome bristles,upper lobe subglabrum,margine ciliatae. Folíola elliptica, 7-11 mm long, lower lobe falcate, 8-13 mm epiphyllolaevi, glabro, hypophyllo villoso, nervis long. Standard 15-25 mm long x 18-27 mm et marginemanifestis. Hypanthium 4.5-16 cm wide, yellow in the center with an orange longum,villosum. Calyx 7-13 mm longus,pilis margin.Fruiting clustered at the collar of the serieeissetulisque nonnullis vestitus. Vexillum 15- plant. Fruits Particulate, pegs horizontal, 25 mmlongum x 18-27 mmlatum, aurantiacum. articles10-18 mmlong x 5-6 mmwide, epicarp Fructusbiarticulatus paxillo horizontāli,articulis smooth, densely villous when young. 2n=20 10-18mm longis x 5-8 mmlatis, pericarpio laevi. chromosomes (Fernández & Krapovickas 1994). Holotype:BRAZIL. Mato Grossodo Sul. Mun. PortoMurtinho, 43 kmE de PortoMurtinho, cami- Additionalmaterial: BRAZIL. Mato Grosso do no a Jardim(cerca del corregoCapibara) en bancos Sul. Mun. Porto Murtinho:44 km E de Porto de arena,paisaje chaqueño, 12-VI-1968, Hammons, Murtinho,BR-267, corrego Capibara, bosque abier- " Langford,Krapovickas & Hemsy565/566 (CEN). tode Schinopsis,150 m, 29-VI-1977, Krapovickas & Isotypes:CTES, US. al. 30134 (CEN, CTES, US); 64 km E de Porto Murtinho,12-VI-1968, Hammons & al. 570 (CEN, Perennial plant, erect. Taproot deep, CTES, US); 82 kmE de PortoMurtinho, 240 m,29- somewhat fusiform,up to 13 mm in diameter VI-1977,Krapovickas & al. 30139(CIES, US); 83 km at the thickest part, branch roots slender, E de PortoMurtinho, 29-VI-1977, Krapovickas & al. without enlargements. Branches wavy, 30141(CTES); 108km E de PortoMurtinho, 12-VI- internodesshort, up to 70 mmtoward the base, 1968, Hammons& al. 573 (CEN, CTES); Faz. usually covered by the stipules, especially Congonha,P7 (RADAM), 22-X-1980,Pires & al. towardthe apex of thebranches, angular, from 17238(CTES).

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Geographic distribution.This species was green,shiny. Flowers small to medium-sized, collected on the westernfoothills of the Sierra along the lengthof the branches. Hypanthium de Bodoquena, in the southwesternextreme of ca. 4 cm long. Standardorange or yellow,with Mato Grosso do Sul, all along the route from red lines on the back. Fruit subterranean, Jardimto Porto Murtinho.The westernmost Particulate; peg, frequentlywith adventitious collectionswere made at the borderof a small roots, short, almost vertical or some 20 cm area of "chaco" which entersinto Brazil, with long,horizontal; isthmus short; articles smooth, the "cerrado." There, it grows in an open covered with a dense layer of hairs. 2n=20 forestof Schinopsis, in shallow, light sandy chromosomes. soil. The remainderof the collections were made in red soils with rocky outcrops in Geographic distribution.All the species of savannas with an abundance of low palms of this section live in an area that includes the the genus Butia. states of Goiás, Tocantins, the centralpart of Mato Grosso and the northernpart of the III. Sect. Extranervosae Krapov. & W.C. Mining Triangle in Minas Gerais. A few Gregory nov. sect species extend beyond the limits of those Fig. 5 states, such as A. prostrata which was also collected in the west of Bahia and A. Burchellii which also lives in the south of Sect. ExtranervosaeKrapov. & W.C. Gregory, Maranhão, in the southeastof Pará and in the in W.C. Gregory,M.P. Gregory,Krapovickas, vicinityof Teresina,in Piauí. Smith & Yarbrough,in Peanuts- Cultureand It has not been possible to establishthe area Uses: 93, 1973. Krapovickas, Agricultural of origin of A. villosulicarpa, which is only Genetics. Selected Topics: 137, 1973, nomen known as cultivatedby indigenous groups of nudum. west-centralMato Grosso. With the exception of A. villosulicarpa, Perennes. Radices ramificationibus none of the wild species was collected west of tubercularibus,concatenatis. Stipulae marginibus the Paraguay River, which from its source liberis usque ad basem. Folia quadrifoliolata constitutesthe western limit of the section epiphylloviridi, nitido. Vexillumaurantiacum Extranervosae . subtus lineis rubescentibusornatum. Fructus The majorityof the species in this section subterraneus, biarticulatus paxillo fere grow on a very special soil type, frequently horizontāli,paulo profundo,saepe radicante, encounteredin the "cerrado," constitutedby a pericarpiofere laevi. thinlayer of soil over a stonysubstrate.

Typussectionis : Arachis prostrata Benth. Obs. The pollen grains of A. prostrata,A. lutescens (Pire 1968) and A. villosulicarpa Perennial plants. Roots with tuberiform (Pire 1974) are tricolpateand syncolpate,that thickenings, frequently concatenated. is, withthe colpi unitedat the poles, similarto Mainstem erect and branches procumbent. those of the genus Stylosanthes,in contrastto Stipules with the margin free to the base. the rest of the genus Arachis where the colpi Leaves tetrafoliolate,upper surface bright are isolated.

Key fordistinguishing the species

A. Leaflets3-4 times longer than wide, upper surface glabrous, lower surface with bristles on themarginal veins. 16.A. setinervosa

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A'. Leafletsless than 3 timeslonger than wide; bristles, if present, are perpendicular tothe margin, not on the veins. B. Upperleaf surface completely glabrous. C. Leafletstwo or more times longer than wide. Fruit articles 10-15 mm long x 6-7mm wide. 17.A. Macedoi C' Leafletsless than 1.6 times longer than wide. D. Plantserect. Leaflets with thickened margins, ca. 40 mmlong x 25 mmwide. Fruit articles 14-16 mm longx 5-8mm wide. 18.A. marginata D'. Branchesprostrate. Leaflets with the margin unmarked, less than 20 mmlong. Fruit articles less than9 mmlong x 6 mmwide. E. On theprostrate branches, leaflets oblong or elliptical to obovate-ovate (LAV 1.4-1.8:1). F. Fruitpegs with adventitious roots. Leaflets elliptical to obovate-ovate (LAV 1.4-1.5:1). 19.A. prostrata F. Pegswithout adventitious roots. Leaflets oblong to obovate-ovate(LAV 1.6-1.8:1). 20.A. lutescens E1.Leaflets suborbicular (LAV 1.0:1). Fruitpegs with adventitious roots. 21.A retusa B' Upperleaf surface pilose, at leaston theyoung leaves. G. Primarybranches ca. 5 cmtall. Lateral branches prostrate with leaflets up to 13 mmlong x 10 mm wide.Fruit articles less than 11x7 mm. 22. A. Burchellii G' Primarybranches more than 40 cmlong. H. Fruitarticles 8 mmlong x 5 mmwide. On thelateral branches, leaflets from 12-23 mm long x 5-10 mmwide. 23.A Pietrarellii H'. Fruitarticles 16-23 mm long x 8-10mm wide. Leaflets 16-44 mm long x 5-16mm wide. 24. A. villosulicarpa

16. Arachis setinervosa Krapov. & W.C. Holotype:BRAZIL. Mato Grosso.Mun. Barra Gregory nov. sp. do Garças,crest of Serrado Roncador,along new road,127 km NNE ofvillage ofXavantina, 29.9 km S Herbaperennis. Radix palaris, tuberosa.Rami of crossingof CorregoTanguru. Herb carpeton prostrati, ca. 20 cmlongi, brachyblastos gerentes. laterite pebble substrate on a gentle slope Caulis pilis rigidis2 mmlongis villosus. Stipulae otherwisecovered with scrub cerrado. Main herb violaceae, villosae. Folio la oblongo-lanceolata, formingthe carpet. 13-XII-1969, Eiten & Eiten9904 rigida, epiphyllo laevi, glabro, viridi, nitido, (US). Isotypes:NY, K. hypophyllopilis longissparsim vestito, nervo medio et margine prominentibus,saepe setulis Perennial plant. Taproot with tuberiform brevibus,crassis, patentibus margine instructo. enlargements of some 7 mm in diameter. Hypanthium2.5-4 cmlongum, violaceum. Calyx 5- Branches prostrate,ca. 20 cm long. Stem an- 6 mmlongus, villosus, sine setulis.Vexillum 8-10 gular, internodes 10-15 mm long, purplish, mm longum,luteum subtus lineis rubescentibus villous, with hairs 2 mm long, more or less ornatum. rigid and perpendicularto the surface of the

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stem. The main branches with very short burned.If thisoccurs, the rockyhillside is left ramificationsor spurs 10-15 mm long with covered by Arachis setinervosa. internodes almost non-existent and leaves densely clustered. Stipules with the fused Obs. Arachis setinervosa is characterized portion3 mmlong and thefree apices 5-10 mm by the presence of bristles on the marginal long X 1 mm wide at the base. Petiole 5-8 mm veins on the back of the leaflets. When there long; rachis 2-3 mm long. Apical pair of are bristles present in other species of leaflets somewhat larger than the basal pair; Arachis, these are on the margin itself and leafletsoblong-lanceolate, the apical 10-14 mm grow in the same plane as the surfaces of the longX 3 mmwide and thebasal 8-12 mmlong x leaflets,and not perpendicularlyas is the case 2.5 mm wide; leaves somewhat smaller in A. setinervosa. towardthe apex of the branches.Basal partof It is verydifficult to determinethe presence the stipuleswith prominent longitudinal veins, of red lines on the dorsal surface of the stan- purplish,or withthe violet color limitedto the dard in thedry material, but theanalysis of the veins, villous, hairs 2 mm long; free portion flowers of the type specimen leaves little slender,almost acicular, rigid,with scarce long doubt of their existence. By its other hairs, more dense on the margin, the latter characteristics,it approaches several of the usually with violet tinge. Petiole and rachis species of the section Extranervosae. By the purplish,canaliculate, canal narrow,back with form of the leaflets, oblong-lanceolate, it prominentveins, villous, hairs similarto those approaches A. Macedoi, from which it is of the stem and also with short adpressed differentiatedby the spur branches that give hairs. Pulvinus villous. Leaflets rigid, upper A. setinervosaa unique appearance. surfaceglabrous, smooth, brilliant green; lower surfacewith scattered long hairs, midveinand 17. Arachis Macedoi Krapov. & W.C. marginalveins prominent,marginal veins with Gregory nov. sp. shortthick most of thetime bristles, perpendi- Figs. 1,17; 15,C-E cular to the underside. Inflorescences few- flowered,axillary, arranged along thelength of the branches.Flowers withbasal bracts 5 mm A. Macedoi Krapov.& W.C. Gregory,in W.C. long,villous. Hypanthium25-40 mmlong, lilac Gregoiy,M.P. Gregory,Krapovickas, Smith & colored, villous with long sparse hairs which Yarbrough,in Peanuts- Cultureand Uses: 120, retaingrains of sand. Calyx bilabiate, 5-6 mm 1973, nomennudum. Refers to specimen GKP long,villous, without bristles, lobes unitedat the 10127. base 1.5-2 mm. Standard 8-10 mm long, yellow, withred lines on the back surface. Herbaperennis. Radix polaris ramificationibus tubercularibus.Caules principaliserectus ramis Additionalmaterial: BRAZIL. Mato Grosso. procumbentibus.Gaulis pilis longis sericeis, Mim. Agua Boa: 26,5 kmN do acesso principala sparsimvestitus. Stipulae plerumque glabrae, pili Agua Boa ao longoda BR-158,1 17 kmN de Nova longiin dorso interdumadsunt. Folíola ovaliter- Xavantina,12°16'S, 360 m,Valls & al. 12516 lanceolata epiphyllolaevi, viridi,nitido, glabro, (CEN,CIES). hypophylloglabro vel pilis brevibus,adpressis, sparsimvestito, nervo medio prominente longos Geographie distribution. This species is pilos caducos gerente, margine ciliato paulo known only fromthe high parts of the Serra manifesto.Hypanthium 2-6 cm longum,villosum. do Roncador, the vegetation of which was Calyx5-7 mm longus pilis ca. 2 mmlongis vestitus. studiedby Eiten (1975), supplyinginteresting Vexillum 7-9 mm longum x 8-10 mm latum, details on the habitat of this species (page aurantiacum,dorso lineis rubescentibus ornatum. 120). Accordingto Vails (12516), it occurs in Fructus biarticulatuspaxillo horizontāliusque sandysoil overa layerof lateriterocks, under a ad 20 cm longo, radices adventitiasemittenti, vegetationdominated by Diectomisfastigiata isthmo6 mmlongo, articulis 10-15 mm longis x 6- (Swartz) Kunth,an annual grass thatis easily 7 mmlatis, apice recurvo,pericarpio laevi.

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Holotype:BRAZIL. MinasGerais. Capinópolis, wide, pericarp smooth and with beak. 2n=20 FazendaSanta Terezinha, margen inundable de pe- chromosomes(Conagin 1964, fig. 1, F, V.776, queña laguna, suelo pedregoso, 4-IV-1961, GKP 10127). Gregory,Krapovickas & Pietrarelli10127 (LIL). Isotypes:CEN, CTES, G, GH,MO, NY, SP,US. Selectedadditional material: BRAZIL. Minas Gerais. Mun. Ituiutaba:Fazenda do Poço, 30-XI- Perennialplant, without rhizomes. Taproot 1944, Macedo 598 (RB 55357, US); Faz. Santa withtuberiform enlargements, also presenton Terezinha,beira da lagoa, lugareshúmidos, 9-II- the lateralbranch roots. Mainstem erect, up to 1949, Macedo 1636 (SP, US); id., 1671 (SP); 45 cm tall, lateralbranches procumbent, some Capinopolis,Faz. Santa Terezinha,22-XI-1971, 50 cm long; stems purple, cylindricaltoward Macedo 5059 (MBM); id., 23-11-1984,Valls & al. the base, quadrangulartoward the apex, with 7533(CEN, CTES). Goiás.Km 3 1da Rod.Itumbiara- scatteredlong silky hairs; internodes3-6 cm RioVerde, 19-IV-1978,Tamashiro&al. 7413 (UEQ. long. Leaves tetrafoliolate.On the mainstem thefused portion of thestipules 1 1 mmlong x 8 Relatedmaterial: BRAZIL. Maranhão.BR-230, mmwide, thefree parts acute, 18 mmlong x 2 12 kmde Carolina,14-XI-1979, Martins & al. 7415 mmwide; petiole some 40 mmlong; rachis 10 (CIES); 11,8 km E de Carolina,13-111-1982, Valls & mm long; the apical pair of leafletsup to 40 al. 6609 (CEN, CTES); 11 kmE de Carolina,13-111- mm long x 10 mmwide, the basal pair 35 mm 1982,Valls & al. 6610 (CEN, CTES); 5 kmE de long x 9 mm wide. Leaves of the lateral Carolina,15-III-1982, Valls & al. 6612(CEN, CTES): branchessmaller; basal portionof the stipules 38 kmE deCarolina, BR-230, 18-111-1985, Valls & al. 5 mm long x 4 mmwide, thefree parts acute, 8 8383 (CEN, CTES). Tocantins. 16 km W de mm long; petiole 10 mm long; rachis 4 mm Filadelfia,na estradapara Araguaina, 13-111-1982, long; theapical pair of leaflets13 mmlong x 5 Valls& al. 6626 (CEN, CTES). Goiás. 20 kmNE de mmwide, basal pair 12 mmlong x 5 mmwide. CamposBelos na estradapara Aurora do Norte,12- Stipules with glabrous surfaces, occasional IV-1986,Valls & al.9952 (CEN, CIES); 13km NW de long hairs on the back and towardthe base of NovaRoma, camino a Teresinade Goiás,600 m,3-11- the fused portion. Petiole and rachis from 1990,Arbo & al. 3556(CTES). villousto glabrescent;pulvinus villous. Leaflets oval-lanceolate;upper surfaceglabrous, shiny Common name, "amendoim do resfriado" green; lower surface glabrous or with very (Macedo 598). sparse, diminutive,adpressed hairs, midvein prominentwith long, sparse, caducous hairs; Geographic distribution. This species margin somewhat marked on the lower grows in small temporaryponds in rockysoil, surface, with long sparse cilia. Spikes 3- in the upper part of the rocky hills in the flowered,with very short axis, disposed all northernpart of Triangulo Mineiro (MG). It along the lateral branches. Flowers with two was also collected in the vicinity of the basal bracts, the lower one acute and entire, Paranaiba River,in the southof Goiás. theupper one bifid,both with long hairson the marginand along the midvein.Hypanthium 4 Obs. 1. In the more northerncollections cm long (2-6 cm), villous. Calyx bilabiate, made near Carolina, in the state of Maranhão, covered with long silky hairs ca. 2 mm long; and at Filadelfia,in Tocantins,the materialis thebroad lobe some 5 mmlong, slightly divided verysimilar to A. Macedoi, so we tentatively at the apex; the narrow lobe subfalcate, 6-7 place theseunder this name. It is possible they mm long. Standard 7-9 mm long x 8-10 mm pertainto anothertaxon, but the available ma- wide, orange, in dry state lilac-violet in the terial does not allow us to arrive at a more center,with red lines on the back side; wings definitiveconclusion. ca. 5 mmlong. FruitParticulate; peg horizontal, up to 20 cm long, frequently with Obs. 2. The specimen Macedo 598 (RB) adventitious roots; isthmus 0.6 cm long; consists of two plants,one verysimilar to the articles rounded, 10-15 mm long x 6-7 mm type and the other with wider leaflets (the

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Fig.15. Arachis marginata : A, schematicofthe plant; B, leaf,lower side (G. 10406). A. Macedoi : C, schematicofthe plant;D, leaffrom mainstem; E, leaf from lateral branch (G. 10127).

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apical pair up to 32 mmx 18 mm,the basal pair long x 5-8 mmwide, withsmooth pericarp. up to 27 mmx 15 mm). Holotype:BRAZIL. Goiás [Tocantins].Sandy We dedicate this species to Prof. Amaro hillMission of Duro. oct. 1839, Gardner 3 103(BM!). Macedo of Ituiutaba, Minas Gerais, who Isotype:P!. collected it forthe firsttime. Additionalmaterial: BRAZIL. Tocantins. 30 km E 18. Arachis marginata Gardner de Dianópolis (presentname of the Mission of Figs. 1,18; 15,A-B Duro),22-IV-1961, Gregoiy & al. 10406(CIES, LIL). Mun. Dianópolis:Fazenda Agua Limpa 11°37'S, 46°36'W,17-111-1982, Valls & al. 6649(CEN, CIES); Gardner,in Hooker,Icon. PI. 5, tab. 500, 1842, id.,4-V-1982, Valls & al. 6759 (CEN, CTES). Mun. "Herb.Bras. n. 3 103. Rarein upland sandy Campos PonteAlta do BomJesus: 54 kmS de Dianópolise near the mission of Duro, provinceof Goyaz, 14 km leste do entroncamentopara Taguatinga, Brazil." estradaDianópolis-Barreiras, 17-III-1982, Valls & al.6652 (CEN, CTES). Erect perennial. Main taproot with numerous branches bearing concatenate Geographiedistribution. This species is from tuberiformenlargements 10-25 mmlong x 5-7 a very restricted area in the vicinity of mm wide, rugose and more or less striated. Dianópolis (Misión de Duro), whereit grows in Stems erect, up to 10 cm long, unbranched, the "cerrado" with sparse herbaceous borne fromthe collar of the plant; internodes vegetation,in whiteand reddishsandy soils. 12-30 mm long, 4-angled, villous, with caducous hairs. Leaves tetrafoliolate. The Obs. At present, Arachis marginata is fused portion of the stipules 7 mm long x 5 known only fromthe type locality and areas mm wide, the freepoints acute, up to 13 mm verynearby. Nevertheless, its name has been long. Petiole very short in the basal leaves, applied to differentspecies which also have some 2 mm long, and up to 30 mm long in the much-thickenedleaflet margins, as in the case apical leaves. Rachis 7-12 mm long. Leaflets of A. Burkartii and of A. Oteroi (Bentham oblong or obovate, theapical pair up to 42 mm 1859: 87). These two species have red lines on long x 27 mm wide, the basal pair up to 34 the frontsurface of the standard,the firsthas mm long x 23 mm wide. Stipules villous on rhizomes and neither has roots with the the fused portion,almost glabrous and with tuberiform concatenate thickenings so prominentveins on the freetips; marginwith characteristicof A. marginata. long dense silky hairs. Petiole and rachis canaliculate, villous on the back. Upper 19. Arachis prostrata Benth. surface of the leaflets glabrous, brightgreen, Figs. 1,19; 16,A-C smooth and with a line of long hairs on the midvein; the lower surface with prominently marked veins, forminga network, covered Bentham,Trans. Linn. Soc. London18(2): 159, with long scattered,adpressed hairs and with 1841, "Ad Trahirasprovinci ae Goyaz Brasiliae. " longer, rigid and more dense hairs along the Pohl (v.s. cumfl. comm.a Mus. Caes. Reg. Vind.). midvein; margin stronglythickened on the Bentham,Martius, Fl. bras.15(1): 87, 1859. lower surface of the leaflets,with long dense silky hairs and some short bristles. Perennialplant. Main root and its branches Inflorescences 3 -flowered, axis very short, withtuberiform enlargements from 9 mm long coveredby the stipules,occurring on thebasal x 5 mmwide, up to 35 mm long x 7 mm wide. nodes of the stems. Hypanthium4 cm long, Primary stem erect, 5-12 cm long, covered villous. Standard completely yellow, with almostentirely by thestipules, internodes short, reddish lines slightly marked on the back. ca. 5 mm long, villous. Lateral branches Fruit subterranean,with articles 14-16 mm prostrate,up to 90 cm long, stems villous

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Fig.16. Arachis prostrata : A, schematic ofthe plant; B, leaffrom lateral branch; C, leaffrom mainstem (G. 10240). A. Pietrarellii:D, schematicofthe plant; E, leaffrom lateral branch; F, leaffrom mainstem (G.9923). toward the apex of the branches and primary branches, stipules with the fused glabrescenttoward the base, internodes10-30 portionsome 8 mm long, the free parts 10-12 cm long. Leaves tetrafoliolate.On the erect mm long x 2 mm wide at the base. Petiole 10-

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25 mm long. Leaflets oblong, at times Niquelandiaa Maranhão,12-IV-1961, Gregoiy&al. somewhatobovate, the apical pair up to 20 mm 10234(BAA, CTES, GH,LIL, MO, NY,US); 3 kmS long X 12 mm wide, the basal pair somewhat del rio Maranhão,camino Niquelandia-Anapolis, smaller,up to 19 mmlong x 1 1 mmwide; rachis 12-IV-1961,Gregory & al. 10240(CTES, GH,LIL, 5-10 mm long. On the prostrate lateral MBM,MO, NY, US); 75 kmN de Corumbáde Goiás, branches,the fused portionof the stipules 3-4 on roadto Niquelandiain valleyof rio Maranhão, mm long, the freeportion 5-9 mm long x 2-2.5 700m, 23-1-1968, Irwin & al. 19055(NY, RB, UB, US); mmwide at thebase, becomingabruptly acute; id.,25-1-1968, Irwin & al. 19196(K, NY, RB, UB, US); petiole 3-5 mm long; leaflets obovate or 14km S deNiquelandia, 21-1-1972, Irwin & al. 34701 elliptical,the apical pair usually 9 mmlong x 7 (K, NY, UB); Serrado rioPreto, 16°S, 47°W, ca. 8 km mm wide, up to 13 mm x 9 mm,the basal pair E de Cabeceiras,1000 m, 18-XI-1965,Irwin & al. being somewhatsmaller, up to 12 mmlong x 8 10469 (NY, RB, UB, US); 86 kmNE de Formosa, mm wide. Stipules villous towardthe base and caminoBrasilia-Barreiras, 19-IV-1961, Gregory &al. usually with the surfaces of the apices 10292 (CTES, LIL, US); 115 kmNE de Formosa, subglabrousand themargin villous, hairs 1.5-2 caminoBrasilia-Barreiras, 25-IV-1961, Gregory & al. mm long; petiole and rachis withvillous back 10449(CTES, G, GH, LIL, MO NY,US); km194, rod. and ventral surface canaliculate, glabrous. Belem-Brasilia(mun. Pirenópolis), 30-1-1976, Upper surfaceof leafletscompletely glabrous, Hatschbach& al. 38197 (CTES, MBM); Rod. BR- shiny, smooth; lower surface densely villous 020, rio Macacos (mun.Alvorada do Norte),9-1- withlong, more or less adpressedhairs; margin 1977,Hatschbach 39383 (CTES, MBM). Mun.Flo- somewhat markedon the back, longly ciliate, resde Goiás: 4 kmN dorio Extrema, BR-020, 14°56'S, rarely with short bristles. Flowers along the 46°59'W,27-III-1981, Valls & al. 6284(CEN, CTES); length of the prostrate branches, in short id.,21 -III- 1982, Valls & al. 6667 (CEN, CTES, NY, axillary spikes. Hypanthium2.5-4 mm long, US). Mun.Campinorte: 5 km N de Campinorte,BR- villous. Calyx villous withoutbristles; lower 153,km 273,2, 14°75'S, 49°9W, 3-IÏÏ-1982, Valls & al. lobe subfalcate, 5 mm long, upper lobe 4 mm 6475 (CEN, CTES, LIL, NY, US). Mun.Porangatu: long. Standard 8 mm long x 9-10 mm wide, BR-153,km 384, corrego Funil, 3-ffl-1982, Vails & al. orange with red-violet lines on the dorsal 6485 (CEN, CTES, G, MO, NY, SI, US). Mun. surface; wings yellow, 5 mm long. Fruit Alvoradado Norte:14°33'S, 46°33'W, 21-III-1982, subterranean,Particulate; peg 1-5 cm long, Vails& al.6662 (CEN, CTES, MO, NY, US); id.,9-V- aerial portion violet and glabrous, densely 1982,Vails & al. 6788(CEN, CTES). Mun.Formosa: pubescent below ground, and with abundant 17km N da vilaJK, BR-020, 15°4'S, 47°5'W, 29-III- adventitious roots which present tuberiform 1983,Vails & al. 7004 (CEN, CTES); 8 kmW of enlargements; isthmus very short; articles MonteAlegre, 600-700 m, 1 1-III-1973, Anderson & independent,rarely united forming a single 2- al. 6847 (UB). Mun.Campos Belos: 13,5km NW de seeded cavity, 5-8 mm long x 4-5 mm wide, Campos Belos na estradaa Arraias,2-XI-1984, with a short beak; pericarp smooth, densely Allem & al. 3070 (CEN, CTES); 84 km NE de covered with a layer of small hairs. Seed 4-6 Formosa,BR-020, 2-II-1990, Albo & al. 3485(CTES, mm long x 3-4 mm wide. 2n=20 chromosomes HRCB);3 kmSW de Posse,GO-446, 3-II-1990, Arbo (Conagin 1963, V.780, GKP 10234). & al. 3515 (CTES, HRCB);9 kmNE de Iaciara,camino a Posse, 3-II-1990,Arbo & al. 3537 (CTES, Holotype:BRAZIL. Goiás. Trahiras [Niquelandia], HRCB); 4 kmSW de NovaRoma, camino a Teresina Pohl1836 (K!). de Goiás,3-H-1990, Albo & al. 3555(CTES, HRCB); 5 kmW de Colinas,camino a Niquelandia,4-II- 1 990, Selectedadditional material: BRAZIL. Bahia. Arbo& al. 3665,3669 (CTES, HRCB); 52 kma leste Mun. Riachãodas Neves: 12 kmN de Riachãodas de São Miguel do Araguaia na estrada para Neves,11°39'S, 44°55'W, l-IV-1983, Vails & al. 7058 Porangatu,28-VIII-1984, Valls & al. 7887 (CEN, (CEN, CTES, MO, US); 3 kmS do entroncamentoCTES); 5,2kmN de Campinone,BR-153, 30-VIII- para SantaRita, na estradade Barreirasa Corrente, 1984,Valls & al. 7895(CEN, CTES); RioTraíras, 10 11°21'S,44°56'W, l-IV-1983, Valls & al. 7068(CEN, kmW deNiquelandia, 30-VHI-1984, Valls&al. 7897 CTES). Goiás. 10 km S del rioTraíras, camino de (CEN,CTES); 5 kmN deCampinone, BR-153, 14-IH-

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1985, Vails & al. 8304 (CEN, CTES). Mun. In A. villosulicarpa and in A. Pietrarellii Porangatú:corrego Funil, BR-153, km 83 1, 14-III- we have also observed adventitiousroots on 1985,Vails & al. 8308(CEN, CTES); 11,5 km NE de the pegs, but not withthe frequencyin which Vale Verdee cercade 18km a SW de São Raimundo, theyare presentin A. prostrata. 19-111-1985,Valls & al. 8418(CEN, CTES); 4 kmSW The species closest to A. prostrata is A. de Acreuna,BR-060, 2-IV-1986, Valls & al. 9858 lutescens, fromwhich is differentiatedby the (CEN, CTES); CachoeiraGrande, 27-Ī-1968, Onishi absence, in the latter,of roots on the pegs, as & al. s/n(NY, SI, UB). Piauí. Mun.Altos: 18 km NE we were able to confirm in the numerous deAltos, BR-343, 23-111-1985, Valls & al. 8481(CEN, collectionsthat we wouldmake. Furthermore,A. CTES); id., Valls & al. 8482 (CEN, CTES). Mun. lutescensprefers gravelly soil and occurs to the CampoMaior: 10 kmNE de CampoMaior, 23-III- west and southof the rangeof A. prostrata. 1985,Valls & al. 8486(CEN, CTES). Mun.Piripiri: 3 kmSW doRio dos Matos, BR-343, 30-IV-1987, Valls Obs. 2. The material from Faz. Piçarrão, & al. 11028 (CEN, CTES). Tocantins.26 kmSE dorio Araguari (MG) (Werneck 3, Valls 5913 & SantaTeresa, estrada Gurupi-Peixe, 12°S, 48°37'W, 7539) has an appearance very similar to A. 5-III-1982,Valls & al. 6522 (CEN, CTES, NY, US). prostrata in the size and the shape of the Mun. PortoNacional: barrancado rio Tocantins, leaves, but has a pilose upper leaf surfacelike 16-III-1982,Valls & al. 6638(CEN, CTES, NY, US); A. Burchelliiand the pegs have no adventitious id., Coradin3736 (CEN). Mun. PonteAlta do B. roots, as in A. lutescens. This material was Jesus:fazenda Morro Branco, a 6 kmde estradade collectedin theextreme southeast of therange of Dianópolisa Taguatinga,na estradapara fazenda section Extranervosae and could correspond Agua Limpa,17-III-1982, Valls & al. 6648 (CEN, to anotherentity, but it would be advisable to CTES, NY, US); Natividade,open sandyplaces, studynew populationsto decide on theposition dec.1 839, Gardner 3104 (G, K, P); Montevidiu,5-10 km of thismaterial. na rodoviapara Palmeirópolis, 23-1-1992, Hatschbach & al. 56351 (CTES, MBM). Minas Gerais. Mun. 20. Arachis lutescens Krapov. & Rigoni 2 kmS de Cachoeira 11 -VIII- Capinopolis: Dourada, Fig. 1,20 1984,Valls & al.7725 (CEN, CTES).

Relatedmaterial: BRAZIL. Minas Gerais.Mun. Krapovickas& Rigoni,Darwiniana 11(3): 452- Araguari:Fazenda Piçarrão, declive acima da area 454, lám.2, abajo,1957. de inundaçaodo ribeirãoPiçarrão, Werneck 3 (CEN, CTES); id.,25-11-1981, Valls&al. 5913(CEN, CIES); Perennial plant. Main root and branches id.,24-11-1984, Valls & al. 7539(CEN, CTES). withelongated tuberiform enlargements, from 10 mmlong x 5 mmwide up to 80 mmlong x 10 Geographie distribution.Arachis prostrata mm wide. Aerial part covered with long soft is a species thatgrows in easternGoiás, in the hairs, except the upper side of the leaflets, south east of Tocantins,in the northof Bahia, which is glabrous. Mainstem erect, 3-10 cm in Piauí, and in the north of the Triangulo long with leaves larger than those of the Mineiro, in compact clayey soils subject to branches.Lateral branchesprocumbent, up to flooding, frequently with gravel, and 60 cm long; internodes1-2.5 cm long, villous. preferentiallynear streamsof water. Leaves tetrafoliolate, rarely with a supernumeraryapical leaflet. On the erect Obs. 1. We collectedA. prostrata near the primaryaxes, stipules with the fused portion type locality where, apparently, only this from6-8 mm long, the free parts 10-16 mm species ofArachis grows. The materialcoinci- long x 2 mm wide at the base, abruptly des verywell withthe holotype and in thefield narrowing; petiole 15-45 mm long; leaflets we were able to confirm the presence of fromoblong to subovate,the apical pair usually abundant adventitiousroots on the pegs. We 15 mmlong x 8 mmwide but up to 24 mmx 13 also observed this detail in all of the other mm,the basal pair somewhatsmaller, up to 23 collectionswe made of this species. mmlong x 10 mmwide; rachis5-7 mmlong but

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up to 13 mm. On the lateral branches, the al. 10176(CEN, CTES, LIL, MO, NY, US); 15km S de fused portionof the stipules 3 mm long, the Ceres,16-IV-1961, Gregory & al. 10258(LIL); id., free portion 7 mm long x 2 mm wide at the Gregory& al. 10259(CEN, CTES, GH, LIL, MO, NY, base, abruptlypointed; petiole 2-6 mm long; US); 200km S ofCaiaponia, on road to Jataí, Serra de from leaflets oblong to obovate, apical pair Caiapó,17°12'S, 51°47W, 31-X-1964, Irwin&al. 7605 usually 8 mm long x 6 mm wide but up to 18 (NY,US). Mun.Jaraguá: km 1 1 1, BR-153, as margins mmx 1 1 mm,the basal pair somewhatsmaller, dorio Saraiva, 15°32'S, 49°27W, Coradin 3466 (CEN); up to 16 mm x 9 mm; rachis 2-5 mm long. id,2-m-1982, Valls & al.6466 (CEN, CTES, US). Mun. Stipules villous towardthe base, the surfaceof Morrinhos:100 m lestedo rio Piracanjuba,entre theapices usuallyglabrous with villous margin; Morrinhose Caldas Novas, 17°43'S, 48°50'W, 23-11- petiole and rachis villous on the back, 1981, Valls & al. 5925 (CEN, CTES); 3 km de canaliculate ventralsurface, glabrous; leaflets Israelandia,rio Claro, 15-VM-1984, Valls & al. 7732 with the upper surface completelyglabrous, (CEN, CTES); 11 kmN de BomJardim de Goiás,16- shining, smooth, the lower surface densely Vm-1984,Valls & al.774 1 (CEN, CTES); id, Valls & al. villous, with hairs ca. 2 mm long, somewhat 7742 (CEN, CTES). Mun.Aragarças: 5 kmE do rio adpressed, and withthe marginprominent on Araguaia,estrada para Goiás Velho,18-VIII-1984, the lower surface, longly citiate. Valls& al.7743 (CEN, CIES). MatoGrosso. Cuiabá, I- Inflorescences along the length of the 1927,Riedel 778 (K); Cuiabá,Riedel 489 (K); Cuiabá, branches,short axillary spikes, ca. 5 mmlong, nov.-dec.1844, Weddell 2922 (K, P); cerradosal N de 4-flowered. Flowers sessile in the axil of a Cuiabá,Krug 17 (SP); VarzeaGrande, Cuiabá, 14-IH- bract and a bifidbractlet. Hypanthium 4-6 cm 1948,Addor (RB). Mun. Varzea Grande: BR-364, 30 km long,villous. Calyx bilabiate,villous; lower lobe NofCuiabá, 5-H-1978, Pedersen 12177 (C, CTES, Herb. subfalcate,ca. 6 mmlong; upper lobe ca. 5 mm Pedersen);Guía, próximo a Cuiabá, 24-111-1982, Prado long. Standard suborbicular, 8-12 mm in &al. 306(UEC);MT-060,17kmSdeBR-070caminoa diameter,orange, with red lines on the dorsal Poconé,20-1-1989, Krapovickas & al. 43052(CTES); surface converging toward the base. Fruit CorregoRibeirão, Cuiabá, 14-111-1948, Addor (RB); subterranean,biarticulate; peg 2.5-5 cm long, 100km SE de Cuiabá,9-III-1959, Gregory & al. 9898 reddish on the aerial portion and densely (CTES,LIL, SI, SP); id., 9900 (LDL, SI, SP); 70 km SE de pubescentbelow ground,without adventitious Cuiabá,camino a Rondonópolis,9-III-1959, Gregory & roots; articles6-9 mmlong x 5-6 mmwide, the al. 9903(LIL); 5 kmE de Coxipóda Ponte(13 kmEde apical article somewhat larger, with beak, Cuiabá),9-IH-1959, Gregory & al. 9905 (LIL, SI, SP); 10 pericarp smooth,densely pubescent; isthmus kmN deCuiabá, 1 1-IÜ-1959, Gregory & al. 9906(LIL, short,1-3 mm long. Seed 5 mmlong x 3.5 mm SI); 37 kmN de Cuiabá,ayo. Mirim,ll-m-1959, wide. 2n=20 chromosomes (Conagin 1963, Gregory& al. 9907 (CTES, LIL, SI, SP); 50 kmN de V.779, GKP 10176). Cuiabá,1 1-IÜ-1959, Gregory & al. 9909 (LIL, SI); 17km NE deRosario Oeste, 12-IH-1959, Gregory & al. 9920 Holotype:BRAZIL. Mato Grosso.10 kmeast of (LIL,SI); 2 kmNW de Cuiabá,13-HI-1959, Gregory & Cuiabá."In shale.Species with tuberous elongated al. 9927(LIL, SI). Mun.Cuiabá: BR-364 near Corrego swellingson roots,"15-III-1948, J.L. Stephens SH Pindaival,350 m, 1 1-II-1975, Anderson 11339 (MBM); 255 (LIL 397456!).Isotype: NA!. St.Antonio de Leverger, 15-25 km S de Cuiabá,300 m, 12-11-1975,Anderson 11352 (MBM); 35 km E de Selectedadditional material: BRAZIL. Goiás. 23 Cuiabá, BR-364, 13-XI-1975,Hatschbach 37492 km E de Caiaponia on road to Montevideu,4-II- (CTES,MBM); 15km W de Cuiabá,camino a Poconé, 1959, Irwin 2570 (NY, US); 30 kmN deJataí, on road 18-m-1976,Krapovickas&al.30028 (CEN, CTES, GH, to Caiaponia,24-X-1964, Irwin & al. 7281 (K p.p.); MO,NY, US); FazendaSangradouro, 22 kmW de Sete entrePiranhas e Bom Jardim,6-IV-1958, A. Lima Porcos,57°15*W, 16°S, 16-XH-1976, Krapovickas & al. 3018 (IPA, K); rioPiracanjuba, entre Morrinhos y 30032(CEN, CTES, G,GH MO,NY, US); 106kmEde CaldasNovas, 6-IV-1961, Gregory & al. 10160(CEN, Cáceres,camino a Cuiabá,57°8'W, 15°56'S, 17-XII- CTES, GH,LIL, MO, NY, US); rioPiracanjuba, entre 1976,Krapovickas & al. 30038(CEN, CTES, GH, MO, CaldasNovas y Piracanjuba, 7-IV-1961, Gregoiy & NY, US); Barradas Garças,13-1-1968, Philcox & al. al. 10174 (CEN, CTES, LIL, NY, US); id.,Gregory & 4013(K, NY, UB); Xavantina,52°14'W, 14°38'S, 30-

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Xn-1967,Philcox & al. 3762(K, NY, UB); 17km N of margine pilis 1 mm longis ciliatae. Folíola Aragarças,road to Xavantina, 23-XH-1967, Philcox & suborbicularia,apice retuso,epiphyllo laeve, gla- al. 3718(K, NY, UB); 13km WSW ofthe Cuiabá river bro, hypophyllo margine et vena media onroad from Cuiabá to Cáceres, 22-VIH-1981, Valls & prominentibus,glabro, pilis paucis 1 mmlongis in al.6332 (CEN, CIES); 126km WSWof Cuiabá, on road nervomedio, margine ciliis 1 mmlongis et setulis toCáceres, 22 km W ofSete Porcos, 22-VI1I-1981. Valls viridibusnonnullis. Hypanthium 45 mmlongum &al. 6338(CEN);Cáceres airport, 31-VÜI-1981, Valls& aliquantum villosum. Calyx 5-6 mm longus, al. 641 1 (CEN,CTES); 2 1km N ofCáceres, on road to villosulus, setulis destitutus.Vexillum 12 mm Barrados Bugres, 31-VDI-1981, Valls&al. 6412 (CEN, longumaurantiacum subtus lineis rubescentibus CTES); 71 kmN de Cáceres,on roadto Barrados ornatum.Fructus biarticulatus, paxillo radices Bugres,3l-Vm-1981, Valls&al. 6414 (CEN); id., Valls adventitias emitente,articulis 8-10 mmlongis x 5- &al. 6415(CEN, CTES); 45 kmS deRosario Oeste (15 6 mmlatis, pericarpio laevi. kmNď Guia),2-IX-1981, Valls& al. 6441 (CEN, CTES, NY, US). Mun.Santo Antonio de Leverger: 8 km W de Holotype:BRAZIL. Goiás. Mun. Terezina de Palmeiras,30-1-1989, Krapovickas & al. 43 138 (CTES). Goiás,margem direita da estradade Terezinapara Mun.Poconé: BR-070, km 63 1, 17-V-1985, Valls & al. CamposBelos, logo após a pontesobre o rio São 8740(CEN, CTES). Mun.Barra dos Bugres: Jauquara, João,flores alaranjadas com estriasno verso do l-VI-1985,Valls&al. 8976 (CEN, CIES); PortoEstrela, estandarte,720 m,13°45'S, 47°14'W, Valls, Pittman 1-VI-1985,Valls & al. 8978(CEN, CTES); 73 kmS de & Silva12883 (CEN). Isotype:CTES. Barrados Bugres, camino a Cáceres,l-VI-1985, Valls & al.8982 (CEN, CTES); 56km E deBarra dos Bugres, Prostrate herb. The main root and its 2-VI-1985,Valls&al. 8989 (CEN, CIES); id,Valls&al. branches with cylindrical tuberiform 8995(CEN, CTES); 32kmE de Cuiabá, BR-364, 3-VI- enlargements.Stem square, villous, withhairs 1985,Valls & al. 9013 (CEN, CTES); 10 kmNW de ca. 1 mmlong. Stipuleswith the fused part 3-4 Poconé,24-1-1985, Valls & al. 9348(CEN, CTES); id., mm long, freepart 7-9 mm long x 2-2.5 mm 9349(CEN, CTES). Mun. N. Sia. doLivramento: 4-XI- wide, the back of fused part villous, the 1986,Valls & al. 10473(CEN, CTES). MinasGerais. surfaces somewhat villous toward the base, Uberaba,Regnell III-414 (K). the rest glabrous or subglabrous, with a few long hairs especially on the veins almost up to Geographiedistribution. This species grows the tip, marginsciliate, with hairs 1 mm long. south of 14°30'S in the state of Mato Grosso Leaves tetrafoliolate;petiole 5-8 mm long; and in thewestern part of the stateof Goiás. It rachis 5-6 mm long, both canaliculate, with was collected in Uberaba, in the Triangulo hairs on the back, the canals glabrous,that of Mineiro.The Paraguay River,in thevicinity of the petiole separated from the canal of the Cáceres (MT), constitutesthe western limit of rachis by a transverseline of hairs. Apical the species. The majorityof the plants which leaflets12-22 mmlong x 12-20 mmwide, basal we collected grew in gravelly soil or in soils leafletsca. 1 1-19 mm long x 11-16 mm wide, consistingof a thinlayer of earththat covered suborbicular, the apex retuse with an a substrate of angular gravel. A few indentation of 1-1.5 mm, upper surface populationsgrew in sandysoils (Gregory& al. smooth,lower surfacewith the margin and the 9898, 10176 and 10259). Almost all ofthe midvein prominent, secondary veins collections were made in flooded places or somewhat marked, both surfaces glabrous, places subject to flooding. with only scatteredhairs 1 mm long on the midvein;the marginwith cilia 1 mm long and 21. Arachis retusa Krapov., W.C. some short bristles. Hypanthium45-55 mm Gregory & Vails nov. sp. long,somewhat villous. Calyx bilabiate,upper lobe 5 mm lowerlobe 6 mm Figs. 1,21; 24,A long, falcate, long, somewhatvillous, without bristles. Standard 12 mm long x 14 mm wide, orange, with reddish Herba prostrata.Caulis pilis ca. 1 mmlongis lines on the outside surface. Fruit villosus.Stipulae glabrae, dorso et base villosae, subterranean,the pegs oftenhave adventitious

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roots,articles 8-10 mm long x 5-6 mm wide, cerradao,14-111-1968, Irwin, Maxwell & Wasshausen pericarpsmooth. Seeds 6 mmlong x 4 mmwide. 21163 (RB). Isotypes:K, LIL, NY, UB, US.

Additionalmaterial: BRAZIL. Goiás. 4 km da Perennial plant. Root with tuberiform Estradade Teresinade Goiás a MonteAlegre de enlargements 20 mm long x 6 mm thick. Goiás. Erva rastejante.Flores amarelas formando Primarystem erect, 3-5 cm long,with the stem relvadoem localpertubado do mataciliar do rioSão covered by the stipules; lateral branches João,l-XII-1991, Pereira & al. 1948(CIES, IBGE);2 km prostrate,0.5-1 m long,internodes usually 2-3 ao nortede Teresinae 34 kmao suldo rioParanã, ao cm but up to 4 cm long, angled, reddish, longoda estradapara Monte Alegre, 590 m, 13°34'S, villous, with short adpressed hairs and long 47°13'W,14-TV-1986, Valls & al. 9950(CEN, CIES). hairs 1.5-2 mm long, more dense toward the Mun.Teresina de Goiás: 13°45,S,47014'W, Valls & al. nodes. Leaves tetrafoliolate.On the primary 12883(CEN). Tocantins.Mun. Paranã: Parana, 270 m, branches,the fused portionof the stipules 6-8 12°37'S,47°53 W, Valls & al. 12939(CEN, CTES); 46 km mm long, the free portion 9-10 mm long; SE de Gurupi,a 12km NW do rioTeresa, Fa z. Taboca, petiole 8-14 mm long; rachis 3-6 mm long; naestrada Gurupi-Peixe, 290 m, 1 1°55'S, 48°44W, 5-EH- leafletsoblong, the basal 16-19 mmlong x 6-8 1982,Valls & al.6517 (CEN, CTES). mmwide, theapical 18-20 mm long x 8-9 mm wide. On the lateral branches, the fused Geographicdistribution. This species grows portionof the stipules 3-5 mm long, the free in the extremenortheast of the state of Goiás portion 7-11 mm long; petiole 3-6 (8) mm; and in the south of the state of Tocantins rachis 3-4 (5) mm long; leafletsusually ovate, (Brazil). occasionally oblong and rarelyobovate, basal leaflets 8-12 mm long x 4-8 mm wide, the 22. Arachis Burchellii Krapov. & W.C. apical pair 10-13 mm long x 5-10 mm wide. Gregory nov. sp. Fused part of the stipules with long hairs on Fig. 1,22 the back, the surfaces subglabrous,with few adpressed hairs, to villous; the free portion triangular,acute, withprominent veins toward Herba perennis. Radieis ramificationes the base and with a single vein toward the tuberculares.Caulis villosus,principalis erectus, apex, externalsurface glabrous or with some rami procumbentes.Stipulae subglabrae dorso short adpressed hairs, and occasionally also pilis longis, margineciliis longis instructae. Folia with a few long hairs, margin longly ciliate. caulis principalisfoliolis oblongis, ilia ramorum Petiole and rachis villous except glabrous on foliolis ovatis vel interdumoblongis aut raro upper, canaliculate surface. Pulvinus villous. obovatis,omnibus epiphyllo laevi, nitido, pilis Leaflets with the upper surface smooth, usque ad 1 mm longis vestito, hypophyllopilis shining,with long hairs, up to 1 mm long, adpressisvix perspicuis saepe pilis longissparsim scattered, at times only present on some immixtis,nervo medio villoso et margineciliis leaves; lower side with midvein and margin longis setulisque brevibus instructo prominent, surface with barely visible prominentibus.Hypanthium 3.3-4.8 cm longum, adpressed hairs and with scatteredlong hairs laxe villosum.Calyx 4-6 mm longus, villosus, or withoutthem, midvein villous, marginwith setulisdestitutus. Vexillum 7-11.5 mmlongum x long cilia and emergenceswith a thick base 10-13 mmlatum, luteo-aurantiacum dorso lineis and a short bristle. Flowers all along the rubescentibusornatum. Fructus biarticulatus prostrate branches, in very short axillary paxillo glabro, 6-10 cm longoradices adventitias spikes, 3-4 flowered.Hypanthium 33-48 mm emittente,articulis 10-11 mm longis x 6-7mm latis, long, softly villous. Calyx villous, without pericarpiolaevi. bristles;upper lobe 4-5 mm long, lower lobe subfalcate,5-6 mm long. Standard7-1 1.5 mm Holotype:BRAZIL. Tocantins. 5 km N de long x 10-13 mm wide, yellow-orange,with Araguaina,300 m,creeping herb, forming extensive reddishor purple lines on the dorsal surface, mats, corolla yellow-orange,cut-over sandy converging toward the base. Pegs purple,

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) glabrous on the aerial portion,pubescent and & al. 7875 (CEN, CTES). Piauí. Km 76 estrada with adventitiousroots below ground, 6-10 Teresina-CampoMaior, 8-III-1968, Andrade-Lima cm long.Fruit articles 10-11 mmlong x 6-7 mm 68-5328(CEN, CTES). Mun.Altos: Altos, km 311, wide, pericarp smooth. 2n=20 chromosomes BR-343,23-III-1985, Valls & al. 8473(CEN, CTES); (Fernández & Krapovickas 1994). km 42 on BR-316 ao sul de Teresina,1 km S de DemervalLobão, 26-III-1985, Valls & al. 8501(CEN, Selectedadditional material: BRAZIL. Goiás. 25 CTES). Tocantins.2 kmS ofGuará, 550 m,sandy kmSW ofMonte Alegre de Goiás,northern spur of cerrado,19-111-1968, Irwin & al. 21448(NY, UB); 24 SerraAtalaia, 600 m,13-111-1973, Anderson 702 1 (K, kmS ofParaíso, 600 m, 24-111-1968, Irwin & al. 2 1732 NY, UB, US). Mun. Cavalcante:entre Campos (HB,NY, RB,UB, US); 28,6km S ofParaíso do Norte Velhos y Alto Paraísodo Norte,km 98, 14°33'S, de Goiáson Belém-Brasilia highway, 10°24'S, 49°W, 47°16'W,510 m,Coradin 3836 (CEN). Maranhão. 280 m,27-XII-1969, Eiten & al. 10051 (K, NY, US); Taquara,5 kmNW ofBalsas, 7°29'S, 46°3,W, 250 m, Estreitopara Tocantinópolis,24-IV-1971, Duarte 15-III-1962,Eiten & al. 3620 (NY, SP); Imperatriz, 13942 (NY); PortoReal (PortoNacional), 17-XI- aeroporto,27-IV-1982, Pires 17643 (CTES). Mun. 1828,Burchell 8443 (K); Natividadeto Porto Real, Imperatriz:"Bananal," 15 kmS ofImperatriz, BR-0 10, btw.Carmo & CorregoFundo, 12-XI-1828, Burchell 5°40'S,47026W, 290 m,29-11-1980, Plowman & al. 8328 (K); id.,Burchell 8358 (K, P). Mun.Presidente 9370(CTES): 33 kmN de Imperatriz,BR-010, 2-III- Kennedy:Road from BR-153 to Itapora 12 kmW of 1982,Valls & al. 6586 (CEN, CTES); 13 kmS de Pres.Kennedy, Fazenda Primaveraalong Riberao Imperatriz,BR-010, 12-III-1982,Valls & al. 6600 Feinho,8°25'S, 48°37'W, 400-500 m, 31-1-1980, (CEN, CTES); 40 kmS de Imperatriz,BR-010, km Plowman& al. 8174 (CTES). MunTocantinópolis: 1303,12-111-1982, Valls & al. 6602(CEN, CTES): 41 BR-153, kin 1 192, 6°38'S, 47034W, 300 m 7-X-1980, kmS de Imperatriz,12-III-1982, Valls & al. 6604 Coradin3650 (CEN). Mun.Araguaina: BR-010, km (CEN, CTES): 48 km3 de Imperatriz,12-III-1982, 1097,7°14'S, 48°15'W, 250 m, Coradin 3715 (CEN), 11 Valls& al. 6605 (CEN, CTES). Mun.Montes Altos: kmde Paraísodo Norte,Coradin 3724 (CEN). Mun. km1302, BR-019, 5°46'S, 47°21'W, Coradin 3659a Natividade:entre Natividade y Dianópoliskm 5, (CEN); BR-010,km 1303, 5°46'S, 47°21'W, 230 m, 7- 11°40'S, 47°47'W, 370 m, Coradin3755 (CEN); X-1980,Coradin 3660 (CEN); BR 010, km 1330, Natividade,3 kmN do entroncamentopara Porto 5°34'S,47026W, 250 m. Coradin 3669 (CEN); 12kmN Nacionalna estradapara Dianópolis,16-III-1982, de Carolina,17-III-1985, Valls & al. 8352 (CEN, Valls& al. 6640(CEN, CTES); Estreito-Marabákm 1, CTES); 17km N de Carolina,17-111-1985, Valls & al. 5-IV-1974,Pinheiro & al. 90 (NY). Mun.Formoso do 8359(CEN, CIES); 9,5km N deCarolina 17-111-1985, Araguaia: 2 km de Formosona saida para o rio Valls& al. 8375(CEN, CTES); 20 kmSW ofRiachão, Javaé,4-III-1982, Valls & al. 6496 (CEN, CTES). BR-230,1 8-ni-l 985, Valls & al. 8390(CEN, CTES); Mun.Cristalandia: 42 kmNdeGurupi, BR-153, 6-III- Balsas,19-III-1985, Valls & al. 8402(CEN, CTES): 6 1982,Valls & al. 6530(CEN, CTES): id.,15-111-1985. kmNE deBalsas, 19-111-1985, Valls & al. 8405(CEN, Valls& al. 8309(CEN, CTES); Rosalandia,BR-153, CTES); f V t SE ofVale Verdeentre Balsas e São 6-III-1982,Valls & al. 6532(CEN. CTES); id.,15-ffl- Raimundo.BR-230, 19-III-1985, Valls & al. 8413 1985,Valls & al. 8321(CEN, CTES). Mun.Miracema (CEN, CTES); km567,5 da BR-316 entreTimon e doNorte, : 1 km N do riodos Bois, km 854, BR-1 52, 7- Caxias,24-III-1985, Valls & al. 8496 (CEN, CTES). m-1982,Valls & al. 6540 (CEN, CTES); BR-153,29 Mato Grosso.Mun. São Felixdo Araguaria: BR-242, kmN deMiranorte, 7-III-1982, Valls & al. 6546(CEN, 24 kmE de BR-158,20-VIII-1984, Valls & al. 7799 CTES). Mun.Tupiratins: 150 mN do rioFeio, BR- (CEN, CTES); 16,4km W de São Felixdo Araguaria, 153,8-III-1982, Valls & al. 6554(CEN, CTES). Mun BR-158,22-Vin-1984, Valls & al. 7805(CEN, CIES). Colinasde Goiás:1 8 kmN de Colinas,BR-1 53, 8-III- Mun.Luciara: 14 km S de PortoAlegre, BR-158, 23- 1982,Valls & al. 6555 (CEN, CTES); 3 km S de Vm-1984,Valls&al. 7821(CEN, CIES). Mun.Santa Araguaina,BR-153, 9-III-1982, Valls & al. 6556 Terezinha:15,2 km do aeroportode SantaTerezinha (CEN,CTES); Araguaina,BR-226, 10-III-1982. Valls na estradapara BR-158, 24-VIII-1984, Valls & al. & al. 6558(CEN, CTES). Mun.Babaçulandia, 18 km 7850(CEN, CTES); SantaTerezinha, 25-VIII-1984, SW ofWanderlandia, BR-153, 6°56'S, 48°05'W, 420 Valls& al. 7868 (CEN, CTES). Pará. 36,3km W de m, 26-11-1980,Plowman & al. 9164 (CTES), CampoAlegre, 9°25'S, 50°39'W, 26-VIII-1984, Valls Vanderlandia,10-III-1982, Valls & al. 6559 (CEN,

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CIES); 12km S deVanderlandia, lO-m-1982, Valls & erectus, rami decumbentes.Stipulae villosae al. 6566(CEN, CIES); 7 kmNE deVanderlandia, 16- usque subglabrae, ciliatae. Folíola oblonga III-1985, Valls & al. 8335 (CEN, CTES). Mun epiphylloviridi-olivaceo, nitido, pilis ca. 1.5 mm Tocantinópolis:56 kmNE de Vanderlandia,BR-226, longis sparsim vestito,margine manifestiore, 10-m-1982,Valls & al. 6574(CEN, CTES); id.,16-ÜI- hypophyllovilloso nervo medio prominente, mar- 1985,Valls & al. 8343(CEN, CTES). Mun.Paraíso do gine denseplus minusveadpresse-ciliato conspi- Norte,: 1 2 kmde Paraísodo Nortena estradaa Porto cuo. Fructus biarticulatus,paxillo ca. 15 cm Nacional,15-ÏÏI-1982, Valls & al. 6633(CEN, CTES); longo, radices adventitiasemittente, articulis 8 id.,6634 (CEN, CTES); id.,6635 (CEN, CIES); 24km mmlongis x 5 mmlatis, pericarpio laevi. SW de Paraísodo Nortena estradapara Porto Na- cional,15-III-1982, Valls & al. 6636 (CEN, CTES). Holotype:BRAZIL. MatoGrosso. 17 kmNW de Mun. PortoNacional: 32 kmW da pontesobre rio RosarioOeste, 12-III-1959, Gregory, Rrapovickas & Tocantins,na estradade Paraísodo Nortea Porto Pietrarelli9923 (CEN). Isotypes:CTES, GH, LIL, Nactional,15-111-1982, Valls & al. 6637(CEN, CIES). MO,NY, SP,US. Mun.Pium: proximo a margemda Ilha do Bananal, emfrente a Santa Terezinha, 25-VIII-1984, Valls & al. Perennial plant. Root slender, with 7863 (CEN, CTES). Mun.Miracema do Norte:BR- fusiformenlargements 2-4 cm long x 0.5 cm 153,km 383 a 13,6km S dorio dos Bois, 27-VIII-1984, wide. Mainstem erect, sometimes reclining, Valls& al. 7880(CEN, CTES). ca. 45 cm long. Four prostratelateral branches ca. 1.20 m long. Internodesangled, villous, Geographic distribution. Occurs in glabrous towardthe base of the branches.On Tocantins and adjacent localities in the states the mainstem,the firstinternode ca. 10 cm of Goiás, Maranhão and Piauí. It grows long, those followingshorter, usually 3-5 cm forminga mat,principally in the "cerrado"and long and towardthe apex theyare reduced to in the gallery forest,in sandy soil, sometimes ca. 2 cm long. On the lateral branches, the withclay subsoil. internodes2-4 cm long. Leaves with4 oblong leaflets. On the mainstem,stipules with the Obs. This species is very closely relatedto fused portion 12-15 mm long x 3 mm wide, A. prostrata and A. lutescens, with which it the freeportion 16-22 mm long x 2 mm wide, formsa group that is differentiatedfrom the acute, with rigid apex, marked longitudinal restof the species of this sectionby the size of veins,margin ciliate, surfaces glabrous except the leaflets, which are the smallest. Despite fora few hairs towardthe base; petiole 24-29 this similarity, we consider them as mmlong, villous; rachis9-13 mmlong, villous; independent entities for having different apical pair of leafletsup to 35 mm long x 12 ranges. Going from southwest to northeast mmwide, basal pair up to 30 mmlong x 1 1 mm through the range of the section, the wide. On the lateral branches, the fused geographic sequence is A. lutescens, A. portionof the stipules 4-9 mmlong x 1.5-2 mm prostrata and A. Burchellii. wide, the freeportions 8-13 mm long x 1.5-2 mmwide, villous; petiole 5-14 mmlong, villous; We dedicate this*species to W.J. Burchell, rachis 4-8 mm long; apical pair of leaflets 13- who collectedit forthe first time in 1828 in the 23 mmlong x 7-10 mmwide, the basal pair 12- vicinityof Porto Real (today, Porto Nacional) 20 mm long x 5-8 mm wide; upper surface on the Tocantins River. olive green, shiny, with long (ca. 1.5 mm) scatteredhairs, marginsomewhat prominent; 23. Arachis Pietrarellii Krapov. & W.C. the lower surface with long hairs similar to Gregory nov. sp . those on the upper surfacebut more dense,the Figs. 1,23; 16,D-F midvein prominent, villous, the surface between the secondary veins with reddish spots,margin pronounced, with abundant cilia, Herba perennis.Radix tenuis,ramificationibus more or less adpressed. Inflorescences in tubercularibus. Caulis villosus, principalis axillary spikes distributedalong the lengthof

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) the branches,2-3-flowered, axes short;bracts 24. Arachis villosulicarpa Hoehne filiformand 5 mm hori- lanceolate, long. Pegs Fig. 1,24 zontal,shallow, growing parallel to thesurface of thesoil, ca. 15 cm long,subglabrous, reddish. Articlessmooth, 8 mmlong x 5 mmwide. 2n=20 Hoehne,Arq. Bot. EstadoSão Paulo,n.s. 2: 16- chromosomes(Smartt 1964, GKP 9923). 18,táb. 5, 1944.Gregory, Res. and Farming 5, Prog. Rep.4: figs.25-26 and 28, 1946,"diploid." Mendes, Additionalmaterial: BRAZIL. Mato Grosso. Bragantia7: 262, 1947,2n=20. Conagin, Bragantia Mun.Nobres: 900 m ao N do CorregoSeco e a 4 km 21: 345-348,est. 1 & 2, figs.2 A, 2B and9, 1962. ao S de Nobres,ao N de RosarioOeste, 25-1-1989, Valls& al. 12085(CEN, CIES); 1,4km S doacesso a Perennial plant; taprooted in the first Nobresna estradade RosarioOeste a Sinop,25-1- months,later with branch roots very much 1989,Valls & al. 12086(CEN, CTES); 15 kmN de thickened,customarily producing vegetative RosarioOeste, 2-VI-1985, Valls & al. 9000 (CEN, sprouts. Mainstem erect, some 60 cm tall, CTES); 17km N de RosarioOeste, 2-VI-1985, Valls normallyproducing only vegetativebranches; & al. 9004(CEN). lateral branches prostrate, 1 m long (in the type), or decumbent up to 35 cm long; Relatedmaterial: BRAZIL. Mato Grosso.Mun. internodesangled, 22-40 mm long, villous. AguaBoa: BR-158,27,2 km ao nortede AguaBoa, Fused portionof the stipules 10-17 mmlong x 6 19-Vm-1984,Valls & al. 7784(CEN, CTES); 40,8km mm wide, the freetips acute, 10-20 mm long. N de AguaBoa, 19-Vm-1984,Valls & al. 7786(CEN, Petiole canaliculate, 19-50 mm long; rachis 5- CTES). Mun.Canarana: 37 kmS de Cascalheira,19- 13 mmlong. Leaflets 4, elliptical,the apical pair Vm-1984,Valls & al. 7793(CEN, CTES). Mun.Santa 20-44 mmlong (up to 50 mmin thetype) x 6-16 Teresinha:Santa Teresinha, 25-VÜI-1984, Valls & mm wide (up to 22 mm in the type), the basal al.7861 (CEN, CTES). pair 16-40 mm long x 5-15 mm wide. Stipules villous toward the base, with subglabrous Geographicdistribution. The typecollection surfaces, prominentveins and ciliate margin. was made in the south central part of Mato Petiole and rachis villous, except on both Grosso in thevicinity of Nobres, situatedsome canals. Pulvinus villous. Upper surface of the 18 kmnorth of Rosario Oeste, at thefoot of the leaflets with sparse, very adpressed hairs Serra do Tombador. It also grows in eastern when young, later glabrous, smooth, bright Mato Grosso, on theeastern slope of the Serra green;lower surfacesubvillous to glabrescent, do Roncador, between Agua Boa and Santa withlong hairs on the midvein;margin barely Teresinha,along the Araguaia River,where it marked,ciliate. Inflorescenceswith very short was collected in August,in the dryseason, for axes, covered by the stipules, distributed whichits identificationremains tentative. principally along the tertiary branches. Hypanthiumusually 4 cm long,varying between Obs. Arachis Pietrarellii is related to A. 2.1 and 7 cm, villous. Calyx bipartite,5-6 mm Macedoi, fromwhich it differsby its larger, long, pilose. Standard 6-10 mm long x 10-15 somewhat more obtuse leaflets,especially in mm wide, orange, with red lines on the back the leaves of the lateral branches,and by the convergingtoward the base. Fruitsubterranean, more marked margin. It is also similar to A. Particulate; pegs inclined 40°-70°, 6-16 cm villosulicarpa, but its fruitsare much smaller. long, commonly13 cm long, striate,glabrous, The pegs of A. Pietrarellii customarilyform frequentlybearing roots; isthmus13 mm long; roots which enlarge into tuberoids, and articles usually 18 mm long x 8 mm wide, plantletscan arise at thejunction of root and varyingfrom 16x8to23xl0 mm,S-shaped, peg- denselyvillous; pericarp smooth; seeds 10-18 mm long, S-shaped, elongated. 2n=20 We dedicate this species to José R. chromosomes(Mendes 1947). Pietrarelli,fellow explorer, who found this species. Holotype:BRAZIL. São Paulo. Cult. Jardim

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Botanicode sementestrazidas pelo Sr. Antonio knowledge of this species in the wild state, Telles como procendentes dos Indios each tribemaintaining its own seed. One proof Nambiquaras,Juruena, M. Grosso,Gehrt s.n., 8-1- of its conditionas a cultivated plant is the 1943 (SP 47535!). Isotype:K!. The chromosome strengthof the peg, which is much stronger count2n=20 (Mendes 1947: 262) was determined than can be foundin the wild species. withseeds of these cultivars (SP 47535). Evidently, we are witnessing a domesticationprocess independentof that of Selected additionalmaterial: BRAZIL. Mato A. hypogaea. Both species belong to different Grosso.Mun. Vila Bela: PostoIndigena Mamainde, sections, between which it has not yet been 82 kmSE deVilhena, e logoca. 20 kmSW da BR-174, possible to obtain any hybrid.In the section 21-V-1985,Vails & al. 8820(CEN, CIES). Rondonia. Extranervosae,the species most similarto A. Vilhena,2 l-V-1985,Valls & al. 8816 (CEN, CIES); villosulicarpa is A. Pietrarellii , for its id.,8818 (CEN, CTES). São Paulo. Campinas,Insti- vegetativecharacter, but havingmuch smaller tutoAgronómico, V-44 (Procedencia:Mato Grosso, fruits(fig. 1, 23 and 24). Vilhena,leg. Pedro de Barros, 2-IX-1938, 1-2288), II- 1960,Conagin 18 (IAC 18675);Cult. V44, 4-VI-1968, Obs. 2. In the archives of the Instituto Krapovickas& al. 14446(CTES); Cult.V 862,5-VI- Agronómico at Campinas, the following 1968,Krapovickas & al. 14445(CTES). collections are recorded, all from Mato Grosso. Common name, "amendoim bravo" (IAC, V.44 (1.2288), Amendoim bravo, Estaçao V.44). "wi-ki-rin-gui,"the name provided in Telegráfica de Vilhena, norte de Mato Vilhena by the native informantFrederico of Grosso. Enviado por Pedro Paes de Barros, the Tauandé tribe(Vails 8818). agrònomodo D.N.P.V., Cuiabá, 20.9.1938. V.80 (1.5893), Mato Grosso. Obs. amostra Geographicdistribution. This is a cultivated de sementes. . . esse materialse asemelha com species about which there is no information o de no. 1.2288. Processo no. 43.372. knownregarding its existencein thewild state. V. 122 (1.7241), AmendoimI, Cultivado pe- The materialwas providedby the indigenous los indios Nambyquaras en Juruena, Mato people thatlive in thevicinity of Juruena,in the Grosso. Material trazido pelo Sr. Athos de west central part of Mato Grosso, and from Souza Lima. Entregue ao Sr. Octacilio Vilhena, in the eastern extremeof Rondonia Ferreyrade Souza. 11 .1. 1945. (Krapovickas &al. 1985: 41). V. 123 (1.7242), AmendoimIII, id. V.125 (1.7244), AmendoimII, id. Obs. 1. Arachis villosulicarpa was only V.131 (1.7746), Mato Grosso. Pequena knownfrom seven fruitsamples fromVilhena amostrade sementesrecebida pelo Sr. Octacilio and Juruena,sent through intermediaries to the F. de Souza, por intermédiodo MonsenhorJoão Institutode Botánica de São Paulo and the Baptistadu DreneufS.J. Administrador Apostóli- Instituto Agronómico de Campinas (SP), co da Prelaziade Diamantino,no estadode Mato between 1938 and 1945. The original Grosso. Esse amendoimfoi obtido com os indios informationdescribed it as a native crop, but de Juruena,tratando-se, talves, da mesmaespe- the name "amendoim bravo" of the only cie que ha tempos, recebi do Snr. Raul surviving sample would indicate a wild DrummondeGolsalves, dessa mesmalocalidade conditionfor the material. It was not clear if e encaminheiao InstitutoAgronómico. O mate- the fruitsused forfood came fromcultivation rial depois de passar pelos tratamentos or ifthe natives were collecting them in the wild. necessários foi devolvido a Secção de Recently, we (Vails, Krapovickas and Oleaginosas,para o Snr. Octacilio F. de Souza. Simpson) obtained, near Vilhena, fruits 19.IX.1945. harvested by members of two indigenous V.284 (1.14538), North Carolina. Sementes tribesand had the opportunityto visit a third trazidas pela Dra. C.H.T. Mendes Conagin. tribe,where we saw theplants in cultivation.In Sementesda var. 118. Cit. 862. Agostode 1952. these three cases the natives had no 862 = V.284

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Accordingto Conagin (1962: 348) theplants Hypanthiumca. 45 mm long. Standardorange of V.125 have completely prostrate lower witha conspicuouspurple spot towardthe base branches, for which they are more like the on the upper surfaceand withfainter red lines originaldescription. The rest of the materials on the lower surface. Fruit subterranean, studiedby Conagin have decumbentbranches. frequentlywith three segments; peg and isthmi Hoehne describes this species with very elongated,horizontal, shallow; articles single- long, prostrate, root-bearing stems. It is seeded; pericarpsmooth, covered witha dense possible that this observation is due to the layer of hairs. 2n=20 chromosomes(Gregory presence of pegs with adventitious roots, &al. 1973). frequentin this species. 25. Arachis triseminataKrapov. & W.C. Obs. 3. Arachisvillosulicarpa is, genetically, Gregory nov. sp. We obtained a veryisolated species. only hybrids, Figs. 2,25; 23,C that were highly sterile, in crosses with A. lutescens,A. Macedoi, A. marginata and A. prostrata,all in thesame section. A. pusilla auct.non Bentham, Gregoiy & al., in W.C. Gregoiy,M.P. Gregoiy,Krapovickas, Smith & Obs. 4. In 1974, Pire studiedthe pollen of Yarbrough,in Peanuts- Cultureand Uses: 121, A. villosulicarpa, findingtri-colpate grains 1973. that are syncolpate,that is, with the colpae united at the poles, similar to those of the Herba perennis. Caulis villosus,principalis genus Stylosanthes. erectus,rami decumbentes.Stipulae marginibus basi connatis tubumbrevem 1-2 mm longum IV. Sect. Trisenúnatae Krapov. & W.C. efficientes,apice subulataepilis longis sparsim Gregory nov. sect vestitae,margine ciliatae. Foliola ovalia usque Fig. 5 obovata, acuta, epiphyllo laevi, glabro, hypophyllovilloso, nervo medio prominente, margineciliato. Hypanthium 4.5 cmlongum, villosum. Sect. TriseminataeKrapov. & W.C. Gregoiy,in Calyx4-5.5 mmlongus, villosus. Vexillum 8-9 mm W.C. Gregoiy,M.P. Gregory,Krapovickas, Smith & longumX 10 mmlatum, aurantiacum. Fructus 2-3- Yaibrough,in Peanuts- Cultureand Uses: 93, articulatus,paxillo horizontāliusque ad 72 cm 1973,nomen nudum. longo,isthmo 20-37 cm longo,articulis 10-12 mm longisX 6-7 mmlatis, pericarpio laevi. Perennes. Radix polaris ramificationibus tenuibusnon incrassatis. Stipulae marginibus Holotype:BRAZIL. Bahia.Juazeiro, Estaçao Ex- connatistubum brevem 1-2 mm longum efficientes perimentalMandacaru, 6-IV-1967, Gregory& apice subulatae.Folia quadrifoliolata.Vexillum Krapovickas12881 (CEN). Isotypes:C, CTES, G, aurantiacumsupra subtusque lineis rubescentibus GH,MBM, MO, NY, US. ornatum.Fructus subterraneus2-3 articulatus paxillo horizontalepaulo profundo,pericarpio Perennialplant. Taproot deep, with slender laevi. branches.Mainstem erect and lateral branches decumbent;stem angular with copper-colored Typussectionis: Arachis triseminata Krapov. & hairs ca. 1.5 mm long, dense, especially on the W.C. Gregory youngparts. Leaves tetrafoliolate,those of the mainstemsomewhat larger than those of the Perennial plant. Taproot, without lateral branches. On the mainstem,the fused thickenings. Mainstem erect and branches portionof thestipules up to 10-12 mmlong, the decumbent. Stipules somewhat fused at the freeportion up to 23 mmlong; petioleup to 60 base forminga very short tube and with the mm long; rachis up to 19 mm long; apical pair apices subulate. Leaf tetrafoliolate.Flowers of leafletsup to 29 mmlong x 17 mmwide, the small, along the length of the branches. basal pair up to 28 mmlong x 16 mmwide. On

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the lateral branches,the fused portionof the BR-407,1 l-IV-1983, Valls & al. 7243(CEN, CTES); stipules 7 mm long,the freeportion 10-17 mm 5 1km S de Juazeiro,BR-407, 1 l-IV-1983, Valls & al. long; petiole usually 25 mm but up to 40 mm 7246 (CEN, CTES); 5 kmof Juazeiro,SUDENE, long; rachis 8-15 mm long; apical pair of ResearchStation, 370 m, 5-II-1972, Pickersgill RU72- leafletsusually 17 mmlong x 9 mmwide butup 81 (CTES,IPA, US); 15kmWdeJuazeiro,8-IV-l%7, to 26 x 17 mm; the basal pair 15 x 17 mmbut Gregory& al. 12922 (BAA, CTES, GH, MO, NY, up to 26 x 14 mm. Stipules withthe fusedpart US); 15km de Juazeirona direçãode Uauá, 18-VHI- papery with margins united toward the base 1965, Andrade-Lima65-43 17a (IPA); Fazenda forminga short tube 1-2 mm long, external Tourão,6 km Leste de Juazeirona estradapara surfacewith long scatteredhairs; freeportion Curaçá,18-VI-1981, Valls & al. 6188(CEN, CTES). subulate,with a singlevein and ciliatemargins. Mun. Xique-Xique:baixios de Irece,10-VI-1986, Petiole with long hairs, ca. 2 mm, and small Pinto61/86 (CTES). Mun.Barra: 15kmEde Javi,BR- adpressed hairs, canal on upper side narrow 242,entre Barreiras e Ibotirama, 12°12'S, 43°36'W, 6- and withsmall hairs,separated from the rachis V-1982,Valls & al. 6772(CEN, CTES); 43,3km W do canal by a transverseline of long hairs, also rio São Francisco(Ibotirama), BR-242, 12°21'S, presentat the juncture of the second pair of 43°38*W,14-IV-1983, Valls & al. 7292(CEN, CIES). leaflets. Leaflets oval, sometimes obovate, Minas Gerais. Mun. Janaúba:15°27'S, 43°27'W, acute toward the tip and with the apex Valls & al. 13080 (CEN). Pernambuco. Mun. apiculate; upper surface smooth, glabrous; Petrolina:ao lado de canaisde irrigaçaodo rioSão lower surface villous, midvein prominent, Francisco,26-11-1978, Leitão Filho & al. 8912 (MG, secondary and marginal veins little marked, UEQ. margin ciliate. Flowers abundant along the length of the branches, in short, 3 -flowered Common name, "mundubi" (Valls & al. spikes. Hypanthium45 mm long (29-74 mm), 7243). hyaline,villous. Calyx 4-5.5 mmlong, villous. Standard 8-9 mm long x 10 mmwide, orange, Geographicdistribution. This species grows yellow in the center,with red lines on both in the state of Bahia, in the south of sides, more intense on the upper surface, Pernambucoand in thenorth of Minas Gerais, wingsyellow, 6-7 mmlong. Fruit subterranean, and in the vicinityof the São Francisco River. 2-3 -articled; peg more or less horizontal, The mostwidely separated localities are found commonly30 cm long but up to 72 cm long, between 40 and 50 km on eitherside of said isthmus20-37 cm long; fullydeveloped articles river. It is a component of the "caatinga" 10-12 mmlong x 6-7 mmwide, coveredwith a woodland vegetation,growing in compacted dense coat of very shorthairs, when the hairs silty soil. Plants typicallyform dense stands fall the surface is completelysmooth; the peg and provide a very palatable feed for is insertedon the extreme basal end of the livestock. articlesat thatthick point, ca. 2 mm wide; the isthmus is borne at the apical end without Obs. Arachis triseminata is a species forminga beak, givingthe article a small dorsal that is genetically very isolated because it hump. Cotyledons oval-shaped, the upper was not possible to obtain any hybridsin the surface with veins very sunken. 2n=20 interspecificcrosses attempted. chromosomes (Gregory & al. 1973). Crosses between accessions of the same species from differentlocations produced Additionalmaterial: BRAZIL. Bahia. Juazeiro, hybridswith 50.7 and 68.2% fertilepollen. III-1912, Löfgren 906 (RB); Juazeiro,Horto This is a very interestingresult, given that Florestal,111-1912, Zehntner 36-906 (R); id., 4-1- the two populations were separated by a 1964,Andrade-Lima 64-4200 (CEN); EstaçaoExperi- distance of some 20 km. Both populations mentaldo Mandacaru,13 km NE ofJuazeiro, 22-VI- showed no appreciable morphological 1981,Valls & al. 6240(CEN, CTES); Juazeiro,2 km a differences, which suggests the lesteda BR-407na estradapara Curaçá, 10-IV-1983, development of some type of intraspecific Valls& al. 7232 (CEN, CTES); 29 kmS de Juazeiro, genetic barrier.

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)

V. Sect. Heteranthae Krapov. & W.C. isthmushorizontal, not deep; articles single- Gregory nov. sect seeded; pericarpsmooth, covered witha dense of hairs. 2n=20 chromosomes. Fig. 6 layer

Sect. AmbinervosaeKrapovickas, Agricultural Geographic distribution.This is a typically northeasternsection. It is in Genetics.Selected Topics: 137, 1973, nomen nudum. present all of the that Sect. PseudoaxonomorphaeKrapov. & W.C. states constitutethe "Nordeste" regionof Gregory, in W.C. Gregory, M.P. Gregory, Brazil and barely exceeds their boundaries, in Krapovickas,Smith & Yarbrough,in Peanuts- towardsthe south Minas Gerais and towards Cultureand Uses: 92, 1973,nomen nudum. the west in Goiás. There is a collection made on Marajó Island, in the mouthof the Amazon River, which we tentativelyidentify as A. Annuaevel biennes. Radix polaris ramificationibus Dardani, and whichis veryinteresting because it tenuibus,non incrassatis.Caulis prinicipalis erectus, is the only known case of a wild species of ramiprocumbentes. Stipulae marginibus liberis usque Arachis havingcrossed the Amazon. ad basemapice aciculares. Folia quadrifoliolataFlo- res dimorphi:parvissimi hypanthio brevissimo, minus Obs. Initially(Gregory & al. 1973: 120), this quam2cm,calice 2-3 mm longo, corolla clausa, etflores sectionwas comprisedby A. Dardani, a species normaleshypanthio extenso, calice majorecorolla whosestandard had redlines on bothsides. Later, expansa Fructussubterraneus, biarticulatus, paxillo with the new explorations,upon analyzingA. horizontāli. pusilla and A. sylvestris,it became evidentthat the threespecies formeda fairlyhomogeneous Typussectionis: Arachis Dardani Krapov. & group,composed of annual taxa adaptedto extre- W.C. Gregory me environmentalconditions, such as are often presentin the northeastof Brazil, where the Annual or biennial herbs. Taproot with "caatinga" predominates. Recently, A. slender branches. Mainstem erect, branches Giacomettiiwas added, whichis also an annual procumbent.Stipules withthe marginsfree to and has dimorphicflowers. the base and with acicular apices. Leaves In A. pusilla and in A. Giacomettiithe stan- tetrafoliolate.Dimorphic flowers,distributed dard exhibitsthe reddish lines on bothsurfaces, along the lengthof the lateral branches.Very but in A sylvestris,said lines are foundon the small flowers are produced, with the lower side, as in the section Extranervosae. hypanthiumfrom 3 to 20 mm long and the We believe, for reasons of geographic calyx and corolla closed, 2-3 mmlong; as well distributionand because these species share as normalflowers with the hypanthiumexten- the presenceof dimorphicflowers, a character ded, thecalyx largerand thecorolla expanded. thatis notpresent in any othersection, that it is Standard orange or yellow, withreddish lines advisable to include A. sylvestris in this on the outside surface or on both surfaces. section, making the standard's coloration no Fruit subterranean, biarticulate; peg and longeran exclusive characterof the group.

Key fordistinguishing the species

A. Stem,stipules and petiole with indumentum composed only of rigid bristles, 2-3 mmlong. Canal ofthe petiolevery narrow, separated from the canal ofthe rachis by a pilosetransverse prominence. Leaflets withthe upper surface glabrous. Standard with reddish lines on bothsurfaces. 26.A. Giacomettii A'. Indumentumcomposed of hairs of different length, if there are long hairs present they are 2 mmlong, soft, undulate,not rigid. The petiole-rachiscanal is wide,separated or not.

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B. Standardorange with reddish lines only on thedorsal surface; wings yellow. Canal of thepetiole separatedfrom the canal ofthe rachis by a pilosetransverse prominence. If there are bristles on the lowerleaf surface,they are notorderly. Pericarp smooth. Leaflets commonly with hairs on both surfaces,rarely with the upper surface glabrous. 27.A. sylvestris B'. Standardwith reddish lines on bothsurfaces. Upper leaf surface glabrous. C. Canal ofthe petiole separated from the canal of the rachis by a prominenceand transverse line of hairs.Frequently bristles on theupper leaf surface, orderly, forming a lineparallel to themargin. Pericarpsmooth. Wings yellow with the apex and internal margin orange. 28.A. pusilla C'. The canal ofthe petiole commonly not separated from the canal of the rachis. Lower leaf surface withoutbristles. Fruit articles reticulate, due to thefalling away of the smooth, villous epicarp. Wingsyellow. 29.A. Dardarti

26. Arachis Giacomettii Krapov., W.C. douradasabundantes em todo o pecíolo; plantas Gregory, Vails & C.E. Simpson com dominanciado eixo centralque crescianna Fig. 2,26 vegetaçãoalta. 17-111-1992, Valls, Pizarro, Valente & Werneck13202 (CEN). Isotype:CIES.

Herba annua caule principali erecto rāmis Annual herb.Taproot, weak. Hypocotyl2-3 procumbentibus.Caulis, stipulaepetiolique indu- cm long x 4 mm in diameter.Mainstem erect, mento setarum tenuum strictarum2-3 mm up to 60 cm long, with two basal vegetative longarumtantum obtectus. Folia quadrifoliolata branches, one or two successive nodes with foliolis apicalibus modice obovatis, foliolis shortinflorescences and withone or two short basalibus ovalibus usque oblongis, epiphyllo apical branches;lateral branchesprocumbent, laeve, glabro,hypophyllo glabro autpilis paucis no secondarybranches, with inflorescences on parvis appressis setisque nonnullis sparsim the basal nodes, internodesup to 6 cm long. vestito,nervo medio manifesto, nervis secundariis Stem, stipules and petiole with indumentum marginequesat perspicuis,hoc et nerviomedio made up solelyof straight,slender bristles, 2-3 pilis 1.5-2mm longis se tis nonnullis hirsutis. Flo- mmlong, scattered. Stipules with the basal part res dimorphi:parvissimi hipanthio brevissimo, 5 fused, 15-18 mm long and narrowwings, 8-11 mmlongo et flores normales hypanthio 2 cm longo, mmlong and 1.5 mmwide at thebase. Leaves pilos longosvalde dispersegerens, calyx 4-5 mm tetrafoliolate, apical leaflets somewhat longus,villosus et largesetulosus, vexillum 8 mm obovate and basal leaflets oblong, apiculate. longum auranti acum, supra subtusque lineis Canal of the petiole very narrow, separated rubescentibusornatum. Fructus subterraneus 2- fromthe canal of the rachis by a transverse (3)-articulatus,paxillo glabro ca. 10 cm longo, pilose prominence.On the mainstem,petioles isthmo2-5 cmlongo, articulis 10-12 mm longis x 6- up to 9 cm long and rachis up to 1 cm long, 7 mmlatís, epicarpio dense pilorumminutorum apical leafletsup to 44 mmlong x 20 mmwide obtectis. and the basal leafletsup to 42 mm long x 18 mm wide. On the branches,petioles up to 4.5 Holotype: BRAZIL. Minas Gerais. Mun. cm long and rachis 0.5 cm long,apical leaflets Montalvania,8.5 kmNW de Montalvania(desde o up to 35 mm long x 18 mmwide and the basal rioCoxá) na estradapara Pitarana, 14°21'S, 44°24W, ones up to 32 mmlong x 15 mmwide. Pulvinus 630 m.Mancha de cercade 50 mde extensão:cerdas villous; upper leaf surface smooth,glabrous;

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) lower surface glabrous or with a few small 188:1511, 1929 (juin). Chevalier, Rev. Int. Bot Appl. adpressed hairs and with some scattered Agrie.Trop. 13(146-147): 770, pl. 16,1933. bristles,with the midveinprominent and with the secondary veins somewhat marked; Herbaceous annual. Taproot, weak. midveinand marginwith long hairs 1.5-2 mm Mainstem erect, 4-15 cm long, without long and some scatteredbristles. Flowers very flowers, with two or four vegetative basal small, with hypanthiumvery short,that later branches, covered by the stipules toward the form pegs; exceptionally,larger flowers are apex. Lateral branchesprocumbent, up to 55 present. Well-developed flowers have two cm long, littleor no furtherbranching. Stems basal bracts 5 mm long, with bristles; glabrescent toward the base, later villous, hypanthium2 cm long, reddish,villous; calyx with long hairs up to 3 mm, more or less reddish, somewhat villous and with long straight,and small curved and wavy hairs; straightbristles, 1 mm long, upper lobe 4 mm internodes 10-40 mm long. Leaves long,lower lobe falcate,5 mmlong. Standard8 tetrafoliolate. Leaves of the mainstem mm long, orange with red lines on both somewhat larger than those of the branches; surfaces,wings yellow. Fruitsubterranean, 2- fused portion of the stipules 8-10 mm long, (3)-articled; pegs glabrous, ca. 10 cm long, the freeportion 9-14 mm long x 1-2 mmwide, isthmus2-5 cm long,articles 10-12 mmlong x pointed; petiole 15-30 mm long, rachis 7-10 6-7 mmwide, covered by a dense coat of hairs mm long; leaflets oblong, those of the basal that stronglyretains soil, withoutbeak, the pair 22-30 mm long x 10-14 mm wide, those isthmusis bornenear the middleof the ventral of the apical pair 30-36 mm long x 13-17 mm surfaceof the basal article. Seed 8-9 mm long wide. On the lateral branches, the fused x 4-5 mmwide, dark in color. portionof thestipules 6-7 mmlong and thefree portion 10-11 mm long x 1.5-2 mm wide, Additionalmaterial: BRAZIL. MinasGerais. 8,5 pointed;petiole 10-20 mmlong; rachis 5-8 mm km NW de Montalvania(desde o rio Coxá) na long; leafletsoblong to obovate,the basal ones estradapara Pitarana, 630 m,plantas da população up to 23 mmlong x 14 mmwide and the apical V. 13202,mas com eixo central ereto muito longo e ones up to 24 mmlong x 15 mmwide. Stipules sem desenvolvimentodos ramos laterais, que villous, withhairs ca. 3 mm long, more or less vegetavamentremeadas com gramíneasaltas. 17- straight,and small, curly hairs. Petiole and m-1992,Valls & al. 13203(CEN, CTES); id.,1-1994 rachis villous with the two types of hairs, Werneck& al. 201 (CEN, CTES); 12 km de sometimeswith bristles on the back, the canal Montalvania,desde o río Coxá, em direção a of bothglabrous. Pulvinuswith long hairs and Pitaranga,1-1994, Werneck & al. 178(CEN, CTES). small curlyhairs. Upper surfaceof the leaflets withuniformly scattered hairs, 1-1.5 mmlong; We dedicate this species to Dr. Dalmo lower surfacewith hairs ca. 2 mm long, more Catauli Giacometti,organizer and ex-director or less adpressed,midrib prominent, with long of the CentroNacional de Recursos Genéticos hairs up to 3 mm and short curly hairs; (CENARGEN/EMBRAPA), Brasilia, who secondary veins and margin barely marked, supported and greatly stimulated the margin ciliate, sometimes thick bristles are explorationsfor Arachis germplasmin Brazil. presenton the underside.Inflorescences few- flowered, axillary along the length of the 27. Arachis sylvestris(A. Chev.) A. Chev. prostrate branches. Hypanthium40-50 mm withscattered hairs Fig. 2,27 long,reddish, villous, long and shortcurly hairs. Calyx bilabiate,4-5 mm long, with long hairs and short curly hairs, Chevalier,Rev. Int. Bot. Appl. Agrie. Trop. 9(96): withoutbristles, after anthesis the upper lobe 486, pl. 13, 1929 (aug). Chevalier,loc. cit. 13(146- usually splits almost to the base. Standard 7 147):766, 1933. mm long, orange, with red lines convergent Arachis hypogaea L. subsp. sylvestrisA. toward the base on the external surface, in Cheval.,Compt. Rend. Hebd. Séances Acad. Sci. dried specimensthe entirestandard acquires a

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 reddishtone; wingsyellow. Fruitsubterranean; Timon,10 km W del puentea Teresina,Fazenda pegs shallow,growing horizontally, some 20-25 Bandeirante,3 1-1-1981, Krapovickas & al. 37215(C, cm long,villous on theaerial part;isthmus 6 cm CEN, CTES, F, G, INPA,IPA, K, LIL, MO, NY, P, long; articles 10-20 mm long, smooth,with a US); 7-8 kmW del puentea Teresina,28-1-1981, small beak, covered witha dense mat of short Krapovickas& al. 37191(CTES, F, G,HGB, IPA, K, hairs. 2n=20 chromosomes (Fernández & LIL, MBM, MO, NY, RB, SP, UB, US); 8 kmde Krapovickas 1994). Teresina,perto do fronteiracom Piauí, 31-1-1981, Pires& al. 17361(CTES, MG); Timon,9,5 km W da Holotype:BRAZIL. Bahia. 1929,leg. Bondar (P!). pontesobre o rioParnaíba, 24-III-1985, Valls & al. "... vallée du rio São Francisco,sur les sables 8491(CEN, CTES); Estereito,16-III-1985, Valls & al. d'alluvion.Elle estconnue comme l'Arachide sous 8345 (CEN, CTES); 9,5 km ao N de Carolinana le nomde Mundubi"(Chevalier 1929b: 486). estradapara Estreito, 17-III-1985, Valls & al. 8373 (CEN, CTES); 60 kmE de Carolina,BR-230, 18-m- Selectedadditional material: BRAZIL. Bahia. 4 1985,Valls & al. 8386 (CEN, CTES); 20 kmSW de kmE de Barreiras,road to Passagem and Ibotirama, Riachão,BR-230, 18-ffl-1985, Valls & al. 8392(CEN, 500m, 6-m-1971, Irwin&al. 31651 (NY, UB); id, 5-V- CTES);6kmNE deBalsas, BR-230, 19-111-1985, Valls 1982,Vails & al. 6767(CEN, CIES); Barreiras,patio & al. 8403(CEN, CTES). Mua Loreto:Ilha de Balsas del HotelBarreiras, 12°19'S, 45°W, 450 m,31-III- region,35 kmS ofLoreto, Veados, 7°23'S, 45°7W, 5- 1983,Vails & al. 7037 (CEN, CTES); Barreiras,bal- IV-1962,Eiten & al. 4005 (NY); 18 kmSW de São neariodo riode Ondas,15-IV-1983, Valls & al. 7294 Raimundo,BR-230, 19-III-1985, Valls & al. 8417 (CEN, CTES); 12 km N de Riachão das Neves, (CEN, CTES); São Raimundodas Mangabeiras,rio 11°30'S, 49°55W, l-IV-1983, Valls & al. 7060(CEN, Neves,19-ffl-1985, Valls & al. 8422(CEN, CTES); 1,2 CTES); 16km N do rioCariparé, estrada Riachão das kmNE da torrerepetidora de televisãoda saidaleste Nevesa Formosado RioPreto, l-IV-1983, Valls & al. de São Raimundo,BR-230, 20-HI-1985, Valls & al. 7065(CEN, CTES); 200 mN da saídapara Santa Rita 8423(CEN, CTES); 70 kmNE de São Raimundo,BR- de Cassia, estrada de Barreiraspara Corrente, 230,20-m-1985, Valls & al. 8435(CEN, CTES); São 11°20'S, 44056*W, l-IV-1983, Valls & al. 7071(CEN, Domingos,BR-230, 20-III-1985, Valls & al. 8437 CTES); 12 kmW de Formosado Rio Preto,1 1°3'S, (CEN,CTES); 27 kmNE dePastos Bons, 21-ffl-1985, 45°17'W,2-IV-1983, Valls & al. 7076(CEN, CTES); Valls& al. 8442 (CEN, CTES); 7 kmW de São João Formosado Rio Preto,2-IV-1983, Valls & al. 7079 dos Patos,BR-230, 21 -III- 1985, Valls & al. 8444 (CEN, CTES); 200 mN do riachoPiripiri, Estrada de (CEN, CTES); 25 kmE de São Joãodos Patos,BR- Formosaa Corrente,10°55'S, 45°14'W, 24-IV-1983, 230,21-m-1985, Valls &al. 8451(ŒN, CTES). Mua Valls& al. 7081(CEN, CTES); 1 kmNdoPindobaçu, Caxias:58 kmNE da pontesobre o rioParnaíba, BR- estradapara Senhordo Bonfim,10°44'S, 40°21'W, 3 16,24-111-1985, Valls & al. 8494(CEN, CTES). Mi- Valls& aL 6180(CEN, CTES); id.,19-IV-1987, Valls & nas Gerais.Rio Jequitaí,just S ofJequitaí, 17°14'S, al. 10891 (CEN, CTES); id., 10892(CEN, CTES). 44°26W,21-IV-1982, Valls&al. 6676(CEN, CTES); Ceará. Sinlocalidad, 1859, Allemao & al. 364(R); sin Pirapora,rio São Francisco,11-1-1989, Krapovickas localidad,1860-61, Allemao (P); 19,7km leste de & al. 42867(CEN, CTES). Paraíba.Em terrenosare- Barbalha,camino a MissãoVelha, 22-IV-1987, Valls nosose xerofíticos,25-V-1959, Moraes 2 127(NY, P, «feal. 10922(CEN, CTES); id, 10932(CEN, CTES); id., US). Pernambuco.Serra Talhada, V-1971, Heringer 10933 (CEN, CTES); 3,9 kma lestedo rioCauipe, 11996 (CTES, US). Piauí. Sinlocalidad, 1883, Netto entreGuarura e Catuana,BR-222, 28-IV-1987, Valls 27 (R); km84, estrada Teresina-Campo Maior, 8-III- & al. 10993(CEN, CTES); 4 kmW dorio Cauipe, BR- 1968,Andrade-Lima 68-5329 (CTES, IPA). Mun. 222, 28-IV-1987,Valls & al. 11000(CEN, CTES); Amarante:BR-343, entre Teresina e Floriano,km 40, Ibiapaba,Unidade de Pesquisade Ibiapaba/EPACE, 25-V-1980,Coradin 2605 (CEN, CTES). Mun. 30-IV-1987,Valls & al. 11020 (CEN, CTES). Goiás. Corrente:500 m S da pontesobre o rioCorrente, BR- Mua Formosa:1,5 km S davila JK (BR-020), 15°12'S, 135,10°27'S, 45°10'W, 460 m,3-IV-1983, Valls & al. 47°10W,10-VI-1981, Valls & al. 6001(CEN, CTES); 7105(CEN, CTES); 11km N do rioCorrente, BR-135, id.,21-111-1982, Valls & al. 6668(CEN, CTES); id.,3- 490 m,4-IV-1983, Valls & al. 7123 (CEN, CTES). V-1982,Valls & al. 6744 (CEN, CTES); id.,29-HI- Mua Barreirasdo Piauí: 39 kmS ofGilbués, BR-135, 1983,Valls & al. 7002 (CEN, CTES). Maranhão. 10°8'S,45°13'W, 4-IV-1981, Valls & al. 7126(CEN,

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CTES). Mun.Gilbués: 5 kmS de MonteAlegre do the locations and naming Glaziou as the Piauíe 7 kmNdeGilbués, BR-135, 520 m, 4-IV-1983, collector.Among the collectors involved in this Valls& al. 7130(CEN, CTES). Mun.Nazaré do Piaui: operation is Fr. Allemao, from whom there 7 kmEde Nazaré do Piauí, BR-230, 300 m, 7-IV-1983, exists a specimen fromCeará in the Museum Valls& al. 7176(CEN, CTES). Mun.Oeiras: 3 kmW National d'HistoireNaturelle, Paris. de Oeiras,200 m,8-IV-1983, Valls & al. 7180(CEN, CTES); acesso ao aeroportode Oeiras,8-IV-1983, Obs. 2. Bondar (1929), in an account of a Valls & al. 7191 (CEN, CTES); Altos,BR-343, km tripto the municipalityof Juazeiro(Bahia) in 311,23-111-1985, Valls & al.8475 (CEN, CTES); 13km December 1925, talks about "mundubi," a NE de Altos,BR-343, 23-III-1985, Valls & al. 8477 leguminous creeper that develops (CEN, CTES); 5,5km N de Itaueira,PI-140, 27-III- extraordinarilywell in the rainy season. The 1985,Valls & al. 8516 (CEN, CTES); 35 kmNE de plant sent to Hoehne in São Paulo was CristinoCastro, BR-135, 27-111-1985, Valls & al. 8520 identified as Meibomia incana Sw. The (CEN, CTES); Piripiri,700 m SW do Riodos Matos, cuttingsplanted in Bahia did not do well; they 30-IV-1987,Valls & al. 11027(CEN, CTES). Rio produced flowers and then died, but it was Grandedo Norte.Mun. Caicó: km 17 da BR-427,a 10 verifiedthat the plant buries its fruitsin the kmde Acari,em direçãoa CurraisNovos, 24-IV- same manneras "amendoim." 1987,Valls & al. 10969(CEN, CTES). Mun.Açu: 8,2 Chevalier (1929b: 496) transcribeda good kmWde rioAçu, a W de Açu,BR-304, 25-IV-1987, portionof this article byBondar, so it is very Valls& al. 10980(CEN, CTES). Tocantins.2 kmS de presumable that the type locality of A. Guarai,BR-153,8°51'S, 48°31'W, 7-ffl-1982, Valls & sylvestrisis Juazeiro, in the state of Bahia, al. 6547 (CEN, CTES). Mun.Tocantinópolis: BR- "on deep soil, formedfrom calcareous silt,that 226,km 1 192 (56 kmNE de Vanderlandia),6°38'S, cracks deeply in the dry season" (Bondar 47°34'W,Valls & al. 6575(CEN, CTES). 1929: 9).

Commonname, "mundubi"(Bondar 1929); Obs. 3. The specimen Vails 6676, from "mundubi do Joazeiro" (Otero 1952: 179); Jequitai (MG), has somewhat different "amendoim do porco" (Valls & al. 7191); flowers, whose standard, instead of having "mandubí do porco" (Valls & al. 7172); reddishlines in thedorsal surface,has greenish "mundubimbravo" (Valls & al. 8458). lines. The specimenKrapovickas 42867, from Pirapora (MG), is verysimilar, but because it Geographic distribution.This is one of the has onlytiny flowers it is notpossible to obser- mostwidely distributed species in the northeast of ve the color of the corolla. It is interestingto Brazil, formingvery small populations (Vails note thatboth specimensgrow in the southern 6180 and 6547), or coveringvery broad areas, extremeof the range of A. sylvestris. with invasive tendencies(Krapovickas 37215 and Vails 7076). It grows in dryplaces, in the Obs. 4. in A. sylvestrisit is commonto find "caatinga" as well as in the "cerrado," in loose very visible, abundant hairs on the upper sandysoils, redor grey,preferring shady spots, surfaceof theleaflets, making its identification especially whenfound in undisturbedareas. easy. Nevertheless, there are some specimens, coming from the vicinity of Obs. 1. There exists a questionable Floriano,on the Parnaiba River,in the central specimen: Brazil, São Paulo, Campo do part of Piauí, thathave a completelyglabrous Bocaina, common name "amendoim bravo," upper leaf surface. In some specimens (Vails 8-IX-1879, Glaziou 10513 (P), which 7144), a littlemore distant, the indumentumof correspondsvery well withA. sylvestris,but the upper leaf surfacevaries fromglabrous to was collected outside the range of that pilose. We identify this material as A. species. It has been pointed out (Wurdack sylvestrisbecause its flowershave a standard 1970: 911) that Glaziou was in the habit of withreddish lines on the dorsal surface. altering the labels of other collectors and using them for other specimens by changing Material studiedwith a glabrousupper leaf

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surface: BRAZIL. Maranhão. Mun. Barão do mm wide. On the lateral branches,the fused Grajaú: 100 m W da pontesobre o rioParnaiba, na portionof the stipules 5-8 mm long, the free estradaBR-230, para Floriano (Piauí), junto ao mar- portion10-12 mmlong; petiole 15-35 mmlong, co inauguraldo ponte,140 m, 6°46'S, 43°1'W, 7-IV- rachis 5-9 mm long; leafletsovate, the apical 1983,Valls & al. 7156 (CEN, CTES, G,K, MO, NY, 15-20 mmlong x 8-1 1 mmwide, the basal 13- US); id.,22-111-1985, Valls & al. 8458(CEN, CTES). 17 mm long x 7-9 mmwide. The stipuleswith Piauí. Mun.Floriano: 1 kma lesteda pontesobre o thickenedback, prominentveins and papery rioParnaiba, 170 m, 7-IV-1983, Valls &al. 7165(CEN, wings, free at the base (without forminga CTES, F, P, US); id.,22-III-1985, Valls & al. 8461 tube), free ends acute, single-veined, (CEN,CTES); 7 kmdaBR-343,a partir da BR-230, 22- narrowingabruptly, with very short hairs and III-1985,Valls & al. 8466 (CEN, CTES); 16km S de long hairs (2 mm) scatteredon theback, wings Floriano,PI-140, 27-111-1985, Valls & al. 851 1 (CEN, subglabrous,margin long ciliate. Petiole and CTES). Mun.Manuel Emiaio: 35 kmNE de Cristino rachis with a very narrow canal, back with Castro,BR-135, 310 m, 8°36'S, 44°W, 5-IV-1983, Valls short, somewhat wavy hairs and scattered & al. 7144 (CEN, CTES, G,K, NY); id.,27-III-1985, long, erect hairs; petiole-rachis canal Valls& al. 851 9 (CEN,CTES). Mun.Nazaré do Piauí: interruptedat the level of the first pair of 300 mN do acessoleste de Nazaré,BR-230, 2 10 m, leaflets by a transverse line of long, 6°50'S,42°41'W, 7-IV-1983, Valls & al. 7172 (CEN, compressed hairs. Upper surface of the CIES). leaflets smooth, glabrous; the lower surface with veins and margin softly marked, with 28. Arachis pusilla Benth. scattered adpressed hairs; in young leaves Fig. 2,28 there are also somewhat more erect, long hairs; margin densely ciliate with the hairs strongly directed toward the apex. Bentham,Trans. Linn. Soc. London18(2): 159, Inflorescences " few-flowered, very short, 1841, In Serra Jacobina provinciae Bahia, along the length of the branches and also Brasiliae,Blanchet 2669." Bentham,Martius, FI. presenton themainstem. Flowers dimorphous, bras.15(1): 86, 1859.Chevalier, Rev. Int. Bot. Appl. with very small flowers 2-3 mm long with Agrie. Trop. 9(90): 99, 1929. Chevalier,loc. cit. hypanthium3-10 mmlong, and normalflowers 13(146-147):765, 1933. Buikart, Darwiniana 3(2), lám. withhypanthium 4-5 cm long, villous. Calyx 21, 1939.Hermann, Agrie. Monogr. USDA 19: fig. villous, includingsome long bristles; narrow 24, 1954.Krapovickas & Rigoni,Darwiniana 1 1(3): lobe subfalcate,6 mmlong, the wide lobe 5 mm 449-450,obs. 1, 1957. long. Standardca. 10 mmlong, yellow-orange, withreddish lines on bothfaces; wingsyellow, Annual plant. Root small, 10-15 cm deep, withorange apex. Fruitbiarticulate, peg hori- withvery thin branches. Mainstem erect, 5-20 zontal, 4-20 cm long, violet and villous upon cm long; lateralbranches (n+1) procumbent,up emergence,later glabrous; isthmus0.5-4 cm to 80 cm long, with two short vegetative long; articles 10 13 mm long x 6-8 mm wide, branches at the base, with very few or no the apical article somewhat larger; pericarp subsequent (n+2) vegetative branches. smooth, covered by a dense cloak of Internodes1-4 cm long, angular, more or less diminutive hairs. Seeds with ocherous villous, with rigid hairs ca. 2 mm long, and integument,completely filling the articles. scatteredshort, somewhat curly hairs. Leaves 2n=20 chromosomes (Fernández & tetrafoliolate,those of themainstem somewhat Krapovickas 1994). larger and with the petioles noticeablylonger than those of the lateral branches. On the Holotype:BRAZIL. Bahia. Serra Jacobina, mainstem,the fused portion of thestipules 7-10 Blanchet2669 (K!). Isotypes:G!, P!. mm long and the freeportion 11-17 mm long; petiole 50-90 mm long; rachis 11-15 mmlong; Selectedadditional material: BRAZIL. Bahia. leafletselliptical, the apical 24-36 mmlong x 9- 38,1 kmNE de Ibotirama,camino a Seabra,12°02'S, 15 mm wide, the basal 22-30 mm long x 7-12 42°57W,16-IV-1987, Vails & al. 10837(CEN, CTES).

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Mun.Barra: BR-242, 1 1 kmW da pontesobre o rio 29. Arachis Dardani Krapov. & W.C. São Francisco,12°12'S,43°20'W, 16-IV-1987, Valls Gregory nov. sp. 10833 Mun. 2 km & al. (CEN,CIES). Paiatinga: 1,2 S Figs. 2,29; 17 de Ibotiramana estradapara Paratinga,12°21'S, 43°13'W,360 m, 14-HI-1981, Valls & al. 61 10(CEN, CTES); id.,flores diminutas, 19-ÜI-1982, Valls & al. Herba annua vel biennis. Caulis angulosus, 6655(CEN, CTES, G,K, LIL, MBM,NY, P, SI,US); villosus,principalis erectus, rami procumbentes. id.,flores normales e diminutas, 14-IV-1983, Valls & Stipulae pilis longis cupreis sparsim vestitae, al. 7287 (BAB, CEN, CTES, F); 42,8 km S de apicibus liberis trinervibusglabris, margine Ibotiramana estradaParatinga, 7-V-1982, Valls & al. longe ciliatae. Folíola ovalia vel aliquanto 6773(CEN, CTES); id.,14-IV-1983, Valls & al. 7289 obovata,utrinque acuta, epiphyllolaevi, glabro, (CEN, CTES); 13km S de Paratingana estradapara hypophyllofere laevi,pilis longissparsim vestito, BomJesus da Lapa,7-V-1982, Valls&al. 6781(CEN, margineciliato. Hypanthium 2-7 cmlongum, pilos CTES). Mun.Bon Jesus da Lapa: 14km a lestedo rio longos valde disperse gerens. Calyx 4-5 mm Correntena estradade SantaMaria da Vitoriaa Bom longus,pilis longis cupreisplus minusvedense Jesusda Lapa, 13°23'S,44°05'W, 450 m,9-V-1982, vestitus.Vexillum 7-9 mm longum x 8-10 mmlatum, Valls& al. 6785 (CEN, CTES). Ceará. Mun.Missão velutino-aurantiacum,supra subtusque lineis Velha:19,7 km a lestede Baibalha,camino a Missão rubescentibusornatum. Fructus biarticulatus Velha,7°15'S, 39°10'W, 22-IV-1987, Valls&al. 10921 paxillo usque ad 30 cm longo, isthmo6-9 cm (CEN, CTES). Minas Gerais.Mun. Coronel Murta: longo,articulis 10-13 mmlongis x 6-7 mmlatis, 8,2 kmS do rioJequitinhonha, na estradade Coro- epicarpio densepilorum minutorum obtectis, iis nelMurta a Araçuai,16°41'S, 42°10'W, 23-IV-1982, decisis reticulatio insignispericarpii conspi- Valls & al. 6709 (CEN, CTES). Piauí. Mun. cua. Piracuruca:margem esquerda do rioGenipapo, no ladonorte da BR-222,km 26,4, 3°59'S, 41°24W, Valls Holotype:BRAZIL. Pernambuco.Mun. São &al. 11022(CEN, CTES). Lourençoda Mata,Engenho São Bento,Tapera, 15- IV-1967, Gregory& Krapovickas12946 (IPA). Common name, "amendoim de caracará" Isotypes:CEN, CTES, EAC, G,GH, K, LIL, MO,NY, (Valls & al. 7287). P,RB, SI, SP,US.

Geographic distribution.This species was Annualor biannualplant. Taproot withweak collected in the state of Bahia on both sides of branchings.Mainstem erect, 5-15 cm long,but the São Francisco River, where it grows in capable of attaining 35 cm in length when open areas in presence of the palm growingin competitionwith tall grasses. Late- Copernicia cerifera. It was also collected in ral branchesprostrate, up to 70 cm long. Stem the north of Minas Gerais, near the angled in the young parts, cylindricaltoward JequitinhonhaRiver, in the southof Ceará and the base, with more or less dense, coppery the northof Piauí. hairs ca. 1.5 mm long. The mainstemwithout flowers.On the primarylateral branches,the Obs. The type specimen, Blanchet 2669, vegetative and reproductive branches consists of various very young individuals, alternate,with a predominanceof the latter, analyzed in detail by Krapovickas & Rigoni which are more abundant on the secondary (1957: 449). We have not personallycollected branches. Leaves tetrafoliolate,those of the thisspecies at thetype locality. At present,the mainstemalways largerthan those of the late- above cited specimens are those that most ral branches. On the mainstem, the fused closely approximatethe type of A. pusilla for portionof the stipules 10 mm long, the free the glabrous upper surface of leaves, with parts acicular, 15-20 mm long; petiole 30-42 orderlybristles on the lower surface,and for mmlong; rachis 10-12 mmlong; leafletselliptic the transverseline of hairs at the firstpair of or somewhatobovate, acute toward the ends, leaflets, characters clearly visible on the theapical pair up to 36 mmlong x 22 mmwide, Blanchet 2669 specimen. thebasal pair up to 33 mmlong x 20 mmwide.

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Fig.1 7. ArachisDardant A, schematic ofthe plant; B, leaffrom mainstem; C, leaf from lateral branch (G. 12946).

On the lateral branches,the fused portionof of hairs, which are presentat the level of the the stipules 5-6 mm long, the free parts apical pair of leaflets. Leaflets with upper acicular, 10-13 mm long; petiole 15-21 mm surfaceglabrous, the lower surfacewith long long; rachis8 mmlong; leafletsrounded, broad, scatteredhairs and withsmall, scarcelyvisible obovate, the apical pair up to 25 mm long x 19 adpressed hairs; veins weakly markedon the mmwide, the basal pair up to 20 mm long x 15 lower surface and with faint marginal vein; mmwide. Stipuleswith scattered long coppeiy marginciliate. Flowers along the lengthof the hairs towardthe base of the fused portion;the prostrate branches, in short 3-4-flowered free portion3 -veined, with glabrous surfaces spikes. Hypanthium2-7 cm long, with very and the margin longly citiate. Petiole scatteredsilky hairs. Calyx bilabiate,with long canaliculate, dorsal surfacewith long coppery coppery,more or less dense hairs,upper lobe hairs, similar to those of the stem; canal 4-5 mm long, lower lobe subfalcate,acute, 4- glabrousor withsome shorthairs, open, almost 4.5 mmlong. Standard7-9 mm longx 8-10 mm flat toward the apex, the separation fromthe wide, yellow-orange,with reddish-violet lines rachiscanal usually withoutthe transverse line on both faces; wings yellow, 6-7 mm long.

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Fruitsubterranean, Inarticulate; peg usually 10- 11001 (CEN, CTES); Itapagé,9,3 km o oesteda 14 cm long but up to 30 cm long; isthmus6-9 estradapara Pitombeiras entre Umirim e Itapagéao cm long; articles well-developed, 10-13 mm longoda BR-222,3°45'S, 39°32'W, 28-IV-1987, Valls long X 6-7 mmwide, coveredwith a dense coat & al. 11006 (CEN, CTES); Sobral,14,2 km a oestedo of diminutive hairs which when removed rioJaibaras e 900 m a lesteda estradapara Jaibaras reveals a somewhat prominentreticulum; the e Cariréao longoda BR-222,3°44'S, 40°28W, 29-IV- peg is insertedon the dorsal side, toward the 1987,Valls & al. 11008 (CEN, CTES); Frecheirinha, base of the articlesand the isthmusis borneat margemoeste do rio Caiçara ao longoda rodovia the apex on the ventralside, forminga small BR-222(lado norte)3°46'S, 40°47'W, 29-IV-1987, beak. 2n=20 chromosomes (Fernández & Valls& al. 11016 (CEN, CTES). Maranhão.Pastos Krapovickas 1994). Bons,27 kma NE de PastosBons e 2 kma SE de Orozimboao longodeBR-230, 21-111-1985, Valls & Selected additional material:BRAZIL. S/l., al. 8440 (CEN, CTES). Pará. Ilha do Marajó,Rio Löfgren77 (R); Löfgren128 (R). Alagoas.Porto Real Camará,fazenda Sta. Rita,Retiro Pau Grande,III- do Colegio,VIII-1929, Deslandes 69 (SP). Bahia. 1950,Lima 72 (IAN,SI, US). Paraíba. Mun.Sousa: Barreiras,3 1-ÏÏI-1983, Valls & al. 7039(CEN, CTES); 900,6°47'S, 38°12'W, 23-IV-1987, Valls & al. 10939 llkmNdeRiachãodasNeves,llo40'S,44o55rW, 1- (CEN, CTES); Sousa,3,7 km W da pontesobre o rio IV-1983,Valls & al. 7055 (CEN, CTES); 12km N de Peixeao longoda BR-230,6°47'S, 38°7'W, 180 m 23- Riachão das Neves, l-IV-1983,Valls & al. 7062 IV-1987, Valls & al. 10940(CEN, CTES); Sousa,400 (CEN, CTES); id.,7063 (CEN, CTES). Ceará. Mun. m W da pontesobre o rio do Peixe, na chegadaa SantaQuitéria: CE-032,km 113, Sobral-Caninde, 15- Aparecida,6°47'S, 38°6'W, 23-IV-1987, Valls & al. VI-1979,Coradin & al. 1963(CEN, CTES); Baturité, 10945(CEN, CTES); Sousa,7,7 km E da pontesobre 11 -IV- 1909, Ducke 1970 (MG, RB); Jucas,6°32'S, o riodo Peixe em Aparecidaao longoda BR-230, 39°32'W,250 m, 1-III-1972,Pickersgill RU72-259 6°46,5'S,3802W, 23-IV-1987, Valls & al. 10946(CEN, (CEN, CTES, IPA, US); Sobral,Praça Fernando CTES); Pombal,3,5 km W da pontesobre o rio Mendes,12-IV-1967, Gregory & al. 12943 (CEN, Piranhasna BR-230, entreAparecida e Pombal, CTES, EAC, GH,MO, NY, US); id., 12945(CTES, 6°43'S,37°50W, 24-IV-1987, Valls & al. 10947(CEN, EAC, G, GH, IPA, LIL, MO, NY, US); 50 kmE de CTES); Pombal,1 1,6 km NE de Pombal,BR-427, Sobral,12-IV-1967, Gregory & al. 12939(CEN, CTES, 6°45'S,37°42'W, 210 m, 23-IV-1987, Valls & al. 10953 EAC, GH,MO, NY, US); 40 kmE de Sobral,12-IV- (CEN, CTES); Paulista,cerca de 700 m da divisa 1967,Gregory & al. 12941(CEN, CTES, EAC, G,GH, Paraiba/RioGrande do Norte,BR-427, 6°40'S, K, MO, NY, P, US); 12km NW ofSenador Pompeu, 37°31 W, 230m, 23-IV-1987, Valls & al. 10960(CEN, 5°40'S,39°25'W, 3-III-1972, Pickersgill & al. RU72- CTES); s/l.,regiões secas, 19-IV-1959, Moraes 2088 282 (CEN, IPA,US); 5 kmfrom Campos Sales on the (NY, P, US); Souza,Posto Agricola de São Gonçalo, road to Crato, 7°10'S, 40°20'W, 19-11-1972, varzea,5-III-1936, Luetzelburg 26889 (CEN, IPA, "amendoimcarcará," Pickersgill RU72-186 (CEN, NY). Pernambuco.Fazenda Cachoeirinha(mim. CTES, US); Brejo Santo,2,8 kmS de BrejoSanto, Mirandiba),km 447 da BR-232,entre Salgueiro e BR-116, 7°3 l'S, 39°00W,350 m, 2 l-IV-1987,Vails & BomNome, 8°1'S, 38°49'W, 20-IV-1981, Valls & al. al. 10912(CEN, CTES); BrejoSanto, 28 kmS deBrejo 6215(CEN, CTES); Tapera,17-VI-1932, Pickel 303 1 Santo,BR-1 16, 7°3 l'S, 38°«)^, 21-IV-1987,Vails & (CEN, P, US). Mun. São Lourençodo Mata: Eng. al. 10913(CEN, CTES);Barro,km461,5daBR-116, São Bento,6-V-1967, Andrade-Lima 67-5006 (IPA); ao longoda zonaurbana de Barro,7°1 l'S, 38°47'W, SerraTalhada, Hotel Planalto, 8-V-197 1, Heringer & 330 m,22-IV-1987, Valls & al. 10936(CEN, CTES); al. 638 (CEN, CTES, IPA,UB); EstradaSalgueiro- Aracati,1,6 km a lestedo inicioda pontesobre o rio Carqueja, Serra de São Gonçalo, 23-V-1971, Jaguaribe,BR-304, 4°34'S, 37°47'W,26-IV-1987, Heringer& al. 865 (CEN,IPA, UB). Piauí.Dry shady Valls& al. 10988(CEN, CTES). Mun.Beberibe: 2,2 places fromBrejo Grande,Ceará, to the Cityof kmNW do entroncamentodas rodoviasBR-304 e Oeiras,m-1839, Gardner 2091 (G,K,NY,P);27kma BR-116, entre Aracati e Cristais,4°35'S, 38°13'W, 26- Lestede Oeiras,BR-230, 8-IV-1983, Valls & al. 7 197 IV-1987,Valls & al. 10992(CEN, CTES); Caucaia,4 (CEN, CTES); km3,3 da BR-407na saida de Picos kmo oestedo rioCauipe entre Guarani e Catuana, paraPetrolina, 9-IV-1983, Valls & al. 7215 (CEN, BR-222,3°41'S, 38°53'W,28-IV-1987, Valls & al. CTES); Corrente,10°26'S, 45°10'W, 3-IV-1983, Valls

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& al. 7086(CEN, CIES); Floriano,9 km a Lestedo rio opportunityto observethe plant in nature. Parnaiba,BR-230, 7-IV-1983, Valls & al. 7166(CEN, CTES); Angical,faixa de dominioleste da BR-343a Obs. Arachis Dardani is, genetically, a 100m ao norteda entradada EstaçaoExperimental very isolated species. Hybrids were only de Angicalda Embrapa,22-III-1985, Valls & al. 8471 obtained when it was crossed with two (CEN,CTES); km15 1,7 da rodoviaBR-3 16 cerca de 8 accessions of A. paraguariensis ssp. kmao nortede ElesbãoVeloso, 26-III-1985, Valls & paraguariensis, of section Erectoides. The al. 8503 (CEN, CTES); 27 kma oese de Oierasao pollen fertilitywas verylow in both cases (0.6 longoda rodoviaBR-230, 26-III-1985, Valls & al. and 2%). 8505 (CEN, CTES); 16 kmao sul de Florianona In crosses with A. Macedoi, of section Rodovia PI- 140 (distancia medida da estaçao Extranervosae,the hybridobtained was also rodoviaria),27-III-1985, Valls & al. 8509 (CEN, highlysterile. CTES); Entradanorte de Correntena BR-135,28-111- A special situationarose in crosses between 1985,Valls & al. 8522(CLN, CTES); Trevode acesso two distant accessions of this species, one a Amarantena rodoviaBR-343, 22-111-1985, Valls & fromSobral (CE) and the otherfrom Tapera al. 8469 (CEN, CTES); 9 kma lesteda pontesobre o (PE). The reciprocal hybrids in both cases rio Parnaibaao longoda rodoviaBR-230, 22-III- gave pollen stains of 20.5%, a verylow result, 1985,Valls & al. 8462(CEN, CTES). Rio Grandedo especially in light of the fact that these two Norte.35 kmfrom Currais Novos on roadto Cerro accessions show no significantmorphological Cora,6°10'S, 36°25'W,400 m,"amendoim bravo," differences. 2 l-m-1972,Pickersgill RU72-363 (CEN, CIES, IPA, US); BR-226,Santa Cruz-Currais Novos, km 4, 2 1- We dedicate this species to Dr. Dárdano de Vn-1980,Coradin & al. 3224(CEN, CTES); Faz. Cas- Andrade-Lima,expert on theflora of the "nor- trobetween Acari and Cruzeta, 36°40'W, 6°22'S, 450 deste,"who accompanied us duringthe search m,23-III-1972, Pickersgill & al. RU72-374(CEN, forthe material that we designatedas thetype. CTES, IPA,US); Caicó,24,7 km a SE de Caico,BR- 427,6°32'S, 36°56W, 200 m,24-IV-1987, Vails & al. VI. Sect. Caulorrhizae Krapov. & W.C. 10963(CEN, CTES); Caicó,km 17 daBR-427 a 10km Gregory nov. secí. de Acari em a Currais direção Novos, 6°21'S, Fig. 7 36°37W,210 m, 14-IV-1987, Valls & al. 10968(CEN, CTES); CurraisNovos, 1 5 kmNE de CurraisNovos, BR-427,6°12'S, 36°24'W,24-IV-1987, Valls & al. Sect. CaulorhizaeKrapov. & W.C. Gregory,in 10972 (CEN, CTES); CurraisNovos, 2,7 kmE de W.C. Gregory,M.P. Gregory,Krapovickas, Smith & Riachuelo,naBR-304, 5°49'S, 35°48'W, 25-IV-1987, Yarbrough,in Peanuts- Cultureand Uses: 92, Valls & al. 10974(CEN, CTES); Mossoró,4,8 ao N 1973, nomennudum. Krapovickas, Agricultural de Mossoró,BR-304, 5°7'S, 37°20'W, 30 m,26-IV- Genetics.Selected Topics: 137, 1973. 1987, Valls & al. 10981 (CEN, CTES). Sergipe. Maroim,X-1948, Moreno (SP 64272). Plantaeperennes. Radix non incrassata.Rami procumbentesad nodos radicantes.Caules cavi. Geographie distribution.This is a species Folia quadrifoliolata. Fructus subterraneus restrictedto the northeastregion of Brazil, biarticulatuspericarpio laevi. where it grows mostly in the wooded "caatinga," frequentlyin low-lyingareas with Typussectionis : Arachis repens Handro "carnaúba" palms. The southernmost collectionwas made in Barreiras (12°10'S), in Perennial plants. Taproot, without the state of Bahia. thickenings.Branches spreading,procumbent, The specimencollected on Marajó Island, in rootingat the nodes. Stems hollow. Stipules the mouthof the Amazon River (Lima 72), we with the margins free to the base. Leaves assign to this species with some reservations, tetrafoliolate.Flowers along the lengthof the due to the fact that we only had access to the branches. Well-developed hypanthium,up to herbarium specimen and did not have an 10 cm long or more. Standard commonly

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) yellow, expanded. Fruit subterranean, 12 mm wide, apex mucronate;the apical pair biarticulate; pegs short,vertical or inclined; always somewhat larger than the basal pair. isthmusshort, up to 5 cm long; articlessingle- Stipules glabrous or with some small hairs, seeded; pericarp smooth. 2n=20 adpressedtoward the base, marginbarely ciliate, chromosomes. withshort hairs. Petiole and rachiscanaliculate, with very scattered adpressed hairs to Geographicdistribution. This sectiongrows subglabrous, very rarely with some bristles. in the borderarea betweenthe Brazilian states Pulvinussomewhat villous. Leafletswith upper of Goiás, Bahia and Minas Gerais, reachingas surfaceglabrous; lower surface with very small far as the Atlantic coast, where the type adpressed hairs barely visible to subglabrous, specimen of A. Pintoi was collected. midveinsomewhat conspicuous, the restalmost smooth,margin very slightly marked, with some adpressedhairs or glabrous,with scarce short Key fordistinguishing the species bristles.Inflorescences axillary, few-flowered, the veryshort axis coveredby the base of the stipules.Hypanthium 3.5-1 1 cm long,with more A. Leafletsmore than 2.5 timeslonger than wide. or less long, adpressed hairs, somewhatmore Plantswithout bristles, except rarely a fewon the villoustoward the top. Calyx bilabiate,ca. 7 mm petiole. long,covered with adpressed hairs and withlong scatteredbristles. Standard 8-13 30.A. repens entirelyyellow, mm long x 16-17 mm wide; wings and keel A'. Leafletsless thantwo times longer than wide. yellow.Fruit subterranean, biarticulate; peg 5 cm Bristlespresent on the stems,stipules, petiole long; isthmus1.7-3.5 cm long; articles8-13 mm and rachis,frequently also on theunderside of long x 5-6 mm wide, witha small apical beak, theleaflets. pericarpsmooth. 2n=20 chromosomes(Conagin 31.A. Pintoi 1962:353).

Holotype: BRAZIL. Minas Gerais. Estaçao 30. Arachis repens Handro EngenheiroDolabela (E.F.C.B.), margemdo Rio Cultivadaem R. Fig. 2,30 Jequitahy. Deodoro,D.F., Janeiro, 20-XI-1941,Ramos Otero 2999 (SP 46152!).

Handro,Arq. Bot. Estado São Paulo 3: 180-191, Selectedadditional material: BRAZIL. Minas táb.47, 1958.Conagin, Bragantia 21: 353,est. 7, figs. Gerais. Lagoa Santa, Warming2998 (G); Faz. 6A, 14, 1962.Conagin, Bragantia 22: 128,fig. 1A Jatoba,rio Jequitaí, 45 kmfrom Pirapora to Jequitai, (2n=20),1963. 21-IV-1982,orange fl., Vails & al. 6673(CEN, CTES); id.,yellow fl., Vails & al. 6674(CEN, CTES). Trailing perennial.Root branched,without enlargements. Branches widely spreading, Cultivatedmaterial: BOLIVIA. Pando. Cobija, with adventitiousroots at the nodes. In one- main plaza, ll-XI-1988, Williams809 (CTES). year-old plants, the mainstem without BRAZIL. Amazonas.Campus Aleixo Manaus, cult, branches on the upper nodes, thin and flexi- in sandysoil for lawn effect, l-X-1973, Bisby 1265 ble, bending toward the soil. Internodes3-5 (K); id.,28-Vm-1976, Davis &al. 60389(UEC). Mato cm long, somewhat angular, with adpressed Grosso do Sul. Mun.Douradina: Bocajá, BR-163, whitish,caducous hairs. Leaves tetrafoliolate. cult. 15-11-1933,Hatschbach & al. 59057 (CTES, Stipules withthe fused portion6-10 mm long, MBM). Minas Gerais. SeteLagoas, Estac. exper., wide, with imbricated margins completely cult.proc. Eng. Dolabela,2-V-1961, Gregory & al. envelopingthe stem; the free portionusually 10538(CTES, LIL). BeloHorizonte, Jardim Botanico, shorter,narrowing abruptly from the base. cult.?,2-II-1978, Krapovickas & al. 33403 (CEN, Petiole 7-40 mm long; rachis 3-7 mm long. CTES, F, G,MO, PAMG,SI, UC, US); Pirapora,cuit. Leafletselliptical to obovate,20-35 mmlong x 8- HotelCanoeiros, 21-IV-1982, Valls & al. 6671 (CEN,

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CTES). Rio de Janeiro. Deodoro, cultivadana Gregory,M.P. Gregory,Krapovickas, Smith & Seção Experimentalde Agrostologia,origem: Yarbrough,inPeanuts - Cultureand Uses: 121, 1973, margemdireita do rioJequitai, Eng. Dolabela,Mi- nomennudum. This refers to the specimen Gregory & nas Gerais, ll-XII-1941, Otero3059 (RIZ). Rio Krapovickas12787. Grande do Sul. São Gabriel,Estac. Experim. cult., proced.:São Paulo,22-V-1968, Hammons & al. 365 Herba perennis. Caulis reptantesad nodos (CTES). São Paulo. Institutode Botanica,cult., radicantes,teretes, cavi, setulascaducas gerentes. procedenciaignorada, 12-III-1957, Handro 686 (SP, Stipulaedorso glabrae, margine ciliatae. Folíola US); Institutode Botanica,cult, (same population obovata, epiphylloglabro margineindistincto, as Handro686), Eiten2549 (K, UB); id.,2746 (K); hypophyllosparsim setuloso, margine pilis sericeis Campinas, InstitutoAgronòmico, cult., proc.: etsetulis paucis brevibus instructa. Hypanthium 3.5- Estac. Exper.Monte Alegre do Sul, SP, V-305,X- 9.5 cm longum,villosum. Calyx 5-6 mm longus, 1955,Conagin 1 (IAC 18127);Matao, Inst. Pesqui- villosus,setulosusque. Corolla luteď, vexillo 11 mm sas I.R.I., cult.,5-VI-1968, Hammons & al. 500 longoX 13 mmlato. Fructus biarticulatus paxillo 5- (CTES); Caraguatatuba,Barra do rioJuqueriqueré, 32 cmlongo, isthmo 1-8.5 cm longo, articulis 11-14 PortoNovo, cult., 2-VI-1968, Hammons & al. 467 mmlongisx 6-7 mm latis, pericarpio laevi. (CTES); Araras,Usina de S. João,en césped,cult.?, 17-XI-1978,Mroginski 796 (CIES, SP). Holotype:BRAZIL. Bahia. Cruz das Almas, IPEAL, cult,procedente de Bahia, rio Jequitinhonha, cerca de Common names, "amendoim rasteiro" Belmonte,Bahia, en bancos de arena,de semillasco- (Otero 3059; Otero 1951: 178); "tepe colom- leccionadaspor GeraldoC.P. Pintoy Paulo D.T. biano" (Williams 809). Alvim,en abril1954, 15°52'S, 39°6'W, 31-III-1967, Gregory& Krapovickas 12787 (CEN). Isotypes: CTES, Geographic distribution.Apparently, the G,GH, K, UL, MO,NY, P, RB, SI, SP, US. typical formof this species occupies a very small range, on the Jequitai River, between Perennial plant. Taproot, without Jequitaiand EngenheiroDolabela, towardthe enlargements. Stems erect at first, later northcentral part of Minas Gerais. Due to its prostrate,rooting at the nodes, cylindrical, ability to compete with weeds and to forma angled whendry, with rigid caducous bristles. compact lawn, its cultivation has been Stems distichous. Leaves tetrafoliolate. promotedand has expanded widely. Stipules with the portionfused to the petiole 10-15 mmlong x 3 mmwide, with rigid bristles Obs. Arachis repens has a stronggenetic on the back; the freeportion 10-12 mm long x affinitywith A. Pintoi, of the same section, 2.5 mm wide at the base, with prominent and they produced hybrids with a high longitudinal veins, both surfaces glabrous, percentage of stained pollen (86.8%). The marginwith silky hairs. Petiole up to 6 cm long, morphological differencesbetween them are canaliculate, with rigid bristles on the back, quite notable, especially between the two glabrous canal with finesilky hairs along the parentsutilized in the crosses. margins.Rachis 10-15 mm long, canaliculate, In crosses between different sections, with a few bristles on the back. Pulvinus hybridswere only obtainedwith A. major and pubescent.Leaflets obovate, apical pair up to A. paraguariensis ssp. paraguariensis, of 50 mmlong x 32 mmwide, basal pair up to 45 section Erectoides . The fertilityof these mm long x 28 mm wide; glabrous upper hybridswas verylow. surface with scarcely marked margin; lower surfacewith scattered bristles, more abundant 31. Arachis Pintoi Krapov. & W.C. on the external semi-lamina (exposed in the Gregory nov. sp. foldedleaf) of thebasal leaflets;margin lightly Figs. 2,31; 18 marked, with silky hairs and some short bristles. Inflorescences4-5 -floweredaxillary

A. Pintoi Krapov. & W.C. Gregory,in W.C. 'Errorin original, appeared as aurantiaca.

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Fig.18. Arachis Pintoi : A, branch; B, schematicofthe plant (G. 12787).

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spikes,very short, covered by thefused portion Berilo, 17-VIII-1982,Werneck 34 (CEN). Mun. of the stipules. Flowers sessile, protectedby Araçuai: 1 km apos o rio Araçuai na estrada two bracts,the basal 12 mmlong x 5 mmwide, Araçuai-Virgemda Lapa, 28-IV-1982,Valls & al. with silky hairs along the midveinand on the 6727(CEN, CTES); 8 kmde Araçuaina estradapara margin,the upper bract bifid, 10mmlongx2.5- FranciscoBadaro, 28-IV-1982, Valls & al. 6728 3 mmwide, with silky hairs along thetwo veins (CEN,CTES). Mun.Presidente Juscelino: 3 km W of and on the margin.Hypanthium 6.5 (3.5-9.5) RioParauna, 85 kmS ofDiamantina, BR-259, 30-IV- cm long,with long silkyhairs. Calyx bilabiate, 1982,Valls & al. 6740(CEN, CTES); id.,Valls 6741 with silky hairs somewhat more rigid than (CEN, CTES). Mun.Unai: ribeirãoLagoa do Mel, those of the hypanthiumand with bristles; afluentedo rio Preto,fazenda Palmital, 16°8'S, largerlobe 5 mmlong, tridentate; upper lobe 6 47°13'W,850 m,Werneck 1 & 2 (CEN); id., 18-11- mm long, narrow, falcate. Corolla yellow. 1981,Valls & al. 5895(CEN, CTES). Standard 11 mm long x 13 mm wide, with yellow veins; wings 8 mm long x 6 mm wide; Geographie distribution.The typical form keel 6-7 mmlong, falcate. Four oblonganthers of this species is known only fromthe type basifixed, four spherical anthers dorsifixed, locality.The foragequalities of thisspecies has and one staminode.Fruit Particulate; peg 5- lead to itsdistribution for cultivation, apparently 32.5 cm long, lightlypubescent toward the with great success. base; basal article somewhatsmaller than the distal segment,11-13 mmlong x 6-7 mmwide; Obs. 1. We have includedin the description isthmus1-8.5 cm long; distal article12-14 mm of this species material regarded as similar, long x 7 mm wide; pericarp smooth,covered albeit with reservations.This is a group that with delicate hairs that retainthe soil. 2n=20 deserves special study. This additional mate- chromosomes (Fernández & Krapovickas rial has some adpressed hairs, barely visible 1994). on the undersideof the leaflets,in additionto the bristles.In contrastthe type specimenhas Additional material: BRAZIL. São Paulo. only the bristles, lacking the trichomes. Campinas,Instituto Agronómico, Fazenda Santa Nevertheless,on the veryyoung leaves of the Elisa,adventicia, 1-II-1984, Valls & al.7529(CEN, type specimen, small adpressed hairs, CTES); id., 21-1-1987,Krapovickas & al. 40986 evidently caducous, are found on the (CIES). undersideof the leaflets.

Selectedrelated material: BRAZIL. Bahia. Mun. Obs. 2. This species is very much like A. Bom Jesusda Lapa: 74 kmW ofrio São Francisco repens forits rootingstems, hollow when dry, on roadfrom Bom Jesusda Lapa to SantaMaria da and for its yellow flowers. They are Vitoria,8-V-1982, Valls & al. 6784 (CEN, CTES). differentiatedbecause in A. Pintoi the leaflets Goiás. 120 km além de Formosa,na base de are larger,with a roundedtip, and there are afloramentocalcareo, prostrada e sendo pastada bristlespresent on the stem, stipules, petiole porgado, 22-X-1965, Heringer 1073 1 (CTES, NY); and undersideof the leaflets. Vale do Paraná,Rio dosMacacos, 5-II-1967, Duarte 10313 (RB). Mun. Formosa: 12 km N-NW of Obs. 3. Crosses with A. repens produced Formosa,then 2 km into fazendaJenipapo, rio highly fertilehybrids (86.8%). In contrast, Bandeirinha,12-V-1982, Valls & al. 6791 (CEN, when crossingA. Pintoi withA. major and A. CTES); id.,Valls & al. 6792 (CEN, CTES). Minas paraguariensis ssp. paraguariensis of the Gerais. 15kmN de Montalvania,14°20'S, 44°20'W, sectionErectoides, and withA lignosa and A. 550 m, edge of lake, 18-111-1972,Anderson & al. Rigonii of the section Procumbentes, the 37175(K, NY, US); SerraCabral, 2 kmNof Joaquim hybridswere highly sterile. Felicio,650 m,5-III-1970, Irwin & al. 26986 (UB). Mun. FranciscoBadaro: beira do córregona saida We dedicate this species to Ing. Agron. da cidade para Berilo, 19-VII-1982,Werneck 17 Geraldo C.P. Pinto, who collected and (CEN); 3 km de FranciscoBadaro na saida para cultivatedit.

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VII. Sect. Procumbentes Krapov. & W.C. extended,horizontal, shallow; articlessmooth. Gregory nov. sect. 2n=20 chromosomes. Fig. 7 Geographic distribution. This section extendsitself along the Paraguay River, from Sect. Erectoidesser. Procumbensae Krapov. & Concepción, on the Tropic of Capricorn, W.C. Gregory,in W.C. Gregory,M.P. Gregoiy, towards the north,flanking the Pantanal in Krapovickas,Smith & Yarbrough,in Peanuts- Mato Grosso on the south and the north,and Cultureand Uses: 93, 1973,nomen nudum. then expands westwardas far as Santa Cruz de la Sierra, in Bolivia. Its species generally Perennes.Radix polaris. Rami procumbentes, live in soils thatare periodicallyflooded. rarissimedecumbentes. Folia quadrifoliolata, saepe glabrae vel subglabrae. Vexillum Obs. The positionof its species is intriguing. aurantiacum vel luteum, supra lineis In termsof theirgrowth habit, A. Rigonii and rubescentibusornatum. Fructus subterraneus, A. lignosa appear to pertain to the section biarticulatus,paxillo perlongo, paulum sub humo Arachis and A. appressipila to the section horizontaliterprocurrenti, pericarpio laevi. Erectoides, but the genetic behavior of these three species (see pages 173 and 177 in Typussectionis : Arachis Rigonii Krapov. & Gregory, M.P. & Gregory 1979) shows a W.C. Gregoiy greater affinitywith the section Erectoides. Nevertheless,it is advisable to considerthis as Perennialplants. Taproot, without thickenings. an independentsection given that the fertility of Branches procumbent, rarely decumbent. the hybridswith species of other sections is Leaves tetrafoliolate,generally glabrous or always verylow. Furthermore,there is a very subglabrous.Stipules with free margins. Flowers notable cytological difference.In the species alongthe length of the branches. Well-developed of section Procumbentes, the presence of a hypanthium.Standard expanded, yellow or pair of chromosomes with a dot-shaped orange, with reddishlines on the front.Fruit satellitehas been detected,which is exclusive subterranean,Particulate; peg and isthmusvery to thissection (Fernández & Krapovickas1994).

Key fordistinguishing the species

A. Leafletson thelateral branches with a length/widthratio less than 2: 1. B. Leaves small,leaflets of the lateralbranches up to 19 mm x 10 mm,glabrous. Stems without adventitiousroots. Standard orange. 32. A. lignosa B'. Leaveslarger. C. Leafletsglabrous, up to 35 mmx 19mm. 33.A. Kretschmeri

C' Lowerleaf surface with diminutive adpressed hairs.

D. Leafletsup to30 mmx 17 mm.Corolla yellow. Stems without adventitious roots. 34.A. Rigonii

D'. Leafletsup to24 mmx 18 mm.Standard orange. Stems with adventitious roots. 35. A. chiquitana

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A'. Length/widthratio greater than 2.5:1. E. Length/widthratio of 2.5-3.5: 1. F. Lateralbranches procumbent, with leaflets up to 43 mmx 13mm, glabrous. Stipules violet at thebase. 36.A. matiensis F. Lateralbranches decumbent, with leaflets up to50 mmx 16 mm,with the upper surface glabrousand theunderside with adpressed hairs. Stipules green. 37. A. appressipila

E'. Length/widthratio from 4 tomore than 7:1. G. Stipuleswith very short bristles. Peg thick,hollow. 38.A Vallsii

G'. Stipuleswith long bristles or without them. Peg slender,compact. 39.A. subcoriacea

32. Arachis lignosa (Chodat & Hassl.) part of the stipulesmeasure 4-6 mm long x 4 Krapov. & W.C. Gregory nov. comb. mmwide, the free tips acute, up to 1 1 mmlong x 1 mm wide at the base. Petiole to 25 mm Figs. 2,32; 19,A-B up long, semi-terete or slightly canaliculate. Rachis 5 mm long. Leaflets obovate or A. prostrata Benth. var. genuina f. lignosa elliptical,obtuse, the apical pair up to 19 mm Chodat& Hassl.,Pl. Hassl.2: 448, 1904,"pr. Para- longx 10 mmwide, thebasal pair up to 17 mm guay,Dec. Hasslern. 6515; in campispr. Concep- long x 9 mm wide. Stipules glabrous or with ción,Sept. Hasslern. 7476." short, soft hairs on the margin; petiole A. marginataGardn. var. lignosa (Chodat & glabrous;pulvinus glabrous or withsome small Hassl.) A. Chev.,Rev. Int.Bot. Appl.Agrie. Trop. hairs; leaflets glabrous on both surfaces, 13(146-147): 762, 1933.In makingthis combination, margin somewhat marked on the back, also Chevalieronly took into accountthe specimen glabrous. Inflorescencesaxillary, 3 -flowered, Hassler7476. bornealong the lengthof the lateral branches, A. prostrata Benth. var. lignosa Chodat & the axis covered by the stipules. Flowers Hassl., Hermann,Agrie. Monogr. USDA 19: 10, appear from between two thin bracts, the 1954,nov. stat. in synom. lower acute and entire, the upper bifid. A. lignosa (Chodat& Hassl.) Krapov.& W.C. Hypanthium5 (3.5-9) cm long,subglabrous or Gregory, in W.C. Gregory, M.P. Gregory, withsilky hairs. Calyx bilabiate,subglabrous or Krapovickas,Smith & Yarbrough,in Peanuts- lightlypubescent, occasionally with very few Cultureand Uses: 121, 1973,women nudum. bristles; wide lobe 7 mm long, narrow lobe falcate,7 mmlong. Standard 1 1 mm long x 12 Perennial plant, taproot without mm wide, yellow in the centerwith a broad enlargements.In youngplants, the collar of the orangemargin, wings 7 mmlong, yellow. Fruit plant is found some 5 cm deep; in mature Particulate; peg horizontal, parallel to the plants, the bases of the branches are buried surfaceof the soil, shallow, up to 30 cm long; some 20 cm, but withoutforming rhizomes and isthmusca. 3 cm long; articlessome 10-12 mm usually withoutadventitious roots. Mainstem long x 6-7 mm wide, denselyvillous, pericarp very short,erect. Lateral branches prostrate, smooth. 2n=20 chromosomes (Smartt & 10-30 cm long,ordinarily with short internodes, Gregory1967, GK 10598). 10-15 mmlong (up to 25 mm) somewhatangu- lar, lightlypubescent, without bristles. Leaves Lectotype:PARAGUAY. Concepción.Hassler tetrafoliolate.On thelateral branches the fused 7476 (G!). Isotypes:BM!, LIL!, NY!, P!, UC!. The

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Fig.1 9. Arachis lignosa: A, schematic ofthe plant; B, leaf(G. 1 0598). A. pseudovillosa-. C, leaf; D, schematicofthe plant (G.9625). otherspecimen mentioned by Chodat& Hassler: Rojas 14074(SI); id.,20-1-1949, Ramírez 63 (SI); id., Paraguary,Hassler 65 13 (see Numericorrigendi) 20-1-1949,Rosengurtt B-5496 (CTES, LIL, MVFA, (G!), has rhizomesand correspondsto A. glabrata SI); EstanciaFonciere, 19-1-1949, Ramírez 363 (SI). var.glabrata. PresidenteHayes. PuertoMilitar, puente sobre el río Paraguay,frente a Concepción,10-XII-1989, Selectedadditional material; PARAGUAY. Con- Vanni& al. 1291(CTES). cepción. Concepción,22-1-1955, Meyer 18729 (LIL); id., 18-V-1961,Gregory & al. 10598 (CEN, Geographicdistribution. This species grows CTES, GH, LIL, MO, NY, US); id., 25-11-1968, along the Paraguay River, between Concep- Krapovickas& al. 14248(CTES, US); id.,16-V-1974, ción and PuertoFonciere, in the departmentof Schinini91 88 (CTES); id.,8-XII-1992, Nicora & al. Concepción, in Paraguay. Also, it is found in 9785 (CTES, SI); PuertoFonciere, 18-11-1948, PuertoMilitar, across fromConcepción, in the Stephens& al. SH 178 (LIL, NY); id., 18-11-1948, departmentof P. Hayes.

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Obs. The crosses of A. lignosa with pair of leaflets 19-21 mm long x 11-15 mm species of the same section made relatively wide, thebasal pair 17-18 mmlong x 9-1 1 mm fertilehybrids, for example, A. appressipila wide. In cultivationthe leaves may have larger (ca. 30%) and A. Rigonii (54.4%). On the parts: the freeportion of the stipulesup to 20 otherhand, the hybrids obtained with A major mm long x 5 mm wide at the base, the apical (0.1%) of section Erectoides and A. Pintoi pair of leafletsup to 35 mm x 19 mm and the (0.2%) of section Caulorrhizae were almost basal pair up to 30 mmx 15 mm.Leaves totally totallysterile. glabrousexcept for a fewhairs on thepulvinus, occasionally a few adpressed and scarcely 33. Arachis KretschmeriKrapov. & W.C. visible hairs on the canal toward the apex of Gregory nov. sp. the petiole. Margins of the stipules somewhat Figs. 2,33; 20 imbricated at the base of the fused part. Leaflets oblong or elliptical, occasionally somewhatwider on the distal third,apiculate, Herba perennis.Rami prostrati, basem versus upper surface smooth, lower surface with radices adventitiasemitientes. Caulis angulosus, somewhatprominent midvein; marginbarely glaber. Folia praeteraliquot pilos in pulvinulis markedor unmarked,usually with some widely omninoglabra, foliolis oblongis vel ellipticis, scatteredbristles. Axillary spikes very short, 3- apiculatis,epiphyllo laevi, hypophyllonervo me- 4-flowered,bracts glabrous. Hypanthium up to dio manifestiore,margine parum vel haud 10 cm long,more or less villous. Calyx almost distincto,setulis nonnullis instruēto. Hypanthium glabrous,may show a veryfew weak bristles; usque ad 10 cm longum,sat villosum.Calyx 5-7 upper lobe 6 mm long, lower lobe 7 mm long, mmlongus, fere glaber, setulas tenues poucas ge- falcate. Standardup to 18 mm long x 22 mm rente.Vexillum usque ad 18 mmlongum x 22 mm wide, witha yellow centerand orange toward latum,aurantiacum. Fructus biarticulatus paxillo the margin,with a few veryweak red lines on horizontāliglabro usque ad 45 cm longo,isthmo the upper surface. Fruit Particulate; peg 8-15mm longo, articulis 10-14 mm longisx6-7 mm glabrous,up to 45 cm long,horizontal; isthmus latis,apice paulo recurvo,pericarpio laevi. 8-15 mm long; articles 10-14 mm long x 6-7 mm wide, witha slightbeak, pericarp smooth Holotype:U.S.A. Florida.Fort Pierce. August21, with a heavy coat of hairs which strongly 1978. (KretschmerJr. s/n) submittedby W.C. retainsparticles of soil. Seed 1 1 mmlong x 5.5 Gregoiy3-30-79 (CTES 34805).The cultivated mate- mm wide. 2n=20 chromosomes(Stalker, data rialcame originally from Brasil, Mato Grosso do Sul, publishedin Kretschmer& Wilson 1988). 36 km fromMiranda on the road to Corumbá, FazendaBandeira, 20°7'S, 56°43'W; it was growing Additionalmaterial: BRAZIL. Mato Grosso do in water 4 to 8 inches deep, May 18, 1976, Sul. Mun.Aquidauana: BR-262, 23 kmW do trevo Kretschmer,Jr. & Rayman,cult. IRFL 2273, P.I. paraAquidauana, 7-IV-1986, Vails & al. 9907(CEN, 446898(Kretschmer & Wilson 1988). CTES); BR-262, 20,5 km W do acesso a Aquidauana,19-IV-1984, Vails & al. 7631 (CEN, Perennialplant. Branches prostrate,60 cm CTES); id.,5-IV-1986, Valls & al. 9889(CEN, CTES); long or more,with abundant adventitious roots id, 30-X-1986,Valls & al. 10404(CEN, CTES); 35km on the basal internodes. Stems cylindrical W de Aquidauana,19-IV- 1984, Valls & al. 7633 toward the base and angular toward the ends (CEN); 8 kmS da BR-262na estradaAquidauana- of the branches; internodes 2-4 cm long, Nioaque,7-IV-1986, Valls & al. 9917(CEN, CTES). glabrous. On the prostrate branches, a Mun. Corumbá:dist. Nabileque, Faz. São Bento, vegetative branch alternates regularly with Campo brejosojunto a mata de Carandá, 17-XI- two reproductive branches. Leaves 1977, Allem & al. 1318 (CEN, US); 40 km de tetrafoliolate. For the most part the fused Corumbá,camino a Porto Manga, 7-XII-1976, portionof the stipules 7-10 mm long, the free Krapovickas& al. 30007(CEN, CTES, GH,MO, NY, portion10-11 mmlong x 2-3 mmwide; petiole RB, SI, SP, US); 5 kmE de Monjolona estradade 20-30 mm long; rachis 7-10 mm long; apical Corumbáa Porto Manga, 12-X-1985, Valls & al. 9129

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Fig.20. Arachis Kretschmeri : plant (IRFL 2273).

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(CEN, CIES). Mun.Miranda: Fazenda Bodoquera, long x 9 mm wide (up to 23 mm x 14 mm). Secção Guaicunis,25-X-1978, Allem & al. 2147 Stipules with very small, adpressed hairs (CEN, CTES); 10 kmS da BR-262no acessoá sede widely scatteredon the lateral surfaces,and a Guaicurusda Faz. Bodoquena,19-IV-1984, Valls & lineof somewhatlonger hairs along theback of al. 7637(CEN, CTES); id.,6-IV-1986, Valls & al. 9904 the fused portion; margin weakly villous. (CEN,CTES). Petiole canaliculate,the back weaklyvillous to subglabrous;pulvinus villous. Upper surface of Cultivatedmaterial: ARGENTINA. Córdoba. the leaflets glabrous, the lower surface Manfredi,Estación Experimental INTA, cult. No. 64, subglabrous with scattered, diminutive de semillasdel holotipo, 14-111-1985, Krapovickas & adpressedhairs, margin and veins veryslightly al. 40053(CEN, CIES, LIL, MO,NY, SI, SP,US). marked,margin weakly villous. Spikes axillary, 2-3-flowered,axis very short,covered by the Geographicdistribution. This species grows base of the stipules.Hypanthium 6.6 cm (4-13 on the southernedge of the Gran Pantanal, cm) long,villous. Calyx bilabiate, 6 mm long, fromthe area around Corumbá to the vicinity pubescent, sometimes with an occasional of Aquidauana, in the stateof Mato Grosso do bristle. Standard entirelyyellow, ca. 10 mm Sul, in Brazil. It growsvery well on land thatis long; wings 5-7 mm long, yellow. Fruit subjectto periodicflooding. Particulate; peg horizontalup to 65 cm long; isthmussinuous, 2-17 cm long; articles 10-13 mmlong x 5-7 mmwide, the apical one slightly Obs. The collector of the is type specimen larger, somewhat flattenedat the apex and trials in Florida conductingadaptation (USA) terminating in a beak; pericarp lightly where the that he calls the species, "pantanal reticulate, almost smooth. 2n=20 shows as a peanut," good promise forageplant chromosomes(Krapovickas & Gregory1960). (Kretschmer& Wilson 1988). Holotype:BOLIVIA. SantaCruz de la Sierra.20- 34. Arachis Rigonii Krapov. & W.C. 11-1958,Krapovickas 9459 (LIL). Isotypes:BAB, Gregory CTES,SI. Fig. 2,34 Additionalmaterial: BOLIVIA. Santa Cruz. San- taCruz de la Sierra,4-IV-1959, Gregoiy & al. 10034 Krapovickas& Gregory,Rev. Invest.Agrie. (BAB,CTES, GH,LIL, MO, NY, SI,US). 14(2):157-160, fig. 1, 1960. Geographic distribution.This species is Perennialplant. Taproot withbranch roots known only fromthe type locality, where it withoutenlargements. Mainstem erect,short; was collectedon a streetsome 400 m fromthe lateral branches procumbent,ca. 50 cm long, centralsquare of the city of Santa Cruz de la stems subquadrangular,young parts villous Sierra,when its sandy streetshad not yetbeen along the angles, the rest with scattered paved. adpressed hairs. Leaves tetrafoliolate.On the mainstem,the fused portionof the stipules 8 Obs. This species occupies an important mmlong, the free tips up to 16 mmlong; petiole position in the genus because of its great up to 45 mm long; leaflets oval-oblong, the capacity for interspecifichybridization. The apical pair ca. 25 mmlong x 14 mmwide (up to hybrids obtained with species of its same 30 mmx 17 mm),the basal pair ca. 22 mmlong sectionhave a highdegree of fertility,as is the x 13 mm wide (up to 27 mmx 14 mm). On the case with A. appressipila (40.2%) and A. lateral branches, the fused portion of the lignosa (54.4%). In hybrids obtained with stipules 8 mm long x 5 mm wide, the freetips species of othersections, the fertilitydeclines up to 1 1 mmlong; petiole up to 30 mmlong; the noticeably. Hybrids were obtained with the apical pair of leaflets 16 mm long x 11 mm following species of section Erectoides: A. wide (up to 26 mmx 17 mm),basal pair 14 mm gracilis (12.2%), A. Hermannii (7%), A.

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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) paraguariensis ssp. paraguariensis (1 to Leaves tetrafoliolate,dimorphous, those of the 5%), A. Benthamii (2.6%), A. major (0.3 to mainstemnotably larger than those of the late- 1%), A. Archen (0.8%), A. Martii (0.3%) and ral branches. On the mainstem, the fused A. Oteroi. It was also crossed withA. Pintoi portionof the stipulesup to 8-10 mm long,the (0.2%) of section Caulorrhizae and with A. freeportion up to 24 mm long x 3 mm wide, glabrata var. glabrata and A. pseudovillosa, acute. Petiole up to 35 mmlong; rachis 1 1 mm tetraploidspecies of sectionRhizomatosae. long; leaflets elliptical, acute, apiculate, the apical pair up to 37 mmlong x 13 mmwide, the 35. Arachis chiquitana Krapov., W.C. basal pair 34 x 12 mm.On thelateral branches, Gregory & C. E. Simpson nov. sp. in well-developedleaves, the fused portionof the 5-7 mm thefree 13-18 Figs. 2,35; 23,A stipules long, portion mm long x 2.5-3 mm wide; petiole 10-17 mm long; rachis 5-6 mm long; leaflets from Herba perennis. Caulis principalis erectus ellipticalto obovate,apiculate to obtuse,apical ramis procumbentibusradices adventitias in pair 20-24 mmlong x 1 1-18 mmwide, the basal internodiisbasalibus gerentibus.Stipulae dorso pair 16-21 mm long x 9-13 mmwide. Stipules et basi pilis longis,raro etiamsetulis munitae, withglabrous surfaces,margins longly ciliate, praeter illos usque glabrae. Folíola foliorum hairs caducous, frequentlywith long hairs on caulis principaliselliptica, acuta, apiculata,illa theback of thefused part, occasionally villous foliorum ramealium ovalia usque obovata, at the base, rarely with very short hairs apiculata usque obtusa,epiphyllo laevi, glabro, scatteredon the surfaceof the fusedpart, very hypophyllonervo medio prominenteet nervis rarely with bristles. Petiole and rachis secundariis sat perspicuis,pilis adpressis vix canaliculate,back laxly villous,margin of both conspicuisvestito glabrove, margine manifestiore canals ciliate,small adpressedhairs barely visi- ciliis caducis setulisque brevibus nonnullis ble in the canal. Pulvinus villous. Upper instructo.Hypanthium 4-7.5 cm longum, villosum. surface of the leaflets smooth and glabrous; Calyx 5-8 mm longus,setulis sparsis immixtis lower surfacewith prominent midvein, margin villosus. Vexillum aurantiacum, ca. 12 mm and secondary veins somewhat marked,with longum.Fructus biarticulatus paxillo horizontāli barely visible scattered adpressed hairs or paulo profundo8-12 cm longo,isthmo ca. 2 cm glabrous, occasionally long hairs on the longo,articulis ca. 10 mmlongis x 5 mmlatis, midvein;margin longly ciliate, cilia caducous, apice recurvo,pericarpio laevi. sometimeswith a fewshort, persistent bristles. Flowers on the centralaxis and along the late- Holotype:BOLIVIA. SantaCruz. Prov. Chiqui- ral branches, in very short, few-flowered tos,30 kmNE de San Joséde Chiquitos,60°47'W, spikes. Hypanthium4-7.5 cm long, villous. 170471g300 m.,puesto San Marcos,en lugarbajo, Calyx bilabiate, villous and with scattered inundado,arenoso; perenne, ramas postradas, co- bristles,lower lobe falcate,6-8 mmlong, upper rolaanaranjada, 27-IV-1980, Krapovickas, Simpson lobe wide, 5-6 mm long. Standardorange, ca. & Schinini36027 (CIES). Isotypes:CEN, G,GH, K, 12 mm long (when dry). Fruit subterranean, LIL, LPB,MO, NY, P, SI, SP,US. Particulate; peg horizontal,shallow, glabrous, 8-12 cm long, isthmusca. 2 cm long; articles Perennialplant. Taproot deep, 10-14 mmin smooth,beaked, 10 mm long x 5 mm wide; diameter,with thin branches. Mainstem erect, seed 7 mm long x 4 mm wide. 10-20 cm long,the intemodesof its upperhalf covered by the stipules. Lateral branches Additionalmaterial: BOLIVIA. Santa Cruz. prostrate,up to 70 cm long, internodes20-40 Prov.Chiquitos: 5 kmN de San José,estancia El mm long, villous toward the end of the Cinco,26-IV-1980, Krapovickas & al. 36025(CEN, branches. Adventitious roots on the basal CTES, GH,LIL, LPB, MO, NY, RB, US); 4 kmN de internodes of the mainstem and along the San José,camino a San Ignacio, 28-IV-1980, lengthof the lateralbranches, especially when Krapovickas& al. 36028(CIES, LPB,MO, NY, US); theygrow submergedor in contactwith water. San José,cult, en la estaciónde gasolina,29-IV-

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1980,Krapovickas & al. 36031 (CEN, CTES, LIL, branchesbut up to 65 mm long in full-grown LPB,MO, NY, US). branches. Leaves tetrafoliolate. On the mainstem,leaves somewhatlarger than those Geographic distribution.This species was on the lateralbranches; stipules with the fused collected in the neighborhoodof San José de portionpurplish, up to 19 mm long, the free Chiquitos, in the "Chiquitano" massif, on the ends up to 25 mmlong x 2 mmwide; petioleup northernedge of the "chaco" region.It occurs to 53 mm long; rachis 18 mm long; leaflets in low places subjectto flooding. elliptical,somewhat asymmetrical at the base, apex acute, the apical pair up to 46 mm long x Obs. Ordinarily,the stipules of this species 20 mmwide, the basal pair 42 x 18 mm. On the have no bristles,but in populations36027 and lateral branches, the fused portion of the 3603 1 a fewplants were foundwith bristles on stipulesusually 8 mmlong, the free ends acute, leaves located toward the end of some 15 mm long x 2 mm wide (up to 19 mm x 2 branches. mm); petiole 18 mm long; rachis 6 mm long; leafletselliptical, acute, apical pair 26 mmlong 36. Arachis matiensis Krapov., W.C. x 10 mm wide (up to 43 mm x 13 mm), the Gregory & C.E. Simpson nov. sp. basal pair 24 mm long x 9 mm wide (up to 38 Fig. 2,36 mm x 1 1 mm), frequentlythe leafletstoward thebase of the stemsare relativelywider, from obovate to obtuse, with acute apex. Stipules Herba perennis.Rami procumbentes.Caulis withmargins somewhat imbricated at thebase, angulosus, viridis, glaber. Stipulae glabrae, usually glabrous, occasionally somewhat violaceae. Folia caulis principalis foliolis villous on theback of thefused portion. Petiole ellipticis,acutis, ramealia foliolis ellipticis usque and rachis glabrous, rarelysomewhat villous obovatis, omnia epiphyllo laevi, glabro, withshort hairs. Upper surface of the leaflets hypophylloglabro nervo medio et margine smooth, glabrous (very rarely with short manifestioribus,margine pilis serieeis caducis scatteredhairs); lower surfaceglabrous, veins setulisque nonnullis persistentibusinstrueto. and margin somewhat marked (sometimes Hypanthium2.5-6 cmlongum, villosum. Calyx 5-8 with barely visible adpressed hairs); margin mm longus setulis tenuibus sparsim immixtis with caducous silky hairs and some short, villosus. Vexillum11-14 mm longumx 16 mm persistentbristles. Flowers along the lateral latum,aurantiacum. Fructus biarticulatus paxillo branches in very short,few-flowered axillary horizontāliusque ad 60 cm longo,isthmo 3-9 cm spikes. Hypanthium2.5-6 cm long, villous; longo,articulis 10-15 mmlongis x 6-7 mmlatis, calyx bilabiate,villous and withscattered thin apice recurvo,pericarpio laevi. bristles,lower lobe falcate,6-8 mmlong, upper lobe 5-6 mmlong; standard1 1-14 mmlong and Holotype:BOLIVIA. Dep. Santa Cruz. Prov. up to 16 mm wide, orange,wings up to 8 mm Sandoval,1 kmN de San Matías,56°26'W, 16°21'S, long, yellow or yellow borderedwith orange. 170 m, Krapovickas,Simpson & Schinini36014 Fruitbiarticulate; peg horizontalup to 60 cm (CTES). Isotypes:G, GH,LPB, MO, NY, SI, SP,US. long, glabrous or subglabrous, rarely pubescent;isthmus 3-9 cm long; articles 10-15 Perennialplant. Taproot deep, 10-15 mmin mm long x 6-7 mm wide, smooth, beaked, diameter.Mainstem erect, ca. 25 cm long,late- covered by a dense coat of diminutivehairs. ral branches prostrate,up to 80 cm long. In 2n=20 chromosomes (Fernández & plants of several years, the mainstem may Krapovickas 1994). become remarkably bulky at ground level owing to numerous short,crowded branches. Additionalmaterial: BOLIVIA. Santa Cruz. Similar branchlets, with dense cataphylls, Prov.Sandoval: Sandoval, San Matías,1 kmW del frequentlyoccur along the branches. Stem aeropuerto,17-IV-1980, Krapovickas & al. 36007 green, angular, glabrous, rarely pilose; (CEN, CTES, G, GH,LIL, LPB, MO, NY, US); id., internodesvery shorttoward the apex of the aeropuerto,17-IV-1980, Krapovickas & al. 36008

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(CEN, CTES, G,GH, K, LIL, LPB,MO, NY, SI,US); ao longo da estradapara Cáceres que parteda id., 5 km SW del aeropuerto, 18-IV-1980, rodoviaPoconé-Cuiabá, 10 kmalem de Poconé,24- Krapovickas& al. 36296 (CTES); 2 kmNW de San X-1985,Valls & al. 9350 (CEN, CTES); BR-070km Matías,20-IV-1980, Krapovickas & al. 36303(CTES, 623, 1,8 kmW do corr.dos Macacos, 17-V-1985, LPB, US); 1 km N de San Matías,camino a San Valls & al. 8736 (CEN, CTES); 126 kmWSW of Francisco,24-VIII-1981, Vails & al. 6345 (CEN, Cuiabá,on roadto Cáceres, 22 kmW ofSete Porcos, CTES); 3,5 kmNNW de San Matías,20-IV-1980, 22-VIII-1981,Valls & al. 6337 (CEN, CTES). Mun. Krapovickas& al. 36010(CTES); id.,Krapovickas & Santo Antonio de Leverger: Sto. Antonio de al. 36012 (CTES). Prov.Ñuflo de Chavez:Estancia Leverger,24-1-1989, Valls & al. 12084(CEN, CTES). Novicia,30 kmde Concepción,62°12'W, 16°29'S, 500 m,2-V-1977, Krapovickas & al. 30094 (CEN, Geographic distribution. Occurs in the CTES, G, GH, K, LPB, MO, NY, SI, SP, US); id., easternpart of the Dep. of Santa Cruz (Boli- Krapovickas& al. 30095(CTES); 24 kmW de Con- via) and in the westernborder areas of Mato cepción, camino a San Javier, 3-V-1977, Grosso (Brazil), where it was collected along Krapovickas& al. 30096(CTES). the 16thparallel South, between 57°37'W and BRAZIL. Mato Grosso. Porto Esperidião 62°14'W. It prefers the "cerrado," called 58°28'W,15°52'S, 155 m,26-VIII-1981, Valls & al. "pampa" in easternBolivia. 6357(CEN, CTES, GH,MO, NY, US); 49 kmWNW of Cáceres,on roadto Pto.Esperidião, 220 m,25- Obs. 1. The specimenKrapovickas 30096 is Vm-1981,Valls & al. 6356(CEN, CTES, US); 14km distinguishedby being somewhatmore villous NW ofPorto Esperidião, 220 m,26-VIII-1981, Valls and by the small hairs on the upper surfaceof & al. 6361 (CEN, CTES, US); 66 kmNW ofPorto the leaflets,the same as Vails & al. 6408a. Esperidião,on roadto Pontes e Lacerda,200 m,26- The firstwas collectedat the westernextreme VIII-1981, Valls & al. 6366 (CEN, CTES, GH,MO, and the second at the eastern extremeof the NY, US); 56 kmNW ofPorto Esperidião, on roadto range of this species. Pontese Lacerda,Estancia Dois Irmãos,160 m, 30- Vm-1981,Valls & al. 6405(CEN, CTES, US); 33 km Obs. 2. In theeastern part of the rangeof A. ofPorto Esperidião, Fazenda Paraísodo Norteon matiensis,at 26 km east of Cáceres (MT), on roadto Pontes e Lacerda,170 m, 30-VIII-1981, Valls highwayBR-174, km 705 (Vails & al. 6340 & al. 6407(CEN, CTES, NY, US); 9 kmENE ofPorto and 8910), is founda populationof an Arachis Esperidiãoon roadto Cáceres, 150 m, 30-VIII-1981, that grows in calcareous soil and is Valls & al. 6408a (CEN, CTES, GH,MO, NY, US); distinguishedfrom A. matiensis by its more Cáceres,cult, at the airport, 3 1 -VIII- 1981, Vails & al. vigoroushabit and by itslarger fruit. This could 6409 (CEN, CTES); Cuiabá,in flowerbeds at the be anothertaxon, but it would be advisable to FederalUniversity of Mato Grosso,cult., 20-VIII- examineadditional material to make a decision 1981,Valls & al. 6324 (CEN, CTES); 9 kmENE de in thatregard. PortoEsperidião, 18-V-1985, Valls&al. 8756(CEN, CTES); 56 kmNW de PortoEsperidião, 28-V-1985, 37. Arachis appressipila Krapov. & W.C. Valls & al. 8890 (CEN, CTES); 12 kmda BR-174, Gregory nov. sp. caminode Porto a 28-V- Esperidião Casalvasco, Figs. 2,37; 21 1985,Valls & al. 8893(CEN, CTES); VarzeaGrande, 4-XI-1986,Valls & al. 10468(CEN, CTES); id.,Valls &al. 10469(CEN, CTES). Herbaperennis. Caulis principalis erectus ramis decumbentibus.Caulis angulosusapicem versus Relatedmaterial (with bristles on thestipules): pilis paucis adpressisvestitus, cetero glabrescens. Mato Grosso.Mim. Cáceres:37 kmE de Cáceres Stipulae glabrae vel pilos paucos parvíssimos caminoa Cuiabá, 17-XII-1976,Krapovickas & al. adpressosbasi gerentes,margine laxe ciliatae. 30033(CTES); 47 kmde Cáceres,camino a Cuiabá, Folíola ovali-lanceolataepiphyllo laevi, glabro, 17-XII-1976,Krapovickas & al. 30040(CTES). Mun. hypophyllonervo medio et margine prominentibus, Poconé:30 kmW de Poconé,camino a Cáceres,16- nervis secundariisinconspicuis, in nervispilis Xfl-1976,Krapovickas & al. 30030(CTES); 33,8km brevibus adpressis vestito, in nervo medio

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Fig.2 1. Arachisappressipila : plant (G. 1 0000).

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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) majoribus. Hypanthium4-9 cm longum, laxe wide, orange towardthe marginand yellow at villosum.Calyx 6-8 mm longus, plus minusve setulis the base, pinkishveins on the upper surface; nonnullistenuibus immixtis villosus. Vexillum 14-17 wings7-10 mmlong, yellow. Fruit Particulate; mmlongum x 16-18mm latum, aurantiacum. Fructus peg up to 30 cm long,horizontal; isthmus 1-9.5 biarticulatuspaxillo horizontāliusque ad 30 cm cm long; articlesca. 13 mmlong x 7 mmwide, longo,paulo profundo,articulis 13 mmlongis x 7 the apical articlesalways somewhatlarger, up mmlatis, pericarpio laevi. to 15 mm long x 7 mmwide; pericarpsmooth. Seed withocherous seed coat, 10-11 mm long Holotype: BRAZIL. Mato Grosso do Sul. x 5-6 mmwide. 2n=20 chromosomes(Gregory Corumbá, aeropuerto, 27-III-1959, Gregory, & al. 9993). Krapovickas& Pietrarelli9990 (SP). Isotypes: CIES, G,GH, LIL, MO, NY, SI,US. Selectedadditional material: BRAZIL. Mato Grosso do Sul. Corumbá,XII-1902, Robert 731 & Perennial plant. Taproot, with slender 761 (BM); 11-1911,Hoehne, Comissão Rondon 2624 branches,without enlargements. Flowers and (R); Corumbá,3-XII-1943, Baldwin 3141 (NY, SP, fruits clustered about the crown. Mainstem US); 28-III-1959,Gregoiy & al. 9993(GH, LDL, MO, erect, up to 50 cm long; lateral branches NY, SI, US); 5-XII-1976,Krapovickas & al. 30002 decumbent,ca. 50 cm long, internodestoward (CEN, CTES, GH,MO, NY, US); id.,Krapovickas & the base of the branches up to 7 cm long, al. 30003(CTES, NY, US); 3 kmde Corumbácamino cylindrical,glabrous; theinternodes toward the a Unicum,29-111-1959, Gregoiy & al. 10002(GH, LIL, apex 1-2 cm long, angular, lightlypubescent, MO, NY, US); 4 kmde Corumbá,camino a Unicum, hairs adpressed. Leaves tetrafoliolate.The 29-111-1959,Gregory & al. 10000(GH, LIL, MO, NY, fusedpart of the stipulessome 12 mmlong x 6 SI, US); 12km de Corumbácamino a Unicum,5-XII- mm wide with the margins imbricatedat the 1976,Krapovickas & al. 30004 (CEN, CTES, GH, base, the freetips acute, some 17 mmlong x 2 MO, NY, US); 15 kmE de Corumbá,9-XII-1976, mm wide. Petiole canaliculate, in well- Krapovickas& al. 30010 (CTES); Faz. Nova developed leaves 1.5-3 cm long. Rachis 0.5-1 Campina,9-XII-1976, Krapovickas & al. 30009 cm long,canaliculate. Leaflets oval-lanceolate, (CEN, CTES, G, GH, MO, NY, US); Estr. do apical pair up to 50 mm long x 16 mm wide, Gaturamo,15-IV-1972, Hatschbach 29502 (CTES, basal pair up to 40 mm long x 13 mm wide. MBM); 5 kmS de Corumbá,22-XI-1977, Allem & al. Basal portion of the stipules glabrescentor 1452(US); id.,Allem & al. 1455(US); Corumbá,9-X- withvery small, scattered,adpressed hairs; the 1985,Valls & al. 9056(CEN, CIES); id.,lO-X-1985, free portion with glabrous surfaces and the Valls& al. 9060(CEN, CTES); Faz. Coqueiro,17 km marginweakly ciliate. Petiole glabrescentor desdea BR-262pela estrada para Forte Coimbra, 10- withscattered adpressed hairs. Pulvinuswith a X-1985,Valls & al. 9077 (CEN, CTES); id.,25-X- few short hairs. Leaflets with upper surface 1986,Valls & al. 10322,10323, 10324 (CEN, CTES). smooth, glabrous, the lower surface with prominent midvein and margin, secondary Common name, "amendoimbravo" (Allem veins somewhatmarked, and the surfacewith & al. 1452). scattered, small, adpressed hairs, and adpressed hairs a littlelonger on the midvein; Geographie distribution.This species is themargin weakly ciliate and occasionallywith restrictedto a small area in the vicinityof some bristles. Inflorescencesclustered at the Corumbá,on the easternbank of the Paraguay collar of the plant and also along the lengthof River. thebranches; the axillary spikes ca. 5 mmlong, covered by the stipules, 5-6-flowered;bifid Obs. Even though A. appressipila is a bractsca. 15 mmlong. Hypanthium7 (4-9) cm decumbent species, the crossing results long,weakly villous. Calyx moreor less villous, demonstratethat its greatestaffinity is withthe withsome thinbristles, bilabiate, the wider lobe species that have spreading branches which 6 mm long, the narrowerlobe falcate, 7-8 mm make up the rest of section Procumbentes . long. Standard 14-17 mm long x 16-18 mm The hybridsobtained with A. lignosa (30.9%)

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and with A. Rigonii (40.2%) have partial be oblong or oblong-lanceolate,apical leaflets fertility.In contrast,the hybrids with species of to 59 mmlong x 8 mmwide, basal leafletsto 50 othersections are highlysterile, as is the case mmlong x 7 mmwide, bothsurfaces glabrous, withA. major (4.8%), A. paraguariensis ssp. upper side smooth,under side with only the paraguariensis (1 to 4.5%), A. Hermannii midvein marked, margin with scarce, very (1.9%), A. Martii (0.4%) and A. Benthamiiof small caducous hairs and some shortbristles. section Erectoides, À. guaranitica (0.7%) of Hypanthium5 cm long, withfew hairs. Calyx section Trierectoides, and A. glabrata var. 5-6 mm long, with few shorthairs and some glabrata, a tetraploid species of section veryshort bristles. Standard ca. 1 1 mm long, Rhizomatosae . orange. Fruit subterranean,Particulate; peg thick,hollow, glabrous; articles 12-15 mmlong 38. Arachis Vallsii Krapov. & W.C. x 6-7 mm wide, beak somewhatpronounced. Gregory nov. sp. Seeds 10-11 mm long x 4.5-6 mmwide. Figs. 2,38; 22 Additionalmaterial: BRAZIL. Mato Grosso do Sul. Mun.Corumbá: 35 kmSE de Corumbá,camino Herba perennis. Radix polaris. Caulis a PortoManga, Fazenda Vale do Paraísoy luego6 principalisreclinatus usque ad 1.60 m longus, kmde la ruta,hacia el N, haciaBaía Negray Banda pauciramosus,rami procumbentes. Folia glabra Alta,57°30'W, 19°05'S, 10-Xn-1976, Krapovickas & stipulisbrevissime setulosis, foliolis lanceolatis Gregoiy30012 (CTES); 38,8km a partirda entrada epiphyllolaevi, hypophyllonervo medio promi- para Mirandaao longo da BR-262 em direçãoa nente,margine paucis pilis brevissimiset setulis Corumbá,23-IV-1985, Valls & al. 8678(CEN, CTES); nonnullisbrevibus munitis. Hypanthium 5 cm id.,31-X-1985, Valls & al. 9481 (CEN, CTES); id., longumparce pilosum. Calyx 5-6 mm longus. Valls& al. 9482(CEN, CTES); id.,6-IV-1986, Valls & Vexillumaurantiacum. Fructus subterraneus, al. 9902 (CEN, CTES); id.,28-X-1986, Valls & al. biarticulatus,paxillo longo,crasso, cavo, glabro, 10357(CEN, CTES). articulis 12-15 mm longis x 6-7 mm latis, pericarpiolaevi. Geographiedistribution. This species grows to the south of the Gran Pantanal, in Mato Holotype:BRAZIL. Mato Grosso do Sul. 37.8km Grosso do Sul. It is known only from the a oeste da saida de Mirandana BR-262 em area collectionsmade at thetype locality and a very inundavelao longoda rodovia,campo graminoso small herbariumspecimen (Krapovickas & al. con "carandá"( Copernicia) proximode mata,19- 30012) collected next to the type specimen of IV- 1984,Valls, Rao, Gerin& Silva 7635 (CEN). A. valida, in verysimilar environments, that is, Isotype:CTES. in periodically flooded grasslands with Copernicia palms. Perennial plant. Taproot, without enlargements.Mainstem scandent,up to 1.60 We dedicate this species to José Francisco m long, with little branching. Stems with MontenegroVails, researcher at the Centro abundant adventitiousroots, toward the thick Nacional de Recursos Genéticos base, to 4 mm in diameter,hollow, with a (EMBRAPA), Brasilia, who collectedmost of vestiture consisting of abundant very short the materialof this species. bristles.Leaves tetrafoliolate.Stipules with the fusedportion to 13 mmlong, the free portion 15 39. Arachis subcoriacea Krapov. & W.C. mm long x 2 mm wide, very short bristles Gregory nov. sp. witha few short uniformlydistributed, margin Figs. 2,39; 23,B caducous hairs. Petiole up to 38 mm long and rachis up to 10 mmlong, both nearly glabrous, with very few hairs and a few shortbristles. Herba perennis.Rami procumbentes.Caulis Pulvinus somewhatpilose. Leafletslanceolate, angulosus, glaber. Stipulae glabrae interdum but towardthe base of the branchessome may setulosae. Folia glabra. Folíola lanceolata,

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Fig.22. Arachis Vallsii : A, base of the plant with pegs (V.9902); B, leaf(V.7635).

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Fig.23. Arachischiquitana : A, branch(K.36027). A. subcoriacea: B, branch(K.30037). A. triseminata: C, leaf (G.12881).

acuta, apiculata, rigida, subcoriacea, epiphyllo longum, aurantiacum. Fructus biarticulatus paxillo laevi, hypophyllo nervo medio et margine horizontālica. 25 cm longo, articulis14-17 mm prominentibus,nervis secundariis manifestioribus , longisX 6-7 mmlatis , apice recurvo,pericarpio margine setulis nonnullis brevibus instructo. laevi. Hypanthium5-9 cm longum,parce plus minusve adpresso-piloso.Calyx 6-10 mmlongus, praeter Holotype: BRAZIL. Mato Grosso. Mun. pilos paucos in apicibusdentium et setulas paucas Cáceres, 106 kmE de Cáceres,camino a Cuiabá, sparsasglaber. Vexillumsuborbi cularet 10-15 mm 57°8'W, 15°56'S, 17-XII-1976,Krapovickas &

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Gregory30037 (CEN). Isotypes:CTES, G, GH,K, 696,6(30 kmE de Cáceres),30-V-1985, Valls & al. LIL, MO, NY, P, RB, SI, SP,US. 8943(CEN, CIES); id, Valls&al. 8941(CEN, CIES); BR-070,km 685 (42 kmE de Cáceres),30-V-1985, Perennial plant. Taproot 8-10 mm in Valls& al. 8937(CEN, CTES); id.,Valls & al. 8935b diameter, deep, strong, without enlarged (CEN,CTES); BR-070,km 652 (75 kmE de Cáceres), branchroots. Branches procumbent, short, 20- 17-V-1985,Valls & al. 8750(CEN, CTES); id.,29-V- 30 cm long,with little subsequent branching at 1985,Valls & al. 8920(CEN, CTES); id.,30-V-1985, the basal nodes. Stem angular, glabrous, Valls& al. 8922(CEN, CIES); BR-174km 680, Oasis, internodes 10-30 cm long. Leaves 29-V-1985,Valls&al. 8916b(CEN, CTES); 5,2kmNE tetrafoliolate.Stipules with the fused portion 7- do RibeirãoFlexas, rodovia BR-364, entre Cáceres e 10 mmlong, the free part triangular, acute, 13- Cuiabá,24-X-1985, Valls & al. 9357 (CEN, CTES); 15 mm long x 1-2 mmwide. Petiole 13-24 mm id, 9358(CEN, CTES); id.,9359 (CEN, CTES); 2-XI- long; rachis 5-6 mm long. Apical pair of 1986,Valls & al. 10445(CEN, CTES); id.,Valls & al. leaflets24-30 mm long x 4-7 mm wide, basal 10447(CEN, CTES). Mun.S. Antoniodo Leverger: pair 21-25 mm longx 3-5.5 mmwide. Stipules 4,6 kmde SantoAntonio na estradapara Barao de glabrous or with bristles,with parallel veins Melgaço,27-X-1985, Valls&al. 9401(CEN, CTES); somewhat marked; toward the base, at the id.,9402 (CEN, CIES); id.,9403 (CEN, CIES); id.,24- unionwith the stem,the margins are somewhat 1-1989,Valls & al. 12080(CEN, CTES). superimposed.Petiole and rachis canaliculate, glabrous;pulvinus with a fewshort hairs on the Geographiedistribution. This species grows upper side. Leaflets lanceolate, acute, to the northof the Pantanal Matogrossense,in apiculate, rigid,subcoriaceous; upper surface Mato Grosso, in places subject to flooding, smooth, glabrous; lower surface glabrous, betweenCáceres and the vicinityof Cuiabá. midvein and margin prominent,secondary nerves somewhat marked; marginwith some Obs. Arachis subcoriacea has a certain short bristles. In cultivated material the similarityto A Diogoi forthe size of the plant margins of the stipules and the leaflets are and for its lanceolate leaflets. They differin softlyciliate and witha fewbristles at thebase the textureof theirleaves, more smoothin A. of the stipules. Flowers along the lengthof the Diogoi, and inthe development of the peg, which branches in short, 4-flowered spikes. inA subcoriacea is horizontaland shallow,and Hypanthium5-11 cm long, somewhatvillous, deep and inclinedsome 45° inA. Diogoi. withfew hairs, more or less adpressed. Calyx Included in this species are various glabrous except forsome small hairs at thetip specimenswith broader leaflets, possibly from of the teethand some short,scattered bristles; growingin shaded places. upper lobe 6-7 mm long,tridentate, lower lobe 8-10 mm long,falcate. Standardsuborbiçular, VIII. Sect. Rhizomatosae Krapov. & W.C. 10-15 mm long, orange; wings yellow, 10-11 Gregory nov. sect. mm Fruit long. subterranean,biarticulate; pegs Fig. 8 thin,horizontal, up to 72 cm long, glabrous above ground, below ground with hairs that retaina littlesoil; isthmusup to 20 cm long; Sect.Rhizomatosae Krapov. & W.C. Gregory,in articles14-17 mmlong x 6-7 mmwide, smooth, W.C. Gregory,M.P. Gregory,Krapovickas, Smith & witha well-developedbeak. Yarbrough,in Peanuts- Cultureand Uses: 92, 1973, nomennudum. Krapovickas, Agricultural Selectedadditional material: BRAZIL. Mato Genetics.Selected Topics: 137, 1973. Grosso. Mun. Cuiabá: sandymarsh and adjacent cerrado,on BR-364near Córrego Pindaival, 350 m, Perennes. Radix polaris, rhizomataadsunt. 11 -11-1975, Anderson 1 1338(MBM); RodoviaMT- Rami procumbentes. Folia quadrifoliolata. 364,35 kmleste de Cuiabá,13-XI-1975, Hatschbach Vexillumaurantiacum, raro luteum. Fructus 37496 (CTES, MBM). Mun Cáceres:106 kmE de biarticulatus paxillo plus minusve verticali, Cáceres,17-XÜ-1976, Allem 750 (CEN); BR-070,km pericarpiolaevi aut reticulato.

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Typussectionis : Arachis glabrata Benth. species of this section occupy a central positionwithin the overall range of the genus Plants perennial, rhizomatous. Taproot, Arachis which correspondsto the range of A. without thickenings.Branches procumbent. glabrata var. glabrata. Leaves tetrafoliolate.Flowers along thelength The diploid species, A. Burkartii, grows of the branches. Hypanthiumwell developed. more to the south and has a range with Standard expanded, orange, rarely yellow. practicallyno overlap withthose of the other Fruit subterranean, Particulate; peg and rhizomatousspecies, except fora small fringe isthmusshort, of moreor less verticalgrowth. in the extremeNE of Corrientesand in the SE of Misiones (Argentina), where they grow, Geographic distribution. The tetraploid nevertheless,in differentenvironments.

Key fordistinguishing the species

A. Leafletscoriaceous, with the margin marked on bothsurfaces. Standard with reddish lines on both surfaces.2n=20 chromosomes. Series Prorhizomatosae 40. A. Burkartii A'. Leafletswith the margin somewhat marked only on thelower surface. Standard with reddish lines on the uppersurface. 2n=40. Series Rhizomatosae B. Plantcompletely prostrate, with the leaves adpressed to the soil. Upper leaf surface shiny, commonly withsmall scattered bristles. Pericarp reticulate. 41. A. pseudovillosa B'. Plantsomewhat taller, with the leaves separated from the soil. Upperleaf surface smooth, without bristles.Pericarp smooth. 42. A. glabrata C. Leafletsmore or less oblong. 42a. var.glabrata C'. Leafletslanceolate. 42b. var.Hagenbeckii

Ser. Prorhizomatosae Krapov. & W.C. Perennial plant, rhizomatous. Taproot, Gregory nov. ser. without thickenings.Branches procumbent. Leaves tetrafoliolate. Leaflets coriaceous, Sect. Rhizomatosae ser. Prorhizomatosae withmargin marked on bothsurfaces. Flowers Krapov.& W.C. Gregory,in Gregory,M.P. Gregoiy, along the length of the branches. Standard Krapovickas,Smith & Yarbrough,in Peanuts- with reddish lines on both surfaces. Fruit Cultureand Uses: 92, 1973,nomen nudum. subterranean,Particulate. Pericarp smooth. 2n=20 chromosomes. Perennes. Radix polaris, rhizomataadsunt. Rami procumbentes. Folia quadrifoliolata, 40. Arachis Burkartii Handro Vexillum insignitermarginata. supra subtusque Fig. 2,40 lineis rubescentibus.

Typus: Arachis Burkartii Handro Handro,Arq. Bot. Estado São Paulo 3 : 177-179,

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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) táb.42-43, 1958. Burkart, in Burkart, Flora Ilustrada long, usually yellow toward the base and deEntre Ríos 3: 623-4,fig. 289, 1987. orange toward the apex. Fruit Particulate; A. marginataauct. non Gardner,Bentham, Fl. pegs 2-4.5 cm long; isthmusca. 5 mm long; bras.15(1): 87,p.p., 1859.Chevalier, Rev. Int. Bot. articlessome 13 mmlong x 7 mmwide. 2n=20 Appl. Agrie.Trop. 13(146-147):762, p.p., 1933. chromosomes(Gregory & al. 1973). Burkart,Darwiniana 3(2): 264, 1939.Hoehne, Flora Brasílica25(2) part.122: 16,p.p., 1940.Hermann, Holotype:BRAZIL. Rio Grandedo Sul. 12 km Agrie.Monogr. USDA 19: 10,p.p., fig. 7, 1954. northof Sant'Anna, 13-XI-1936, W.A. Archer 4439 A. marginataforma submarginata Hoehne, Flo- (SP 45487!).Isotypes: IAN!, K!, NY!, US!. raBrasílica 25(2) part.122: 16,1940, nomen p.p. Selectedadditional material: ARGENTINA. Co- Perennial plant, rhizomatous; rhizomes rrientes.Dep. Ituzaingó:ruta 38 y ríoAguapey, 2 1- shallow, branched,rooting, elongated, 2-3 mm III-1981,Tressens 1238 (CTES); ruta34, 12km E de in diameter,the young parts thinner,scaly. ruta12, 15-XII-1984,Tressens & al. 2975 (CTES). Stems very short,upright or reclining,almost Dep. SantoTomé: Garruchos, 25 -IV- 1969, Pedersen completely covered by the stipules. Leaves 9147 (CTES, Herb. Pedersen); 20-IX-1974, tetrafoliolate.The fusedportion of the stipules Krapovickas& al. 25779 (CTES); 15 km E de up to 7 mm long x 7 mm wide, the free tips Virasoro,5-II-1972, Krapovickas& al. 21038 acute, up to 12 mmlong. Petiole usually short, (CTES); SantoTomé, 1 -11-1976, Krapovickas 29082 10-20 mmlong, but may reach 75 mmin length (CTES); Ea. Timbó,ayo. Ciriacoy ruta40, 27-11- in some cases. Rachis ordinarilyca. 5 mmlong 1983,Schinini & al. 23420(CTES). Dep. SanMartín: (rarely up to 10 mm). Leaflets obovate, Torrent,13-XII-1944, Ibarrola 1662 (LIL); TresCe- rounded or cuneate, sometimes obovate- rros,14-11-1979, Schinini & al. 17203 (CTES); lanceolate, apex obtuse, rounded or acute, Burkart8166 (SI). Dep. Paso de los Libres:Campo mucronulate(in driedspecimens the apex may Gral.Avalos, 7-IV-1972, Carnevali 3141 (CTES); be split), coriaceous, with marked margin, Bonpland,27-III-1964, Krapovickas & al. 11300 especially on the lower surface;the apical pair (CTES); paradaPucheta, 17-11-1979, Ahumada 2502 always somewhatlarger than the basal pair,up (CTES); ayo.Ayuí, 17-11-1979, Schinini & al. 17277 to 25 mm long x 1 1 mm wide, in cultivated (CTES). Dep. Mercedes:Solari, 8-III-1945, Ibarrola material up to 33 mm long x 17 mm wide. 2590(NY, SI); 26-III-1964,Krapovickas & al. 11285 Stipules withthe surfacesalmost glabrous, the (CTES); 61 kmN de Mercedes,5-III-1973, Quarin & fused portion villous on the back, margin al. 932 (CTES); 38 kmE de Mercedes,19-V-1968, ciliate. Petiole somewhatcanaliculate, usually Hammons& al. 355 (CTES); Mercedes,1 1-III-1953, villous, sometimes with scattered hairs, but Krapovickas7967 (LIL, US); 11 kmS de Mercedes, always withhairs ca. 2 mm long, occasionally cerroPajarito, 23-11-1984, Tressens & al. 2435 with some bristles present. Leaflets with (CTES); ayo.Yuquerí, camino a ItáCorá, 21-11-1984, brilliantupper surface, usually glabrous or with Tressens& al. 2313 (CTES). Dep. MonteCaseros: long sparse, caducous hairs; the lower surface Colonia Libertad,27-III-1964, Krapovickas & al. usually villous, sometimessubglabrous, with 11301 (CTES); CampoGral. Avalos, ayo. Curupi, 23- scattered adpressed hairs, rarely with some 11-1979,Schinini & al. 17598(CTES); MonteCase- bristles;margin weakly ciliate and sometimes ros,27-III-1947, Nicora 4728 (SI); id.,29-X-1950, with some short bristles. Inflorescences Nicora5584 (SI); id.,28-111-1964, Krapovickas & al. axillary,few-flowered, axis short,covered by 11306 (CTES); Mocoretá,rio Mocoretá, lO-XI-1980, the stipules.Hypanthium 6-12 cm long,pilose. Ahumada& al. 3879 (CTES); San Francisco,33 km Calyx bilabiate,villous, with long silky hairs and SW de MonteCaseros, 12-IX-1979, Schinini & al. some long thinbristles; the wide lobe 6-8 mm 18803(CTES). EntreRios. Dep. Federación:rincón long, the narrow lobe falcate, 7-9 mm long. del Mocoretáy el ríoUruguay, Burkart 2 1775 (SI); Standard 14-17 mm long x 17-20 mm wide, Federación,20-XII-1946, Meyer 1 1 102 (LIL). Dep. yellow at the base and orange toward the Concordia:San Carlos,12-XII-1946, Meyer 10946 margin (rarely completely yellow), with (LIL); Concordia,6-II-1927, Burkart888 (SI); reddishlines on both surfaces;wings 8-1 1 mm Chajarí,Burkart 8167 (SI). Misiones.Dep. Apósto-

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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 les: ayo. Chimiray, Ruta 40, lO-IX-1978, 17-V-1983,Valls & al. 7335(CEN, CTES); 1 kmS do Krapovickas34141 (CTES); Azara,1,5 km E de Ruta arroioMatapí, BR-472, km 2 16,17-V-1983, Valls & al. 40, caminoal puerto,2-II-1976, Krapovickas & al. 7337(CEN, CTES); 21 kmE deUruguaiana, BR-290, 29116 (CTES); 4 kmN de San José,20-III-1983, 18-V-1983,Valls & al. 7344 (CEN); 57 km E de Cristóbal& al. 1899(CIES). Uruguaiana,BR-290, km 568, 18-V-1983, Valls & al. BRAZIL. Rio Grandedo Sul. Isabelle1833 (K); 7345 (CEN, CTES); Alegrete,rio Ibirapuitan, BR- Rio Grande, 1833, Gaudichaud 1991 (P); 290,18-V-1983, Valls & al. 7347(CEN, CTES); 30km Montenegro,Deslandes s/n (SP); Tupancireta,11- E de Alegrete,BR-290, 18-V-1983, Valls & al. 7349 XI-1936,Archer 4429 (IAN, K, LIL,NY, SP,US); 26- (CEN). Mun. São Sepe: 1 kmW do entroncamento 1-1942,Rambo 9173 (PACA); 28-1-1942, Rambo 9613 BR-290/BR-392,19-V-1983, Valls & al. 7357 (CEN, (PACA); 28-1-1942,Rambo 9973 (LIL); 30-1-1942, CTES). Mun.Encruzilhada do Sul,BR-290, rio Irai, Rambo 9980 (PACA); Juliode Castilhos,11-XI- 19-V-1983,Valls & al. 7359(CEN, CTES). Mua Dom 1967,Hagelund 5482 (CTES); Giruá,18-XI-1974, Pedrito:BR-293, rio Ibicui da Armada,28-XI-1979, Hagelund83 17 (CTES); Alegrete,Rio IbiraPuitan, Pedersen12565 (CTES, Herb.Pedersen); 39 kmS de 13-III-1948,Palacios & al. 1816 (LIL); Alegrete,ayo. CruzAlta, rio Ivaí, BR-158, 30-XI-1980, Krapovickas Capivari,14-III-1948, Palacios & al. 1758(LIL); ca- & al. 37085(CEN, CTES); Capão Seco,ruta a Pelo- mino de Alegretea S. Francisco,8 km de Passo tas,12-XI-1969, Rosengurtt & al. 9241 (MVFA). Novo,Nicora 4695 (SI); 10km SE ofAlegrete, 21-V- URUGUAY.Banda Oriental, Saint Hilaire 2364 1968,Hammons & al. 364 (CTES); id.,18-XÜ-1981, (K, P). Artigas.Cuareim, 2-V-1901, Berro 1501, fl. Albo& al. 2449(CTES); 55 kmW deRosario do Sul, naranja(MVFA); Cuareim, 2-V-1901, Berro 1897, fl. 8-XII-1978,Krapovickas & al. 34237(CTES); 14km amarillo azufre (MVFA); Artigas,5-XII-1943, W de Rosariodo Sul, 19-1-1973,Krapovickas & al. Bartlett21028 (US); Sarandi,22-111-1984, Pedersen 22792 (CTES); 11 kmNW de Minas do Camaqua 13896(CTES, Heib.Pedersen); ayo. Catalancito, 30- (mun.Caçapava do Sul),6-XÜ-1978, Krapovickas & I-1948,Castellanos (LIL 15979);Bella Unión,23-1- al. 34203(CIES); S. Gabriel,Faz Sta.Cecilia, 1-1944, 1942,Rosengurtt B 3715 (MVFA, SI); id., 29-III- Rambo25703 & 25829(PACA); São Luiz,Caaió, 13- 1962,del Puerto1949 (MVFA); TresCruces, l-III- VIII-1954,Rambo 53309 (PACA); São Pedroad. fi. 1944,Rosengurtt B 4587 (MVFA, SI); ColoniaRive- Ibicui,II-1946, Rambo 34446 (PACA); SantaAna do ra,18-VI-1950,RosengurttB6025 (MVFA); Santa Livramento,Cerro do Registro,24-11-1947, Castella- Rosa, m-1922, Schroeder(Osten 16703) (MVM); nos (LIL 15981); Livramento,Morro Vigia, 12-1- ayo.Yucutuyá y rioCuareim, Ea. El Ombúde Mallo, 1944, Rambo3910 (PACA); Uruguaiana,Rambo 13-IV-1978,dei Puerto & al. 15363(CIES, MVFA); 63224 (PACA); Uruguaiana,24-1-1948, Palacios & ayo.Cuaró, próx. rio Cuareim, 21 -XI- 1976, dei Puer- al. 232 (LIL); 14 kmE de Uruguaiana,21-V-1968, to 13078 (MVFA); Tomás Gomensoro,9-1-1971, Hammons& al. 362 (CTES); Uruguaiana,10-XII- Marchesi10037 (MVFA); Catalán,ruta 30, 4-XII- 1945, Swallen 7659 (US); Camino Itaqui a 1957,Rosengurtt B 6842 (MVFA); Ruta30, km 679, Uruguayana,17-V-1963, Rosengurtt & al. 9467 4-II-1966,dei Puerto & al. 5961(MVFA); Ruta 30, km (MVFA);Quarai, Faz. doJarau, 1-1945, Rambo 26031 83,2-IV-1986, dei Puerto & al. 18171(MVFA); Cam- (LIL, PACA); Torres,Campo Bonito,26-X-1978, po de La Estanzuela,rata 3 y bifurcacióna T. Waechter1021 (ICN); id.,10-11-1983, Krapovickas & Gomensoro,21-X-1961, Millot 545 (MVFA). Rivera. aL38473 (CEN, CTES); id., 21-V-1983, Valls& al.7363 Tranqueras, Arechavaleta, Herb. Osten 4105 (CEN, CTES); Caçapavado Sul,BR-153, km 64,6, 4 (MVM); id., XI-1899,Arechavaleta 5369, 5369a kmN do rioCamaquá, 30-1-1982, Valls & al. 6898 (MVM);id., 19-111-1913, Osten 6525 (MVM); id.,18- (CEN); São Luiz Gonzaga,BR-285, km 447, 15-V- II-1945,Legrand 4161 (MVM); id., 30-IV-1968, 1983,Valls & al. 7317 (CEN, CTES). Mun. Santo Hammons& al. 27 (CTES); Tranqueras,declives de Antoniodas Missões: BR-285,a 400 m N do rio la CuchillaNegra, 1-1941, Legrand 2409 (MVM); 74 Icamaquá,15-V-1983, Valls & al. 7320(CEN, CTES). kmN deTacuarembó, Ruta 5, 29-IV-1968, Hammons Mun.Itaquí: BR-472, km 53, 16-V-1983,Valls & al. & al. 24 (CTES); Rivera,10-1-1944, Legrand 3447 7330(CEN, CIES); BR-472,5,5 km S deItaquí, 16-V- (MVM, SI); Rivera,Archer 4449 (SP, US); Cerro 1983, Valls & al. 7333 (CEN, CTES). Mun. Aurora,10-11-1961, Arrillaga& al. 1008(MVFA); Uruguaiana:BR-472, km 15 1, 16-V-1983,Valls & al. Paso Platón,15-III-1962, del Puerto 1795 (MVFA). 7334(CEN, CTES); BR-472,20 kmS de Uruguaiana, Tacuarembó. Tacuarembó,1-1899, Arechavaleta

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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)

5370(MVM); id.,1-1940, Legrand 1847 (MVM); ca- This series is differentiatedfrom the series minoa Borracho,3 leguasde Tacuarembó,1 1 -III- Prorhizomatosae principally because its 1945,Rosengurtt B 4742 (MVFA, SI); Luján,24-1- species are tetraploidand due to its different 1977,Pedersen 1 1618 (CTES, Herb.Pedersen); 30 genetic behavior such that, unlike A. km NE de Tacuarembó,Ruta 5, 29-IV-1968, Burkartii,its species can cross and produce Hammons& al. 23 (CTES); EstaciónExperimental hybrids with diploid species from other del Norte,22-XI-1976, del Puerto13039 (MVFA); sections.Also, the rangesof thetwo series are Grutade los Heiechos,10-III-1990, Izaguirre & al. different(fig. 8). 19730(MVFA); Grutade los Cuervosa CerroTra- vieso,3-XI-1990, Bayce & al. 19970(MVFA); arro- 41. Arachis pseudovillosa (Chodat & yoLaureles, 18-11-1968, Lema 6959 (MVFA).Salto. Hassl.) Krapov. & W.C. Gregory nov. Salto,Osten 5374 (CORD, MVM, SI, US); Mirador comb. Las Hammons& al. Lavanderas,Salto, 28-IV-1968, Figs. 2,41; 19,C, D 16(CTES); 4 kmS de Salto,28-IV-1968, Hammons & al. 17 Ruta km 6-VI-l & (CTES); 4, 106, 967,Izaguirre A. prostrataBenth, var. pseudovillosa Chodat 21003 al. (MVFA); Termasdel Arapey,15-1-1967, & Hassl.,PI. Hassl.2: 449, 1904. Rosengurtt& al. 10543(MVFA). Paysandú. Arroyo Osorio13949 Sacra,1-II-1948, (MVM). Rhizomatous perennial; rhizomes shallow, branched, rooting, elongated, 2-3 mm in distribution.This Geographie species grows diameter, the younger parts more slender, over a of the stateof Rio Grande do greatpart densely pubescent, scaly. Stems extremely Sul in the northof in (Brazil), Uruguay,and short, almost subterranean,with the leaves near the River in in places Uruguay Argentina: completelypressed against the soil. Leaves the south of the east of Corrientes Misiones, tetrafoliolate.The fusedportions of thestipules and the northeast of Entre Rios. The 5-9 mm long x 4 mmwide, the freetips acute, southernmostlocalities are made ofthe up arroyo 6-14 mm long. Petiole usually short,4-16 mm a littleto thesouth of the of Sacra, city Paysandú, long, rarelyup to 3 1 mm long. Rachis 3-7 mm in and near the of Uruguay, Capão Seco, city long, rarely 10 mm. Leaflets usually broadly in thesoutheast of Rio Grandedo Sul. Pelotas, obovate, rounded, apex obtuse, rounded or This with species prefers high places, somewhatacute, mucronate,coriaceous, veins hard and with shallow, soils, frequently rocky and marginmarked only on the lower surface; One to this is the outcroppings. exception the apical pair always somewhat larger and collectionof Bonito Torres,Campo (Waechter morerounded than the basal pair,apical pair up in the extremenortheast of Rio Grande 1021), to 35 mmlong x 23 mmwide, thebasal pair up do near the borderwith Santa Sul, Catarina, to 3 1 mmlong x 19 mmwide. Stipuleswith the whereit in soil of in a grows nearlypure sand, surfacessubglabrous, villous on theback of the of Butia population capitata. fused portion,margin ciliate (cilia caducous), occasionally some bristlespresent only on the Obs. Arachis with its 20 Burkartii, margin of the free portions. Petiole is the rhizomatous chromosomes, only diploid canaliculate, usually villous, sometimeswith and species presents a quite remarkable bristles;rachis somewhatless villous than the such that no were genetic isolation, hybrids petiole. Leaflets with the upper surface shiny obtained from the numerous crosses dark green, commonly with small scattered be with within the attempted, they species bristles; the lower surface usually villous, same section or with from other species occasionally subglabrous, but always with sections M.P. & (Gregory, Gregory1979). plantresidue and particlesof soil clingingto the surface, margin with scattered bristles, and frequentlyin young leaves some short hairs. Ser. Rhizomatosae Inflorescences axillary, few-flowered,with short axis, covered by the stipules. Arachis Bentham Type species; glabrata Hypanthium5-10 cm long, villous, with long

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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 silky hairs. Calyx bilabiate, villous and caminoa Amambai,17-11-1959, Gregory & al. 9678, frequentlywith some long delicatebristles; the 9679 (LIL, US); 20 kmW de Amambai,camino a broad lobe 7 mmlong, the narrow lobe falcate, AntonioJoão, 17-11-1959, Gregory & al. 9672 (LIL, 7.5-8 mmlong. Standard 13-15 mmlong x 14- US); 5 kmde Amambai,camino a PontaPorã, 19-11- 16 mm wide, with reddishlines on the upper 1959,Gregory &al. 9688,9703 (LIL, US); 13kmaoN surface, yellow toward the center with an do rio Amambai na estrada de Ponta Porã a orange margin,occasionally entirelyyellow; Amambai,24-IV-1984, Valls & al.7695 (CEN, CIES); wings8 mmlong, yellow. Fruit Particulate; peg 8,5km ao Sul do rioAmambai ao longoda estradade 3-5 cm long, covered toward the base with a PontaPorã a Amambai,24-IV-1984, Valls & al. 7701 short tomentum; isthmus ca. 1.5 cm long; (CEN,CIES); a 3 kmda praçacentral de Rio Brilhante articles 11-15 mm long x 6-7 mm wide, e a 1,5km do acesso sul aBR-163, 26-IV-1984, Valls & reticulate when mature, with beak, covered al.7710 (CEN, CTES); 23 kmaSEde Antonio João e 11 when young with a tomentumof shorthairs kmNE do aceso a Maracajúna estradapara Ponta that retain particles of soil. 2n=40 Porã,8-IV-1986, Valls & al. 9932(CEN, CTES). Mun. chromosomes (Fernández & Krapovickas Guia Lopes da Laguna: on road (BR-267) from 1994). Maracajúto GuiaLopes, about 12 kmW ofErvania, 24-X-1986,Pedersen 14710 (CTES, Herb. Pedersen). Lecto-holotype:PARAGUAY. "Herba 0,05- PARAGUAY.Amambay. Finca Elvira, 20kmNW 0,15 mpétala citrina,in campispr. Ipé hú,Sierra de PedroJuan Caballero, 13-V-1961, Gregory & al. de Maracayu, oct. (Hassler) n. 5069" (G!). 10559(LIL, US); EstanciaSan Luis,20 kmNW de Isotypes:BM!, K!, NY!, P!, UC!. PedroJuan Caballero, 13-V-1961, Gregory & al. We selectedthis specimen as thelectotype due 10560,10561 (LIL, US); PedroJuan Caballero, Rojas to the abundanceof materialand because of its 6481 (SI); 17-11-1951,Schwarz 11931 (LIL); Río being distributedto a largernumber of herbaria Ipané,18-VIII-1980, Schinini & al. 20434 (CTES); " than the other syntype:petula aurantiaca, in ¿str. CapitánBado, San Fernando,31-X-1986, campopr. flumenCarimbatay, sept., (Hassler) n. Pedersens/n (CTES). 4511" (BM!, G!). Geographiedistribution. This species grows Selected additionalmaterial: BRAZIL. Mato inthe Sierra de Amambay,along itstotal length Grossodo Sul. 15kmNdeNioaque, BR-419, 11-11- and in adjacent areas, in the border regionof 1993,Hatschbach & al. 58909(CTES, MBM); 5 km Mato Grosso do Sul and Paraguay. It prefers SW deRio Brilhante, 12-V-1961,Gregoiy&al. 10555 open fieldswith short grass vegetation. (LIL, US); 28 km SW de Dourados,12-V-1961, Gregory& al. 10558(LIL, US); 9 kmW de Caarapó, Obs. 1. In some localities, we observed 19-1-1979,Krapovickas & al. 34334 (CEN, CTES); plants of a size intermediate between A. entrePonta Porã e Dourados,24-IV-1939, Otero 362 pseudovillosa and A. glabrata, which were (SP): 35 kmE de PontaPorã, 15-V-1976, Hatschbach foundgrowing together in: Brasil, Mato Grosso 38670 (CTES, MBM); PontaPorã, Herb. Hassler do Sul, 8-9 km northof Ponta Porã, Feb. 1 1, 9886 (G); 7-10 km N de PontaPorã, 11-11-1959, 1959, Gregory& al. 9627, 9628 (LIL); 1 km Gregory& al. 9625,9626, 9627a, 9630 (LIL, US); id., north of the Rio Amambai, 13-VI-1968, 26-VI-1977,Krapovickas & al. 30130(CTES); 22 km Hammons & al. 576 (CTES) and 577 (CTES, de Ponta Porã camino a Bela Vista, 19-11-1968, US); 5 km from Amambai, 19-11-1959, Krapovickas& al. 14055 (CTES); 40 kmNW de Gregory& al. 9706 (LIL, US). Because of the PontaPorã, camino a Bela Vista,13-11-1959, Gregory presenceof bristleson theupper surfaceof the & al. 9634, 9635 (LIL, US); FazendaNestor Qué, leaves, these specimens most resemble A. Campanario,14-IV-1939, Otero & al. 283 (SP); 1 km pseudovillosa. N delRío Amambai, camino Ponta Porã a Amambay, 17-11-1959,Gregory & al. 9664(LIL, US); id.,14-V- Obs. 2. In general,the genetic behavior of 1961, Gregory & al. 10565(LIL, US); id.,10566 (LIL, A. pseudovillosa is very similar to A. US): id., 13-VI-1968,Hammons & al. 575 (CEN, glabrata, butbecause we had a smallernumber CTES, US): 1km E de AntonioJoão (Capitan Bado), of sampledlocations available, the results for the

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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) formerare morelimited than in the latter species. taproot. Rhizomes at a depth of 5-20 cm, Hybrids were obtained between A. ramified,rooting, very extended; in year-old pseudovillosa and A. glabrata var. glabrata plants the rhizomes originate on the basal in two cases, but withoutdata on thefertility of nodes of the cotyledonarybranches. Aerial pollen. stems to 40 cm long, decumbent,scandent, Hybrids were also obtained with annual pubescent to glabrous. Leaves tetrafoliolate. species of section Arachis such as A. Stipules subulate, villous to glabrescent, Batizocoi and A. duranensis, withA. Rigonii sometimeswith some bristles.Leaflets oblong, of section Procumbentes, and with A. ellipticalor obovate,with the margin somewhat paraguariensis ssp. paraguariensis in markedon the underside.Upper leaf surface section Erectoides, but the hybridsnever did usually glabrous, but younger leaves may flower.A surprisingcase is thebehavior of the exhibit some very short, scattered hairs. hybrid between A. pseudovillosa and A. Lower leaf surface with adpressed hairs to Hermannii of section Erectoides, with46.1% subglabrous, and frequently with hairs pollen stain (p. 178 in M.P. Gregory & al. somewhat longer on the midvein. Spikes 1979). pauciflorous,very short, axillary. Hypanthium well developed,villous. Calyx villous and with Illustration:Otero, 19, 1941: photographof abundant bristles. Standard orange, rarely specimenOtero 362, fromPonta Porā. yellow, with red lines on the upper surface. Fruit subterranean,Particulate; peg 5-10 cm 42a. Arachis glabrata Benth. var. glabrata long, with short isthmus; pericarp smooth. 2n=40 Fig. 2,42a chromosomes(Gregory & al. 1973).

Holotype:BRAZIL. Mato Grossodo Sid. Rio Bentham,Trans. Linn. Soc. London18(2): 159, Pardo,Riedel 107 Hb. Mus. Petr. 1837 (K!). 1841,"Ad Rio Pardo Brasiliae.(v.s. comm.a Mus. Acad.Petrop. cum fl.)." Selectedadditional material: ARGENTINA. Co- Bentham,Martius Fl. bras.15(1): 87, 1859,excl. rrientes.Dep. SantoTomé: ayo. Chimiray y ruta40, fig. Chevalier,Rev. Int. Bot. Appl. Agrie.Trop. 12-IV-1974,Krapovickas & al. 25225(CTES). Dep. 13(146-147):759, 1933,pro parte.Hoehne, Flora Ituzaingó: 25 km E de Ituzaingó, 15-1-1966, Brasílica25(2) part.122: 19, táb. 14,fig. 1, 1940. Krapovickas& al. 12000(CIES, NY, US). Dep. San Hermann,Agrie. Monogr. USDA 19: 8,fig. 4, 1954. Miguel: Colonia Madariaga,17-11-1946, Ibarrola Conagin,Bragantia 21: 355, est. 8, figs.6B & 15, 4229 (CEN, CTES, G, LIL, MO); Loreto,25-1-1959, 2n=40,1962. Gregory& al. 9553 (LIL, US). Dep. GeneralPaz: A. prostrataBenth. var. genuina Chodat & Puisoyé,25-X-1945, Ibarrola 3610 (CIES, LIL). Dep. Hassl.,PI. Hassl.2: 448, 1904,Hassler 8439 (G!). Empedrado:Empedrado, 3-ÜI-1950, Schwarz 9973 A. prostrata Benth. var. pseudomarginata (CEN,CTES, LIL); Rincóndel Madrejón, 2-IÜ-1972, Chodat& Hassl.,Pl. Hassl. 2: 449, 1904,lectotype: Carnevali 2978 (CTES). Dep. Mburacuyá: Paraguay,Atira, Hassler 3445 (G!). Mburucuyá,3/5-III-1945, Hunziker 6683 (CORD, A. glabrataBenth. var. membranifolia [Benth.?] LIL); Estancia Santa Teresa, 26-XII-1976, ex A. Chev.,Rev. Int. Bot. Appl. Agrie. Trop. 9(90): Krapovickas& al. 30044(CEN, CIES, GH,K, MO, 100, 1929,nomen nudum, "A. de SaintHilaire no. NY, US). Dep. Saladas:Saladas, laguna Soto, 23-IX- 367." São Paulo,Ribeirão Corrente (KJ). 1944,Schwarz 4 (LIL). Dep. Concepción:Tabay, 1- A. glabrataBenth. forma major Hoehne, Flora XI-1965, Krapovickas & al. 11609 (CEN, CTES, G, Brasílica25(2) part.122: 20, táb. 15, 1940,nomen US). Misiones.Dep. San Martín:Puerto Tabay, 30- nudum. XI-1945,Schwarz 1578 (K, LIL). Dep. Cainguás: A. glabrataBenth. forma minor Hoehne, Flora CampoGrande, l-XI-1946, Pierotti 5297 (LIL). Dep. Brasílica25(2) part.122: 20, táb. 14,fig. 2, 1940, San Ignacio:Colonia Corpus, 20-X-1948, Schwarz nomennudum. 6456 (CTES, LDL); San Ignacio,27-1-1947, Meyer 11727(LIL). Dep. San Javier:San Javier,14-11-1947, Perennial, rhizomatous plant with robust Huidobro5182 (LIL). Dep. Concepción:Concep-

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ciónde la Sierra,1-II-1948, Schulz 6930 (CIES); 16 Anastacio:Palmeiras, 17-11-1970, Hatschbach 23729 km NW de Concepciónde la Sierra,11 -III- 1969, (CIES, MBM); 41 kmS deAquidauana, Faz. Varzea Krapovickas& al. 15107(CTES). Dep. Apóstoles: Alegre, 30-VI-1977, Krapovickas& al. 30143 Azara,7-IX-1977, Fernández 363 (CTES); Pindapoy, (CTES). Mun.Aquidauana: Aquidauana, Faz. San- 13-1-1937,Clos 5981 (BAB); Apóstoles,26-1-1926, ta Cruz, 17-VII-1969,Hatschbach 21924 (CTES, Clos 1918(BAB). Dep. Candelaria:Picada Güemes, MBM). Mun.Miranda: estrada Miranda-Capao, 15- 21-11-1947,Huidobro 4872 (CTES, LIL); Ayo.Má- XII-1976,Shepherd & al. 4103(CEN). Mun. Jardim: 8 quina,10 km E de SantaAna, ruta 105, 1 l-XI-1972, kmW de Jardim,10-VI-1968, Hammons & al. 563 Mroginski781 (CTES); Bonpland,Jörgensen 191 (CEN). Mim. PortoMurtinho: 51 km E de Porto (BAB). Dep. Capital:20 kmW de San José,cerca del Murtinho,12-VI-1968, Hammons & al. 567 (CTES); arroyoPindapoy, 8-III-1953, Krapovickas7934 id.,Hammons & al. 569 (CTES). Mun.Nioaque: 27 (CEN, CTES, LIL, US); Posadas,La Granja,12-XI- km S de Nioaque,camino a Jardim,10-VI-1968, 1907,Ekman 1719 (NY). Krapovickas& al. 14425 (CEN, CTES). Mun. BRAZIL. Goiás.Mun. Jatai: Balsamo, l-XI-1950, Maracaju: Maracaju, 3-II-1952, Kuhlmann(SP Macedo2694 (LIL, US). Mun.Rio Verde: Rio Verde, 69920). Mun. Nova Andradina:Faz. Douradinho, 17-111-1959,Gregory & al. 9947(GH, LIL, MO, NY, ll-XI-1975, Hatschbach37397 (CTES, MBM). SI, US). Mun. Alagarças:Alagarças, 28-1-1945, Mun. Rio Brilhante:Rio Vacaria, 24-X-1970, Shimoya3593 (US). Mun.Bom Jesus: BR-452, km Hatschbach25 177 (CTES, MBM, NY); 10 kmE de 140W do acessoa SantaRosa, 23-IV-1980, Vails & Entroncamento,1-III- 1959, Gregory & al. 9797(LIL, al. 51 15 (CEN); 33 kmde Bom Jesus,18-III-1959, MO,NY, US); 5 kmE deEntroncamento, ll-V-1961, Gregory& al. 9951 (GH,LIL, MO, NY, SI, SP, US). Gregory& al. 10550(LIL, US). Mun.Caarapó: 9 km Mun.Itumbiara: Itumbiara, 18-III-1959, Gregory & W de Caarapó,19-1-1979, Krapovickas & al. 34593 al. 9954 (LIL, MÒ, NY, SI, SP, US). Mun.Goiatuba: (CEN,CIES, G,K, LIL,MBM US). Mun.Bela Vista: Goiatuba,5-IV-1961, Gregory & al. 10139(LIL, NY, BelaVista, 8-V-1939, Otero & al. 387(SP); BelaVista US). Mun.Parauna: Rio Verdao, BR-060, km 172, 24- 14-11-1959,Gregory & al. 9645(LIL, GH,MO,NY, SI, IV-1980,Vails & al. 5 153(CEN). Mato Grosso. 70 km SP, US). Mun. AntonioJoão: 10 km W de Penso N de Cuiabá, caminoa Rosariodo Oeste, 11-III- [nowAntonio João], 13-11-1959, Gregory & al. 9637 1959,Gregory & al. 9918 (LIL, NY, SI, US). Mun. (GH, LIL, MO, NY, US). Mun. PontaPorã: Ponta Barra do Garças: Barra do Garças, 4-X-1968, Porā, 18-IV-1939,Otero 327 (RBR, SP). Mun. Fonseca& al. 1094(NY, UB). Mun.Rondonópolis: Amambay:Antonio João [now Coronel Sapucaia, Rondonópolis,18-XI-1975, Hatschbach37674 acrossfrom Capitán Bado, Paraguay],18-11-1959, (CTES, MBM); Rondonópolis,8-III-1959, Gregory Gregory&al.9681 (LIL, MO, NY, US); 1kmNdelrio & al. 9893(GH, LIL, NY, MO, US). Mun.Pedra Preta. Amambay,17-11-1959, Gregory & al. 9667(LIL, GH, 32 kmSSE de Rondonópolis,BR-364, 3-IX-1981, MO, NY, US). Minas Gerais. Mim. Ituiutaba: Valls & al. 6442 (CEN, CTES); Birro, entre Ituiutaba,27-X-1944, Macedo 568 (LIL). Mun. Rondonópolisy AltoGarças, 15-III-1959, Gregory Prata:100 kmN de Frutal, 19-111-1959, Gregory&al. & al. 9935(LIL, GH,MO, NY, SI,US). Mato Grosso 9964(GH, LIL, MO,NY, SI,US). Mun Frutal:10 km do Sul. Mun. Ribas do Rio Pardo:Rio Pardo,X- S de Frutal,19-III-1959, Gregory & al. 9966 (LIL, 1826,Riedel 604 (K p.p.,P, US); Ribasdo Rio Pardo, MO, NY, SI, US). Mun.Uberaba: 28 kmde Uberaba 3-HI-1959,Gregory & al. 9827(GH, LIL, MO, NY, SI, na estradaa Belo Horizonte,15-X- 1 966, Duarte 772 US); id.,Gregory & al. 9830(LIL, GH,MO, NY, SI, (NY). São Paulo. Faz. Guanabara,Ilha Seca, 2-VIII- US). Mun.Campo Grande: Campo Grande, 25-XII- 1936,Hoehne & al. (SP 45842);Piraçununga, Emas, 1932,Otero 10 (RBR, SP); CampoGrande, Capao 13-IX-1946,Kuhlmann (SP 80244); Matao, 12-1- Bonito,6-IX-1936, Archer & al. 98 (NY,US); Campo 1980,Krapovickas & al. 35249 (CEN, CTES, US). Grande,Fazenda Experimentalde Criaçao,24-11- Mun.Tanabí: VilaMonteiro, 21-VIII-1941, Gehrt(SP 1959, Gregory& al. 9813 (LIL, US); 70 km S de 45842). Mun. Barbosa: SP-425, 25-IX-1975, CampoGrande, l-m-1959, Gregory & al. 9806(CEN, Hatschbach& al. 37125(CTES, MBM). Mun.Santa CTES, GH, LIL, MO, NY, SI, SP, US). Mun. Rio Cruz do Rio Pardo: 2 km E del río Pardo,Posto Verde:Rio Verde,13-XII-1976, Krapovickas & al. Taquarí,SP-280, km 186, 1-II-1987, Krapovickas & 30020(CTES, NY, US); 52 kmN deRio Verde, 7-III- al. 40989(CIES). Prov:S. Paulo,St. Hilaire 991 bis (P). 1959,Gregory & al. 9882(LIL, MO, NY, US). Mun. PARAGUAY. Alto Paraná. 20 km N de

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Hernandarias,10-1-1974, Schinini 8055 (CTES). plazas in the centerof Asunción. Towards the Amambav.Pedro Juan Caballero, 1-1934, Rojas 6666 north,it extends throughthe states of Mato (SI); PorteraOrtiz, 7 kmN de P.J.Caballero, 1 l-II- Grosso do Sul and Mato Grosso, along the 1959,Gregory & al. 9629 (LIL, GH,MO, NY, US); edges of the Mato Grosso Pantanal. Its cursussuperioris fl. Apa, 11-1902,Hassler 8439 (G, northernlimit is foundat Rosario Oeste, some K, NY, P, UC); CerroCorá, 16-VÜI-1980, Schinini & 70 km northof Cuiabá. Towards the east, its al. 20178 (CTES, G, MO); Bela Vista,14-11-1959, rangeextends through southeastern Goiás and Gregory& al. 9643 (GH, LIL, MO, NY, SI, US). the MiningTriangle to the state of São Paulo, Caaguazú. Km 90, rutaAsunción-Oviedo, 25-1- where its easternlimit is foundat Emas, near 1949,Rosengurtt B-5557 (SI); km 116, ruta Asun- Pirassununga.In Brazil, it grows in the "cerra- ción-Caaguazù,3-II-1959, Gregory & al. 9591 (LIL, do," and in Paraguay and Argentinain open GH, MO, NY, US). Central.Yaguarón, 1-II-1966, field formations.It is a good natural forage, Krapovickas& al. 12290(CTES, NY, US); San Lo- much desiredby livestock,for which reason it renzo,XII-1936, Archer & al. 7470 (SI); Asunción, is always foundheavily grazed and therefore SantísimaTrinidad, 5-1-1937, Archer 4904 (K, LIL, difficultto identify. SP, UC, US); Luque,1-1949, Rojas & al. 14152(SI). Concepción.Fluminis Aquidaban, X-1901, Hassler Obs. 1. Arachis glabrata is very variable, 7664(G, UC); SanSalvador, 11-1917, Rojas 293 1 (LIL, even withina singlepopulation, and it was not SI); PuertoRizzo, 17-1-1955,Meyer & al. 18693 possible to establish intraspecifictaxa, except (LIL); 10km Wde Horquetas,17-V-1961, Gregory & forthe var. Hagenbeckii, which has a more or al. 10596(LIL, MO, NY, US). Cordillera.Cordillera less definedrange and rathernarrow leaflets de Altos,1902, Fiebrig 263 (G, SI); fluminisYacá, pr. thatfacilitate its identification. Valenzuela,11-1900, Hassler 71 15 (G); 12 kmN de Paraguarí,camino a Piribebuy,7-II- 1 982, Schinini & Obs. 2. Despite themorphological variation al. 21947 (CTES). Guairá. Villarica,XII-1874, exhibited by A. glabrata var. glabrata, its Balansa1526 (G, K, P). Itapúa.San Miguel Cunizù, 3 behavior in intraspecific crosses is fairly kmNde Villa Encamación, 14-XÜ-1936, Archer 4655 uniform.Of eighthybrids obtained, in six the (IAN,NY, SP, US); arroyoPorá, 7 kmN de Encarna- fertilityfluctuated between 68.6 and 92.9% ción,28-1-1959, Gregory & al. 9567 (LIL, NY, US); and in one 45.2% was obtained. One very Trinidad,29-1-1959, Gregory & al. 9568 (LIL, NY, interestingcase is the resultof the cross 10 x US); id.,Gregory & al. 9570 (LIL, NY, US). Misio- 84, in whichwas found 12.9% stained pollen. nes. San JuanBautista, X-1931, Rojas 5722 (SI); This value is verylow, especially if one takes Santiago,estancia La Soledad,20-X-1967, Pedersen into account thatboth parentsgrow verynear 8664 (CTES, K, US, Herb.Pedersen). Paraguarí. to one another,given thatTrinidad (no. 10) is Paraguarí,4-IV-1885, Kurtz 225 (CORD); Paraguaiy, situatedsome 20 kmto thenorth of thelocality XII-1900,Hassler 65 13 (G); rioTebicuari, frente a of no. 87 (7 km N of Encarnación) in Florida,4-II-1966, Krapovickas & al. 12399(CTES). southeasternParaguay. It is evidentthat some San Pedro. Lima, estanciaCarumbé, 8-X-1967, typeof geneticbarrier has establisheditself at Pedersen8547 (CTES, K, Herb. Pedersen);San theintravarietal level. Estanislao,VHI-1900, Hassler 4261 (G,K, P). In the crosses with species from other sections, 12 populations of A. glabrata var. Commonname, "mendoimdo campo baixo" glabrata were used, and eight hybridswere (Otero 1951: 175). obtained with A. duranensis, eight with A. Batizocoi, and one with A. stenosperma, all Geographic distribution.Ťhe range of this annual species fromsection Arachis ; fivewith species, which is very polymorphic,has its A. paraguariensis ssp. paraguariensis, four southernlimit in the north of Corrientesand the with A. Hermannii, four with A. major, and south of Misiones, in Argentina.It apparently two with A. Benthamii from section does not cross the Paraguay River and is very Erec toides^ and two withA. appressipila and common in eastern Paraguay where, for two with A. Rigonii from section example, it forms part of the lawns of the Procumbentes. The great majority of these

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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 hybridsnever flowered,except forone hybrid Corrientes,"Hassler 5863 (G!), ß longicalyx, withA. major with29.3% pollen staining,one "flumenCarimbatay," Hassler 45 12 (G!). Lectotype: withA. appressipila with22.2%, and one with Hassler5863 (G!). A. Batizocoi with 0.03% (see p. 195 and A. marginata Gardner subsp. Hagenbeckii Gregory,M.P. & Gregory1979). (Harmsex Kuntze)A. Chev.,Rev. Int.Bot. Appl. The hybridsbetween rhizomatous species Agrie.Trop. 13(146-147): 763, 1933. and those fromsect. Arachis did notproduce a A. prostrataBenth. forma Hagenbeckii (Harms single flower, except for cross 19 (A. ex Kuntze)Hoehne, Flora Brasílica 25(2) part.122: Batizocoi) X 94 (Hammons & al. 569) which 11,1940. flowered.The rhizomatousparent is a plant A. prostrataBenth. subsp. Hagenbeckii (Harms that clings to the ground,with very brilliant ex Kuntze)Hoehne, Flora Brasílica 25(2) part.122: dark green leaflets, somewhat differentfrom 15,tab. 11, fig. 2, 1940. the rest of the A. glabrata material. A. angusti/olia(Chodat & Hassl.) Killip ex It is interestingto ratifythat the four species Hoehne,Flora Brasílica 25(2) part.122: 12, 1940, of section Erectoides are sympatricwith A. excl. exiccatum, based on A. prostrata var. glabrata, but the species from sections pseudomarginataforma angusti/olia Chodat & Arachis and Procumbentes that participated Hassl. in the crosses are allopatìe withA. glabrata A. prostrataBenth. var. angusti/oliaChodat & and among themselves. Hassl.ex Hoehne, Flora Brasílica 25(2) part.122: 7 & 12, 1940.nov. stat. pro syn. Obs. 3. Arachis glabrata has the characteristicsof a good forage plant and is Perennial, rhizomatous plant. It is being used for this purpose in Florida (USA) differentiatedfrom the var. glabrata by its where various cultivarshave been developed, more elongated and pointed leaflets and by like 'Arb,' selected froma collectionmade by being more glabrous. 2n=40 chromosomes W. Archer,on 9 Nov. 1939, in Campo Grande, (Fernández& Krapovickas 1994). Mato Grosso do Sul, Brazil (PI 1 18457) (Prine 1964), or 'Arbrook,' developed frommaterial Lecto-holotype:PARAGUAY. Dep. Paraguari, collectedin Trinidad,Paraguay (Gregory& al. "Caapucú, Süd Paraguay,IX-1892, herb. Otto 9570, PI 262817) (Prine & al. 1986). Kuntze"[Hagenbeck] (NY!).

42b. Arachis glabrata Benth. var. Selectedadditional material: ARGENTINA Co- Hagenbeckii (Harms ex Kuntze) F.J. rrientes. Dep. Ituzaingó: Ituzaingó, 1-1947, Herm. Spegazzini10051 (BAB, SI); Ea. Valle,24-XI-1950, Bertoni5189 (CTES, LIL); 30 kmE de Ituzaingó,5- Hermann,Agrie. Monogr. USDA 19: 9, figs.5 & ni-1953,Krapovickas 7904 (CIES, LIL, US); Puerto 6, 1954,based on ArachisHagenbeckii Harms ex Valle,30 kmE deItuzaingó, 2-X-1978, Schinini & al. Kuntze. 15679(CTES); 20 kmE de Ituzaingó,20-IX-1970, A. HagenbeckiiHarms ex Kuntze,Revis, gen. pl. Krapovickas& al. 16077(CTES, GH); 50 kmE de Itá 3(2): 52-53, 1898,"Süd Paraguay:Caapucu; auch Ibaté,ruta 12, 15-1-1966,Krapovickas & al. 11946 aus dem Gran-Chacogebietebekannt (gesammelt (CTES, NY, US). Dep. Berónde Astrada:Laguna von Hagenbeck)."Type: Gran Chaco, Hagenbeck Torpy,21-X-1949, Schwarz 8395 (LIL). Dep. General (B, destroyed).Photo F2255 (reproducedin Burkart, Paz: 15km S de GeneralPaz, 4-ŪI-1953, Krapovickas Darwiniana3(2): 266,lám. 20, 1939.Hoehne, Flora 7855(CIES, LIL, US). Dep. SanMiguel: 21 kmS de Brasílica 25(2) part. 122, táb. 11 fig. 2, 1940. Loreto,7-III-1974, Schinini & al. 8227(CTES); 6 km Hermann,Agrie. Monogr. USDA 19: 19,fig. 5, 1954. SW de San Miguel,3 1-III-1974,Krapovickas & al. Conagin,Bragantia 21: 356-357,est. 9, figs.7, A & 24703(CIES). Dep.Mburucuyá: Ea SantaMaría, 26- 16,1962, 2n=40. XII-1976,Krapovickas & al. 30043 (CTES, NY, US). A. prostrata Benth. var. pseudomarginata PARAGUAY. Caaguazú. Pastoreo,5-II-1959, Chodat & Hassl. formaangusti/olia Chodat & Gregoiy& al. 9610a& b (LIL, US); Ruta2, 9 kmdel Hassl.,PI. Hassl.2: 449, 1904,a brevicalyx,"flumen cruceCnel. Oviedo-Villa Rica, 20-IX-1970, Maruñack

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127(CTES, G,MO). Canindeyú.Fl. Carimbatay,IX- inW.C. Gregoiy,M.P. Gregoiy,Krapovickas, Smith 1898,Hassler 45 12 (G); fi. Corrientes, XII-1898, Hassler & Yarbrough,in Peanuts- Cultureand Uses: 92, 5863(G); 46 kmS deKatueté, 3 km N delrío Itambeiy, 1973,nomen nudum. rutaPuerto Stroessner-Saltos deiGuairá, 18-III-1982, Sect.Axonomorphae ser. Amphiploides Krapov. Schillini23201 (CTES). CentralSan Lorenzo, 1-1937, & W.C. Gregory,loc. cit. 1973,nomen nudum. Rojas7566a (SI); SanLorenzo, 13-X-1981, Schinini& Sect. Axonomorphaeser. Annuae Krapov. & al. 21285 (CEN, CTES, G, GH); Itá, 29-1-1966, W.C. Gregory,loc. cit.1973, nomen nudum. Krapovickas& al. 12142(CIES, G,NY, US); Itá,30-1- Sect. Axonomorphaeser. PerennesKrapov. & 1966,Krapovickas & al. 12209(CTES); Ayo.Mboiy, W.C. Gregory,loc. cit.1973, nomen nudum. ruta2, 1-1-1973,Schinini 5700 (CTES). Cordillera. Valenzuela,20-XII-1950, Sparre & al. 1153 (LIL); Plantsannual or perennial,without rhizomes EusebioAyala, 3-Ū-1959, Gregory & al. 9580 (GH, LIL, or stolons. Taproot, without thickenings. MO,NY, US); id.,Gregoiy & al. 9587(GH, LIL, MO, Mainstem erect, branches procumbent, NY, US); 1 kmN delayo. Piribebuy, camino a Arroyos spreading.Stipules with open margins.Leaves yEsteros-Tobatí, 7-II-1959, Gregory & al. 9618(GH, tetrafoliolate.Flowers distributedalong the LIL,MO, NY, US); Caacupé,25-V-1964, Krapovickas length of the branches. Hypanthium well & al. 11483 (CEN, CTES, G,MO); Emboscada,XII- developed. Standardalways expanded,orange 1980, Bordas 1247 (CTES). Guairá. Villa Rica, or yellow, withoutreddish lines on the dorsal Jorgensen3637 (LIL, NY, US); VillaRica, XI-1941, surface or with a pinkish purple tinge (A. Rojas9242 (SI); VillaRica, 16-XÜ-1936, Archer 4664 Batizocoi). Fruit subterranean,generally with (IAN,K, NA,NY, SP,UC, US); Jatatú,8 km de ViUa two single-seededarticles or one multi-seeded Rica,16-XII-1936, Archer 4670 (IAN,NY, SP, US). article(A. hypogaea)-,peg verticalor inclined, Misiones.San Ignacio, 3 1-1-1959,Gregory & al. 9575 nothorizontal; pericarp smooth or reticulate. (GH,LIL, MO, NY, US). Paraguarí.Caapucú, 31-1- 1959,Gregory & al. 9576 (GH, LIL, MO, NY, SI, US); 2 Type species: Arachis hypogaea L. kmNde Caapucú,16-VI-1977, Krapovickas & al. 30 107 (CTES,G, US); ColoniaPiraretá, camino a Valenzuela, Geographic distribution.The range of the 2 l-XI-1950,Sparre & al. 453(CTES, LIL). San Pedro. sectionArachis has a peculiar shape (fig. 9). It Primavera,5/8-III-1954, Woolston 117 (K, SI); 4 kmE presentsan axis that coincides more or less de SanEstanislao, 14-V-1974, Schinini 8864 (CTES); with the 57th and 58th meridians, that Ruta3, 10 kmS del rioAguaray Guazú, 14-V-1974, encompasses the watershedsof the Paraguay Schinini8891 (CTES); 4km NE deSan Estanislao, 19-V- and Uruguay riversand ends at the La Plata 1974,Schinini 9103 (CIES, MO); SanEstanislao,ruta 3, River. In theextreme northern part of thisstrip 14-XÜ-1983,Vanni&aL 217 (CEN, CIES, G,MO). is located the Chapado dos Parecis, which separates the basin of the Paraguay River Geographic distribution. Among the fromthat of the Amazon and whichappears to polymorphismsof A. glabrata, the variety be an obstacle that,due to its height,does not Hagenbeckii is easily recognizable by its permitthe rangeto extendfurther to the north narrow leaflets. It occupies a defined area at this place. On thiscentral axis, the perennial withinthe range of the varietyglabrata. The species prevail,where almost all of them are var. Hagenbeckii grows in the north of the foundalong watercoursesand some of themare province of Corrientes(Argentina) and in the adapted to periodicflooding, like A. Diogoi, A. southof Paraguay. Its northernlimit is foundin helodes and A. Kuhlmannii,as well as annual the departmentsof Canindeyúand San Pedro, species, likeA. Hoehnei and A. valida, thatlive in Paraguay. in the Mato Grosso Pantanal where they are permanentlyexposed to largefloods. IX. Sect. Arachis The two arms that extendtoward the north Fig. 9 correspondto the basin of the Tocantins River to the east and to the Mamoré and Guaporé riversystem to thewest, between Trinidad and Sect.Axonomorphae Krapov. & W.C. Gregory, Guayaramerin, in Bolivia. In these two

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northward expansions, the species" character is exclusive to this section and is encounteredare annuals and most are adapted present in a few perennial species, like A. to conditionsof prolongedinundation, as is the villosa, fromthe Uruguay River, in the ex- case of A. benensis, A. trinitensis and A. treme southern part of the range of the Williamsii in the Beni and of A. palustris in genus, and A. microsperma, from the Apa the Tocantins. Isolated at the easternextreme River, on the border of Mato Grosso do Sul is A. stenosperma,an annual species growing with Paraguay. On the otherhand, thereis a on the sands of the Atlantic coast, evidently greater number of annual species with this carriedthere by the action of humans. kind of fruit,like A. valida, fromCorumbá, Expansion toward the southwest is in the Mato Grosso Pantanal, or A. constituted by annual species, like A. glandulifera, A. magna, A. ipaënsis and A. Batizocoi, A. duranensis and A. ipaënsis, Williamsiifrom Bolivia, or A. montícola, the which are adapted to conditions of periodic only tetraploid wild species, from drought. Two of these extend from the dry northwesternArgentina. "chaco" up to the firstfoothills of the Andes: A. Batizocoi (300 to 950 m elev.) and A. Obs. 2. In a first attempt to put the duranensis (250 to 1250 m elev.). These two species of section Arachis in order (Gregory species, togetherwith A. montícola (1350 to & al. 1973), it was clear that there were 1560 m elev.), also an annual, are theones that diploid perennial species (series Perennes), grow at the highestelevations of all theknown diploid annuals (series Annuae) and wild species of the genus Arachis. tetraploid annuals (series Amphiploides). Today, in light of the results of the experi- Obs. 1. In the genus, annual species occur mental crosses conducted, this division is no only in section Arachis and in section longer so clear cut. There is a greater Heteranthae. In the latter, its four genetic affinity between the perennial constituentsgrow in the northeastof Brazil species A. Cardenasii, A. villosa, A. where the droughts are proverbial. The correntina and A. Diogoi and the annuals appearance of all of these annual species is A. stenosperma and A, duranensis, with very similar, with flowers distributedalong whom they share the 'A' genome, than the length of spreading and very long between these annuals and A. Batizocoi branches. Evidently,this biological type has ('B' genome), or A. glandulifera ('D' shown itself to be an adaptive advantage genome), which are also annuals. There is that has made possible the occupation of evidence of other genomes, such as in the new areas with extreme conditions of case of A. ipaënsis, which is closely related flooding or drought,expanding the range of to A. magna, both of which are diploid and its species toward the Amazon basin or to very similar to A. montícola, a tetraploid, arid zones such as the Brazilian northeastor both in plant habit and in the shape of the the foothills of the Andes, to eventually fruit.It is evident that new studies are still reach as much as 1500 m above sea level. necessary to evaluate the characters that Also present in section Arachis are fruits could be useful for organizing the species withhighly reticulate pericarps (fig. 3). This withinsection Arachis.

Key fordistinguishing the species

A. Biarticulatefruit, articles separated by an isthmus.Peg fragile. B. Plantsannual or biennial. C. Pegvillous and with bristles.

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D. Fruitarticles reticulate. 43. A. glandulifera D' Fruitarticles smooth. 44.A. cruziana C. Peg glabrousor with few hairs, without bristles. E. Fruitarticles markedly reticulate. F. Undersideof leaf subglabrous, with very small adpressed hairs and with a fewlong hairs on themidvein and on themargin. G. Stipuleswith bristles. H. Fruitarticles up to 2 1 mmx 9 mm.2n=40. 45. A montico!a H1.Fruit articles up to 17mm x 9 mm.2n=20. 46.A. magna G' Stipuleswithout bristles. Articles up to 17mm x 10mm. 2n=20. 47. A. ipaënsis F1.Underside of leaf with hairs ca. 2 mmlong on themidvein and on themargin. I. Stipuleswithout bristles. Fruit articles up to 17mm x 10 mm. 48. A. valida I'. Stipuleswith bristles. Articles up to 12 mmx 7 mm. 49.A. Williamsii E' Fruitarticles smooth or slightly reticulate. J. Standardyellow, with pinkish purple tinge on the dorsalsurface. Stipules with bristles. Undersideof leaf with hairs ca. 2 mmlong. 50.A. Batizocoi T. Standardorange or yellow, without purple tinge. K. Undersideof leaf with hairs ca. 2 mmlong, scattered. L. Stipuleswithout bristles. Upper side of leaf glabrous. 51.A duranensis L' Stipuleswith bristles. Frequently also with long hairs on the upper surface of young leaves. 52.A. Hoehnei K1.Underside of leaf glabrous. M. Stipuleswithout bristles. N. Fruitarticles 14-22 mm long x 5-7mm wide. 53.A. stenosperma N1.Fruit articles less than 13 mm long x 6 mmwide. O. Calyxand hypanthiumglabrous to subglabrous.Leaves glabrous. 54.A. praecox 0' Calyxand hypanthium pilose. P. Calyxwith a fewsilky hairs, without bristles. Leaflets glabrous. 55.A. palustris P1.Calyx villous and withbristles. Underside of leaf glabrous or with a fewlong hairson themidvein.

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Q. Stemwith only a fewhairs on theyoung parts. 56.A. benensis Q' Stemvillous. 57.A. trinitensis M' Stipules,petiole and rachiswith numerous bristles. 58.A. decora B' Plantsperennial. R. Undersideof leafwith hairs ca. 2 mmlong, upper side glabrous. Stipules without bristles. Fruit articlessmooth. 59.A. Herzogii * R' Undersideof lea glabrousto villous, with hairs no greaterthan 1 mmlong. S. Fruitarticles noticeably reticulate. T. Articlesup to9 mmx 4 mm.Upper leaf surface glabrous, underside with adpressed hairs. 60.A microsperma T' Articles10-15 mm x 7-8mm. Leaflets with both surfaces villous. 61. A. villosa S' Articlessmooth or with reticulation barely marked. U. Leafletsglabrous, with some short bristles on themargin. 62. A. helodes U' Leafletswith hairs, at leaston theunderside. V. Leafletmargins with two kinds of trichomes: short adpressed hairs and long cilia (up to2 mmlong) and frequently with some bristles. 63. A correntina V. Leafletmargins with one or two kinds of trichomes. W. Fruitarticles small, 7-11 mm long x 4-6mm wide. X. Leafletswith margin ciliate, outstanding on bothsurfaces. 64. A. Simpsonii X'. Leafletswith margin marked only on theunderside. Y. On thelateral branches, leaflets always rounded, obtuse, large (L/W=1.4-1.7: 1); marginciliate and with some bristles. 65. A. Cardenasii Y' Leafletssmaller (L/W= 1 .6-2.9: 1); marginwith adpressed hairs and some bristles. 66.A. Kempff-Mercadoi W' Articleslarger, 10-14 mm long x 5-7mm wide. Z. Leafletsfrom lanceolate to oblong-lanceolate(L/W=2.5-4.8:l); upper leaf surface withvery small hairs. 67. A. Diogoi Z' Leafletsoblong-lanceolate toobovate (L/W= 1 .4-2.3 : 1); upperleaf surface glabrous. 68. A. Kuhlmannii A' Fruitnot articulated, with 1-5 seeds. Peg tenacious. 69. A. hypogaea

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43. Arachis glandulifera Stalker long x 20 mmwide; wings yellow, sometimes with half Figs. 3,43; 24,B-D upper orange and lower half yellow, 11 mm long x 7 mm wide. Fruit biarticulate;peg 1 1-18 cm long, villous, with Stalker,KT., Amer.J. Bot. 78(5): 633-634, figs. 1-4, short hairs and some bristles; isthmus 3 cm 1991. long; articles 15-20 mm long x 10-12 mm wide, epicarp reticulate.2n=20 chromosomes Taprooted annual. Mainstem 50-120 cm (Stalker 1991). long, growingerect at firstbut later trailing, branched and with flowers toward the base. Holotype: U.S.A. North Carolina. Moore Lateral branches trailing, 80-100 cm long. County;Sandhills Research Station, field D2, plot Stems semi-cylindricalto quadrangular,angu- number90 WSN 40, 12-IX-1990,Stalker 90-40 lar, villous, withtwo kindsof trichomes:short (withoutindication of the accession or itsoriginal wavy hairs, long hairs more or less rigidfrom locality)(NCSU). Isotypes:NA, US. 1.5-2 mmlong, and scarce broad-basedbristle- like emergences. Internodesof the mainstem Materialstudied: BOLIVIA. Santa Cruz. Prov. ca. 4 cm long, those of the lateralbranches 5- Ñuflode Chavez:Novicia, ca. 30 kmS de Concep- 10 cm long. Leaves tetrafoliolate. On the ción, 16°29'S, 62°12'W, 500 m s.m., l-V-1977, mainstem,the fused base of the stipules 5-6 Krapovickas& al. 30091 (CEN, CTES, G, GH, K, mm long x 4 mm wide, the freetips 12 mm x LPB,MO, NY, SI, SP,US). Prov.Velasco: 21 kmS de 3 mm wide; petiole 3-5 cm long; pulvinus 2 San Ignacio,camino a San Miguel,16°33'S, 61°W, mm long; rachis 10-15 mm long; apical 400m s.m., 7-V-1977, Krapovickas & al. 30098(CEN, leaflets usually 36-43 mm long x 18-21 mm CTES, GH, LPB, MO, NY, US); 27 km S de San wide, but up to 52 mm x 26 mm, basal Ignacio,7-V-1977, Krapovickas & al. 30099 (CEN, leaflets usually 30 mm long x 34 mm wide, CTES,GH, LPB, MO, NY, US); SanMiguelito, 23 km but up to 44 mm x 22 mm. On the lateral N de San Ignacio,8-V-1977, Krapovickas & al. branches, the fused portionof the stipules 4- 30100 (CTES); comunidadSan Miguelito,100 km al 5 mm long x 3-3.5 mm wide, the freeportion N de San Ignacio, 20-XI-1987, Bruderreck20 7-9 mm long; petiole 10-16 mm in lengthup (CIES). to 20-30 mm long toward the tip of the stem; rachis 7-10 mm long; apical leaflets 25-35 Geographiedistribution. This species grows mm long x 12-18 mmwide, basal leaflets 17- in the centralpart of the departmentof Santa 25 mm long x 7-14 mmwide. Stipules with6- Cruz (Bolivia), in the Provinces of Ñuflo de 8 longitudinal veins along the fused portion Chavez and Velasco. and a single prominentvein along the free portion,with two classes of trichomes:short, Stalker (1991) with the accessions K more or less wavy hairs, long, somewhatrigid 30091, 30098, 30099 and 30100, made hairs more abundant toward the base, and intraspecific crosses that produced proge- scattered bristles from the base almost to nies with a high percentage of stained pollen the apex of the tips, margin ciliate; petiole and normal meiotic behavior. In the semi-cylindrical,canaliculate, villous, with interspecific crosses he found that the short hairs, long hairs and bristles; leaflets hybrids of A. glandulifera with A. eliptical, upper surface glabrous, lower duranensis (genome A) and with A. surface villous with short hairs, long hairs Batizocoi (genome B), all diploids, showed a along the midvein, and scattered bristles; high degree of sterility,with very irregular margin somewhat thickened,ciliate and with meiosis. The attempts to cross A. some bristles. Calyx tube 10-55 mm long, glandulifera with A. hypogaea failed. In villous withlong silkyhairs. Calyx bilabiate, light of these results, Stalker proposed the villous with a few bristles, the broad lobe 7 genome D for A. glandulifera, which is mm long, 3 -toothed,the narrow lobe curved, different from the genomes A and B of 8 mm long. Standard orange, up to 17 mm section Arachis.

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Fig.24. Arachis retusa : A, branch (Pereira 1948). A. glandulifera : B, leaffrom the mainstem; C, leaf from lateral branch; D, fruitarticle (K.30091).

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44. Arachis cruziana Krapov., W.C. proximal leafletsca. 29-30 mm long x 14-15 Gregory & C.E. Simpson nov. sp. mm wide. Hypanthiumvillous. Calyx villous and with bristles. Standard Fruit Fig. 3,44 orange. subterranean,Particulate, peg 10-17 cm long, villous on the aerial part and with bristles, Herba annua. Radix polaris, debilis. Caulis articles 14-17 mm long x 7-8 mm wide, with principalis50-85 cm longusramis prostratis. Caules beak, pericarpsmooth. Seed 11-15 mm long x villosiusque glabrescentespilis ca. 2 mmlongis 5-8 mmwide. setulisquebase ampliatissparsim vestiti. Stipulae setuliset pilis longissparsis vestitae. Folíola oblon- Additionalmaterial: BOLIVIA. Santa Cruz.Prov. ga usque obovata epiphyllo glabro, laevi, Chiquitos:8-10 km NE de San José (60°47'W, hypophyllonervis et margine perspicuis pilis sparsis 17047.5)ca 300 m s m.En pequeña"pampa", suelo ca. 2 mmlongis setulisque paucis vestito,margine arenablanca; postrada,corola anaranjada,anual, pilisca. 2 mmlongis ciliato. Hypanthium villosum et 27-IV-1980,Krapovickas & al. 36026 (CTES, LPB, setulosum. Vexillum aurantiacum. Fructus MO,NY, US). subterraneus,biarticulatus paxillo 10-17cm longo, parteaerea villosaet setulosa,articulis 14-17 mm Geographic distribution.It grows to the longisX 7-8mm latis, apice recurvo, pericarpio laevi. south of the Serranía de Chiquitos, in the vicinity of San José de Chiquitos, in the Holotype:BOLIVIA. Santa Cruz. Prov. Chiquitos, 2 departmentof Santa Cruz (Bolivia), in places kmW deSan José (60°47W, 17°47'S),ca. 300ms.m.,en wherethe soil is almost pure sand. el km263 del ferrocarril, enpastizal, anual, ramas postradas,25-IV-1980, Krapovickas, Simpson & Schinini 45. Arachis montícola Krapov. & Rigoni 36024 US. (CIES). Isotypes:CEN, LPB, MO, NY, Fig. 3,45

Annual plant.Taproot, weak. Mainstem50- 85 cm long,erect when young, later spreading; Krapovickas& Rigoni,Darwiniana 11(3): 441- lateralbranches prostrate, ca. 70 cm long; stem 445,1957. subcylindrical to quadrangular, villous, A. pusillaauct. non Benth., Burkart, Darwiniana glabrescent,hairs ca. 2 mm long,and scattered 3(2): 276,fig. 19, h, 1939. wide-based bristle-bearing emergences; internodes4-5 cm longon themainstem and 5-10 Annual plant, taprooted. Mainstem erect, cm long on the lateral branches. Leaves ca. 30 cm long, lateral branches procumbent, tetrafoliolate,distal leaflets from oblong to up to 1.10 m long. Stems brownish-violetin obovate, the proximalones oblong,upper leaf partsexposed to the sun, angular,more or less surface glabrous, smooth, underside with villous,with hairs ca. 2 mmlong; internodeson scattered hairs 1.5-2 mm long and some the mainstemup to 3 cm long, on the lateral bristles, veins and margins marked, margin branches 2-6 cm long. Leaves tetrafoliolate. ciliate,hairs ca. 2 mm long and a few bristles; On the mainstem,the fused portion of the stipules with bristlesand with scatteredlong stipules 11-14 mmlong, the free portion 23-25 hairs,more abundant on thedorsal surface.On mm long, subfalcate; petiole 34-43 mm long the mainstem,stipules with the fused portion and rachis 12-14 mmlong; leafletsoval-oblong, up to 15 mm long,the freeportion 15 mmlong the apical pair 39-47 mm long x 20-23 mm X 3 mmwide; petiole 35-45 mmlong; rachis 7- wide, the basal pair 36-42 mm x 16-20 mm 15 mmlong; distal leaflets 51-58 mmlong x 23- wide. On the lateralbranches the fusedportion 25 mmwide, proximal leaflets 41-55 mmlong x of the stipules 7-10 mm long, the free portion 19-22 mm wide. On the lateral branches, 13-19 mmlong; petiole22-30 mmlong and the stipuleswith the fused portion 5 mmlong, free rachis 8-10 mm long; leaflets from oval to portion 13-15 mm long x 2-2.5 mm wide; obovate,obtuse, the apical pair 29-34 mmlong petiole 27-30 mmlong; rachis ca. 10 mmlong; x 20-23 mmwide, the basal pair 27-30 mmlong distal leafletsca. 30 mm long x 18 mm wide, x 16-20 mm wide. Stipule surfaces glabrous,

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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 but may show some long hairs on the back of MO, NY, P, SI, US); ApeaderoReyes, 5 km S de thefused portion; wings, back and basal halfof Yala, 1261m, 15-111-1982, Schinini & al. 21768(CEN, the free tips with numerous rigid bristles; CTES,G, LIL, MO, NY, SI,US); SanPablo, 3 kmS de marginciliate. Petiole and rachis canaliculate, Yala, 1300m, 15-III-1982, Schinini & al.21769 (CEN, subvillous and with a few scatteredbristles, CTES, G, GH, LIL, MO, NY, SI, US). Dep. canal glabrous. Leaflets with the upper Humahuaca:Sierra de Zenta,2800 m, 3-III-1929, surface smooth, glabrous; the lower surface Venturi8354 (K, LIL,NY, SI,US). with midvein and margin barely marked, subglabrous, with barely visible diminutive Cultivatedmaterial: ARGENTINA. Salta. Esta- adpressed hairs, frequentlywith some long ciónExperimental Agricola, INTA, Cerrillos, culti- hairs on themidvein, rarely a fewlong hairson var INTA LN 83-A,procedente de Jujuy,Lozano, the rest of the surface; margin longly ciliate 1l-m-1982, Schinini & al. 21765(CIES). with one or two shortbristles. Flowers along the lengthof the lateralbranches, rarely at the Geographicdistribution. Arachis montícola tip of the mainstem. Spikes axillary, 5- occurs in the province of Jujuy (Argentina), flowered,of veryshort axis but may reach up whereit was collectedalong theRio Grande,in to 5 cm in lengthat the base of the n+1 basal a small area some 10 kmlong, in the ravinesof branches.Hypanthium villous, usually 5-6 cm said river,between the localities of Lozano and long (3-7 cm). Calyx bilabiate,villous and with Apeadero Reyes, where it forms very scattered bristles, upper lobe 6-7 mm long, localized but dense populationsbetween 1260 lower lobe falcate 7-10 mm long. Standard and 1565 m in altitude. suborbicular,up to 15 mmlong x 17 mmwide, The collectionVenturi 8354, in the Sierrade yellow at the base, otherwiseorange withred Zenta at 2800 m altitude,is doubtfulsince at radial lineson theupper surface; wings yellow, thiselevation the vegetationchanges radically up to 10 mm long. Fruit subterranean, relativeto the places where we observed this Particulate; peg 7-14 cm long, subvillous;. species. Our collections were made in an isthmus 1.5-7 cm long; articles cylindrical, environmentthat fitsin with the vegetational somewhat flattened at the apex, beaked, group "Distrito Chaqueño Serrano," of the usually 15.5 mmlong x 8.5 mmwide, the apical "Chaqueña" vegetational province (Cabrera articlessomewhat larger, up to 2 1 mmx 9 mm, 1976: 27), which reaches some 1800 m above pericarp tough, reticulate, with prominent sea level where the dominant vegetation is longitudinalveins. Seed ca. 13.5 mmlong x 6.5 xerophytic forest alternating with grassy mm wide; 2n=40 chromosomes(Krapovickas steppe. & Rigoni 1951 and 1957). The elevationnoted by Venturicorresponds better with the "Prepuneña" vegetational Holotype:Specimen cultivated in Manfredi (Cor- province, that in Jujuy occupies altitudes doba, Argentina)from seeds collectedin Yala between2000 and 3400 m (Cabrera 1976: 34), (Jujuy,Argentina), III- 1954, Krapovickas8012 withprecipitation below 200 mmannually and (BAB). Isotypes:AS, BAA, C, CTES, F, G, GH,K, withvegetation consisting of shrubbysteppe, LIL, MBM,MO, NY, P, SI, US. stands of thistle, small dwarf forests and bromeliadcushions. Both the "Chaqueña" and Selected additional material:ARGENTINA. the "Prepuneña"formations belong to "Domi- Jujuy.Dep. Dr. M. Belgrano:Yala, 1450m, 14 km nio Chaqueño" vegetationaldomain. NW de Jujuy,23-V-1950, Báez & al. 7264 (seeds It is interestingto note thatBurkart crossed fromthis collection were grown to produce the type the Sierra de Zenta in February1940 on a trip specimen) (BAB, CTES, SI); Yala, 27-1-1940, by mule from Ledesma to Humahuaca, Schreiter 11063 (LIL); Yala, 29-III-1977, without encounteringa single specimen of Krapovickas& al. 30062 (CTES, G, GH,MO, NY, Arachis. US); Jujuy,7-IV-1945, 0'Donell 2776(LIL, NY, SI); Nevertheless, there is a very important Lozano, 20 km NW de Jujuy, 30-111-1977, historicalfact: Salas (1945: 30) indicatesthat at Krapovickas& al. 30063(ŒN, CTES,G, GH, K, LIL, the time of the Conquest, Omaguaca was a

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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) provinceof uncertainextent and thatwithin it diploid species A. duranensis and A. was included the llaman los and withthe A. valley" "...quele stenosperma perennialspecies españoles Del Mani. ["...that the Spaniards Cardenasii and A. Diogoi. In the hybrid call Peanut Valley."] and mentions a obtainedwith A villosa, 45.7% stainedpollen documentdated June 15, 1596 in which it is was observed. " clearly stated ...quebrada de tumbayaques la quebrada del many" ["...Tumbaya Creek 46. Arachis magna Krapov., W.C. which is the creek of the peanuts"]. Tumbaya Gregory & C.E. Simpson nov. sp. is foundabout 45 kmnorth of San Salvador de Figs. 3,46; 26,C-D Jujuy,at 2094 m altitude.Some 20 km to the north,in the Quebrada Purmamarcain Ántigal de Ciénega Grande, Salas himself(1945: 258) Herba annua. Caulis angulosusdum juvenis found some peanut shells (.Arachis hypogaea villosus.Stipulae dorso et base villosae setulis L.) in three graves, which indicates the longis sparsimimmixtis, margine ciliatae. Folia cultivationof peanuts in times prior to the caulis principalisfoliolis oblongisvel interdum Conquest in places higherthan the area known subovatis,acutis, ilia ramorumovalibus usque for A. montícola. obovatis,acutiusculis usque obtusis,epiphyllo glabro,hypophyllo breviter et sparsim adpresso-piloso, Obs. 1. A. montícola is the only species of nervo medio et marginepilis longis vestito. section Arachis with the same number of Hypanthium5-7.5 cm longum,villosum. Calyx 5-6 chromosomes (2n=40) as A. hypogaea and mmlongus,villosus setulis paucis immixtis. Vexillum with which it is possible to obtain fertile 10-14mm longum x 13-16mm latum, aurantiacum, hybrids. Without a doubt, A. montícola is raro luteum.Fructus subterraneus, biarticulatus linked to the historyof the cultivatedpeanut paxillo3-17 cm longo, isthmo usque ad 5 cmlongo, and it is possible that it has had some articulis13-17 mmlongis x 7-9 mmlatis, apice participationin theorigin of thiscultigen. recurvo,pericarpio reticulato nervis manifestis. The material fromLozano (30063) differs somewhat fromthe type specimen in that the Holotype:BOLIVIA. SantaCruz. Prov. Velasco, plant and fruits are somewhat smaller. San Ignacio,60°58'W, 16°22'S, 370 m s.m„ en el Evidently,it pertainsto an isolatedpopulation, bordeS de la población,plaza del Ex Combatiente, despite the short distance (6 km) that plantashasta 2.50 m diám.,eje centralhasta 35 cm separates Lozano fromYala. alt.,ramas procumbentes, 5-V-1977, Krapovickas, Both populations also differ in the Gregory,Schinini & Simpson 30097 (CTES). morphologyof chromosome"B," carrierof the Isotypes:G, GH,K, LIL, LPB, MO, NY, SP, US. satellite.The materialfrom Yala has the "B" pair similarto ,4. hypogaea ssp. fastigiata; on Annualplant. Mainstem erect, 30-55 cm tall, the otherhand, the Lozano materialhas a "B" withoutflowers; lateral branchesprocumbent, pair similar to that of ssp. hypogaea up to 1.30 m long. Stemangular, villous on the (Fernández & Krapovickas 1994). young parts, to subvillous, hairs ca. 1.5 mm Pickett (1955) studied the oils of A. long, occasionally bristlespresent; internodes montícola (Yala) and of A. correntina. The of the mainstemup to 55 mm long, of the analyses show a great similaritywith those lateral branches 30-90 mm long. Leaves generallypublished forA. hypogaea. The oil tetrafoliolate. On the mainstem, the fused ofA. correntinais similarto thatof the runner portionof the stipules 14-19 mm long,the free peanuts (ssp. hypogaea)-,on the other hand, tips 25-37 mmlong x 2-4 mmwide at thebase; the oil of A. montícola resembles the petiole38-52 mmlong, rachis 10-17 mmlong; "Spanish" types (ssp. fasti giata) leafletsoblong, occasionally somewhatovate, (Krapovickas & Rigoni 1957: 441). acute, the apical pair 53 mm long x 20 mm wide but up to 75 mm x 30 mm (length/width Obs. 2. Arachis montícolaproduced highly ratio2. 1-2.6:1), thebasal pair 46 mmx 16 mm sterilehybrids when crossed with the annual but up to 66 mm x 28 mm (length/widthratio

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2.0-2.8:1). On the lateral branches,the fused Obs. Because of the large size of the plant portionof the stipules 8-9 mm long, the free and the formand size of the leaves and the tips 14-21 mm long x 2-3.5 mm wide at the fruit,Arachis magna much resembles A. base; petiole 1 1-25 mm long; rachis 7-11 mm montícolaand it is verydifficult to distinguish long; leafletsoval to obovate, slightlyacute to betweenthem. They differprincipally by the obtuse,apical pair 25-45 mmlong x 17-25 mm level of ploidy, A. magna is diploid, 2n=20 wide (length/widthratio 1.4-1.8:1), basal pair chromosomes,and A. montícola is tetraploid 20-40 mmlong x 13-21 mmwide (length/width and occurs in NW Argentina.It is possible to ratio 1.6-1.9: 1). Stipuleswith the fused portion differentiatethem because inA montícolathe frommore or less villous to subglabrous,with back of the leafletsis subglabrouswith barely long hairs moredense along thedorsal line and visible diminutiveadpressed hairs and long toward the base, thereare also long scattered hairs only on the midvein. In contrast,in A. bristles;free portion with surfaces glabrous or magna, while the back of the leaflets has a with some long hairs and bristlestoward the similarindumentum, there are also some long base, margin ciliate. Petiole and rachis hairs,ca. 2 mmlong, on therest of the surface. canaliculate, villous. Upper surface of the The youngplants ofA. magna show greater leaflets glabrous, smooth; lower surface differences,since the cotyledonarybranches midvein and margin somewhat marked,with develop afterthe firstflowers have produced small, scattered,adpressed hairs and withlong fruitsand at this stage the plants resemble hairs along the midvein and on the margin. species of the sections Trierectoides and Flowers along the length of the lateral Erectoides. branches,in very short,5-7-flowered axillary Arachis magna is veryclose to A. ipaënsis, spikes. Hypanthium5-7.5 cm long, villous. as muchfor its growth form as forits fruit.We Calyx bilabiate, villous and with a few long believe it properto separate thembecause A. bristles; upper lobe 4-dentate, 5 mm long, ipaënsis is a less vigorous plant and all of its lower lobe falcate,6 mmlong. Standard 10-14 parts are smaller. The species can be mm long x 13-16 mm wide, orange, rarely separated by the presence of bristles on the yellow, wings yellow, 6-8 mm long. Fruit stipules, which is constant in all of the subterranean,Particulate; peg 3-17 cm long, collections of A. magna and lacking in A. with long hairs very scattered on the aerial ipaënsis. Furthermore, the latter species part; isthmusup to 5 cm long; articlesusually occurs in the departmentof Tarija in southern 13 mm long x 7 mmwide but up to 17 mmx 9 Bolivia, near Villa Montes, in thickets of mm, with marked beak, pericarp reticulate Bromelia serra Griseb., a typically withprominent veins. Seed 10-11 mmlong x 5- "chaqueña" species. 6 mm wide. 2n=20 chromosomes(Fernández & Krapovickas 1994). 47. Arachis ipaënsis Krapov. & W.C. Gregory nov. sp. Additionalmaterial: BOLIVIA. Santa Cruz. Figs. 3,47; 25 Prov.Ñuflo de Chavez:El Carmen,ca. 42 kmSSW de Concepción, 62°26'W, 16°38'S, l-V-1977, Krapovickas& al. 30092(CTES, US); Embocadadel A. ipaënsisM.P. Gregory& W.C. Gregory.J. Carmen(ca. 40 kmSSW de Concepción),2-V-1977, Hered. 70: 192. 1979, nomennudum. Refers to Krapovickas& al. 30093 (CEN, CTES, G, GH, K, "Parentno. 70" (Krapovickas& al. 19455). LPB,MO, NY, P, SI,US). Herba annua. Caules glabrescentes.Stipulae Geographic distribution. To date, this dorso paucis pilis longis vestitae, margine species has been collected in the northeastof ciliatae, setulisdestitutae. Epiphyllum glabrum, the departmentof Santa Cruz, Bolivia, to the hypophyllumnervo mediopilis 1.5 mm longis, northof the Chiquitano massif, in an area of ceterumjuventute pilis brevioribusadpressis savannahs and rollinghills. It grows in deep sparsimvestitum, deinde glabrescens,margine humus-bearingsoils. non incrassatociliatum et setulisraris brevibus

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Fig.25. Arachis ipaënsis : A, plant; B, apicalarticle; C, basalarticle of the fruit (K. 19455). instructum.Hypanthium villosum 5-15 cm internodesof the mainstemup to 75 mm long, longum. Calyx 5-6 mm longus, villosus et internodesof the lateralbranches up to 55 mm setulosus. Vexillumaurantiacum1 , 11-13 mm long. Leaves tetrafoliolate, leaflets from longumX 14-15mm latum. Fructus subterraneus, obovate to elliptical. On the mainstem, the biarticulatuspaxillo glabro 5-22 cm longo, fused base of the stipules 7-13 mm long, the isthmo1-15 mm longo, articulis 14-17 mm longisx free tips 17-20 mm long x 2 mm wide; the 7-10 mmlatis, pericarpio reticulato. petiole usually 45 mm long, varyingbetween 35-53 mm; the rachis 10 mm long; the apical Holotype:BOLIVIA. Tarija. Ipa, Quebradade leafletscommonly 38 mm long x 18 mm wide Thaiguate(30 kmN de VillaMontes) 63°25'W, 2 1°S, but up to 45 mm x 25 mm, the basal leaflets 650 m s.m.,3-VI-1971, Krapovickas, Mroginski & somewhat smaller, ca. 33 mm long x 15 mm Fernández19455 (CTES). Isotypes:CEN, G, K, LIL, wide but up to 42 mm x 20 mm. Leaves on the LPB,MO, NY, P, SI, US. lateral branches with the fused portionof the stipules3-5 mm long,the freeportion 9-10 mm Taprootedperennial. Mainstem up to 45 cm long x 2 mm wide. The petioles usually 20 mm in length,with few branches at the base. The long (up to 32 mm); rachis 7-10 mm long. The firsttwo cotylendonarybranches begin with apical pair of leafletsca. 23 mm long x 18 mm numerous clustered inflorescences,that fruit wide (up to 28 mm x 18 mm), the basal pair rapidlyand laterelongate to formprocumbent some 20 mm long x 14 mm wide (up to 27 mm branches ca. 50 cm long. Stems glabrescent, x 16 mm). Stipules slightlyimbricated at the the youngerportions partly villous, withmore base, surfaces glabrous, except for some long or less erect long hairs ca. 1.5 mm long; hairs on the back of the fused portionand cilia along the margin.Petiole, rachis and pulvinus 'Errorin original, appeared as luteum(yellow). with some long scattered hairs. Leaflets

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glabrous above and glabrescentbelow, when pilorum,dorso pilis longis instructae,margine youngwith very short, sparse, adpressedhairs, ciliatae.Folia caulisprincipalis foliolis oblongis, midveinwith some long hairs (1.5 mm long), acutis ilia ramorumfoliolis oblongis,epiphyllo margin not thickened,somewhat wavy when laevi,glabro, hypophyllo nervo medio prominen- dry, ciliate, rarely with an occasional short te,nervis secundariis manifestis, praeter pilos ca. bristle. Flowers occur along the lengthof the 2 mmlongos in nervomedio glabro, margine vix lateral branches and are also clusteredat the conspicuobreviter ciliato. Hypanthium 7-10.5 cm base of theplant. Hypanthium villous, 5-15 cm longum,villosum. Calyx 7-8 mmlongus, villosus. long, the longest ones near the base of the Vexillumca. 15 mmlongum, aurantiacum. Fructus plant. Calyx bilabiate, the wider lobe 5-6 mm biarticulatuspaxillo ca. 12 cmlongo, isthmo ca. 6 long, the narrowerlobe falcate, 6-7 mm long. mmlongo, articulis 12-17 mmlongis x 8-10 mm Corolla orange (see footnote7),standard 11-13 latis,apice recurvo,pericarpio valde reticulato. mmlong x 14-15 mmwide, wings 6-8 mmlong. FruitParticulate; pegs glabrous,5-22 cm long, Holotype:BRAZIL. Mato Grossodo Sul. 35 km superficial;isthmus 1-15 mm long; articles 14 SE ofCorumbá. Fazenda Vale do Paraisoon roadto mm long x 7 mm wide but up to 17 mm x 10 PortoManga, and then6 kmN towardBaía Negra mm, epicarp reticulate, with more or less andBanda Alta, 19°1 l'S, 57°29'W,80 m,in a palm prominentveins. 2n=20 chromosomes(M.P. gladeof carandaí (Copernici a sp.),in darklow soil Gregory&W.C. Gregory,1979). weatheredfrom limestone and probablyinundated intimes of high water, 10-XII-1976. Krapovickas & Additionalmaterial: From the same locationas Gregory3001 1 (CEN). Isotypes:CTES, G, GH, K, theholotype: 12-IV-1977, Krapovickas & al. 30076 MO, NY, P, RB, SI, SP,US. (CTES,NY,US). Annualplant, very vigorous. Mainstem 5-60 Geographic distribution.This species has cm tall, withoutflowers, much branched;late- been found, up until now, only in the type ral branches procumbent,up to 1.5 m long, locality,where it grows on the upper parts of branched; internodes3-6.5 cm long, more or streamravines, among bromeliads. less villous. Leaves tetrafoliolate. On the mainstem,stipules with fused portion 13-17 Obs. Arachis ipaënsis formedhybrids only mm long, the freeends 32-34 mm long x 2-3 withannual diploid species of sectionArachis, mm wide at the base; petiole 38-55 mm long; such as A. duranensisand A. Batizocoi and with rachis 10-15 mm long; leafletsoblong, acute, A. hypogaea var. hypogaea, producingin all theapical pair 50 mmlong x 18 mmwide, up to cases very sterilehybrids with less than 3.7% 75 x 24 mm (length/widthratio 2.7-3.1:1), the pollen staining. These results show that A. basal pair ca. 42 mmlong x 16 mmwide, up to ipaënsis,which lives nearthe edge of theranges 63 x 18 mm (length/widthratio 2.6-3.5:1). On ofA duranensisand ofA. Batizocoi,is an entity the lateral branches,the fused portionof the thathas developedsome veryeffective genetic stipules 7-8 mm long, the freetips 16-19 mm isolationmechanisms, comparable to thosethat long x 3-5 mm wide at the base; petiole 8-31 separateA. duranensisfrom A. Batizocoi. mm long; rachis 6-10 mm long; leaflets Morphologically,A. ipaënsisis verysimilar to eliptical,apical pair ca. 24 mm long x 13 mm A. magna. The latterspecies occursin theextre- wide,up to 5 1 x 26 mm(length/width ratio 1.8- me NE of thedepartment of Santa Cruz in Boli- 2.25: 1), basal pair 21 mm long x 10 mmwide, via. up to 42 x 18 mm(length/width ratio 1.9-2.3: 1). Stipules with long hairs on the back of the 48. Arachis valida Krapov. & W.C. fusedportion, surfaces glabrous or witha line Gregory nov. sp. of hairs along the center of the free parts, Figs. 3,48; 26,A-B marginciliate. Petiole and rachis canaliculate, withthe back villous and the canal glabrous. Pulvinus villous. Leaflets with the upper Herba annua.Stipulae glabrae aut medioserie surface smooth and glabrous, the lower

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Fig.26. Arachis valida : A, leaffrom mainstem; B, leaffrom lateral branch (K.3001 1). A. magna: C, basalarticle; D. apicalarticle of the fruit (K.30093). A. Hoehnei: E, leaffrom mainstem; F, leaf from lateral branch (Sc.21450).

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surface with prominentmidvein and marked Holotype:BOLIVIA. Dep. Beni. Trinidad,Uni- secondary veins, glabrous except for some versidadTécnica del Beni, 15 m E deledificio labo- longhairs along themidvein; margin somewhat ratorios;fruto reticulado, 21-III-1990. Williams & marked on the underside and shortlyciliate. Claure11 18 (CTES). Flowers along the length of the lateral branches,in axillaryspikes up to 10 mmlong, Annual plant. Mainstem erect, ca. 10 cm 3 -5-flowered; bracts ciliate on the margin. tall, lateralbranches procumbent, 35 cm long; Hypanthium7-10.5 cm long, villous. Calyx stem quadrangular,villous, hairs 1.5-2 mm bilabiate, villous and with dispersed bristles, long. Leaves tetrafoliolate,stipules villous upper lobe 4-toothed,7 mm long, lower lobe towardsthe base and withbristles over almost falcate, 8 mm long. Standard orange, ca. 15 the entiresurface, petiole and rachis villous, mm long; wings yellow, ca. 7 mm long. Fruit upperleaf surfacesmooth, glabrous, underside Particulate; peg ca. 12 cm long, somewhat withmargin and veins slightlymarked, small villoustoward the base; isthmusca. 6 mmlong; very adpressed hairs and long hairs (1 mm articles 12-17 mm long x 8-10 mm wide, the long) on the midveinand margin,where there apical article somewhatlarger than the basal, are also a few shortbristles. On themainstem, covered by a thin coat of diminutivehairs; stipuleswith the fusedportion 10-12 mm long pericarpstrongly reticulate, with a pronounced and thefree portion 15-20 mmlong x 2.5-4 mm beak. 2n=20 chromosomes (Fernández & wide, petiole 30-35 mm long, rachis 10 mm Krapovickas 1994). long,leaflets oblong, the distal ones 32-34 mm longx 16-18 mmwide, the proximal ones 28-38 Additionalmaterial: BRAZIL. Mato Grossodo mm long x 12-16 mm wide. On the lateral Sul. Fromthe type locality: l-VII-1977, Krapovickas branches,stipules with the fiisedportion 8-11 & al. 30147(CEN, CIES, NY, US); 12-X-1985,Valls mmlong, freeportion ca. 16 mm long x 3 mm & al. 9153(CEN, CIES); id.,Valls & al. 9157(CEN, wide, petiole 12-20 mm long, rachis 6-10 mm CIES); id.,Valls & al. 9162(CEN). long, distal leafletsobovate, 23-35 mm long x Geographie distribution.This species is 15-21 mmwide, proximalleaflets oblong, 21- known only fromthe type locality, where it 29 mmlong x 13-16 mmwide. Hypanthium30- grows in inundatedgrasslands with palms of 75 mmlong, villous. Calyx villous,with bristles the genus Copernicia. on the margin,upper lobe 5 mm long, lower lobe falcate,7 mmlong. Standard 12 mmlong, 49. Arachis Williamsii Krapov. & W.C. orange. Fruit subterranean,Particulate; peg Gregory nov. sp. 10-15 cm long, glabrous or with some short Fig. 3,49 hairson theexposed parts,articles 1 1 mmlong x 6.4 mmwide, stronglyreticulate, with beak.

Herba annua.Radix polaris. Caulis pilis 1.5-2 Additionalmaterial: BOLIVIA. Beni.Prov. Cer- mmlongis villosus, principalis erectus ca. 10 cm cado,Trinidad, Campus of the Universidad Técnica longus,rami procumbentes. Stipulae basi villosae del Beni, 155 m, 14°48'S, 64°52'W, in recently et setulislarge instructae.Folia caulisprincipalis cultivatedmaize field 400 m SE ofUniversity buil- foliolis oblongis,ilia ramorumfoliolis distalibus ding. Soil dark grey sandy clay, 25-XI-1988, obovatis, proximalibus oblongis. Epiphyllum Williams867 (CTES); id.,500 m al SE del edificio laeve, glabrum,hypophyllum margine et nervis laboratorios,22-IÜ-1990, Williams 1120 (CTES). paulo manifestis,pilis parvisperderne adpressis, nerviomedio et marginemajoribus (1 mmlongis) Cultivatedmaterial: ARGENTINA. Córdoba. vestitum.Hypanthium 30-75 mm longum, villosum. Manfredi,E.E.A. (INTA). Cult.66, origin:Williams Calyx5-7 mmlongus, villosus, margine setulosus. 1118,21-m-1991, Krapovickas 43797 (CIES); id,43798 Vexillumaurantiacum, 12 mm longum.Fructus (CIES); id,27-111-1992, Krapovickas44092 (CIES). subterraneus,biarticulatus paxillo 10-15cm longo, subglabro,articulis 11 mmlongis x 6.5 mmlatis, Geographic distribution. This species is apice recurvo,pericarpio notabiliter reticulato. known only fromthe type locality: Trinidad,

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departmentof Beni (Bolivia). hairs and some bristles.Upper surface of the We dedicate this species to Dr. David leaflets glabrous, smooth, and the lower Edison Williams who recentlydiscovered it surface with veins and margin littlemarked, and is the author of the doctoral thesis with hairs 1.5-2 mm long, scatteredover the "Peanuts and Peanut Farmersof the Rio Beni: whole surface,margin longly ciliate and with Traditional Crop Genetic Resource some short bristles. Spikes axillary, 5- Managementin the Bolivian Amazon." flowered, with axis ca. 5 mm long. Hypanthiumvillous, 5 cm long (2.5-8 cm). Commonname, "manicillo" (Williams 867). Calyx bilabiate,with some long silkyhairs and with bristles,upper lobe 4-7 mm long, lower 50. Arachis Batizocoi Krapov. & W.C. lobe falcate, 5-7.5 mm long. Standard 10-12 Gregory mm long x 10-14 mm wide, lightyellow with Fig. 3,50 tenuousradial, reddish lines, more pronounced on the dorsal surface and the spaces between the lines with violet tinge; wings yellow, 7-8( Krapovickas& Gregory,Bonplandia 3(1 1): 129, mmlong. Fruitbiarticulate; peg 5-12 cm long, 1974. villous on the aerial portion;isthmus 0.5-4 cm A. BatizocaeKrapov. & W.C. Gregory,in W.C. long; articles 10-15 mmlong x 7-10 mmwide, Gregory,M P. Gregory,Krapovickas, Smith & with pronounced beak, pericarp smooth or - Yarbrough,in Peanuts Cultureand Uses: 96, lightlyreticulate. Seed 8-12 mmlong x 5-6 mm 1973,nomen nudem. wide. 2n=20 chromosomes (Krapovickas, Fernández& Seeligmann 1974). Perennialor biennialherb. Taproot, deep but frail. Mainstem 0.30-1 m long, erect when Holotype:BOLIVIA. SantaCruz. Prov. Cordille- young, then recumbent,little branched, with ra,Parapetí, en suelode arena,cerca de la estación flowers toward the apex. Lateral branches del ferrocarrilSanta Cruz-Yacuiba.24-11-1958, (n+1) usually only 4, procumbent,little or no Krapovickas9505 (LIL). Isotypes:CTES, F, G, branching,up to 4 m long. Stems angular, MICH,MO, SI. villous, especially on the young parts; internodes4-10 cm long. Leaves tetrafoliolate, Additionalmaterial: BOLIVIA. Chuquisaca. L. those of the mainstem notably larger than Clavo,El Salvador-CIMBOC,20°34'S, 63°8'W, 7-IV- those of the lateral branches. On the 1993,Saravia Toledo & al. 11442. Santa Cruz. Cor- mainstem,the fused portionof the stipulesup dillera,Parapetí, 24-11-1958, Krapovickas 9484 to 12 mm long, the freeportion up to 21 mm (CIES, LIL),9495 (CTES, LIL), 94% (LIL),9497 (LIL), long; petioleup to 70 mmlong; rachis up to 22 9498(LIL), 9503 (LIL), 9504 (LIL); Paja Colorada, 28 km mmlong; apical pair of leafletsoval, usually 50 S deCamiri, 14-IV-1977, Krapovickas & al. 30079 (CEN, mm long x 35 mm wide (up to 60 mm x 43 CIES, G,GH, K, LIL, LPB, MO, NY, P, US); Ipatí (37 km mm), basal pair oblong, 40 mm long x 23 mm N de Camiri),16-IV-1977, Krapovickas & al. 30080 wide (up to 59 mm x 33 mm). On the lateral (CEN,CTES, G, GH, LIL, LPB, MO, NY, US); 5 kmS de branches,the fused portion of thestipules up to Lagunillas,16-IV-1977, Krapovickas & al. 30081 (CEN, 8 mmlong, the free portion up to 12 mmlong x CTES, G,GH, K, LIL, LPB,MO, NY, US); 3 kmN de 5 mmwide at thebase; petiole 15-35 mmlong; Ipati,17-IV-1977, Krapovickas & al. 30082 (CEN, rachis 10-15 mm long; apical pair of leaflets CTES, G, GH, K, LPB, MO, NY, US); 22 kmN de fromsuborbicular to obovate, usually 27 mm Gutierrez,17-IV-1977, Krapovickas & al., 30083 (ŒN, long x 25 mm wide (up to 35 mm x 32 mm), CTES, G, GH, K, LIL, LPB, MO, NY, P, SI, US); basal pair oval, 26 mm long x 20 mmwide (up Lagunillas,1000 m, 11-195 1,Cárdenas 4741 (LIL, US). to 3 1 mmx 26 mm). Stipules pilose towardthe PARAGUAY.Boquerón (ex NuevaAsunción). base and on theback of thefused portion, with RutaTrans-Chaco, 25 km S de Nueva Asunción, scattered bristles, more abundant near the 20°50'S,61°55'W, 12-XÜ-1987, Schinini &al. 25658 union withthe petiole, marginciliate. Petiole (CTES); RutaTrans-Chaco, 20°40'S, 62° W, 12-XII- and rachis canaliculate, with long scattered 1987,Schinini & al. 25704(CTES).

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Geographic distribution.This species was destitutae.Folia caulis principalisfoliolis ovali- collectedin the SW of the departmentof Santa oblongis, ilia ramorumfoliolis rotundatis, Cruz (Bolivia) at altitudesvarying between 700 epiphylloglabro, hypophyllopilis ca. 2 mm and 1000 meters above sea level, in open longis,serieeis sparsim, in nervomedio densius places with a soil of almost pure sand, or on vestito,margine vix manifesto pilis longissparsim low sandy hills. The southernlimit of its range instructae.Hypanthium 3-6 cm longum,pilosum. is foundin El Salvador, near Carandayti,in the Calyx7 mmlongus, pilis longissericeis setulisque department of Chuquisaca. It was also vestitus.Vexillum suborbiculare, 10 mmlongum x collected in the vicinityof Nueva Asunción,in 9 mmlatum, aurantiacum, raro luteum.Fructus the Paraguayan "chaco," near the Bolivian biarticulatuspaxillo 3.5-14 cm longo, glabro, border. isthmo4-11 cm longo, articulis 10.5-15.5 mm longisx 5-8 mmlatis, tenuiterreticulato paene Obs. Within section Arachis, A. Batizocoi laevi. crosses with the annual species A. duranensis, A. ipaënsis and A. stenosperma Holotype:ARGENTINA. Córdoba.Manfredi, and withthe perennialspecies A. Cardenasii, cult,de semillasprocedentes de Campo Durán, A. correntina, A. Diogoi and A. villosa, prov.Salta, Argentina (Krapovickas 29-V-1953), III- producing,in all cases, highlysterile hybrids. 1954,Krapovickas 8010 (CTES). Isotypes:BAB, With species from the other sections, A. CEN,F, G,GH, K, LIL,MO, NY, P, SI,UC, US. Batizocoi has only crossed withthe tetraploid rhizomatousspecies, A. pseudovillosa and A. Annualplant. Taproot tetrarchial. Mainstem glabrata var. glabrata, producing hybrids erect,20-30 cm tall, withnumerous vegetative thatnever flowered or are highlysterile. branchesgrowing from the lowerhalf and with Smartt, Gregory & Gregory (1978a,b) flowerstoward the apex; internodesup to 1.5 proposed the B genome for A. Batizocoi, as cm long. Lateral branchesprocumbent, up to opposed to the A genome found in at least 9 1.40 m long, the basal ones horizontal,those other species in section Arachis, due to the originatinghigher up on themainstem bending low genetic compatibilityof A. Batizocoi as towardthe soil describinga recliningelongated demonstrated by the production of highly S; along the cotyledonary branches, two sterilehybrids. vegetativeand two reproductiveaxes alternate more or less regularly, on the remaining 51. Arachis duranensis Krapov. & W.C. branches there is a predominance of Gregory nov. sp. reproductivebranches; stems green but witha the the Fig. 3,51 violet tinge on parts exposed to sun; internodesusually 3 cm long,ranging from 1.5 up to 4 cm long, villous on the young parts. A. argentinensisSpeg. Hoehne. Genero Arachis. Leaves tetrafoliolate.On the mainstem,the FloraBrasílica 25(2) part.122: 7 & 14,1940, nomen fusedportion of the stipules8 mmlong x 4 mm nudumin syn. sub. A. villosa.(USNA 16177,Argen- wide, the freetips 14-18 mm long x 2-3 mm tina,Salta, Spegazzini) wide, acute; petiole 23-36 mm long; rachis 8- A. duranensisKrapov. & W.C. Gregory,in W.C. 13 mmlong; apical leafletsusually 30 mmlong Gregory,M.P. Gregory,Krapovickas, Smith & x 22 mm wide (up to 36 mm x 24 mm), basal Yarbrough,in Peanuts- Cultureand Uses: 96, leaflets26 mm long x 18 mm wide (up to 32 1973,nomen nudum. This refersto the specimen mm x 21 mm). On leaves of the lateral Krapovickas7988. branches,the fused portionof the stipules4-5 A. SpegazziniiM.P. Gregory& W.C. Gregory.J. mm long x ca. 3 mm wide, the freepart 7-11 Hered.70: 192. 1979,nomen nudum. "Parent no. 8" mm long x 2 mmwide; petiole 7-11 mm long; (Gregory& al. 10038). rachis ca. 4 mm long; apical pair of leaflets usually 14 mm long x 13 mm wide (up to 17 Herba annua. Stipulaesubglabrae, dorso pilis mmx 16 mm),basal pair of leaflets12 mmlong longis vestitae, margine ciliatae, setulis x 10 mmwide (up to 15 mmx 13 mm). Stipules

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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) withsurfaces subglabrous to lightlypubescent, Dep. Ledesma:ruta 34, 4 kmS de FrailePintado, 9- withlong hairs on theback of thefiised portion Xn-1970,Schinini & al. 19539(CTES); id.,9-V-1983, and ciliate margin, without bristles. Petiole Krapovickas& al. 38905(CTES). Salta. Dep. Capi- somewhatcanaliculate, with long hairs on the tal:Salta, 7-III-1905, Spegazzini (BAB 13765);Río back and pubescent in the canal. Leaflets of Arias,Camping Municipal, 4-V-1980, Krapovickas the mainstemoval-oblong, those of the lateral & al. 36036(CEN, CTES, G,GH, K, LIL, MO, NY, P, branchesrounded, occasionally weaklyretuse, SI, SP, US); SportingClub, 10-111-1982, Schinini & mucronulate;upper surface glabrous, lower al. 21763(BAB, CTES, F, G,LIL, MO, NY, P, SI,UC, surfacewith long, scattered,silky hairs, ca. 2 US); San Luis, 11-1-1949, Filipovich 404 (LIL); El mm long, more abundant on the midvein, Prado,5-II-1949, Legname 946 (LIL); El Prado,río marginslightly marked, with sparse long hairs. Arenales,7-V-1959, Gregory & al. 10038(CTES, GH, Axillaryspikes up to 5-flowered,axis up to 1 LIL, MO,NY, SI, SP,US); SanLuís, río Arenales, 10- cm long. Hypanthium3-6 cm long, pilose. V-1975,Krapovickas & al. 28458(C, CTES,LIL, UC, Calyx bilabiate, covered withlong silkyhairs US); San Luís, 11-III-1982,Schinini & al. 21764 and withbristles; upper lobe 7 mm long. Stan- (CEN, CTES, GH,K, MEXU, MO, NY, SI, US); La dard suborbicular, yellow, bordered with Loma,21-111-1979, Fernández613 (CTES); La Loma, orange or yellow, 10 mm long x 9 mm wide, GrandBourg, lO-IV-1980, Krapovickas & al. 36006 upper surface striatedwith radial red lines; (CEN, CTES, G, GH,LIL, MO, NY, SP, US); Aero- wings 7 mm long x 3.5 mm wide, yellow; keel puertode Salta,9-IV-1980, Krapovickas & al. 36005 falcate, 8 mm long, yellowish white. Fruit (CEN,CTES, G,GH, K, LIL, MO, NY, RB,UC, US). Particulate, peg 3.5-14 cm long, almostverti- Dep. La Caldera:estancia La Despensa,16-IV-1943, cal; isthmus7 (4-1 1) cm long,glabrous, violet Hunziker1635 (CORD, LIL, SI). Dep. Metán:14 km on the aerial part, pilose on the subterranean WNWde El Tunal,río Juramento, flor amarilla, 4-IV- part near the soil surface; articlescylindrical, 1980,Krapovickas & al. 36002(CEN, CTES, G, GH, flattenedat the apex and withbeak, 13 (± 2.5) K, LIL, MO, NY, SI, SP, US); id.,flor anaranjada, mm long x 6.5 (± 1.5) mm wide; pericarp Krapovickas& al. 36003(CEN, CTES, G,GH, K LEL, weakly reticulate; epicarp densely covered MO, NY, SI, SP,US). Dep. Anta:La Lagunita(Coro- with thinhairs that stronglyretain a layer of nel Olleros),9-III-1958, Morello & al. 356 (CTES, soil. Seed 10.5 (± 2) mm long x 5 (± 1) mm LIL); Palermo,Rio del Valle,64°14'W, 24°41'S, 23- wide. 2n=20 chromosomes (Krapovickas & III-1980,Vorano s/n (CTES, MCNS,US); 5 kmN de Rigoni 1957: 438, sub. A. pusilla, Smartt& Palermo,21-III-1982, Schinini & al. 21772 (CTES, Gregory1967, GKP 10038). MO, NY); ríodel Valle, 5 kmN de Las Lajitas,24-IV- 1983,Krapovickas & al. 38900 (CTES); 1 kmS de Additionalmaterial: ARGENTINA. Jujuy. Dep. Anta,2 l-ffl-1982,Schinini & al.2 1770(CTES, G, GH, Capital:2 kmS de Palpalá,flor anaranjada, 29-III- LIL,MO, NY, SI,US); id.,1 l-V-1983, Krapovickas & 1977,Krapovickas & al. 30061-A(CTES, GH,MO, al. 38906 (CTES); Anta,24°55'S, 64°25'W, 21-III- NY, US); id.,flor amarilla, Krapovickas & al. 30061-B 1982,Schinini & al. 21771 (CTES, LIL, MO, US). (CTES, GH, MO, NY, SI, US); id., población, Dep. Chicoana:Los Los,Filipovich 415 (LIL). Dep. Krapovickas& al. 30061(CEN, CTES, F, G, GH,K, Gral. Joséde San Martín:Piquirenda, Quebrada LIL, MO, NY, P, SI, SP, UC, US); id., 8-IV-1980, Yacuí,3-II-1925, Schreiter 3841 (LIL); id.,Schreiter Krapovickas& al. 36004(CEN, CTES, G, GH,LIL, 4107 (LIL); CampoDurán, 22-1-1930, Horovitz (SI); MO, NY, RB,SI, US); 8,6km SE de S.S. deJujuy, ruta id.,Parodi 9189 (SI); id.,29-V-1953, Krapovickas 66, 14-m-1982,Schinini & al. 2 1767(CEN, CTES, G, 7988 (CTES, G, LIL, SI, US); id.,Krapovickas 7993 GH,K, LIL, MO, NY, SI, SP, US). Dep. El Carmen: (CTES, SI); id.,6-VI-1972, Krapovickas & al. 19490 Perico,17-1-1906, Spegazzini (BAB 15508);Perico, 1 (CTES, G, MO, NY, UC, US); id., 8-V-1983, kmN de ríoPerico, 28-III-1977, Krapovickas & al. Krapovickas& al. 38903 (CTES); 3 kmE de Senda 30060(CTES, G, GH,LIL, MO, NY, SI, US); Perico, Hachada,río Seco, 2-IV-1977,Krapovickas & al. aeropuertoEl Cadillal,3 1-III- 1977, Krapovickas & 30065(CEN, CTES, F, G, GH, K MO,NY, P, SP,UC, al. 30064(CEN, CTES, GH,K LIL,MO, NY, US); rio US); 5 kmWde Dragones,2-IV-1977, Krapovickas Perico,aeropuerto El Cadillal,12-III-1982, Schinini & al. 30066(CEN, CTES, G,GH, K LIL, MO, NY, P, & al. 2 1766(CEN, CTES, G,GH, LIL, MO,NY, US). RB, SI, UC, US); id., 9-V-1983,Krapovickas & al.

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38904 (CTES); Dragones, 12-III-1992,Saravia Andes. Its range is tied to the rivers Toledo & al. 10766 (CTES); Senda Hachada,río Pilcomayo,Bermejo (withits tributaryGrande Seco,4-IV-1977, Krapovickas & al. 30067(CTES, G, or San Francisco) and Pasaje or Juramento GH,LIL, K, MO, NY, SI, SP, US); 16km W de Gral. (with its tributariesArenales and Arias). The Ballivián,en arroyoafluente del rio Seco, 4-IV-1977, species grows on a stripof land at the base of Krapovickas& al. 30068(CEN, CTES, G, GH, K, LIL, the first foothills of the Andes, from the MO, NY, P, SI, UC, US). departmentof Chuquisaca (Bolivia) to the BOLIVIA. Tanja. Prov.Gran Chaco: Tatarenda, provinceof Salta (Argentina),between 250 m 25-III-1 902, Fries 1465 (G); 18 kmN de Yacuiba, (Dragones, Salta, Argentina) and 660 m caminoa Villa Montes,ayo. Coloradito del Palmar (Caraparí, Tarija, Bolivia). Along the San (quebradade Tatarenda),7-IV-1977, Krapovickas & Francisco River,passing throughFraile Pinta- al. 30070(CEN, CTES, G, GH, K, LIL,LPB, MO, NY, do, its range reaches as far as Palpalá (Jujuy P, SI, US); 30 kmN de Yacuiba,Campo de La Tapia, province)at 1 100 m altitudeand extendsalong Caiza, 600 m,7-IV- 197', Krapovickas& al. 30069 the rivers Arias and Arenales as far as the (CEN,CTES, G, GH, K, LIL,LPB, MO, NY, P, SI, UC, area surroundingthe city of Salta (El Prado, US); 2 kmWde Saladillo,rio Caraparí, 1000 m, 8-IV- 1250 m s.m.). It grows in deep, sandy soils, in 1977,Krapovickas & al. 30071(CTES, G, GH,LIL, the vicinityof watercourses. LPB,MO, NY, US); 14km N deCaraparí, 870 m, 8-IV- 1977,Krapovickas & al. 30072(CEN, CTES, G, GH, Obs. 1. Priorto 1977, we had only two live K, LIL, LPB, MO, NY, P, SI, US); 32 km N de accessions, one from the Arenales River Yacuiba,540 m, 10-IV-1977, Krapovickas & al. 30073 (parentno. 8, GKP 10038) at 1250 m elevation (CEN,CTES, G, GH, K, LIL,LPB, MO, NY, P, SI,SP, and 10 km to the W of the city of Salta, with US); 2 kmN de PalmarGrande (38 kmS de Villa yellow flowers, and the other from Campo Montes),400 m, 10-IV-1977,Krapovickas & al. Duran (parent no. 39, K 7988) at 500 m 30074(CEN, CIES, G,GH, K, LIL,LPB, MO, NY, P, elevationand ca. 400 km to the NE of the city SI, SP,US); VillaMontes, XI-1910, Herzog 1 1 10 (G); of Salta, with orange flowers. As these Simbolardel Carmen, 20 kmE deVilla Montes, 28-V- accessions had somewhatdifferent behaviors, 1971,Krapovickas & al. 19408(CTES, LPB,US); id., we regardedthem as independentspecies: the 2-IV-1971,Krapovickas & al. 19435(CEN, CTES, G, first A. Spegazzinii and the second A. LEL,LPB, MO, NY, SI, US); 5 kmS de VillaMontes, duranensis. The hybrids obtained between caminoa Yacuiba,450 m, 10-TV-1977,Krapovickas these two populationsproduced between 31.3 & al. 30075(CEN, CTES, G,GH, LIL, LPB, MO, NY, and 36.7% pollen staining.Nevertheless, the P, SI, US); 7 km N de Villa Montes,6-V-1983, behavior of the two parents (8 and 39) in Krapovickas& al. 38901 (CTES); 18km S de Villa crosseswith other wild specieswas verysimilar. Montes,7-V-1983, Krapovickas & al. 38902(CTES); Beginningin 1977, withthe intensification of 3 kmS de VillaMontes, 1 l-XII-1990, Saravia Toledo theexplorations and thanksto thecollaboration 2738(CTES). Chuquisaca.Prov. Acero: Carandayti, of Ing. Agr. AlfredoVorano of the Estación III-1935,Rojas 7387 (G, LIL, SI); EstaciónExperi- ExperimentalAgropecuaria (INTA) of Salta, mentalZootécnica "El Salvador,"27 km W de the collectionsof wild peanuts fromJujuy and Carandayti,500 m, 13-IV-1977,Krapovickas & al. Salta were increasedconsiderably, allowing us 30077(CEN, CTES, G,GH, K, LIL,LPB, MO, NY, P, to cover the range between those old SI, US); El Salvador,6-VI-1991, Saravia Toledo 2989 populationswith new accessions. The color of (CTES); 7 kmW de Carandayti,500 m,13-IV-1977, the flowersdid not turnout to be significant, Krapovickas& al. 30078(CTES, MO, NY, US). given thatpopulations were encounteredwith PARAGUAY.Alto Paraguay (ex Chaco).Mayor bothkinds of flowers,both in the cityof Salta Pedro Lagerenza,60°45'W, 20°5'S, 16-IV-1978, as well as furtherto the east. In Palpalá (Jujuy Schinini&al.15101 (CTES, G, SI,US). province), the yellow flower was associated with green stems and the orange flowerwith Geographicdistribution. Arachis duranensis somewhat purple stems. Nevertheless,in El is a species associated with the watershedof Tunal (Salta province),where specimenswith the Rio de la Plata, at the foothillsof the yellow flowers(K 36002) and orange flowers

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(K 36003) growtogether, this association does glabrata and A. pseudovillosa. notexist. It is remarkable the ease with which A. With the increase in collections, it was no duranensis crosses with other species, longer feasible to differentiatethe two initial especially whentaking into account its margi- populations, so we decided to consider all of nal positionat the extremewestern edge of the thediploid annual material from NW Argentina overall range of the genus Arachis. as pertainingto a single species, characterized by the presence of long hairs (ca. 2 mm long) 52. Arachis Hoehnei Krapov. & W.C. on theunderside of theleaves, stipuleswithout Gregory nov. sp. and fruitswith a smooth bristles, pericarp. Figs. 3,52; 26,E-F

Obs. 2. Both the morphology of the chromosomes (Fernández & Krapovickas Herba annua.Stipulae dorso subvillosae setulis 1994) and the electrophoreticanalysis of seed sparsimimmixtis, margine ciliatae. Folia caulis proteins(Bianchi-Hall & al. 1993) show great principalisfoliolis lanceolatis, ilia ramorumfoliolis variability that is not associated with obovatis,apiculatis, supra subtusque pilis ca. 2 mm exomorphological characters nor with longis sparsim vestitis, subtus quam supra geographicdistribution. densioribus,margine longe ciliatis setulisque nonnullisinstructis. Hypanthium 4-10 cm longum, Obs. 3. In the extreme NW of the villosum.Calyx 7-8 mm longus, setulis lutescentibus Paraguayan "chaco," on the banks of the immixtisvillosus. Vexillum aurantiacum, 15 mm Timane River, a temporary stream, at the longumX 17 mmlatum. Fructus biarticulatus paxillo localityof Mayor Pedro Lagerenza (60°45'W, basisubvilloso 4-10 cm longo, isthmo ca. 2 cmlongo, 20°5'S), grow two materials that were articulis10-12 mm longis x 6-8mm latis, pericarpio collectedby Schininiand Bordas in 1978. One laevi. resemblesĀ Cardenasii (15219 B), but is an annual, and the otherresembles A. duranensis Holotype:BRAZIL. Mato Grossodo Sul. Mun. (15101 and 15219 A), but with some Corumbá,Baia Vermelha,Fazenda Santa Teresa, differencessuch as, for example, the very 57°28'W, 18°15'S, 6-XII-1976,Krapovickas and villous peg. From seeds takenfrom no. 15101, Gregory30006 (CEN). Isotypes:CTES, G, GH, K, we obtained parents250 withyellow flowers, LIL, MO, NY, RB, SI, SP,US. aff. A. duranensis, and 251 with orange flowers, aff. A. Cardenasii. This locality is Perennial plant. Taproot without very interestingand deserves to be explored enlargements.Mainstem 10 cm long without again to resolve the taxonomie position of inflorescences,the stemcompletely covered by these two materials. the stipules; lateral branches procumbent, spreading,ca. 1 m long,with flowers along their Obs. 4. In section Arachis, A. duranensis length;internodes 30-40 mmlong with two kinds produces highlysterile hybrids with the annual of hairs: small whitish,more or less adpressed, species A. Batizocoi and A. ipaěnsis, and with otherslong, yellowish, ca. 2 mmlong, and some 2 1.2% pollen staining with A. stenosperma. yellowishbristles. Leaves tetrafoliolate.On the When crossing it with perennialspecies such mainstem,the fusedbase of the stipulesca. 12 as A. Cardenasii, A. correntina,A. Diogoi mmlong x 2 mmwide, the free tips 17 mmlong x and A. villosa, thepercentage of stainedpollen 1.5 mmwide; petiole45 mmlong; rachis 12 mm increases noticeably, reaching as much as long;the apical leaflets45-55 mm longx 13-15 71.2% in the hybridwith A . Cardenasii. mmwide and thebasal leaflets4 1 mmlong x 1 1 With representativesof other sections, it mmwide. Leaves ofthe lateral branches with the producedhighly sterile hybrids with A. gracilis fusedportion of the stipules5-6 mm long x 3.5 and A. Hermannii, of section Erectoides, and mmwide, the free portion 12-14 mm long x 2.5-3 hybrids that never flowered with the mmwide; petioles10-14 mmlong; rachis7 mm rhizomatous tetraploids A. glabrata var. long;the apical pair of leaflets26-28 mmlong x

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13-15mm wide, the basal leaflets2 1-23 mmlong Geographiedistribution. This species grows X 11-12 mm wide. Stipules puberulent,with in Mato Grosso do Sul, in Brazil, along the abundantbristles on thefused basal portionand Paraguay River, from Amolar to Porto da with some bristleson the freeportion, margin Manga. The species was also collected in Pa- longlyciliate. Petiole withtwo kinds of hairs, raguay,near the ArroyoNegla, a tributaryof similar to those of the stem: some small and the Aquidabán River, which drains into the whitish,others long and yellowish,and some Paraguay River. yellowish bristles. Leaflets of the mainstem narrowlyelliptical (length/width ratio 3:1); leaflets We dedicate this species to the botanist of the lateral branches elliptical(length/width Federico Carlos Hoehne, author of several ratio 1.8:1); upper leaf surfaceglabrous, young contributionsto the knowledge of the genus leaves usually have sparse hairs 2 mm long; Arachis, and who would collect it forthe first lower leaf surfacewith scattered, more or less timeduring his participationin the "Comisión erecthairs 1.5 mm long, and diminutivehairs, Rondon." adpressedand visible,more numerousyet still allowingthe epidermis to be seen; midveinwith 53. Arachis stenosperma Krapov. & W.C. longhairs somewhat more dense than on therest Gregory nov. sp. not with of thelower surface; margin thickened, Figs. 3,53 and 53a; 27 long scatteredcilia and withsome shortbristles. Inflorescences4-5-flowered, axis short,covered by the stipules, bracts rigid, longly ciliate. A. stenospermaM.P. Gregory& W.C. Gregory.J. Hypanthium4-10 cm long. Calyx bilabiate,the Hered.70: 192,1979, nomen nudum. "Parent no. 42" widerlobe 6-7 mm long,tridentate, with apical (Hammons& al. 4 10). incisionsof 0.5 mm,the narrowerlobe 7-8 mm long, subfalcate with silky whitishhairs and Herba annua vel semiperennis.Caule badii, numerous,somewhat longer yellowish bristles, pilis sericeis 1-2 mmlongis villosi. Folia caulis ca. 1 mmlong. Standard orange, 15 mmlong x 17 principalis foliolis oblongo-lanceolatis, ilia mm wide; wings yellow, 10 mm long x 7 mm ramorumoblongis usque ovatis, epiphyllo glabro, wide. FruitParticulate; peg 4-10 cm long,wine- hypophyllonervo medio pilis nonnullissericeis, colored and somewhatvillous towardthe base; cetero glabro, margine haud incrassato pilis isthmusca. 2 cm long;articles 10-12 mm long x sericeis setulisque nonnullis instructo. 6-8 mmwide; epicarpsmooth without prominent Hypanthium5-9 cm longumpilis sericeissparsim veins,villous. 2n=20 chromosomes(Fernández & vestitum.Calyx 6-7 mm longussetulis immixtis Krapovickas1994). villosus. Vexillumluteum, raro aurantiacum, suborbiculare,usque ad 15 mmlongum x 20 mm Additional material: BRAZIL. Mato latum.Fructus biarticulatuspaxillo 10-17 cm Grosso do Sul. Mun. Corumbá: Amolar, longo, isthmo 1.5-2.5 cm longo, articulis YIII-1908, Hoehne 19 (R 64955); Corumbá, cylindraceis1 7-22mm longis x 5-7 mmlatis apice ll-X-1985, Vails & al. 9094 (CEN, CTES); notabile,pericarpio papyraceo, laevi. id., Valls & al. 9095 (CEN); Corixo do Areiao, 15 kmW do Portoda Manga, 12-X-1985,Valls Holotype:BRAZIL. Paraná.Paranaguá, en sue- & al. 9140 (CEN, CTES); 3,7 km W do Porto los arenososdel puerto,Vila dos Portuarios[Porto da Manga, 12-X-1985,Valls & al. 9146 (CEN, Pedro II], 28-V-1968, Hammons,Langford & CTES). Krapovickas410 (CTES). Isotypes:CEN, G, LIL, PARAGUAY. Amambay. Ruta 3, arroyo MO,NY, SI,US. Negla, 35 km S de Bella Vista, 24-VIII-1980, Schinini& al. 20563 (CTES); id., 21-X-1981, Perennialplant, without rhizomes. Taproot, Schinini 21450 (CTES); id., 16-XII-1983, with thinbranches, hypocotyl ca. 5 cm long, Vanni & al. 327 (CTES); id., 345 (CTES); id., withsmall adventitiousroots. Mainstemerect, 8-IV-1986, Valls & al. 9923 (CEN, CTES); id., 5-10 cm long,with vegetative branches; basal 25-11-1994,Krapovickas & al. 45023 (CTES). internodes5-8 mm long, brownon those parts

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Fig.27. Arachis stenosperma : A, plant; B, branch(HLK.410).

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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 exposed to the light,with silky hairs 1-2 mm cylindrical,from 1 7 mmlong x 5 mmwide to 22 long; internodescovered by the stipulestoward mmlong x 7 mmwide, withpronounced beak, the apex. Secondarybranches procumbent, 80 pericarppapery, epicarp smooth,villous, with cm long, on which two vegetative branches very small hairs. Seeds 14 mm long x 5 mm alternate with two reproductivebranches in wide but up to 17 mm long x 6 mm wide, succession; internodesca. 2 cm long, brown, cylindrical, apex acute, somewhat curved, withlong silkyhairs. Leaves tetrafoliolate.On seed coat pink. 2n=20 chromosomes (M.P. the mainstem,the fusedportion of the stipules Gregory& W.C. Gregory1979). 10 mm long x 4 mmwide withca. 7 veins, the free portion 15-25 mm long x 3 mm wide, Selectedadditional material: BRAZIL. Mato acute; pulvinus2 mmlong, furrowed, with long Grosso.Mun. Barra do Garças:km 38 da BR-158, silkyhairs; petiole up to 5 cm long; rachis 12 Barrado Garças-NovaXavantina, 450 m, ll-III- mmlong; apical leafletsup to 43 mmlong x 16 1982,Allem & al. 2796 (CEN, CTES); id.,yellow mmwide, basal leafletsup to 40 mmlong x 14 flower,17-VIII-1984, Vails & al. 7762(CEN, CTES); mm wide. On the lateral branches,the fused id, orangeflower, Vails &al. 7764(CEN); 45 kmNde portion of the stipules 10 mm long x 3 mm Barra do Garças, Faz. Mocambo, 16-1-1989, wide, the free portion 12 mm long x 3 mm Krapovickas& al. 42957 (CEN, CTES); 32 kmE de wide, acute; petiole 10 mm long; rachis 4 mm Cuiabá,BR-364, 3-VI-1985, Valls & al. 9010(CEN, long; apical leafletsup to 20 mmlong x 13 mm CTES); id.,Valls & al. 9012(CEN, CTES); 6,6km W wide. Stipules withsilky hairs on the back of do rioBambá, BR-364, en murundús,3-VI-1985, the fused portion,surfaces glabrous, margin Valls & al. 9017 (CEN, CTES); 6,7 kmW do rio ciliate. Petiole canliculate, the back and the Bambáe 46 kmE do rioCoxipó, BR-364, 2 l-X-1985, marginof the canal withlong silkyhairs, canal Valls& al. 9306(CEN, CTES). Mun.Rondonópolis: itselfglabrous or with some very shorthairs. BR-364,ca. 200 mW do rioVermelho e 300 mW do Leaflets oblong-lanceolate,more or less acute acesso a Rondonópolis,22-X-1986, Valls & al. on themainstem, oblong to ovate withrounded 10309(CEN, CTES); id.,Valls & al. 10310(CEN, apex on the lateral branches; upper leaf CTES). Paraná. Mun.Paranaguá: Porto Dom Pedro surfaceglabrous, lower leaf surfaceglabrous, II, l-Xn-1911, Dusen 13472(US); Vila dosPortua- with some silky hairs on the midvein,margin rios,28-V-1968, Hatschbach & al. 19240 (MBM); not thickened,with silky hairs and scarce, Costeira, 1-1-1969,Hatschbach 20679 (CTES, intercalated bristles. Axillary spikes, 4- MBM). Mun. Antonina:Ponta da Pita,21-1-1966, flowered, axis very short, covered by the Hatschbach& al. 13605(CTES, MBM, K, NY, SI, stipules. Flowers protectedby two bracts; on US); Pontada Pita,28-V-1968, Hammons & al. 408 thefirst flower, the basal bractis entire,single- (CEN, CTES, G,MO, NY, P, US); Antonina,20-11- veined, 10 mm long x 3 mm wide, with long 1965,Saito & al. 1268 (SI); PortoAntonina, 24-V- hairs on the vein and hyalinewings, the upper 1983,Valls 7377 (CEN); Rua da acesso a praiade bract bifid, 1 1 mm long, withtwo prominent, Pontade Pita a cerca200 mda praia,24-V-1983, Valls pilose veins; theupper flowers with both bracts 7379(CEN, CTES). Rio de Janeiro.Miers (P); Riode bilabiate and 2-veined. Hypanthium5-9 cm Janeiro,Gaudichaud 864 (G, P); Rio de Janeiro, long, with scattered, silky hairs. Calyx Botafogobay, Miers 3871 (K); Rio de Janeiro, bilabiate, with long silky hairs and some Gaudichaud53 (P). São Paulo. Mun.São Sebastião: scatteredbristles; the wide lobe tridentate,6 en dunacosteira, lO-XI-1976, Gibbs & al. 3506(NY, mm long, the narrow lobe subfalcate, 7 mm UEC); Pontalda Olaria,27-V-1983, Valls & al. 7382 long. Standardsuborbicular, up to 15 mmlong (CEN, CTES); São Francisco,27-V-1983, Valls & al. x 20 mm wide, yellow, withfaint red lines on 7384(CEN, CTES). Mua Cananeia:Cananeia, 22-V- the upper surface;wings up to 1 1 mm long x 9 1986,Valls & al. 10229(CEN, CTES); Butantan,27- mmwide, yellow; keel 10 mmlong. Anthers8, XII-1920, Gehrt4744 (BM, NY, SP) (specimen dimorphic:4 large, oblong, basifixed; 4 small illustratedby Hoehne 1940, táb. 9, sub A. spherical,dorsifixed; and 1 staminodium.Fruit prostrata); Juqueriqueré,Caraguatatuba, Edwall Particulate;peg usually 10 cm longbut up to 17 (SP 1541).Mun. Peruibe: Peruibe, 21-III-1987, Veiga cm; isthmus 1.5-2.5 cm long; articles 66 (CIES, IAC); id.,29-XÜ-1987, Veiga 258 (CTES,

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IAC); Ubatuba,14-V-1987, Romero B-77 (CTES, highlysterile hybrids with A. Batizocoi and A. IAC). montícola; morethan 20% pollen stain with>4. duranensis,A. villosa, A. correntina and A. Cultivatedmaterial: BRAZIL. Mato Grossodo Cardenasir, and up to 88.5% withA. Diogoi. Sul. Campo Grande,EMBRAPA, cult.2. no. 15 Outside sectionArachis it produced a hybrid (introduc.no. 71, procedente de BR-158,45 kmN de with A. glabrata var. glabrata, in section Barrado Garças,leg. Rainer Schultze-Kraft), 30-1- Rhizomatosae, but the hybrid plants never 1979,Krapovickas 34563 (CTES). flowered.

Geographic distribution.This is the only 54. Arachis praecox Krapov., W.C. species of section Arachis that grows on the Gregory & Vails nov. sp. Atlantic where it is found in soils of coast, Fig. 3,54 almost pure sand, from Rio de Janeiro to Paranaguá in the state of Paraná (Brazil). It was also collected in the southeastpart of the Herba annua. Radix polaris, debilis. Caulis state of Mato Grosso, between the area principalis brevis, 2-3 cm longus, ramis around Barra do Garças to near Cuiabá and procumbentibus. Caulis pilis 1 mm longis Rondonópolis. The material from the two subvillosus, deinde glabrescens. Stipulae areas is very similar and represents a glabrae, margineciliis sparsis 0.5 mm longis separationof morethan 1000 kmwhich is very instructae.Folia glabra, ilia caulis principalis difficult to interpret. The topographic foliolis oblongis,ilia ramorumfoliolis obovatis, characterof the territorythat separates these epiphyllolaevi, hypophyllonervis et margine two areas presentsgreat difficultiesfor long manifestioribus,margine pilis paucis brevibus distance dispersion by the usual methods setulisquenonnullis ornato. Hypanthium 4-6 cm accepted for the genus Arachis in which longumglabrum vel subglabrum.Calyx glaber, 4 propagationby means of subterraneanfruits mmlongus. Vexillum aurantiacum, 12 mmlongum. representsa great limitation.In the case of A. Fructusbiarticulatus, paxillo 12 cm longo,isthmo stenosperma, we do not discard human ca. 3 cmlongo, articulis 13-16 mm longis x 6-7mm activityas an agentof dispersal,given its good latis,apice recurvopericarpio laevi. yieldof relativelylarge fruits. It is interestingto note that on the Atlanticcoast it was always Holotype:U.S.A. Texas. Stephenville,cult., foundin raderaihabitats, while in Mato Grosso, originallyfrom Brazil, MT, 71 kmN ofCáceres on at least one population grew in a relatively roadto Barrado Bugres,Valls & al. 6416, 10-IX- undisturbedenvironment (Vails 9017). 1985,Simpson s/n (CEN). Isotypes:CTES, K, US. Accordingto a communicationfrom J.F.M. Vails, there once existed a precolonial road Annual plant withweak taproot.Mainstem called "Peabira," that departedfrom São Vi- short, 2-3 cm long, covered by stipules. cente,near São Paulo, and unitedthe Atlantic Procumbentbranches 40-50 cm long, stem coast with the Rio Paraná and permittedthe subvillous,with hairs 1 mm long, scatteredon indigenous people to communicate with the the younger parts. Leaves tetrafoliolate, mountainregion, going along the edge of the stipules glabrous, marginwith scatteredcilia pantanal to the north (Prous 1992: 374-5; 0.5 mm long; petiole and rachis glabrous, Donlato 1985: 28-31). except fora few hairs on the pulvini and on a transverserib at the heightof the firstpair of Obs. 1. This species is characterizedby the leaflets; leafletswith both surfaces glabrous, long, narrow, cylindrical fruit articles and upper surface smooth, lower surface with seeds. Frequently, there is a small veins and margin slightly evident, margin supernumeraryleaflet presenton the leaves, glabrous or with a few shorthairs and some located betweenthe two basal leaflets. small bristles.On the mainstem:stipules with thefused portion ca. 6 mmlong, free portion 14 Obs. 2. Arachis stenosperma produces mmlong, petiole 17 mmlong, rachis 4 mmlong,

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leafletsoblong, the distal ones ca. 16 mmlong glabervel paucis pilis brevibusinstruētus, setulae X 6 mm wide, the proximal ones ca. 15 mm desunt.Vexillum 7-8 mmlongum x 10 mmlatum, long X 5 mm wide. On the lateral branches: velutino-aurantiacum.Fructus biarticulatus stipuleswith the fused portion 4 mmlong, free paxillo 3-15 cm longo, articulis 9.5-12.5 mm portion7 mmlong x 3 mmwide, petiole 12 mm longisx 5.5-6mm latis, pericarpio laevi. long, rachis 5 mm long, leaflets obovate, the distal ones ca. 14 mm long x 10 mmwide, the Holotype:BRAZIL. Tocantins.Mun. Miracema proximal ones ca. 14 mm long x 9 mm wide. do Norte,13 kmdo riodos Bois, BR-153,km 840, Hypanthium 4-6 mm long, glabrous or 9°29'S,48°36rW, 250 malt., anual, na áreainundada subglabrous. Calyx withthe wider lobe 4 mm e na aguacorrente; mata ciliar aberta, dominada por long,glabrous, the narrow lobe subfalcate,4.5 "babaçu" e Heliconia,solo orgánicoencharcado. mm long. Standard ca. 12 mm long x 13 mm As folhascon sintomasde ataque de Cercospora wide, orange. FruitParticulate, peg ca. 12 cm arachidicola.Flor aniarelo-alaranjada. 7-III-1982. long,isthmus 2.5-3 cm long,articles 13-16 mm Valls, Krapovickas,Rao & Silva 6536 (CEN). long x 6-7 mm wide, with parrot's beak, Isotypes:CTES, G, GH,LIL, MO, NY, US. pericarpsmooth. Annual plant. Root small, 1-2.5 mm in Additionalmaterial: BRAZIL. Mato Grosso. diameter (hypocotyl 2-3 mm in diameter), Mun.Barra do Bugres:7 1 kmN de Cáceres,camino shallow. Mainstem ca. 15 cm long, erect, a Barrado Bugres,31-VIII-1981, Valls & al. 6416 opposing branches of the first two nodes (CEN); id, 3 l-V-1985,Valls & al. 8965(CEN, CIES); separated2-4 cm; apical internodesvery short, 14,1 kmNW do rioCachoeirinha e 5,7km antes do covered by the stipules. Lateral branches rioSalobra na estradade Cáceresa Barrado Bugres, procumbent,up to 85 cm long. Stems glabrous 26-X-1985,Valls & al. 9397(CEN, CTES); id.,3-XI- or withscattered 1 mmlong hairs, occasionally 1986,Valls & al. 10467(CEN, CTES). more dense toward the end of the branches; internodeswith abundant adventitious roots on Cultivatedmaterial: U.S.A. Texas. Stephenville, the submerged parts. Leaves tetrafoliolate, originallyfrom Brazil, MT, Valls & al. 6416,15-Vni- those of the mainstemvery similar to those of 1985,Simpson s/n (CTES); id.,21-IX-1985, Simpson the lateral branches; leaflets elliptical, s/n(CTES). somewhatmore lanceolate on the centralaxis and occasionally somewhat obovate on the Geographie distribution.This species is lateralbranches. The fusedpart of the stipules knownonly fromthe type locality,located 71 usually 7 mm long, up to 12 mm,the freepart km to the north of Cáceres, on the road to 13 mmlong, up to 25 mm; petiole20 mmlong, Barra do Bugres, in the state of Mato Grosso up to 45 mm long and rachis 5 mm long,up to (Brazil). It grows in a fieldsubject to flooding 10 mm; apical pair of leaflets 18 mm long x withsmall rises ("murundus")covered by "ce- 10.5 mmwide, up to 42 x 17 mm,basal pair 16 rrado" vegetation, on poorly drained, mm long x 9 mm wide, up to 33 x 12 mm. compacted sandy clay soil. Stipuleswith glabrous faces or withsome hairs toward the base, margin with not too dense 55. Arachis palustris Krapov., W.C. cilia; the freepart acute 2-2.5 mm wide at the Gregory & Vails nov. sp. base. Petiole and rachis canaliculate with Fig. 3,55 some 1 mm long hairs on the back, canal and marginsof the canal glabrous. Pulvinus more or less villous (villous on veryyoung leaves). Herba annua. Stipulaeglabrae velpilis paucis Leaflets apiculate, both faces glabrous; upper basi instructae,margine subciliatae. Folíola side smoothand lowerside almost smoothwith elUptica,in caule principale latius lanceolata. somewhat marked veins; margin barely Epiphyllumglabrum. Hypophyllum glabrum, mar- noticeable, lightly ciliate and with some gine ciliato paucisetuloso. Hypanthiumtenue, scattered,weak bristles.Flowers in veryshort villosum,15-50 mm longum. Calyx 4-6 mm longus, spikes, few-flowered,all along the lateral

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branches.Bracts weak, ca. 10 mmlong, ciliate villosus.Vexillum ca. 8 mmlongum, aurantiacum. on the margin and with a few very weak Alae 5 mm longae, luteae. Fructusbiarticulatus bristlestoward the apex. Hypanthiumslender, paxillo 2-12 cm longo,glabro, isthmo ca 15 mm 15-50 mm long, villous. Calyx with some longo,articulis 10-12 mmlongis x 5-6 mmlatis, scatteredsilky hairs, withoutbristles; upper laevibus,apice recurvo. lobe 4-5 mm long, lower lobe falcate, 6 mm long. Standard 7-8 mm long x 10 mm wide, Holotype:BOLIVIA. Beni.Trinidad, en césped, orange-yellowwith some verydelicate reddish cercade la pistaprincipal del aeropuerto, 9-IV-1979, lines on the upper face. FruitParticulate; peg Krapovickas,Gregory, Simpson, Pietrarelli& 3-15 cm long, glabrous; articles 9.5-12.5 mm Schinini35005 (CTES). Isotypes:CEN, G, GH, K, long x 5.5-6 mm wide, covered by a delicate LPB,MO, NY, RB, SI, P, US. coat of very small caducous hairs; pericarp almost smooth with an open and scarcely Annual plant. Taproot weak, 3-4 mm in markedreticulation that can be seen onlywhen diameter,shallow. Mainstem erect, up to 45 the hairs fall off;beak accentuated.Seed 6-1 1 cm long, internodesup to 45 mm long, lateral mm long x 4-5 mmwide. branches procumbent,pressed closely to the soil, ca. 40 cm long; stem angular, somewhat Additionalmaterial: BRAZIL. Maranhão.Mun. villous on the young parts, hairs white,ca. 2 Carolina:12 km E de Carolina,BR-230 para Balsas, mm long, internodes15-30 mm long. Leaves 7°22'S, 47°25'W, 170 m alt., local encharcado tetrafoliolate.On the mainstemthe fused part periodicamente,13-III-1982, Valls & al. 6611 (CEN, of the stipules 8 mm long, free part 14 mm CTES, GH,K, MO, NY, P, SI, US); 10-12km E de long; petiole up to 35 mm, rachis 7 mm long; Carolina,18-111-1985, Valls & al. 8377(CEN, CTES); apical pair of leafletsup to 3 1 mmlong x 8 mm 38 kmE de Carolina,BR-230, 18-III-1985, Valls&al. wide,basal pair up to 26 mmlong x 7 mmwide. 8381 (CEN, CTES). Tocantins.Mun. Miracema do Fused portion of the stipules on the lateral Norte:14,6 km S do riodos Bois, 27-VIII-1984, Valls branches6 mmlong, free part ca. 12 mmlong; & al.7881 (CEN, CTES); id, 7883(CEN, CIES). Mun. petiole up to 25 mm long, rachis 7 mm long; Filadelfia:7025'S,43°37W, Valls&al. 13023(CEN). apical pair of leafletsup to 22 mm long x 1 1 mm wide, basal pair up to 19 mm x 8 mm. Geographic distribution.This is the wild Stipules subglabrous with some long hairs species of sectionArachis thatgrows farthest toward the base and on the back near the north.The area extendsfrom 7°22'S to 9°29'S, insertionof the petiole; the freepart linear, ca. foundon both sides of the Tocantins River. It 2 mm wide at the base, 3-4 veined, margin was collectedin totallyflooded soils, wherethe softlyciliate. Back of petiolesubglabrous, with submergedplants were in fullflower. some long hairs, upper face canaliculate, marginsof the canal pilose; pulvinusglabrous 56. Arachis benensis Krapov., W.C. or somewhatpilose; rachis narrow,margins of Gregory & C.E. Simpson nov. sp. the canal pilose. On the mainstem, leaflets on the lateral Figs. 3,56; 28,A-B lanceolate,acuminate; branches, leaflets lanceolate to oval; upper surface glabrous, smooth; lower surface glabrous or Herba annua. Caulis angulosus, novellus with some long hairs on the midvein,margin subvillosus.Stipulae subglabrae, dorso et basi pilis somewhat prominentwith a few long fragile longisnonnuliis instructae. Folia caulisprincipalis hairs and some short bristles. Flowers along foliolislanceolatis, acuminatis, illa ramorumfoliolis the lateral branches, in very short, few- lanceolatisusque ovalibus,omnia epiphyllo laevi, floweredspikes. Hypanthium20-50 mm long, glabro,hypophyllo glabro vel pilis longisnonnuliis villous. Calyx 4-5 mm long, bilabiate, villous innervo medio, in margine submanifesto paucis pilis and with some bristles. Standard ca. 8 mm longis caducis setulisque brevibus nonnuliis long, orange; wings 5 mm long, yellow. Fruit instructo.Hypanthium 20-50 mm longum, villosum. subterranean,biarticulate; peg 2-12 cm long, Calyx 4-5 mm longus,setulis nonnuliis immixtis aerial part glabrous; isthmusca. 15 mm long;

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Fig.28. Arachis benensis : A, portion of branch (K.35005); B, leaffrom mainstem (K.35006). A. helodes : C, leaffrom mainstem;D, leaffrom lateral branch (G.9926). A. correntina: E, leaffrom lateral banch; F, leaffrom mainstem (K.7830). articles10-12 mmlong x 5-6 mmwide, smooth, Station,250 m,9-XI-1965, Booke 48 (K); Trinidad, with strongbeak; the peg is insertedon the en matorralesvecinos al aeropuerto,14-IV-1979, back at the basal end. Seed 7-8 mm long x 4-5 Krapovickas& al. 35006(CEN, CTES, G, GH,LPB, mm wide, with the distal end pointed and MO, NY, US); Guayaramerín,aeropuerto, 18-IV- somewhat curved. 2n=20 chromosomes 1979;Krapovickas & al. 35007(CEN, CTES, GH,K, (Fernández & Krapovickas 1994). LPB, MO, NY, US); Guayaramerín,aeropuerto, 21- XI-1988,Williams 860 (CTES); id.,861 (CTES). Additionalmaterial: BOLIVIA. Beni. Trinidad, savanna,1-1965, Braun 7 & 8 (US); Trinidad,S.A.I. Geographie distribution.This species was

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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) collected in Trinidad and in Guayaramerin, mainstem,stipules with the fused portionca. localities on the Mamoré River in the 10 mm long and the free portion ca. 16 mm departmentof Beni and in northernBolivia. It long x 2-3 mmwide, petiole 25-40 mm,rachis grows in savannahs among grasses or in the 8-12 mm long, leafletsoblong-lanceolate, the borderof shrubbybrush in clayeysoil. distal ones 30-35 mm long x 11-18 mm wide, theproximal leaflets 26-29 mmlong x 9-14 mm 57. Arachis trinitensisKrapov. & W.C. wide. On thelateral branches, stipules with the Gregory nov. sp. fused portion ca. 5 mm long and the free Fig. 3,57 portion10-11 mmlong x 3-4 mmwide, petiole 9-10 mm long, rachis 6-7 mm long, leaflets oblong to obovate, the distal ones ca. 17 mm Herba annua. Radix polaris. Caulis apicem long x 10-14 mm wide, the proximal ones ca. versuspilis 1.5-2 mm longis villosus. Stipulae 15 mmlong x 8-1 1 mmwide. Hypanthium20- setulis destitutae,glabrae vel pilis parvissimis 50 mm long, villous. Calyx 5 mm long, lobe adpressissparsim vestitae, dorso villosae. Caulis subfalcate,6 mm long, villous, withabundant principaliserectus, foliolis oblongo-lanceolatis, bristles. Standard ca. 10 mm long, orange, rami procumbentesfoliolis oblongis usque wings yellow. Fruitsubterranean, Particulate; obovatis. Epiphyllum laeve, glabrum. peg 5-10 cm long, glabrous, articles 9-11 mm Hypophyllumfere laeve, infoliis junioribus pilis long x 5-6 mmwide, pericarpsmooth. parvis adpressis et nonnullisca. 2 mm longis sparsimvestitum in foliis vetustioribus fere solum Additionalmaterial: BOLIVIA. Beni. Trinidad, pili longi nervo medio adsunt, marginepilos UniversidadTécnica del Beni, 20 mNE del edificio paucos adpressos ca. 1 mm longos gerentes. laboratorios.Fruto liso, 21 -III- 1990, Williams 1117 Hypanthium20-50 mm longum, villosum. Calyx 5- (CIES). 6 mmlongus, villosus et largesetulosus. Vexillum 10 mm longum, aurantiacum. Fructus Cultivatedmaterial: ARGENTINA. Córdoba. subterraneus,biarticulatus paxillo 5-10 cm Manfredi,E. E. A. (INTA), cult. no. 42, origin: longo,glabro, articulis 9-11 mmlongis x 5-6 mm Williams 1117, 21-III-1991,Krapovickas 43795 latis,pericarpio laevi. (CTES); id. 27-III-1992, Krapovickas 44091 (CIES). Holotype:BOLIVIA. Beni.Prov. Cercado, Trini- dad, Campusof the Universidad Técnica del Beni, Geographic distribution. This species is 2.5 kmN ofTrinidad. Procumbent. Mainstem erect, knownonly from the type locality, in Trinidad, to40 cmhigh. Orange standard, yellow wings. Pegs capital of theBeni department(Bolivia), at 236 short,white. Dark greysandy clay, 25-XI-1988, m elevation above sea level. Williams866 (CTES). 58. Arachis decora Krapov., W.C. Annual plantwith taproot. Mainstem 10-40 Gregory & Vails nov. sp. cm covered branches long, erect, by stipules, Fig. 3,58 procumbent.Stem villous on theyounger parts, hairs 1.5-2 mm long. Leaves tetrafoliolate, Herba annua.Radix palaris. Caulis violaceus, with surfaces or with stipules glabrous very apicem versus villosus, deorsum versus small sparse adpressed hairs, back villous, glabrescens.Stipulae dorso et basi villosae,mar- without and margin ciliate, bristles; petiole gine ciliatae. Setulae in stipulis,petiolis et in rachis hairs 2 mm surface villous, long, upper rachidiadsunt. Caulis principalis erectus, foliolis and lower surface glabrous smooth, nearly ovato-lanceolatis.Rami procumbentesfoliolis leaves with small smooth,younger dispersed obovatis usque oblongis. Epiphyllum laeve, hairs and a few hairs ca. 2 mm adpressed long glabrum. Hypophyllumnervis et margine vix older leaves with long, commonly only long conspicuis, glabrum vel pilis parvissimis, hairs on the with midvein; margin few adpressis,nervo medio paucis pilis majoribus, hairs ca. 1 mm On the adpressed long. marginepilis parvissetulisque nonnulis vestitum.

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Hypanthium3 cm longum,partim villosum. Calyx Additionalmaterial: BRAZIL. Goiás. Nova 4 mmlongus, labium majore subglabro,labium Roma, entradada Fazenda Ouro na estradade minoresubfalcato villoso et setuloso. Vexillum Nova Roma a Iaciara, Skorupa& al. 644 (CEN); aurantiacum.Fructus subterraneus paxillo gla- CamposBelos, 20 kmao NE de CamposBelos na bro,articulis ca. 14 mmlongis x 7 mmlatis, apice estradapara Aurora do Norte,Allem 3485 (CEN); recurvo,pericarpio laevi. id., Valls & al. 9953 (CEN). Mun. Teresina de Goiás: 13°26'S,47°08'W, Valls & al. 12900(CEN). Holotype:BRAZIL. Goiás.Mun. Campos Belos, Mun. MonteAlegre de Goiás: 13°18'S, 46°42'W, 20 kma nordestede CamposBelos na estradapara Valls & al. 13290 (CEN); id., 13°14'S, 46°44'W, Aurorado Norte, 13°0rS, 46°42'W, 690 m s.m, plan- Valls & al. 13348 (CEN). Mun. Simolandia: tas vigorosas crescendo em local perturbado 14°28'S, 46°29'W, Valls & al. 13371 (CEN). recentementeem beira de barrancoao longo da Tocantins.Mun. Combinado:12°55'S, 46°35'W, estrada,vegetaçao baixa secundaríaem antigaarea Valls& al. 12893(CEN). de mata,12-IV-86, Valls, Simpson & Werneck9955 (CEN). Isotype:CTES. Geographie distribution.This species is known only from places close to the type Annual plant,with weak taproot.Mainstem locality, in northeasternGoiás and southern erect, ca. 15 cm long, procumbentbranches Tocantins(Brazil). ca. 50 cm long. Stems purple,villous toward the apices, glabrescent, bristles present. 59. Arachis Herzogii Krapov., W.C. Leaves with4 leaflets,stipules villous toward Gregory & C.E. Simpson nov. sp. the base and on the dorsal numerous surface, Fig. 3,59 bristlestoward the base, ciliate margin,short hairs. Petiole and rachis partiallyvillous and with abundant bristleson the dorsal surface. Herbaperennis. Radix polaris. Rami prostrati. Leaf upper surface glabrous, smooth, lower Caules novelli villosi, deinde glabri. Stipulae surface with veins and margin subdued, subglabrae, basi villosae. Caulis principalis glabrous or with very small adpressed hairs foliolis lanceolatis, rami foliolis oblongis and a few largerhairs on the midvein,margin obovatisve.Epiphyllum glabrum, laeve, margine withsmall sparse hairs and withshort bristles. sat incrassato,hypophyllum villosum, pilis ca. 2 On the mainstem: stipules with the fused mm longis sparsim immixtis,margine sat portionca. 10 mmlong, free portion ca. 20 mm incrassatoet nervisperspicuis. Hypanthium 40-45 long x 1.5 mm wide, petiole ca. 40 mm long, mm longum,villosum. Calyx 5-6 mm longus, rachis ca. 8 mm long, leaflets ovate- villosus, setulosus. Vexillum11 mm longum, lanceolate, the distal ones ca. 40 mm long x aurantiacum. Fructus subterraneus, 14 mm wide, the proximal ones ca. 32 mm biarticulatus,articulis 8-10 mmlongis x 5-5.5 mm long x 12 mm wide. On the lateral branches: latis,pericarpio laevi. the fused portion of the stipules ca. 5 mm long, the freeportion ca. 12 mm long x 2 mm Holotype:BOLIVIA. SantaCruz. Prov. Chiqui- wide, petiole ca. 18 mm long, rachis ca. 5 tos,18 km N de San José60°47rW, 17°47'S, camino a mm long, distal leaflets obovate, ca. 21 mm San Ignacio, ca. 300 m s.m., perenne,ramas long x 13 mm wide, proximal leafletsoblong, prostradas, corola anaranjada, 28-IV-1980, ca. 17 mm long x 10 mm wide. Hypanthium Krapovickas& al. 36030 (CTES). Isotypes:CEN, 3 cm long, partiallyvillous. Calyx 4 mmlong, GH,K, LIL, LPB,MO, NY, US. larger lobe subglabrous, witha few hairs and an occasional bristle at the apex, smaller Perennial plant, with taproot. Branches lobe subfalcate, villous and with various prostrate,young shoots quadrangular,villous, bristles. Standard orange. Fruit withhairs 2 mmlong, cylindrical and glabrous subterranean, peg glabrous, articles ca. 14 towards the base; internodes2-3 cm long. mm long x 7 mm wide, with parrot's beak, Stipules with the fused portion5-7 mm long smooth. and the freeportion 14-17 mm long x 4-5 mm

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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) wide, dorsal surface villous, surfaces villous obovata, epiphyllo laevi, glabro, hypophyllo toward the base, the rest subglabrous,with a adpresso-piloso,nervo medio sursum pilis longis, few long hairs especially on the veins and marginepilis parvis adpressis,eiliis 1 mmlongis almost to the apex. Leaves tetrafoliolate, etiamsetulis brevibus munito. Hypanthium ca. 7 petiole 3-10 mmlong, somewhatlonger on the cm longum,villosum. Vexillum 10 mm longum, leaves of the mainstem;rachis 3-7 mm long, aurantiacum.Fructus subterraneus, biarticulatus both with villous dorsal surface, a glabrous articulisparvis usque ad 9 mmlongis x 4 mmlatis, petiole-rachiscanal divided by a transverse pericarpiovalde reticulato. line of hairs at the firstpair of leaflets;distal leaflets 18-33 mm long x 8-18 mm wide, Holotype:BRAZIL. Mato Grossodo Sul. Mun. proximal leaflets 15-26 mm long x 7-13 mm Bela Vista,junto a alfandegade Bela Vista em wide, oblongor obovate on thelateral branches gramadosornamentais, area urbanizadapróxima a • and lanceolate on the mainstem,upper side margemdo rioApa, 22-IV-1984,Valls & al. 7681 glabrous and smooth with the margin (CEN). Isotype:CTES. somewhatthickened, the underside villous with hairs ca. 2 mm long, scattered,allowing the Perennial plant, ramose; with taproot. epidermal surface to be seen, margin Branches prostrate,internodes 1-4 cm long, somewhat thickened and marked veins. shoot somewhatvillous on the youngerparts. Hypanthium40-45 mmlong, villous. Calyx 5-6 Leaves with four leaflets. Stipules with the mmlong, villous and withbristles. Standard 1 1 fusedportion 4-5 mmlong and the freeportion mm long, orange. Fruit subterranean, 10-12 mm long, dorsal surface villous with Particulate,articles 8-10 mmlong x 5-5.5 mm adpressed hairs and long bristles.Petiole 8-15 wide, smooth. mmlong, rachis 4-5 mmlong, villous with some bristles, upper surface with small adpressed Additionalmaterial: BOLIVIA. Santa Cruz. hairs; leafletsfrom oblong to obovate, slightly Prov.Chiquitos, 6.5 kmN de San José,60°47'W, apiculate, upper surfaceglabrous and smooth, 17°47'S,camino a SanIgnacio, ca. 300m s.m., 28-IV- lower surface with adpressed hairs and long 1980,Krapovickas & al. 36029(CIES, GH,LPB, MO, hairson themidvein, margin with small adpressed NY, SI, SP,US). hairs,cilia 1 mmlong and shortbristles; distal pair 12-18mm long x 6-10 mmwide, proximal pair 12- Geographicdistribution. This species grows 15 mmlong x 5-8 mmwide. Hypanthium ca. 7 cm at 17°47'S, near San José de Chiquitos,in the long,villous. Calyx 6-7 mmlong, villous and with southernpart of the departmentof Santa Cruz bristles.Standard orange, ca. 10 mm long. Fruit (Bolivia), in patches of light-coloredsand and subterranean,Particulate, peg glabrous,articles gravel, in a not verydense forest. up to 9 mmlong x 4 mmwide, pericarp with very We dedicate this species to Theodor markedreticulation. Herzog, botanical explorer and author of various works on the vegetation of eastern Additionalmaterial: BRAZIL. Mato Grosso do Bolivia. Sul. Bela Vista,entrada de Bela Vistadesde Jardim, 24-IV-1985,Valls & al. 8711(CEN, CTES). 60. Arachis microsperma Krapov., W.C. Gregory & Vails nov. sp. Geographic distribution. This species is known fromthe Bela Fig. 3,60 only typelocality, Vista, on theApa River in the SW part of the stateof Mato Grosso do Sul (Brazil). Herba perennis.Radix polaris, valida. Rami procumbentes.Caulis internodiis1-4 cm longis, 61. Arachis villosa Benth. villosiusculis.Folia junioribus parva. Stipulae Fig. 3,61 adpresso-pilosae, dorso villosiusculae, basi setulaslongas gerentes. Petiolus rachisque villosi setulisquenonnullis instruēti. Folíola oblongaad Bentham,Trans. Linn. Soc. London 18: 159,1841,

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"Ad Rio Grande Brasiliae Meridionalis. thickened,evident on both surfaces in dried Tweedie." Benth., FI. bras. 15(1): 87, 1859. specimensbut principally on thelower surface; Arechavaleta, Anales Mus. Nac. Montevideo the upper surface smooth,pubescent to more 3(20): 355, 1901. Chevalier,Rev. Int.Bot. Appl. or less villous, the lower surface usually Agrie.Trop. 13(146-147): 760, 1933.Parodi, Revista denselyvillous, margin ciliate and occasionally Argent.Agron. 1(2): 136,1934. Burkart, Darwiniana with some short bristles. Inflorescences 3(2): 267,fig. 19, g, i,j, 1939.Hoehne, Flora Brasílica axillarywith short axis, coveredby thebase of 25(2) Part.122: 14,táb. 8, 1940.Burkart, Proc. 8th the stipules, 1-3-flowered.Hypanthium ca. 5 Amer.Sc. Congr.3: 175,fig. 1, a-h, 1942.Hartley, (3-6.4) cm long, pilose. Calyx bilabiate, with Div. Pl. Ind.,Melbourne, Div. Rep. 7: 34,fig. 2, 1949. longsilky hairs and longbristles, the wider lobe Hermann,Agrie. Monogr. USDA 19:11,fig. 9, 1954. 6-6.5 mm long,the narrowlobe falcate,7 mm A. prostrataBenth. var. villosa (Benth.) A. long. Standard 13-14 mm long x 13 mm wide, Chev.,Rev. Int. Bot. Appl. Agrie. Trop. 9(90): 100, yellow at the base and orange toward the 1929. margin; wings 8 mm long, usually yellow, Stylosanthes leiocarpa auct. non Benth., occasionally withthe upper half orange. Pegs Grisebach,Symbol, fl. Argent. 106, 1879.Lorentz, occurringalong the lengthof the branches,up Veg.Nordeste Entre Rios, 38, 62, 133,171, 1878. to 20 cm long, reddish,with a few very small scatteredhairs; fruit Particulate, articles 10-15 Perennialplant. Taproot thick,reaching 1.5 mm long x 7-8 mm in diameter,the terminal cm in diameter,lateral roots slender,without article always somewhat larger; pericarp enlargements.Mainstem erect, 10-35 cm tall, prominentlyreticulate, beak manifest. Seed little-branchedtoward the base in year-old oval-elliptical, 10-11 mm long x 6 mm wide, plants,with inflorescences on theupper nodes; testa brown, very thin. 2n=20 chromosomes internodesangular, villous, 10-20 mmlong. La- (Krapovickas & Rigoni 1951). teral branchesprocumbent, spreading, up to 1 m long,non-rooting, little-branched; internodes Holotype:Brazil. Rio Grande, Tweedie, 1837 (K!). angular, villous, up to 40 mm long. Leaves tetrafoliolate. On the mainstem the fused Selectedadditional material: ARGENTINA. Bue- portionof the stipulesup to 9 mmlong x 6 mm nos Aires. Facultadde Agronomíay Veterinaria, wide, the freetips acute, up to 20 mmlong x 2 JardínBotánico, adventicia, 19-11-1934, Parodi mm wide at the base; petiole 20-35 mm long; 11885 (BAA); id.,15-VI-1937, Clos 6135 (BAB); id., rachis 10 mm long; apical leafletsusually 23 4-VI-1937,Burkart (SI 391); id.,12-11-1938, Burkart mm long x 1 1 mm wide; basal leaflets2 1 mm 9050(SI); id.,27-XI-1936, Archer 4569 (K, US): Insti- long x 9 mm wide. On leaves of the lateral tutoDarwinion, cult. 18-XII-1942,Bartlett 19244 branches,the fused portion of thestipules from (NY, US). Corrientes.Dep. Paso de los Libres: 3-5 mm long x 5 mm wide, the freepart 7-15 Paso Troncón,16-XII-1946, Huidobro 3712 (CTES, mm long x 2.5-3 mm wide at the base, acute; LIL); Paso de los Libres,12-1-1945, Ibarrola 2075 petiole ordinarily10 mm long (from 3 to 18 (LIL, NY); Paso de losLibres, Hotel de Turismo,27- mm), rachis 3-7 mm long; the apical pair of ffl-1964,Krapovickas & al. 11292 (CTES, MO); id., leafletsusually 15 mmlong x 9 mmwide (up to 19-V-1968,Hammons & al. 357 (CTES); Pasode los 18 mmx 1 1 mm); thebasal pair of leaflets13.5 Libres,Laguna Mansa, 18-11-1972,Krapovickas & mmlong x 8.5 mmwide (up to 16 mmx 9 mm). al. 21697 (CTES, G, NY); Bonpland,21-11-1945, Stipules with surfaces almost glabrous, more Ibarrola2148 (LIL, NY); Estancia "El Recreo", or less villous on theback of thefused portion, Bonpland,8-XI-1973, Lourteig & al. 2760(CTES, P). withoutbristles, margin with long, more or less Dep. Monte Caseros: Juan Pujol, 10-11-1945, adpressed cilia. Petiole and rachis somewhat Ibarrola2391 (LIL, NY); Mocoretá,13-11-1945, canaliculate, villous on the back. Pulvinus Ibarrola2406 (LIL, NY); El Ceibo, frentea isla densely villous. Leaflets of the mainstem Correntina,22-1-1942, Rosengurtt B-3676 (CTES, ovate, more or less acute at the apex, those of MVFA, SI); MonteCaseros, Cambai, 10-11-1957, thelateral branches tend to be morerounded to Nicora5699 (SI); ríoMiriñay y rioUruguay, 28-III- obovate,the apex usually mucronulate,margin 1964,Krapovickas & al. 113 1 1 (CEN, CTES, LIL,

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MO); CampoGral. Avalos, ayo. Curupí, 21-11-1979, NuevaPalmira, 8-III-1964, Arrillaga 1973 (MVFA). Schinini,& al. 17524(CTES, G, MO). EntreRíos. San José.Arazatí, 24-XII-1939, Legrand 1696 (SI); Federación,Mandisobí, 17-11-1876, Lorentz 743 id., 17-III-1957,Arrillaga 644 (MVFA); id.,24-IV- (BAF);Federación, 20-XII-1946, Meyer 1 1092 (LIL); 1960,Arrillaga 898 (MVFA); Boca del Cufré,10-1- SantaAna, 24-IX-1961, Burkart 22585 (SI). Dep. Co- 1992,Baycé & al. 20780(CTES, MVFA); desembo- lón:Colón, III- 1927, leg.? (BAB 51612); Colón,28- cadura del ayo. Cufré,28-11-1967, Lema 6505 XI-1946,Meyer 10631 (LIL). Dep. Uruguay:estan- (MVFA); Barradel arroyoSan Gregorio,2-1-1969, cia La Seimira,30-III-1967, Pederson 8198 (K). Lema& al. 8234(CTES, MVFA); Rincón del Pino, 3- URUGUAY.Banda Orientaldel Uruguay,Saint XI-1971,Reu 10868 (MVFA). Montevideo. Montevi- Hilaire2369 (K, P); id.,363 & 366 (K); Sauce Pep. deo,Sello d.883 (K); Montevideo,III-1900, Berro s/ Colonia?],11-1900, Arechaveleta 5368a (MVM). n (CORD); entreLa Coloraday PuntaEspinillo, 4- Artigas.Bella Unión,XII- 1928, Herter 84489 (G, ffl-1983,Izaguirre & al. 17115 (MVFA). MVM, SI); id.,28-1-1948, Castellanos 15980 (LIL); San Gregorio,3 1-III-1962, del Puerto 2 103(MVFA); Geographiedistribution. To date,the species ayo.Guaviyú (Paredón), 19-XII-1977, del Puerto & is known only fromArgentina and Uruguay, al. 14787(MVFA); id.,13-III-1978, del Puerto & al. where it grows in the dunes and sandy soils 15170 (MVFA); ayo. Sauce, 27-1-1964,Marchesi along the Rio Uruguay and on the Uruguayan 1096(MVFA). Salto. Belén, 12-XII-1905, Berro 5886 coast of the Río de la Plata to the edge of the (MVFA); ríoUruguay y ayo. San AntonioGrande, cityof Montevideo.In thecity of Buenos Aires 27-1-1937,Rosengurtt B-941 (CTES, MVFA). (Parodi 1934), it was foundin sand transported Paysandú. Chapicuy,22-11-1941, Rosengurtt B- fromthe Uruguayan coast of theRío de la Plata. 3280 (MVFA); Chapicuy,Santa Sofía, 15-XI-1942, It has beenmentioned as existingin Rio Grande, RosengurttB-3280'/2 (CTES, MVFA). Río Negro. butit has notbeen possible to confirmthis. Ayo.Caracoles, 22-XI-1913, Berro 7092 (MVFA); rio Uruguayy ayo. Negro, Chebataroff(CTES Obs. 1. A problem persists regardingthe 16279);San Javier,2-V-1969, del Puerto & al. 8388 type locality of A. villosa. On the holotypeis (MVFA); ríoUruguay y ayo.Guaviyú, Saladero, 17- read: "Rio Grande, Tweedie, 1837." In his I-1967,Rosengurtt & al. 10587(MVFA); Nuevo Ber- originaldiagnosis, Bentham ( 184 1: 159) writes: 17-111-1989,Davies & al. 19538 "Ad Rio Grande, Brasiliae Meridionalis, lín, (CTES, MVFA); " id., 5-VI-1992,Izaguirre & al. 20991 & 20995 Tweedie. Later, Benthamhimself (1859: 87) Soriano.Arenal Grande, 29-XI-1932, Ca- adds: in arena ad oras Rio (MVFA). "copioso " flum. brera2585 (SI, NY). Colonia.Puerto del Sauce,23- Grande et Uruguay: Tweedie, indicatingits II-1900, Berro 1117 (CTES, MVFA, SI); Nueva existencein two places: on the Uruguay River Helvecia,11-11-1916, Berro (MVFA); Artilleros,17- and in Rio Grande. XI-1949, Buikart 18159 (CIES, SI); Artilleros,26-IV- It is veryprobable thatTweedie' s collection 1968,Hammons& al. 8, 10y 14(CTES); Conchillas, was made only on the banks of the Rio Uru- 19-XII-1921,Castellanos (BA); Colonia, 1-1917, guay, on the westernlimit of the state of Rio Hauman(BA); Colonia,Clos 3225 y 6292 (BAB, Grande do Sul. According to Castellanos CTES); PlayaFomento, 3 l-XII-1968,Rosengurtt B- (1945: 6), Tweedie embarked from Buenos 11001 (MVFA); id., 10-1-1962,del Puerto1039 Aires in 1832 in search of plants,ascended the (MVFA); id., 4-V1-1992,Izaguirre & al. 20993 Rio Uruguay and continuedon to Rio Grande (MVFA);Balneario Fomento, 10-1-1992, Baycé & al. do Sul, Santa Catarina and Rio de Janeiro, 20763(CTES, MVFA);Playa Real de S. Carlos,7 km returningfrom there to his point of departure. N dePuerto Colonia, 17-1-1937, Archer 4952 (K, NY, Three years later he attempted a second US); Ayo.Pintos, Artilleros, near Puerto Platero, expeditionto Rio Grande, but upon leaving 17-XII-1943,Bartlett 21201 (NY, US); coastof the Montevideothe boat on whichhe traveledwas Uruguay,between arroyo Del Vancis & Colonia, shipwrecked. Tweedie 350 (K); Punta Gorda, 19-1-1962, All of the material studied by us was Rosengurtt& al. 8641(MVFA); Indaré,Rosario, 23- collected in sandy places on the banks of the III-1963,del Puerto& al. 2464 (MVFA); Carmelo, Rio Uruguay,from Paso de los Libres to the PlayaSere, Izaguirre & al. 19665(CTES, MVFA); river'smouth, and along the Uruguayancoast

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of the Río de la Plata, which supports the portionof the stipulesup to 9 mm long and the supposition that the holotype had been freeparts up to 15 mm long; petiole up to 43 collectedat some pointalong theRio Uruguay. mmlong; rachisup to 13 mmlong; leafletsoval Accordingto informationsupplied by Vails, to lanceolate,the apical pairup to 34 mmlong x in 183 1 the southernboundary of theBrazilian 13 mmwide, thebasal pair up to 30 mmlong x state of Rio Grande do Sul used to be the 10 mm wide. On the lateral branches, the Arapei River, thereby including within the fusedportion of the stipulesup to 5 mm long, municipalityof Alegretethe entiredepartment thefree portion up to 13 mmlong; petioleup to of Artigasand part of the departmentof Salto, 10 mm long; rachis up to 6 mm long; leaflets both of which currentlypertain to Uruguay obovateto suborbicular,the apical pair usually (Trindade 1985: 148 map and 176). 1 1 mmlong x 8 mmwide (up to 17 x 15 mm), Therefore,it is possible that Tweedie had thebasal pair 10 mmlong x 7 mmwide (up to collected the typeof A. villosa on the banks of 16 x 12 mm). Stipules glabrous, free ends the Uruguay River, betweenthe mouthof the subfalcate,with the marginsomewhat ciliate; Arapey River (departmentof Salto) and Bella petiole and rachis canaliculate, glabrescent, Union (departmentof Artigas), in present-day with villous back when young; leaflets with Uruguay. both surfaces glabrous, upper side smooth, lower side withmidvein and marginsomewhat Obs. 2. In the Museum of Paris (P) thereis marked,margin sometimes with the occasional a specimenlabeled "Brasil: Rio Grande, 1833, shortbristle. Flowers along the lengthof the Guadichaud 883," a specimen actually lateral branches in short4-5 -flowered spikes. collected by Sello in Uruguay. One proof of Hypanthium30-75 mm long, villous. Calyx thisis the existenceof the specimen"Montevi- bilabiate, villous and with scatteredbristles, deo, Sello d.883" (K). upper lobe broad, 4-toothed,5-6.5 mm long; lower lobe thin,falcate, ca. 7 mm long. Stan- Obs. 3. Arachis villosa crossed only with dard up to 15 mm long x 16 mm wide, other species of section Arachis. With A. commonlyorange with a yellow center,but Batizocoi it produced highly sterile hybrids, can also be completelyyellow; wings up to 10 with2.9% stainedpollen. Hybrids of more than mm long, yellow. Fruit subterranean, 20% stained pollen were obtained with A. Particulate;peg glabrous,5-8 cm long; articles stenosperma,A. Diogoi, A. duranensis and 8-12 mm long x 5-7 mm wide, with marked A. montícola, and with 70 to 73% stained beak, pericarp fragile, somewhat reticulate. pollen withA. Cardenasii and A. correntina. 2n=20 chromosomes(Smartt 1964, Smartt& Arachis villosa also produces hybridswith A. Gregory1967, GKP 9926). hypogaea var. fastigiata. Holotype: BRAZIL. Mato Grosso. In 62. Arachis helodes Martius ex Krapov. palustribusad Cuyabá, 1839, leg. S. Manso, & Rigoni MartiiHerbar. Florae Brasil 588 (G!). Isotypes: Figs. 3,62; 28,C-D NY!,P!. Selectedadditional material: BRAZIL. Mato Krapovickas& Rigoni,Darwiniana 11(3): 451- Grosso.Cuiabá, 8-III-1946, Burkart 15715 (CTES, 452,lám. 2, 1957. SI); id., 15-III-1948, Stephens262 (NA); id., Stephens& al. 256 (LIL, NA); id., 13-III-1959, Perennial plant. Taproot deep, without Gregory& al. 9926(CIES, G, GH,LEL, MO, NY, SI, enlargements.Mainstem erect,ca. 5 cm long, SP, US). Mun.Poconé: Cotia, rio Bento Gomes, 60 withoutflowers; lateral branches procumbent, kmSW de Cuiabá,camino a Poconé,16-XII-1976, ca. 50 cm long. Stem glabrescent,somewhat Krapovickas& al. 30029(CEN, CTES, GH,MO, NY, villous and angular in the young parts; US). Mun.Cáceres: fazenda Sangradouro, 57°15rW, internodes 10-30 mm long. Leaves 16°S,16-XII-1976, Krapovickas & al. 30031 (CEN, tetrafoliolate. On the mainstem, the fused CTES, GH, LIL, MO, NY, SP, US); 101 km de

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Cáceres, camino a Cuiabá, 17-XII-1976, covered with soil towardthe base of the plant Krapovickas& al. 30036(CEN, CTES, GH,LIL, MO, and weak adventitious roots are frequently NY, RB, US); SantoAntonio de Leverger,20-VIII- found on the buried parts. Stems villous, 1981,Vails & al. 6325 (CEN, CTES); id.,Valls & al. occasionallywith some bristles;internodes 15- 6326(CEN, CTES); id.,24-1-1989, Valls & al. 12083 35 mm long, angular when young, later (CEN,CTES); BR-070,km 660, 1,2km W do ribeirão cylindrical. Leaves tetrafoliolate. On the Flexas,30-V-1985, Valls & al. 8934(CEN, CTES); 13 mainstem,the fused porton of the stipules 6 kmS ofPoconé, 2 l-Vm-1981, Valls & al. 6331 (CEN, mmlong, the free portion up to 20 mmlong x 3 CTES). Mun. N. Sra. do Livramento:4-XI-1986, mmwide, triangular; petiole up to 45 mmlong; Valls& al. 10470(CEN, CTES); id.,Valls & al. 10471 rachis 12 mm long; leaflets elliptical- (CEN, CTES); id.,Valls & al. 10476(CEN, CTES); lanceolate,the apical pair up to 39 mm long x 25,3 kmda BR-070camino a Poconé,22-X-1985, 15 mm wide, the basal pair up to 36 mm x 14 Valls& al. 9318 (CEN, CTES). mm. On the lateralbranches, the fusedportion of thestipules 4-7 mmlong, the freeportion 7- Geographie distribution.The range of this 10 mmlong x 2-3 mmwide, triangular;petiole species is ratherlimited in thatit extendsonly usually 5-10 mmlong, up to 20 mm; rachis 3-7 some 100 km to the west and some 60 km to mm long; leaflets elliptical to obovate, the the south of Cuiabá, Mato Grosso (Brazil). It apical pair commonly 12 mm long x 10 mm grows in open places subject to flooding. wide, up to 23 x 1 1 mm,the basal pair some 1 1 mm long x 9 mm wide, up to 20 x 9 mm. Obs. Arachis helodes is very uniformwith Stipules somewhat villous toward the base respectto the shape of the leaflets,which are withglabrous surfacesand the marginciliate; always glabrous. Nevertheless, it shows a on the surfaces of the fused portionthere are certain amount of polymorphism. The frequentlylong rigid bristles present. Petiole population Krapovickas & al. 30030 had and rachis canaliculate,back villous withlong individualswith orange flowers and individuals silky hairs and short,more or less adpressed with yellow flowers. In the population hairs, with bristles frequentlypresent; canal Krapovickas & al. 30029, most of the plants with short scattered hairs. Pulvinus densely had orange standards, but about 1% had villous. Upper surface of the leaflets smooth, yellow flowersand 1% had whiteflowers. sometimesglabrous, or withvariable hairiness, from short scattered hairs to more or less 63. Arachis correntina (Burkart) Krapov. dense short hairs; lower surface with & W.C. Gregory nov. comb. prominentmidvein and margins,the surface Figs. 3,63; 28,E-F withshort adpressed hairs or withmore or less raisedhairs, on themidvein, more or less dense hairs 2 mm long, whichmay be presenton the A. villosa Benth. var. correntinaBurkart, rest of the surface as well; marginwith two Darwiniana3(2): 269, 1939.Krapovickas & Rigoni, kindsof trichomes:short, whitish, more or less Rev. Invest.Agrie. 5(3): 289-293 (2n=20), 1951. adpressed hairs, long (1-2 mm) yellowish Conagin,Bragantia21(21): 357, est. 10, figs. 7B, 17, curvedhairs, or some perpendicular,and short 1962. Conagin,Bragantia 18(5): 51-70,figs. 1-5, scatteredbristles. Flowers along the lengthof 1959. the lateral branches in short, 3-4-flowered A. correntina (Burkart) Krapov. & W.C. spikes. Hypanthium6-11 cm long, villous. Gregory, in W.C. Gregory, M.P. Gregory, Calyx villous and withthin scattered bristles, Krapovickas,Smith & Yarbrough,in Peanuts- lowerlobe falcate,6-8 mmlong, upper lobe 5-7 Cultureand Uses: 97, 1973,nomen nudum. mm long; standardorange, 18-23 mm wide x 11-16 mm long, wings yellow or yellow with Perennialplant. Taproot deep, 5-15 mm in theupper half orange. Fruit Particulate; peg 5- diameter.Mainstem erect 10-25 cm long, late- 10 cm long, somewhatvillous near the base, ral branches prostrate,up to 1 m long. In laterglabrous; isthmus3-4 cm long; articles8- mature plants, the branches are usually 13 mm long x 5-7 mm wide, smooth,beaked,

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covered with dense diminutivehairs. 2n=20 Ituzaingó,5-III-1953, Krapovickas 7890 (CTES, LIL, chromosomes (Krapovickas & Rigoni 1949, US); 10km W de Ituzaingó,5-III-1953, Krapovickas 1951). 7897(C, CTES,F, G,LIL, UC,US); Ituzaingó,1-1947, Spegazzini10071 (BAB, SI); ayo.Las Casuarinas,8- Holotype:ARGENTINA. Corrientes, dep. Capi- XII-1946,Pierotti 6208 (LIL); 30 kmW de Ituzaingó, tal,camino de Corrientesa San Cosme,entre km 10 y 16-1-1970,Krapovickas & al. 15680(CTES, K, MO, 11,muy abundante, flores amarillas, 21-XII-1935, P, SI); VillaOlivari, 18 km Wde Ituzaingó,2-X-1978, Clos5930 (BAB 59065!). Schinini& al. 15669(CTES, UC, US); EstanciaSan Pedro,56°52'W, 27°45'S, 10-XI-1976,Arbo 1088 Selectedadditional material: ARGENTINA. Co- (CTES, G, K, US). Dep. Berónde Astrada:Campo rrientes.Corrientes, XI-1921, G.T. Bertoni s/n (LPS YaguáCuá, 24-IV-1960, Pedersen 5550 (K, US, Herb. 23 125).Dep. Capital:en céspedde plaza,XII-1975, Pedersen).Dep. Concepción:Médano el Carmen, MartinezCrovetto 10137 (CTES); mataderoayo. 6-1-1960,Pedersen 5326 (K, US, Herb.Pedersen); Pirayuí,8-1-1966, Krapovickas & al. 11905 (CTES, Ea. San Justodel Palmar, 8-III-1967, Pedersen US); ayo. Riachuelo,puente Pesoa, 2-III-1953, 8084 (CTES, Herb.Pedersen); Tabay, 7-X-1973, Krapovickas7830 (CTES, LIL, MO, NY, US); ayo. Schinini7453 (CTES); El Batel,Paso Crucecita, Riachueloy ruta12, 23-1-1959, Gregory & al. 9530 11 -II- 1968, Krapovickas & al. 13800(CTES, MO); (GH, LIL, MO, NY, SI, SP, US); id.,Gregory & al. 6 km E de Santa Rosa, estanciaMillán, 27-III- 9531 (GH,LIL, MO, NY, SI,US); rutaProv. 5,4 km E 1975,Arbo 839 (CTES); ruta17, 9 kmNE de Santa de ruta 12, camino a Laguna Brava, 14-1-1966, Rosa, 30-III-1974, Krapovickas & al. 24580 Krapovickas& al. 11919(CIES, NY,US); lkmWde (CTES, G, UC); entreSanta Rosa y el río Santa Laguna Brava, 28-XII-1976,Krapovickas & al. Lucía, 5-II-1968,Krapovickas & al. 13780 (CEN, 30048(CTES, US); entreLaguna Brava y ruta 1 2, 28- CTES, F, US); 11 kmNW de SantaRosa, 16-XII- XII-1976, Krapovickas & al. 30049(CTES); ruta12, 4 1977, Tressens 1023 (CTES, G, NY). Dep. kmNE delaeropuerto "Cambá Punta", 28-XII-1976, Mburucuyá: 32 km E de Saladas, estancia Krapovickas& al. 30050 (CTES); id.,31-III-1980, Pindapoy,26-XI-1970, Carnevali 2271 (CTES); Krapovickas& al. 36000 (CTES, GH,NY, US); id., Loma Alta,4-1-1962, Pedersen 6420 (K, US, Herb. Krapovickas& al. 36001(CTES, MO, NY, US). Dep. Pedersen); 10 km W de Mburucuyá,5-II-1968, San Cosme:between San Cosmeand Ramada Paso, Krapovickas& al. 13782(CTES, G). Dep. Lavalle: 9-XII-1947,Stephens & al. SH 16 (LIL); ruta12, Puntadel Rubio,Anzótegui 1210 (CTES, SI, US). accesoa Paso de la Patria,12-IV-1964, Krapovickas PARAGUAY. Central. Cerro Però, entre & al. 11350 (C, CTES, G. F, MO, P, SI). Dep. Itati: Ipacaraiand Pirayú, 10-11-1966, Krapovickas & al. RamadaPaso, 24-1-1959,Gregory & al. 9548 (GH, 12593(CEN, CTES, G, MO, NY, P, SI, US); id.,16- LIL,MO, NY, SI, SP,US); ruta12, km 995, 24-1-1959, VI-1977, Krapovickas& al. 30108 (CTES, F, G, Gregory& al. 9549(LIL, US): ruta12, km 1015, 24-1- GH,K, LIL, MO, NY, UC, US). PresidenteHayes. 1959,Gregory & al. 9551 (LIL, US); ruta12, 4 kmE Villa Hayes,4 km W del río Paraguay,13 -XII- del acceso a Itati,23-1-1972, Krapovickas & al. 1979,Schinini 19616 (C, CTES, F, G,LIL, MO, NY, 20797(C, CEN, CTES, F, G,K, MBM,MO, SI,UC); P, UC, US); rutaTrans-Chaco, entre B. Aceval y ruta 12, 15 km del acceso a Itati, 23-1-1972, Villa Hayes,18-V-1981, Krapovickas & al. 37459 Krapovickas& al. 20854 (BAB, CEN, CTES, F, G, (CTES, G, IPA,MO, SI). LIL, LPB, MEXU, RB, SI, UC); ruta12 a 5 kmde Yacaré,23-X-1957, Sayago 2933 (SI). Dep. San Luis Cultivatedmaterial: ARGENTINA. Cordoba. delPalmar: ayo. Riachuelo, 7 kmSE de San Luisdel Manfredi,13-V-1950, Krapovickas 7201 (CTES), Palmar,14-1-1966, Krapovickas & al. 11941(CTES, (proc.:Corrientes, entre Itati y Tuyutí,leg. Báez y UC, US); San Luisdel Palmar, Luisa A. Loma,4-XI- Rigoni). U.S.A. Georgia. Tifton,PI 262135, PI 1968,Krapovickas 14488 (BAA, CTES, F, G,K); San 258943,Hammons & al. TiftonAcc. A-28, 2-XI-1979 LuisDel Palmar,28-X-1969, Burkart 27595 (CTES, (CTES), proc.Argentina, Corrientes, ruta 12, río SI). Dep. San Miguel:Loreto, 25-1-1959, Gregory & Empedrado,leg. Clos & Nieva,21-VI-1947, D.E.I.P. al. 9557(LIL, US); id, Gregory& al. 9558(US); 12km 6519. N de Loreto,8-III-1974, Schinini 8370 (CTES, G). Dep. Ituzaingó:ruta 12, km 1134,entre Itá Ibatée Geographicdistribution. This species occurs

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in the northwesternpart of the province of 64. Arachis Simpsonii Krapov. & W.C. Corrientes(Argentina), in deep sandy soils. Gregory nov. sp. similar were foundin the Very plants proximity Fig. 3,64 of Asunción (Paraguay) near Ipacaray, to the east of the Paraguay River and in the neighborhoodof Villa Hayes, to the west of Herba perennis.Caulis villosus.Stipulae pilis said river. In this last place it grows in fine brevibus,adpressis vestitae,margine ciliatae, sandysoils, on small hillsof redsoil withrocky illae foliorum apicalium setulosae. Folíola outcrops. ovalia, mucronulata,epiphyllo pilis brevibus sparsim vestito,hypophyllo pilis ca. 1.5 mm Obs. 1. Arachis correntinais very close to longis,villoso, nervo medio et margineciliato A. villosa Benth., from which it is prominentibus.Hypanthium 45 mm longum, distinguishedprincipally by its smoothfruit and villosum.Calyx 6-7 mmlongus, setulis nonnullis the lightlymarked margins of its leaflets.In A immixtisvillosus. Vexillum suborbiculare, ca. 15 villosa the articles of the fruitshow a very mm longum,aurantiacum. Fructus biarticulatus marked reticulum, similar to that of A. paxillo 7-20cm longo,articulis 8-10 mmlongis x hypogaea. 4-5 mmlatis, apice recurvo,pericarpio laevi. The type specimen has leaflets with the upper surfacecompletely glabrous, but in ma- Holotype: BOLIVIA. Santa Cruz. Prov. terialcollected in thetype locality, at 10 kmNE Sandoval,5 km SSW de San Matías (58°26'W, of Corrientes and 4 km NE of the airport 16°21'S),cerca de unalaguna, suelo de aréna,16-IV- (Krapovickas & al. 30050, 36000 and 36001), 1980, Krapovickas,Simpson & Schinini36009 the young leaves have shorthairs scatteredon (CTES). Isotypes:CEN, G, GH, K, LIL, LPB, MO, the upper surface. None of these specimens NY, P, SI, SP,US. has bristleson the stipules,but in the material thatgrows in the vicinityof the type locality, Perennial plant. Taproot thick, obconical, the presenceof bristlesis variable. 10-15 mm in diameter.Branches procumbent, ca. 80 cm long, stemangular, villous withtwo Obs. 2. Arachis correntina was one of kinds of hairs: long, ca. 1.5 mm long, easily the parents used in the first interspecific falling off, and small, adpressed, more hybrid obtained in the genus Arachis, persistent hairs; internodes 2-5 cm long. producingan F, triploidupon crossing it with Leaves tetrafoliolate.On the lateral branches, A. hypogaea var. fastigiata (Krapovickas stipuleswith the fused portion 5-7 mmlong, the & Rigoni 1949, 1951). freeportion triangular, falcate, 15-20 mmlong Arachis correntina only produces x 4 mmwide. Petiole 8-20 mmlong; rachis 5-8 hybrids when crossed with species of mm long. Leaflets oval, mucronulate,apical section Arachis. When crossed with the pair sometimesbroad, 25 mm long x 16 mm annual species, it produced highly sterile wide, up to 36 x 13 mm; basal pair commonly hybrids.with A. Batizocoi, and hybrids of 21 mm long x 12 mm wide, up to 28 x 12 mm. greater fertility with A. stenosperma Stipules with 6-7 prominentveins, surfaces (37.6%) and with ./4.duranensis (54.3%). It with small, adpressed, scatteredhairs, villous also produced hybrids with the perennial toward the base and on the back of the fused species A. Diogoi (35.85%), A. Cardenasii portion,margins citiate; toward the apex of the (63.8%) and A. villosa (73.15%). (see pp. branchesthe stipules have few bristleson the 197-198 and M.P. Gregory& W.C. Gregory fused portion.Petiole canaliculate, villous on 1979). thedorsal line and on themargins of the canal, On the basis of cytogeneticstudies, Murty the remainderwith scatteredadpressed hairs, and Jahnavi (1986) conclude that A. canal withsome small adpressed hairs. Rachis correntina also could have been onç of the narrowtoward the base, with pubescence si- species that contributedthe 'A' genome of milar to that of the petiole; pulvinus villous. A. hypogaea. Leafletsvillous; uppersurface with short hairs,

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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 uniformlyscattered, margin prominent; lower from 7 to 8 mm long x 5 mm wide, with a surfacevillous, hairs ca. 1.5 mmlong, midvein reticulatepericarp. These reticulate articles and marginal veins very prominent;margin are reminiscentof those of A. villosa, but are ciliate. Flowers along the length of the much smaller. branches, in very short few-floweredspikes. Hypanthium 45 mm long, villous. Calyx Obs. 2. Arachis Simpsonii is similar to A. bilabiate,villous and withsome bristles;upper villosa, fromwhich it is distinguishedby its lobe 6 mm long, lower lobe 7 mm long, largerleaves and because inA. villosa thefruit subfalcate. Standard suborbicular,ca. 15 mm articles are larger and noticeably reticulate long, orange. Fruit subterranean,Particulate; (Fig. 3, 62 and 65). Both species have villous peg 7-20 cm long, villous on the aerial part; upper surfacesof the leaflets. articles8-10 mmlong x 4-5 mmwide, smooth, Due to the presence of hairs on the upper with beak, covered with a dense coat of side of the leaflets,it can also be comparedto diminutivehairs. Seeo ca. 7 mmlong. A. Diogoi; but in the latter the leaflets are muchnarrower, with differently colored upper Additionalmaterial: BOLIVIA. Santa Cruz. and lower surfaces and the margins almost tProv. Sandoval: at northedge of San Matías,on unmarked. roadto San Francisco, 24-VÜI-1981, Vails & al. 6346 (CEN, CIES). BRAZIL.Mato Grosso.Campo cerra- We dedicate this species to Dr. Charles E. do atkm 135 on the road from Cáceres to S ofSierra do Simpson with whom we collected the type Aguapei,4-m-1977, Kiikbride & al. 3042(INPA, US). specimen. Relatedmaterial: BRAZIL. Mato Grosso.Mun. Cáceres:23 kmda BR-174,camino P. Esperidiãoa 65. Arachis Cardenasii Krapov. & W.C. Casalvasco, 28-V-1985,Vails & al. 8896 (CEN, Gregory nov. sp. EstanciaVilla Isabel e Rest.Serra- CTES); (Posto Fig. 3,65 no), km 55 da MT-265 (camino de Porto Esperidiãoa Casalvasco), 28-V-1985,Vails & al. 8900(CEN). ArachisCardenasii Krapov. & W.C. Gregory,in W.C. Gregory,M.P. Gregory,Rrapovickas, Smith & Geographic distribution. This species Yarbrough,in Peanuts - Cultureand Uses: 98, 1973, inhabitsthe border area between Santa Cruz nomennudem. This refers to the speimen GKP 10017. (Bolivia) and Mato Grosso (Brazil). In addition to the type collection, a herbariumspecimen Herba perennisi Caulis angulosus,villosus, was also made in the Sierra do Aguapei, fereviolaceus. Stipulae basi villosae.Folia caulis located some 90 km WNW of San Matías. It principalisfoliolis ovalibusvel subobovatis,ilia grows in the "cerrado," which is the type of ramorum foliolis suborbicularibus usque vegetationcalled "pampa" in northeasternBo- obovatis,epiphyllo laevi, glabro, hypophyllo pilis livia. adpressisvix conspicuis vestito, nervo medio pilis ca. 2 mmlongis gerente prominente, folia juniora Obs. 1. The specimen Kirkbride& Lleras subtus longe pilosa, marginepaulo manifesto 3042 is very similar to the type, but has a longeciliato parcius setuloso. Hypanthium 2-6 cm greater number of bristles on the stipules, longum,villosum. Calyx 6-8 mm longus,pilis which are also foundon the petiole and on the longissetulisque vestitus. Vexillum aurantiacum, rest of the rachis. It also has some bristleson raroluteum, 10-14 mm longum x 12-16mm latum. the margin of the leaflets. In the type Fructus biarticulatuspaxillo 10-15 cm longo, specimen, very few bristles are found on the isthmus3 cm longus,articuli laeves, 7-11 mm stipulesof the apical leaves. longix 4-6 mmlati, pericarpio tenui. The specimens Vails & al. 8896 and 8900 resemble A. Simpsonii in the shape of the Holotype:BOLIVIA. SantaCruz. Prov. Chiqui- leaves and in the tomentum,but they are distinguishedfor having fruitswith articles ®Errorinoriginai, appeared as annua.

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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) tos,2 kmSW deRoboré, ca. 300m del río Urasibiqui, bi-ovulate.Fruit biarticulate; peg glabrous on 21-IV-1980,Krapovickas, Simpson & Schinini36015 the aerial part, 10-15 cm long; isthmus3 cm (CTES). Isotypes:CEN, G, GH, K, LIL, LPB, MO, long; articles 7-11 mm long x 4-6 mm wide, NY, P, SI, SP,US. beak somewhat developed, pericarp thin, smooth,with short hairs thatretain a thinlayer Perennial plant. Taproot deep, without of soil. 2n=20 chromosomes (Smartt 1964, enlargements.Mainstem erect, 10-90 cm long, Smartt& Gregory1967, GKP 10017). withlong branchesusually on thebasal 10 cm, later continuingwith very short vegetative Additionalmaterial: BOLIVIA. Santa Cruz.Prov. branches and reproductivebranches. Lateral Chiquitos:Roboré, X-1934. Cárdenas 2988 (G); id.,8- branchesprocumbent, ca. 60 cm long,but up to n-1959,Meyer 20287 (LIL); id., 3 l-HI-1959,Gregory & 1.2 m long,with the base partiallyburied in the al. 10017(LIL, MO, NY, US); id.,l-IV-1959, Gregory & soil; stemangular, villous, hairs ca. 2 mmlong, al. 10026(CTES, GH, LIL, MO, NY, SI, SP, US); id.,21- internodes3-4 cm long, usually violaceous. IV-1980,Krapovickas & al. 36016 (CEN, CTES, G, GH, Leaves tetrafoliolate. On the mainstem, K, LIL, LPB, MO, NY, US); id., 22-IV-1980, stipuleswith the fused portion 10-13 mmlong, Krapovickas& al. 36017 (CEN, CTES, G, GH, LPB, MO, the free tips 20-30 mm long x 3 mm wide; NY, US); id.,Krapovickas & al. 36018(CEN, CTES, petiole 40-45 mmlong; rachis 10-12 mmlong; LPB, NY, US); id.,Krapovickas & al. 36019 (CEN, leafletsoval or somewhatobovate, the apical CTES, LPB, GH, K, MO, NY, US); 24-IV-1980, pair up to 55 mmlong x 28 mmwide, thebasal Krapovickas& al. 36022(CEN, CTES, LPB, NY, US); pair up to 46 mm long x 21 mm wide. On the id.,Krapovickas & al. 36023(CEN, CTES, G, GH, LPB, lateralbranches, stipules with the fused portion MO,NY, US); Roboré,21-XI-1989, Nee 37826 (CIES); 5-7 mm long, the freetips 12-17 mm long x 3 6 km de Roboré,camino a Santiago,23-IV-1980, mmwide; petiole 5-12 mmlong; therachis 5-7 Krapovickas& al. 36020(CEN, CTES, GH,LPB, MO, mm long; leaflets suborbicular or obovate, NY, P, SI, US); Santiago,12-11-1958, Krapovickas obtuse, apical pair up to 35 mm long x 25 mm 9412(CTES, LIL); id.,23-IV-1080, Krapovickas & al. wide, thebasal pair up to 32 mmlong x 22 mm 36021 (CEN,CTES, G, GH, K, LPB,MO, NY, P,US); wide. Stipules villous toward the base, with Santiago de Chiquitos,23-XI-1989, Nee 37846 subglabrous surfaces, usually with some (CTES); 15 kmS de San José,camino a Natividad, adpressed hairs on the surface of the fused 29-IV-1980,Krapovickas & al. 36032(CEN, CTES, G, portionand witha line of long hairs on each of GH,K, LIL, LPB, MO, NY, SI, US); 17km S de San the two lateral veins on the back; the free José,20-IV-1980, Krapovickas & al. 36033 (CEN, portionacute, withciliate margin.Petiole and CTES, G, LPB, MO, NY, US); 21 kmSSW de San rachis canaliculate, the back with short José,29-IV-1980, Krapovickas & al. 36034 (CEN, adpressed hairs and scatteredhairs ca. 2 mm CTES, G, GH,LIL, LPB, MO, NY, US); 2 kmW de long, margins of the canal ciliate; pulvinus Natividad,29-IV-1980, Krapovickas & al. 36035 villous. Leaflets with the upper surface (CEN, CTES, K, LPB, K, MO, NY, P, US). Prov. glabrous, smooth, with the margin little- Sandoval:San Matías, 58°26'W, 16°21'S, 20-IV- marked; lower surface with some barely visi- 1980, Krapovickas & al. 36011 (CTES); id., ble adpressed hairs, midveinprominent, with Krapovickas& al. 36013(CTES). hairs ca. 2 mm long, frequentlyalso with PARAGUAY.Alto Paraguay (ex Chaco).Mayor sparse long hairs on the rest of the lower PedroLagerenza, 60°45'W, 20°5'S, río Timane, 5-IV- surface of young leaves; marginlongly ciliate 1978,Schinini & al. 15219B (CTES, G,MO, SI, US) and with some bristles. Axillary spikes very (see Obs. 2 inA. duranensisy,Cerro León, 60°15' W, short,5 -flowered,bracts with long hairs on the 20°26'S,laguna Tareyí, 26-VIII-1981, Schinini & al. veins. Hypanthium2-6 cm long, pilose. Calyx 21198 (CTES). bilabiate, covered with long hairs and with bristles,upper lobe 4-dentate,6-8 mm long, Geographic distribution. This species is lower lobe falcate, 7-9 mm long. Standard foundin the departmentof Santa Cruz (Boli- orange,rarely yellow, 10-14 mm long x 12-16 via), mainly on the southern edge of the mmwide; wings yellow, 7-10 mm long. Ovary Chiquitana mountains.It was also collected in

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San Matías, on the borderwith Mato Grosso longisX 5-6 mm latis, pericarpio laevi. (Brazil), and in the northof the Paraguayan "chaco." It grows in deep, sandy soils. Holotype:BOLIVIA. Dep. Santa Cruz. Prov. Gutierrez,10 km W de Portachuelo(camino a Buena Obs. Smartt,Gregory & Gregory(1978a), Vista),63°24'W, 17°20'S, ca. 350 m s.m.Junto con based on cytogenetic studies, proposed A. Paspalumnotatum, cerca de arroyo,también entre Cardenasii and A. Batizocoi as the possible Bromelia,borde de bosque de Cecropia,Cassia, donors of the 'A' and 'B' genomes, Attalea, Psidium, Samanea etc., 20-IV-1977, respectively, of A. hypogaea and that this Krapovickas& al. 30085 (CTES). Isotypes:CEN, species could have originatedfrom a sterile GH,K, LIL,LPB, MO, NY, US. hybridsimilar to thatproduced by said diploid species. Perennial plant, with a strong and deep Arachis Cardenasii crossed only with taproot,ca. 10 cm in diameterat the collar. species of its own section. It producedhighly Stem angular with small adpressed hairs and sterile hybrids with the annual taxa A. long hairs up to 1.5 mm long, aligned over the Batizocoi, A. montícola, A. hypogaea var. angles, or nearly glabrous. Lateral branches hypogaea and A. hypogaea var. fastigiata. prostrate ca. 1.20 m long. Leaves Hybrids were obtained with more than 48% tetrafoliolate.On the mainstem,stipules with pollen stained with the annual species A. the fused portion 14 mm long and the free stenosperma and A. duranensis and with the portion20 mm long; petiole 45-60 mm long; perennial species A. Diogoi, and with even rachis 10-12 mm long; leafletslanceolate, the greater fertilitywith A. correntina (63.8%) distal ones 40-45 mm long x 14-17 mm wide, and A villosa (70.4%), also perennial(p. 198, the basal ones 35-42 mm long x 12-15 mm and Gregory,M.P. & Gregory1979). wide. Leaves of the lateral branches: stipules Resslar & Gregory (1979) studied the withthe base fused4-10 mm long and the free meiosis of A. Cardenasii, A. duranensis and tips 15-17 mm long x 3 mm wide, glabrous or A. villosa and theirhybrids, and concludedthat withsome small adpressedhairs, on themargin these three species have a common genome, hairs somewhatlarger, also adpressed; petiole which theydesignated genome 'A.' 5-25 mm long; rachis 4-7 mm long; distal leaflets20-35 mmlong x 12-13 mmwide, basal 66. Arachis Kempff-Mercadoi Krapov., leaflets18-19 mmlong x 10 mmwide, oblong, W.C. Gregory & C.E. Simpson nov. sp. sometimes obovate, upper surface smooth, Figs. 3,66; 29 fromvillous to glabrous on the same branch, lower surface with small adpressed hairs or almost glabrous, margin and veins little Herbaperennis. Radix polaris. Coulis principalis marked, hairs more or less adpressed and erectus.Rami prostrati usque ad 1.20m longi. Caulis some short bristles on the margin. pilis parvisadpressis cum pilis ca. 1.5 mmlongis Inflorescencesshort. Hypanthium70-90 mm immixtisvestitus, aut fere glabro. Stipulae glabrae long,villous. Calyx 5 mmlong, villous and with vel pilis nonnullisparvis, adpressis, margine bristles.Standard 12 mm long x 15 mm wide, majoribus vestitae. Folia caulis principalis wings 8 mm long x 5 mm wide. Fruit foliolislanceolatis, ilia ramorumfoliolis oblongis subterranean,Particulate; peg 13 cm long; vel interdumobovatis. Epiphyllum laeve, villosum isthmus4.5 cm long; articlessmooth, 9-10 mm usque glabrum. Hypophyllumpilis parvis longx 5-6 mmwide. Seed 6.5-8.5 mmlong x 4- adpressis vestitumvel fere glabrum,margine et 4.5 mm wide, testa uniformlytan, darkerand nervispaulo prespicuis,margine pilis plus minusve browner in old seeds. 2n=20 chromosomes adpressissetulisque nonnullis brevibus instructo. (Fernández& Krapovickas 1994). Hypanthium70-90 mm longum, villosum. Calyx 5 mmlongus, villosus setulosusque. Vexillum 12 mm Additionalmaterial: BOLIVIA. Santa Cruz.Bue- longum.Fructus subterraneus, biarticulatus, paxillo na Vista,20-1-1915, Steinbach 1000 (LIL); id., 11- 13 cmlongo, isthmo 4.5 cmlongo, articulis 9-10 mm XII-1920,Steinbach 5211 (NY); id.,Steinbach 6673

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Fig.29. Arachis Kempff-Mercadoi : plant (K.30084).

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(K); SantaCruz, 22-VI-1965, Adolfo M. 443 (US). littleto thenorth of thecity of Santa Cruz de la Prov. Gutierrez:orillas del río Piray,8 km E de Sierra,to the west of the Piray River,between Portachuelo,80 kmN de Sta.Cruz, 63°1 8W, 17°20'S, Portachueloand Buena Vista, in deep, sandy 20-IV-1977,Krapovickas & al. 30084 (CTES, GH, soils. It was also found in Ascensión de LPB, MO, NY, P, SI, SP, US); 13,1 km W de Guarayos, in the same department. Portachuelo,camino a Buena Vista, 7-IV-1979, Krapovickas& al. 35003 (BAB, CEN, CTES, GH, Obs. We dedicate this species to our lost LIL, LPB, MO, NY, US). Prov.Sara: 2 kmENE of friend,Noel KempffMercado, who promoted Portachuelo,on highwayto Montero,ca. 17°21'S, the cultivationof this species, for its colorful 63°17'W,21-V-1991, Nee 40482 (CTES); caminoa and prolonged flowering,in the parks and NuevaMoca, 2,5 kmN de la rutaPortachuelo-Bue- gardensof his native Santa Cruz de la Sierra. na Vista, 10 km W de Portachuelo,7-IV-1979, Krapovickas& al. 35002 (BAB, CTES, GH, LPB, 67. Arachis Diogoi Hoehne 20 km MO, NY, SI, SP, US); cercade Caranda, SW Fig. 3,67 de Portachuelo,7-IV-1979, Krapovickas & al. 35004 (CTES, GH,K, LIL, LPB, MO, NY, SP, US). Prov. AndrésIbañez: near rio Los Ajos, 0,5 km N of Hoehne,Comm. lin. telegr. Bot. 8: 71-72, 1919. Terevinto,17°43'S, 63°22'30"W, 30-XI-1988, Nee & "Exempl.s.n. do Dr. JulioCesar Diogo. Estampan. al. 37008 (CTES); Carandá,500 m, 25-IV-1966, 147. Colhida nas margensarenosas da bahia da Brooke108 (K); betweenPortachuelo yBuena Vis- Gahyba, em Matto Grosso; florescendo em ta,14-111-1964, Badcock 19 (K). Setembro." A. villosa Benth. subsp. Diogoi (Hoehne) A. Relatedmaterial: BOLIVIA. Santa Cruz. Prov. Chev.,Rev. Int. Bot. Appl. Agrie. Trop. 13(146-147): Ñuflode Chávez:Ascensión de Guarayos,2-3 km al 761,1933. N caminoa Centinela,26-IV-1977, Krapovickas & al. A. chacoenseKrapov. & W.C. Gregory,in W.C. 30086(BAB, C, CEN,CTES, GH LPB,MO, NY, US); Gregory,M.P. Gregory,Krapovickas, Smith & id.,aeropuerto, 27-IV-1977, Krapovickas & al. 30087 Yarbrough,in Peanuts- Cultureand Uses: 73, (CTES, NY, US); id.,3 kmal S, arroyoSan Joaquin, 1973, nomen nudum.Name proposed for the 27-IV-1977,Krapovickas & al. 30088(CEN, CTES, specimenGregory & Krapovickas10602 from Puer- GH,K, LPB,MO, NY, SI,US); id.,1 km al W,27-IV- to Casado,Paraguay. 1977,Krapovickas & al. 30089(CTES, G, GH,LIL, LPB, MO, NY, SP, US); bordesud de Ascensiónde Perennial plant. Taproot 5-10 mm in Guarayos,28-IV-1977, Krapovickas & al. 30090 diameter, deep, without enlargements. (CTES, GH,LPB, MO, NY, US); 2 kmNE ofAscen- Mainstem erect. Branches prostrate, sionde Guarayos,23-DÍ-1988, Williams 699 (CTES); somewhat decumbent,30-60 cm long. Stem 2 km N of Surucusi,on road to Ascensionde cylindrical,somewhat angular on the young Guarayosbefore El Puente,22-IX-1988, Williams parts, villous, at times with some bristles 697(CIES). present; internodes30-60 mm long. Leaves tetrafoliolate.Stipules with the fused portion 6- Cultivatedmaterial: BOLIVIA. SantaCruz. Prov. 8 mm long,the freeparts ca. 1 1 mm and up to AndrésIbañez: Santa Cruz de la Sierra(procede del 20 mm long. Petiole 12-17 mm long on lateral camino Buena Vista-Portachuelo),20-IV-1977, branches and up to 35 mm long on the Krapovickas& al. 30103 (CTES, NY, US); id., mainstem.Rachis 5-8 mmlong. Apical pair of Krapovickas& al. 30104 (CTES, US); id., 24-IV- leaflets commonly26 mm long x 6 mm wide 1977,Krapovickas & al. 30105(CTES, NY, US); id., but up to 39 mmx 8 mm; basal pair commonly 6-IV-1979,Krapovickas & al. 35001(CEN, CTES, G, 22 mm x 5 mm but up to 30 mm x 7 mm. GH,LIL, LPB, MO, NY, US); id.,23-11-1988, Williams Stipules violet,with the fused portionalmost 691 (CTES); id.,Valls 13250 (CEN). smooth,the veins slightlymarked, softly villous with short, dense, silky hairs; free portion Geographiedistribution. This species grows narrowlytriangular, 1.5 to 3 mm wide, acute, in the departmentof Santa Cruz, in Bolivia, a subfalcate, with veins somewhat prominent;

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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, Arac hi s (Leguminosae) hairs short, adpressed, scattered; margin id.Valls & al. 9148(CEN). ciliate. Petiole and rachis canaliculate, canal PARAGUAY. Alto Paraná. PuertoCasado, I- and back pubescent,long hairs on the margins 1917,Rojas 2923 (LIL); id.,Rojas 2928 (SI); id.,18-11- of the canal and on the dorsal line; pulvinus 1948,Stephens & al. SH 115 (LIL, NY); id.,25-11- densely villous. Bristles occur frequentlyon 1950,Ramírez 630 (SI); id.,19-V-1961, Gregory & al. the stipules,petiole and rachis. Leaflets from 10602(CTES, GH,LIL, MO, NY, SI,US). Paraguarí. narrowlanceolate to oblong-lanceolate,acute, ArroyoTebicuary y frentea Villa Florida,15-11- apiculate; upperside smooth,somewhat villous 1950,Rosengurtt B-5762 (CTES, MVFA,SI); id.,4- with short,more or less erect scatteredhairs; 11-1966,Krapovickas & al. 12404(CTES, G,NY, SI, underside villous, midrib and margins SP, US); id., 16-VI-1977,Krapovickas & al. 30106 somewhatprominent, with violet tinge; margin (CEN,CTES, F, GH,K, LIL,MO, NY, US). ciliate and with some bristles. Flowers along the length of the branches in short 3-4- Cultivatedmaterial: ARGENTINA. Misiones. floweredspikes. Hypanthium4.5-7 cm long, PuertoIguazú, cult., formando césped, procedente villous. Calyx bilabiate, denselyvillous, with de PuertoCasado, 12-1-1972,Mroginski & al. 360 long silkyhairs and some bristles;upper lobe (CIES). broad, 7 mm long, tridentate, lower lobe subfalcate, 8 mm long. Corolla frequently Geographiedistribution. This species grows entirelyyellow, occasionally with an orange in thewatershed of the Rio Paraguay, fromthe standard;standard up to 16 mm long x 18 mm shores of Lagoa Gaiba and the Pedro II canal wide, wings up to 11 mm long. Fruit which unitesthe formerwith Lagoa Uberaba, subterranean,Particulate; peg ca. 15 cm long, on the border of Mato Grosso do Sul with aerial partvillous, violettinged; articles 10-11 Bolivia, to Puerto Casado, in Paraguay. It is mm long x 5-6 mm wide, withbeak, pericarp adapted to the Pantanal where it tolerates reticulate.2n=20 chromosomes(Smartt 1964; annual floods up to 3 m deep (Vails & al. Smartt& Gregory1967; GK 9901, 10602). 9147). We ascribe to this species, with certain Holotype: BRAZIL. Mato Grosso do Sul. doubts, material collected in San Miguelito, Margensda Lagoa Gahiba,IX- 1908, Diogo 3 17 (R near San Ignacio (Santa Cruz, Bolivia), a 4925!). locality that while pertainingto the Amazon basin, is not very far fromthe headwaters of Selectedadditional material: BOLIVIA. Santa the Rio Paraguay. By the same token, we Cruz.Prov. Velazco: San Miguelito, 23 kmN de San included specimenscollected between Cuiabá Ignacio,8-V-1977, Krapovickas & al. 30101(CTES, and Rondonópolis(MT) and othersfrom as far G,GH, LPB, MO, NY, SI,US); id.,Krapovickas & al. south as the arroyo Tebicuary, in the 30102(CEN, CTES, G, GH, K, LIL,LPB, MO, NY, SI, departmentof Paraguarí (Paraguay). US). BRAZIL. Mato Grosso. 70 km SE de Cuiabá, Obs. 1. Arachis Diogoi presents certain estradaRondonopolis-Cuiabá, 9-III-1959, Gregory polymorphismsthat we do not find to be & al. 9901(CTES, GH,LIL, MO, NY, SI, SP,US); 62 related to one another.In the two collections km SW ofCuiabá on roadto Poconé,at rio Bento thatwe made at Lagoa Gaiba, thetype locality, Gomes,21-VÏÏI-1981, Valls & al. 6330(CEN, CTES). the majorityof the plants had yellow flowers, Mato Grosso do Sul. PortoIndio, orilla del canal and about 1% showed flowerswith an orange PedroII que unela lagoa Gaibay la lagoa Uberaba, standard. 57°55'W, 17°42'S,6-XII-1976, Krapovickas & al. On the other hand, all of the individuals 30001(CEN, CTES, G, GH,LPB, MO, NY, SI, US); observed from the collection from Pedro II Jaguaribe,orilla sud de Lago Gaiba, 57°40'W, canal (30001) had bristles on the stipules, 17°48'S,6-XII-1976, Krapovickas & al. 30005(CEN, petiole and rachis. The type specimenhas not CTES,GH, K, LIL,MO, NY, SP,US). Mun.Corumbá: one bristle.In contrast,in the populationfrom 1 km W do Porto da Manga, na estradapara Jaguaribe(30005) there are individuals with Corumbá,12-X-1985, Valls & al. 9147(CEN, CTES); bristlesand individualswithout them.

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Obs. 2. The interpretationof A. Diogoi has with species of section Arachis, producing had some problemsthat have led to confusion. very sterile hybridswith the annual taxa A. It was described by Hoehne in 1919 and the Batizocoi, A. montícola, A. hypogaea var. type specimen was deposited in the Museo hypogaea and A. hypogaea var. peruviana, Nacional de Rio de Janeiro. Later, upon withmore than 28.9% stained pollen withA. preparing his revision of the genus Arachis villosa, A. corrientina and A. Cardenasii, (1940), work done at the Institutode Botánica and up to 88.5% with A. stenosperma (pp. de São Paulo, Hoehne was not able to 197-198 and Gregory,M.P. & Gregory1979). reexaminethe collectionsdeposited in theMu- seo Nacional de Rio de Janeiro (Hoehne & 68. Arachis Kuhlmannii Krapov. & W.C. Kuhlmann195 1: 230). Surely,it is owingto this Gregory nov. sp. thatHoehne would in his circumstance assign, Figs. 3,68; 30 monograph,this name to a differentspecies fromthe area aroundCampo Grande(MGS), a species that we name A. Archeri. Both Herbaperennis.Radix polaris. Coulis pilis ca. 1.5 species have the form of the leaflets in mm longis villosus usque glabrescens.Stipulae common,but they differ in habit,prostrate in A villosaebasi setulosaeque,apicibus liberis glabrae, Diogoi and erect or decumbentin A. Archeri, margineciliatae. Coulis principalis erectus foliolis in root system,with enlarged branchings in A. oblongo-lanceolatis.Rami procumbentesfoliolis Archeri, and in the position of the flowers, obovatis.Epiphyllum laeve, glabrum, hypophyllum along the length of the spreading stems in glabrumvel in foliis novellispilis parvissimis Hoehne's species and clusteredaround the base adpressisvestitum, margine haud incrassatociliis of theplant in thespecies fromCampo Grande. setulisquenonnullis instructo. Hypanthium 20-60 The name A. Diogoi appears repeatedlyin mmlongum, villosum. Calyx 6-7 mm longus, villosus the cytogeneticsliterature to designateplants setulosusque.Vexillum aurantiacum, raro luteum, cultivated at the Instituto Agronòmico de 10 mmlongum. Fructus subterraneus, biarticulatus, Campinas (São Paulo, Brazil) accordingto the paxillo 8-12 cm longo,aliquanto villoso, isthmo 3 concept that Hoehne would develop in his cmlongo, articulis 10-14 mm longis x 5-6 mmlatis, monographof 1940. In accordance with the pericarpiosubtiliter reticulato, fere laevi. criteria upon which we now support our determinations,the material designated V.85 is Holotype:BRAZIL. Mato Grosso.70 km E de A. major and V.84 and V.128 are A. Archeri Cáceres,camino a Cuiabá,Serra de Araras,16°08'S, (Mendes 1947, Conagin 1962, Pompeu 1977). 57°18'W,200 m s.m.,17-XII-1976, Krapovickas & In 1961, when we collected the specimen Gregory30034 (CEN). Isotypes:CTES, G, GH,K, 10602 in Puerto Casado, on the Paraguay LIL, MO, NY, P,RB, SI, SP,US. River, we found ourselves with a very differentplant fromthe one that at that time Perennialplant. Taproot, thick, up to 23 mm was called A. Diogoi and that grows in the in diameter,deep, withoutenlarged branchings higherparts of the Sierra de Maracaju, near or tuberoids. Mainstem erect, 20-25 cm in Campo Grande,in Mato Grosso do Sul. Thus, height,without flowers, branched at the base, we gave this plant the name A. chacoense, withinternodes shorter than the stipules, villous mentionedin the literaturebeginning in 1973. on the younger parts. Lateral branches Later, in 1976, we had the opportunityto prostrate,up to 80 cm long,with few branches, collect thetrue A Diogoi in thetype locality of but the secondary branches vegetative and this species. The great similarity between with inflorescences along their length, collections 10602, 30001 and 30005, and their internodes 3-6 cm long, from villous to similargenetic behavior, induced us to treatall glabrescent,with long hairs ca. 1.5 mm long. of these collections as pertainingto a single Leaves tetrafoliolate.On the mainstem,the species. fused base of the stipules 1 1-15 mm long, the freetips 25-30 mm long x 2-3 mm wide; the Obs. 3. Arachis Diogoi has been crossed petiole 37-45 mm long; the rachis 10-16 mm

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Fig.30. Arachis Kuhlmannii: A,leaf from mainstem; B, leaffrom lateral branch (K.30034).

long; theapical leaflets40-60 mmlong x 12-27 hairs,sometimes with bristles; on the leaves of mm wide, the basal leaflets 37-54 mm long x the mainstemthe petioles are more glabrous. 10-22 mm wide. Leaves on the lateral Leaflets of the mainstem oblong-lanceolate, brancheswith the fused portionof the stipules those of the lateral branches obovate, usually 5-7 mm long,the freeportion 13-25 mm long x rounded, sometimes somewhat acute, upper 3-4 mmwide; thepetiole 9-25 mm long; rachis leaf surface glabrous, lower leaf surface 7-10 mmlong; apical pair of leaflets28-37 mm glabrous, in young leaves with very small longx 13-23 mmwide, thebasal pair26-32 mm adpressed hairs, margin not thickened,with longx 1 1-21mm wide. The stipulesvillous with cilia and some bristles.Inflorescences axillary, bristleson the fused portion,the free portion 4-5-flowered, axis short, shorter than the with surfaces glabrous and margin ciliate, stipules. Hypanthium20-60 mm long, villous. stipules of the mainstem usually more Calyx villous and withbristles, the wider lobe 6 glabrous. Petiole villous or with a few long mm long, the narrow lobe subfalcate, 7 mm

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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 long. Standardorange, 10 mmlong, wings and 24 km NE do rio Taboco, estradaRio Negro- keel yellow; in some populations(30034) some Aquidauana,29-X-1985, Valls & al.9479 (ŒN, CIES); flowersentirely yellow. Fruitbiarticulate; pegs id.,30-X-1986, Valls & al. 10420(CEN); Faz Rancharla, 8-12 mm long, somewhatvillous on the aerial 20°02,S,55°57W, 24-XI-1989, Pott&al. 5451 (CTES). portion.At the beginningof its development, Mun. Corumbá:Faz. Nhu Mirim,Pantanal de when the peg has not yet enteredthe soil, a Nhecolandia,4-X-1976, Allem 120 (CEN); Faz Nhu zone of dense, adpressed hairs ca. 4 mm long Mirini,150 kmEde Corumbá, 7-XÜ-1976, Krapovickas appears some 2 mm back fromthe tip. 2n=20 & al.30008 (CEN, CIES, G,GH, K, LIL, MO, NY, SP, chromosomes(Fernández & Krapovickas1994). US); id.,17-X-1985, Valls & al. 9230& 9231 (CEN, CTES); Faz. Guanandi,18°52'S, 56°11W, 18-X-1985, Selected additionalmaterial: BRAZIL. Mato Valls& al. 9235& 9236(CEN, CTES); 11 kmda Faz. Grosso.Mun. Cáceres: 94 kmde Cácerescamino a Guanandina trilha para Nhu Mirim, 19-X-1985, Valls & Cuiabá,17-XÜ-1976; Krapovickas & al. 30035(CEN, al.9243 (CEN, CTES); Faz Ipanema,19°04'S, 56°32W, CTES, GH,K, MO,NY, RB, SI,US); Cáceres,cult, at 17-X-1985,Valls&al. 9214 (CEN, CTES). Mun. Miran- theairport, 31-VIII-1981, Vails & al. 6410 (CEN, da:Miranda, campo de futebol, 19-IV-1984, Valls & al. CTES); 1, 1 kma NW da pontedo rioParaguay a W 7639(CEN, CTES); id.,5-IV-1986, Valls & al. 9894 de Cáceresna saídapara Porto Velho (BR-354), 25- (CEN,CTES); inicioda rodoviaMiranda-Bonito, 6-IV- X-1985,Valls & al. 9375 (CEN, CTES); 47 kmE de 1986,Valls&al. 98% (ŒN, CTES). Cáceres,BR-174, km 680, 29-V-1985,Valls & al. 8916a(CEN, CTES); id.,Valls & al. 8918(CEN); 42 Relatedmaterial: BRAZIL. Mato Grosso.Pe de kmE deCáceres, BR-174, km 685, 30-V-1985, Valls & Anta, 20 km NW of Cáceres on road to Porto al. 8935a (CEN, CTES); 23,7 kmalem do ribeirão Esperidião,25-VIII-1981, Valls & al. 6351 (CEN, Flexas,entre Cuiabá e Cáceres,restaurante Oasis, CTES); Vila Bela, 27-VIII-1981,Valls & al. 6380 24-X-1985,Valls & al. 9354(CEN, CTES); id.,Valls & (CEN, CTES); 71 kmS ofVila Bela, 27-VIII-1981, al. 9355(CEN, CTES); Faz. SantoAndré, 20 kmSSW Valls& al. 6389(CEN); 75 kmS ofVila Bela, 27-Vm- de Cáceres,camino a San Matias(Bolivia), 24-VIII- 1981,Valls & al. 6390 (CEN); Casalvasco,29-VIII- 1981,Valls & al. 6344(ŒN, CTES); Caiçara,3 kmN 1981,Valls & al. 6396(CEN, CTES). Mun.Cáceres: 9 ofroad from Cáceres to Porto Esperidião (20 kmNW kmENE de PortoEsperidião (BR-174, km 91,9), 30- ofCáceres), 25-VIII-1981, Valls & al. 6352 (CEN, Vm-1981,Valls&al. 6408b(CEN);18-V-l 985, Valls CTES); 18,6km NW de Cáceres,BR-174, 25-VIII- & al. 8758(CEN, CTES); 45 kmN ofCáceres, on road 1981,Valls & al. 6355(CEN, CTES); id.,18-V-1985, toBarra do Bugres,31-VIII-1981, Valls&al. 6413 Valls & al. 8753 (CEN, CTES); 56 kmNW de Porto (CEN,CTES); id.,3 l-V-1985,Valls & al. 8964(CEN, Esperidião,BR-174, km 148,3, 30-VÜI-1981, Valls & CTES); 57 km S de Barra do Bugres,camino a al.6404 (CEN, CTES); id.,18-V-1985, Valls&al. 8763 Cáceres,l-VI-1985, Valls & al. 8979(CEN, CTES); 73 & 8764(CEN, CTES); id.,28-V-1985, Valls & al. 8887, kmS de Barrado Bugres,camino a Cáceres,1-VI- 8888 & 8889 (CEN, CTES). Mun. N. Sra. do 1985,Valls & al. 8980(CEN, CTES); 55,1km NE de Livramento:rio Piraim, 4-XI-1986, Valls & al. 10506 Cáceres,na estradapara Barra do Bugres,26-1-1985, (CEN, CTES); id.,Valls & al. 10507(CEN). Mun. Valls& al. 9394& 9395(CEN, CTES). Poconé:Várzea Comprida, 10-11-1978, Allem & al. 1663(US). Mato Grossodo Sul. BetweenCorumbá Common name, "amendoimbravo" (Allem and Cuiabá, V-1927, Smith 107 (K). Mun. & al. 1663). Aquidauana:Aquidauana, 27-11-1959, Gregoiy & al. 9824 (LIL); 25 kmW de Aquidauana,BR-262, 12- Geographie distribution. Arachis XII-1976,Krapovickas & al. 30017(CEN, CTES, G, Kuhlmannii grows in the northern and GH,K, MO, NY, P, SI, SP,US); 12km W do trevode southernedges of the PantanalMatogrossense, acessoa Aquidauana,BR-262, 7-IV-1986, Valls & al. in the statesof Mato Grosso and Mato Grosso 9912(CEN, CTES); id.,30-X-1986, Valls & al. 10405 do Sul (Brazil) and is one of the few species (CEN, CTES); Faz. RioNegro, 30-X-1978, Allem & collected in the very center of the Pantanal al. 2269 (CEN, CTES); 14,3km NE da Faz. Proteção (Pantanal da Nhecolandia). na estradaRio Negro-Aquidauana, 29-X-1985, Valls & al. 9470(CEN, CTES); 2,2km NE da Faz. Pontale Obs. 1. The specimen Krapovickas & al.

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30017 represents a population somewhat 104).Perù, rio Marañón. differentfrom the typeby havingleaves of the Mandues, Schmidel,1567. Derroteroy viaje a mainstema bit smallerand morelanceolate. In Españay las Indias(ed. 1947:54, 75, 80, 82). Para- this population, abundant in individuals,the guay. great majorityof the plants had flowerswith Tlalcacahoatl,Molina, 1571. Vocabulario en len- orange standards,but between 1 and 5% had gua castellanay mexicana,1: 64. Hernández,1615. totallyyellow flowers. Quatrolibros de la naturaleza(por Ximenez), ed. 1790,2: 159,ed 1946,3: 159.Sahagun ?, 1831. Histo- Obs. 2. From material of this species ria generalde las cosas de la Nueva España (MS, (Gregory& al. 9824) collectedin Aquidauana, 1575)ed. 1956,3: 173.México. Carranza & Lindquist(1962) describeda new Manobi,Lery, 1578. Histoire d'un voyage fait en fungus, Thecaphora Frezii Carr. & Lindq. la terredu Brésil(ed. 1960, 162). Brasil,Bahia de This basidiomycete attacks the fruits, Guanabara. deformingthem by eliminatingthe isthmus that Truffede l'Amérique,Dalechamp, 1587. Hist, separates the two articles, therebygiving the general,pl. 2: 791, chap. CXXIX. Tomado de affectedfruits an appearance similarto those Monardes(1565). (Chevalier 1933: 729). of Arachis hypogaea (loc. cit. fig. l,a). Amendoes,amendois, [Soares de Souza], 1589. Fortunately,experimental attempts to innoculate Noticiado Brasil.Soares de Souza, 1851. Tratado thecultivated peanut were notsuccessful. descritivodo Brasilem 1587 (ed. 1971: 184-185). We dedicate this species to João Geraldo Brasil,Bahia. Kuhlmann,who collected the plants in Mato Macara, Alvaresde Almada,1594. Tratado bre- Grosso as a member of the "Comissão ve dos rios de Guiné (segúnFicalho, 1884: 133, Rondon" or "Comissão de Linhas Telegráfi- Chevalier1933: 735). Archipiélagode Bissagos, cas, Estratégicasde Mato-Grosso ao Amazo- Africa. nas." Nucesquaspian, Clusius, 1601. Rar. pl. hist.,lib. IV: 79. Costade Guinea. 69. Arachis hypogaea L. Inchic,Inca Garcilaso de la Vega,1609. Comenta- riosReales (ed. 1943,2: 179).Perù. Linnaeus,C. Sp. pl.2: 741. 1753."Hort, cliff. 353. Chocopa,Bertonio, 1612. Vocabulario de la len- Hort.ups. 228. Roy. lugdb. 390 ...Habitatin Brasilia, guaaymara: 306 (reimpr. 1984). Bolivia. Peru." Mandues,carios populi edunt, Bauhin, G. 1623. Arachidnaquadrifolia [Plumier] Trew, PI. rar., Pinax,lib. III, sect.1: 90-91. tab.3, 1764. Manobi, Laet, 1630.Beschrijvinghe van West- Arachidnahypogaea (L.) Moench,Methodus, Indien510, firstillustration of thefruit (ed. 1633, 122.1794. NovusOrbis seu Descriptionis Indiae Occidentalis, Arachis hypogaea subsp. oleifera A. Chev., lib.XVm, cap. XI: 586).(Hammons 1973: 23, fig. 2). Rev.Int. Bot. Appl. Agrie. Trop. 13(146-147): 770, Guerte, Nieremberg,1635. Historianaturae, 1933."(~A. hypogaeasensu stricto) ." maximeperegrinae, lib. XVI, C. 103(Chevalier 1933: 728). Maní, Oviedo,1535. Historia General de las In- Arachus [HypogajosJ americanus, dias,lib. 7, cap. 5, folio75. "yslaespañola." undergroundcicheling of America, indian Lirén ?, Oviedo, 1535. HistoriaGeneral de las earthnuts, Parkinson, Jn. 1640. Theatrum Indias,lib. 7, cap. 12,folio 76. "yslaespañola y en botanicum,chap. XI: 1069-1070."...in mostplaces tierrafirme." of America...."Illustrates two open fruits,one of Mandibí,Nuñez Cabeza de Vaca, 1555.Naufra- themwith two seeds, next to a plantof Viciasativa giosy Comentarios(ed. 1942: 129, 162,201, 205, L. subsp. amphicarpa (Dorthes) Ascherson& 212). Oviedo,1851. Historia General de las Indias, Graebner(Hammons 1973: 25, fig. 3). lib.23, cap. 12.Paraguay. Mandubí, Marcgrave,1648. Historiaererum La frutaque se cria debaxode tierra,Monardes, naturaliumBrasiliae (ed. 1942: 37, fig.). First 1565.Historia medicinal de las cosasque se traende illustrationof the plant (Hammons 1973: 26, fig.4). nuestrasIndias Occidentales (ed. 1574, 3a. parte: Piso,1658. De Indiaeutriusque re naturali et medica,

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256-7,fig. (Hammons 1973: 27, fig. 5). Herbariumof Stockholm(S, 307-19) and that Fructus peruanas amygdaloides sub terra of the Clifford Herbarium (BM) are two nascens,Bauhin, J., 1650. Hist. pl. 1(3): 292,cap. 30. syntypesto take intoaccount fortypification. Cap. 3 1,Manobi Lerii. The specimen LINN, 909-1, "HU," that Cacaguate, Cobo, 1653. Historiadel Nuevo Krapovickas and Rigoni (1960: 214) Mundo:359, nombre mexicano. consideredas the type and Verdcourt(1971: Pistache,Dutertre, 1654. Histoiregénérale des 442) as a syntype,should not be discarded isles S. Christophe,de la Guadeloupe, de la because in Hortus Upsaliensis (1748) it is an Martinique,et autresdans l'Amerique(ed. 1667, element that appears in the protologue of Histoiregénérale des Antilles2: 121). Species Plantarum. Aracus [Hypogajos]Americanus, Ray, 1686. Nevertheless,it would be safer to choose Hist.pl.1:919. the specimen from the Clifford Herbarium Arachidna Indiae utriunque tetraphylla, (BM) as the lectotype, being the one that Hermann,1689. Paradisibatavi prodromus, 314. offersa greaterdegree of certaintyin termsof Sloane,1696. Cat. pl. Jamaica, 72. itsauthenticity and forbeing linked to theorigi- Senna tetraphyllas. Absi congenerhirsuta nal idea that Linnaeus had regarding this Maderaspatensisflore flavo siliquis punctatis species, given that Hortus Cliffortianus scabris,folículos sub terrantcondens, Plukenet, (1738) is the oldest referenceby Linnaeus that 1691.Phytographia, tab. LX, fig.2. Plukenet,1696. appears in the protologue. Almagestum,341-2. Madras (India) (BM!). The CliffordHerbarium specimen and that Pindals,Sloane, 1696. Cat. pl. Jamaica, 72. of theLinnean Societyare verysimilar and are Arachidnaquadri/olia, villosa, flore luteo, Plu- bothprepared in the same manner,with all the mier,1703. Nova pl. amer. 49, fig. 37. leaves well extended. They are lateral Lupinusquadrifolius exoticus, Barrelier, 1714. branches,distichal, with very similar, glabrous Plantae per Galiam, Hispaniam et Italiam leaves. In the firstof these specimens (BM), observatae,tab. 1215. the typeof ramificationcannot be seen; but in Arachidnoidesamericana, Nisolle, 1723. Mém. the second (LINN), two nodes with Acad.Sci. (Paris):387-392, fig. inflorescencesare followedby two nodes with Arachis,Linnaeus, 1735. Syst.nat., Diadephia shortvegetative branches and two nodes with Decandria,nomen. Linnaeus, 1737. Gen. pl. ed. 1, inflorescences.The specimen S, 307-19 has 592. Linnaeus,1737. Hort. Cliff. 353-4, Brasilia & leaves with the leaflets folded and an Perú(BM!). Linnaeus,1748. Hort, upsal. 1 : 228-9. alternatingramification pattern can be seen: Chamaebalanus japónica, Rumphius,1747. two inflorescences, one vegetative branch, Herb,amboin. 5: 426,tab. 156. two inflorescencesand an apical vegetative Earthor Groundnut,Miller, 1754. Gard. Diet abr. branch. In his Species Plantarum, Linnaeus ed.4, 116.Aírica. does not mentionthe numberof seeds, but he Cacahuete,Terreros, 1765-83. Diccionario Cas- does in his Genera Plantarum, ed. V (1754: tellano. 329) where he says that Arachis has fruits Pea-nut,Gray, Asa. 1856.The flowers of the pea- with two seeds. All of these characteristics nut (Arachis hypogaea L). Amer.Jour. Sci. and correspond to the Virginia-type peanut, Arts,2nd Ser. 22: 435-436. commonly cultivated in the eastern United States. Lectotype: In the Linnean herbaria, Arachis hypogaea is represented by three First reports of the peanut: The first specimens, very similar to one another.The publishedmention of the peanut thatwe know one at theLinnean Society (LINN, 901-1) only of is owed to Gonzalo Hernández de Oviedo y has the annotation"HU" (Hortus Upsaliensis) Valdês (1478-1557), in his "HistoriaGeneral y and could have been incorporatedafter 1753 Natural de las índias, Islas y Tierra Firme del because it lacks the no. "1" indicating its Mar Océano," thatappeared forthe firsttime position in the text of Species Plantarum in Seville in 1535. This chroniclerarrived in the (1753). The specimen of the Linnean Americasfor the first time in 15 14 and resided

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in those lands untilhis death,except fora few theytake themto sell cooked, and with their tripsthat he made to Spain. His account is the coveringsremoved they are verywhite inside principalresult of the experienceshe acquired and of good flavor.I have not seen in Spain nor during his travels and work in Tierra Firme anywherea fruitor flavorthat compares with (northernColombia and Venezuela), Panama, these lirenes. But nevertheless,this fruitis of Nicaragua and the Antilles. good flavor and there is plenty of it on this In "La Historia General de las Indias," islandof Hispaniola and on themainland and in which is thetitle that appears on the firstpage many parts of these Indies." of the princeps edition, Oviedo expressly Oviedo says the same in the second edition mentionsthe peanut in Book 7 "De la agricul- (Oviedo 1547). In the editionof José Amador tura," ChapterV (fol. LXXV), and writes: de los Rios (Oviedo 1851, 1: 279-280) he contextuallyrepeats this description of lirenes, "Of themaní [peanut]which is anotherfruit with the additional statementsthat "that vein and ordinarysustenance that the Indians have that attaches the liren is no thickerthan a on thisisland of Hispaniola. One fruitthat the commonneedle" and "and removingthat little Indians have on this island of Hispaniola is skincovering, which is thinnerand softerthan called mani, whichthey plant and harvestand the skin of a walnut, and inside remains the is a very ordinary plant for them in their whiteliren and it is of good flavor." gardens,and is sized like pinenutswith shells, Evidently,when Oviedo speaks of lirenes, and they hold them to be healthful,but the he is describing the plant of Arachis Christians take little comfortin them,being hypogaea. By virtue of his details that the eatenmostly by lowlymen and boys and slaves branches extend themselveslike those of the and by people who do notpardon their taste for ajes ( Dioscorea trífida L.f.) and batatas anything.It is of mediocreflavor and of little ( Ipomoea batatas (L.) Lam.), fromwhich are substance, but is very common with the suspended the fruits that are produced Indians,and is foundin greatquantities on this underground,there remains almost no doubt island and others." regardingthis supposition, given the unique Except for the name and the fact that the fruitingcharacteristics of Arachis hypogaea. maní is "sized like pinenutswith shells," it is Moreover, by this descriptionwe are able to difficultfor us to recognize this plant as the recognizea runner-typepeanut. peanut given that Oviedo adds that "it is of It is currentlyaccepted thatleren or lairen mediocreflavor and of littlesubstance." is a name applied to Calathea allouia (Aubl.) On theother hand, it is quite likelythat when Lindl. (Marantaceae), an erect monocot with discussing lirenes, Oviedo provides a good edible starchytubers (León 1987: 106). descriptionof Arachis hypogaea. In the same . Bartolomé de Las Casas (1474-1566), a Book 7, ChapterXII (folio LXXVI), it reads: contemporaryof Oviedo, mentionsmaní (Las "Of the plant and fruit called Lirenes. Casas 1909: 29), whose fruitcompares with Lirenes is a fruitthat is borne on a plant that that of broad beans, field peas and chickpeas the Indians cultivate and even today the and the seeds with shelled hazelnuts. In this Christians on this island grow them in their work of Las Casas, which went unpublished fieldsand gardensand inheritedlands. This is until 1909, maní can be recognizedas Arachis an herbthat extends itself and lays itsbranches hypogaea and it was in thisform that its name on the ground and theysow it fromthe plant was disseminated throughout Spanish itselfas I have been told is done withthe ajes America, replacing many local vernacular or batatas and under the ground they bear names. theirfruit which is whiteand of thesize of thick The first illustration of the fruit was dates and some larger and smaller also of published by Laet (1630) and reproducedby whitecolor, and each fruitof these is attached Hammons(1973: 23, fig.2). It depictsa "Virgi- or tiedby a thinstem with which it is connected nia" type peanut, that we can identifyas to the branch. The Indians cook these and Arachis hypogaea L. var. hypogaea, for these days the marketsabound withthem and whichLaet does not indicatethe place of origin

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of theillustrated material. villosa which produced a sterile hybrid Hammons (1973: 22, fig. 1) reproducesa (Krapovickas & Rigoni 1957: 438). figurefrom Clusius (1605) as a possible first Then, Krapovickas & Rigoni (1954, 1957) illustrationof the seeds of Arachis hypogaea, successfully crossed A. hypogaea with the butthis interpretation, which Sloane (1696: 72) new species A. montícola (2n=40), obtaining indicatedas doubtful,is not precise since the fertile offspring and demonstrating A. markedveins and the pronouncedthread more montícolato be the wild species most closely closely resemble allspice ( Myristicafragrans relatedto the cultivatedpeanut. Houtt.). Thereafter,numerous crosses were made The firstillustration of thepeanut plant was involvingmore species, all fromthe section published by Marcgrave (1648, 1: 37), Arachis (Gregory, M.P. & W.C. Gregory reproducedby Hammons (1973: 26, fig. 4). 1979, Smartt1990). Both the descriptionas well as the illustration It is possible that A. montícola is a wild permitthe identificationof the "mandubí" of ancestor of A. hypogaea, but the alternative Marcgrave as the peanut called "crema" in possibilitythat it is a derivativeor feralform of Bolivia and "guaycurú"or "guanaco" in Para- the cultivatedpeanut should not be discarded. guay and in NE Argentina, and whose A mechanismof reversionto thewild condition cultivationwe knowfrom Bolivia, northeastern is outlinedby Krapovickas & al. (1974), when Argentina, Paraguay and, Brazil. This describing A. Xbatizogaea Krapov. & Av. "mundubi" of Marcgrave, therefore,would Fernández,as the resultof the recuperationof belong to the var. hypogaea. fertilityin a sterileinterspecific hybrid, with the participationof A. Batizocoi and A. hypogaea. Origin of the cultivated peanut: Arachis Arachis hypogaea is an amphidiploid hypogaea has 2n=40 chromosomes whose ancestors have still not been (Kawakami 1930 and Husted 1931) and its conclusivelydetermined. Various species have polyploidcondition was establishedby Husted been proposed as possible ancestors of the (1936) in a classic work on themorphology of cultivatedpeanut. Gregory& Gregory(1976, its chromosomes. 1979) expressed the possible participationof Numerous attempts at interspecific species belongingonly to the sectionArachis, hybridization were made to establish the proposingA. Cardenasii and A. Batizocoi as relationships between A. hypogaea and the the probable diploid progenitors of the wild species of the genus Arachis. The first cultivatedpeanut (Smartt & al. 1978). Arachis such cross was made by Gregory(1946, fig. duranensis and A. ipaënsis have even better 29) using the cv. 'N.C. Bunch' x Arachis possibilities(Fernández & Krapovickas 1994), glabrata, "polyploid," obtaining only but the participationof morethan two species undeveloped seeds. Later, the cross of A. shouldnot be discarded,taking into account the hypogaea with A. Archeri was attempted differencesbetween the two subspecies of A. (Smith 1956 and Johansen& Smith 1956, sub hypogaea. A. Diogoi) in which,although fertilization had The rangeof the diploid species involvedin begun, there were failures in the embryo's the originof the cultivatedpeanut is restricted formation. to NW Argentinaand SE Bolivia. The first hybrid (Krapovickas & Rigoni The characters of these wild species that 1949, 1951) was achieved by crossing A. we can findin A. hypogaea are: runnerhabit, hypogaea var. fastigiata (2n=40) with A. small fruits with marked constriction and correntina(2n=20), obtaininga sterileF, with evident but subdued reticulation,with two 30 chromosomesand irregularmeiosis. Later, seeds, and these demonstratingdormancy. these authorsattempted to cross A. hypogaea This suite of characters, which we can with the diploids A. duranensis (sub A. considerprimitive, is foundonly in the subsp. pusilla), A. villosa, A. villosulicarpa and the hypogaea (Simpson & al. 1986). tetraploid A. glabrata var. Hagenbeckii, In South America,the subsp. hypogaea has obtaining a satisfactoryresult only-with A. itsmost important center of variationin Bolivia

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and it is in SE Bolivia, on the firstfoothills of Anotherdifference between the subspecies is the Andes, in the departmentsof Tarija and the presence of flowers on the mainstem in Chuquisaca, where we collected samples of subsp. fastigiata and theirabsence in subsp. cultivatedpeanut withthe greatestamount of hypogaea , also being a very constant those charactersthat we considerprimitive. In character,but which does not apply to the this area are found only the annual diploid "Cruceño" peanutwhich pertains, evidently, to species A. duranensis, A. ipaënsis and A. subsp. fastigiata and is grown in NE Bolivia Batizocoi. On the otherhand, A. Cardenasii, fromSanta Cruz northward. also a diploid,is a perennialand lives furtherto The facts suggest that subsp. hypogaea the east, in the Chiquitana hills. The other had its origin in SE Bolivia and that subsp. species involved,A. montícola,is an annual, fastigiata had differentiateditself furtherto tetraploid,and occurs in a very small area the north,possibly in Peru, where it shows its between1 0 and 20 kmNW of thecity of Jujuy, greatest variabilitywith the presence of the where peanut is not presently cultivated, vars. fastigiata , peruviana and althoughthere are historicand archeological aequatoriana, withoutdiscarding the possible evidences of its pre-Columbianexistence (Sa- participationof some otherwild species. The las 1945). antiquityof peanut cultivationin the Andean A notable difference between subsp. region is manifestedin the diverse ways it is hypogaea and subsp.fastigiata is the color of used. In Bolivia it is a commonelement of the the leaves, dark green in the formerand light daily diet, being boiled in soups, chopped, green in the latter. It is a very constant mixed in cornbreadsor cakes (Rusby 1901) character, except for a few accessions of and used to produce a fermentedbeverage, subsp. hypogaea from south-central Peru. "chicha de maní" (Weddell 1853).

Key for distinguishing the taxa of A. hypogaea

A. Mainstemwithout flowers and n+1 brancheswhich alternate regularly: two vegetativeand two reproductive(alternate branching). 69a. subsp.hypogaea B. Leafletswith underside glabrous or with a fewhairs on themidvein. 1. var.hypogaea B'. Leafletswith hairs 1-2 mm long on theunderside, dispersed across entire surface. 2. var.hirsuta A'. Mainstemwith flowers and lateral branches on whichthe reproductive and vegetative branches present themselvesin no specificorder (sequential branching). 69b. subsp.fastigiata C. Fruitswith more than two seeds. Extended fruiting. D. Leafletswith the underside glabrous or with hairs only on themidvein. E. Fruitswith subtle or slightlymarked reticulation, without pronounced longitudinal ribs. Reproductivebranches usually short and slender. 1. var.fastigiata E'. Fruitsalways with sharply marked reticulation and withoutstanding longitudinal ribs. Reproductivebranches long, 5-10 cm, robust,both on the mainstemand on the lateral branches. 2. var.peruviana

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D'. Leafletswith hairs on theunderside 1-2 mm long, dispersed across entire surface. Reproductive brancheslong, primarily on thelateral branches. Mainstem commonly with inflorescences or shortreproductive branches. 3. var.aequatoriana C. Fruitsusually with two seeds. Fructification clustered around the base ofthe plant. Frequently with compoundspikes. 4. var.vulgaris

69a(l). A. hypogaea subsp. hypogaea var. A. hypogaeasubsp. africana Bois, 1927.Plantes hypogaea aliment.1: 96. 1927. Based on A. africanaLour., illegitimatename. A. Lour.,Fl. Cochinch.2: 430. 1790. A. hypogaeasubsp. africana var. communisA. africana " "Habitatin variis locis Africaeorientalis, non Chev.,Rev. Int. Bot. Appl. Agrie. Trop. 9(91): 193. Arachis africana Burm. F. 1768 = Voandzeia 1929. Lectotype:"arachide du Cayor" (subvar. subterranean (L.) Thouars. According to pallida). Adam, L'Arachide, 32, fig. 13. 1908. Loureiro'sdescription, it is a runner-typepeanut, Bouffil,Biologie de l'arachide,54, fig. 7, 1. 1947. withglabrous leaflets and 2-3-seededfruits. A. hypogaeasubsp. africana var. robustiorA. A. hypogaeavta. glabra DC. Prodr.2: 474. 1825. Chev.,I.e., 193. 1929."Dotiga (bambara de Ségou), A. africanaLour, is mentionedas a synonym. Lotiga ou Loséno (bambarade Haute-Guinée)." A. hypogaeavar. aegyptiacaHassk., Retzia 1: Adam,L'Arachide, 34. 1908. 190. 1855. "In hortobogoriensi coluntur specima A. hypogaeasubsp. africanavar. microcarpa ex Aegyptointroducta ." A prostrateform that A. Chev.,I.e., 193. 1929.Type: "Fila tigaou Dion Hasskarlsupposes to be perennial,with fruits 1-2 gouassidu Soudan."Adam, L'Arachide, 35. 1908. (rarely3) -seeded. A. hypogaeasubsp. asiatica var. macrocarpa A. hypogaea var. indica Kurtz, Verh. Bot. A. Chev.,I.e., 194. 1929. "Tigakoumba,Sokoba, VereinsProv. Bradenburg: 45. 1875.Latin name for Sokoba tiga, Tiga ba (en bambara),Sama tiga the"Arachide de l'Inde"that Cordemoy (1886: 249) (Arachidede 1'Elephant),Faréba (en Kassonké). describesas prostrate. Les fruitset les grainspar leurtaille rappellent la A. guaranianaBertoni, Catálogo espec. o var. var.robustior, mais la planteest dressée." In 1933, plant,cult. Estac. Agron. Pto. Bertoni: 6. 1912.The Chevalierincludes the North American "Virginia descriptionis verybrief, and may referto the Bunch" and "VirginiaRunner" peanuts in this "guaycurú"peanut accordingto the specimen variety.Adam, L'Arachide, 33-34, fig. 18. 1908. "PuertoBertoni, Alto Paraná, leg. T. Rojas 8157, A. rasteiroA. Chev.,Rev. Int.Bot. Appl. Agrie. junio 1938"(SI). Trop.9: 487. 1929.Chevalier creates a Latinname for A. hypogaea subsp. procumbens Waldron, theplant whose fruits he describesand illustrates Contr.Bot. Lab. MorrisArbor. Univ. Pennsylvania inthesame volume [9(91): 196-197,pl. 5,no.l andpl. 4(2): 3 12. 1919.Includes the cultivare "North Caroli- 6, no.3], Native to Santa Catharina, Brazil, its fruits na," "VirginiaRunner," "Virginia Bunch" and have 3 and up to 4 seeds witha bi-coloredseed "Jumbo." coat,and itis similarto the "Nambyquarae" peanut A. hypogaeavar. africanaGiróla, Cult. Mani butwith somewhat smaller fruits: 4-5.5 cm long and Argentina,17. 1922. Based on A. africanaLour, 15-18mm in diameter. (illegitimatename). Non Kurtz. A. hypogaea subsp. rasteiro (A. Chev.) A. A. nambyquaraeHoehne, Com. lin.telegr., Bot. Chev.,Rev. Int. Bot. Appl. Agrie. Trop. 13(146-147): Annexo 5, 12: 21, táb. 190. 1922. "N° 2052, 772. 1933.Based on A rasteiroA. Chev. Kuhlmann, Pimenta Bueno, Rondonia, Matto A. hypogaeasubsp. nambyquarae (Hoehne) A. Grosso,em Abrilde 1919 (Cultivadade sementes Chev.,Rev. Int. Bot. Appl. Agrie. Trop. 13(146-147): trazidasdesta localidade, onde a plantaé cultivada 77. 1933.Based on/1,nambyquarae Hoehne. pelos indiosNambyquaras, na malocado cacique A. hypogaea var. nambyquarae (Hoehne) Abaitora)"(R!). Buikart,Darwiniana 3(2): 281. 1939.

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A. hypogaea formamacrocarpa (A. Chev.) ded, sinuous, fromprocumbent to decumbent. Hoehne,Flora Brasílica 25(2)122: 18. 1940. On the principal lateral branches (n+1), two A. hypogaea formamicrocarpa (A. Chev.) vegetative branches alternate regularlywith Hoehne,I.e., 19. 1940. two reproductivebranches. Leaves medium- A. hypogaeaforma typica Hoehne, I.e. : 18. 1940. sized; leafletswith the upper surfaceglabrous A. glabrataBenth. subsp. rasteiro (A. Chev.)A. and withhairs 1-2 mm long spaced across the Chev.,Rev. Int.Bot. Appl. Agrie.Trop. 28: 515. underside.Reproductive spikes simple, 1-2 cm 1948. long. Fruitswith up to three,rarely four seeds; A. hypogaeaforma communis (A. Chev.) F.J. pericarp strongly reticulate, with uniform Herm.,Synop. Arachis, 14. 1954. reticulation. A. hypogaeaforma nambyquarae (Hoehne) F.J. Herm.,I.e. 14.1954 Geographic distribution. This peanut is frequentlyencountered in the archaeological Herbaceous annual, usually late-fruiting. deposits fromthe coast of Peru. In modern Mainstem erect,without inflorescences. Late- timesit is stillcultivated in theAmericas, on the ral branches usually procumbent,sometimes coast of Peru and in centralMexico, as well as decumbent. On the basal lateral branches in thebasins of the Pacific and Indian Oceans. (n+1), two vegetative branches alternate This is the peanut that Plukenet (1691) regularly with two reproductive branches. indicated as coming from Madras, in Medium-sizedleaves; leafletsglabrous on both southeasternIndia, and thatRumphius (1747) sides, or with some hairs on the underside called Chamaebalanus j aponi cus. It is midvein. Reproductive spikes simple, short, knownfrom the Philippines(Blanco 1878, 2: rarelyup to 5 cm long. Fruitstypically with 2 362), Java (Dubard 1906, fig. 2), China (3) seeds and in some Bolivian landracesup to (Dubard 1906, fig. 3; Skvortzow 1920: 144), 4 seeds; pericarpmoderately reticulate. Madagascar (Dubard 1906, figs. 4 & 5), and fromthe burials of Ancón, in Peru (Dubard 69a(2). A. hypogaea subsp. hypogaea var. 1906, fig. 1). hirsuta Köhler 69b. A. hypogaea subsp. fastigiata Köhler,Med.- Pfl. 3, 42. 1898. Based on A. Waldron asiatica Lour,(the illustration does notcorrespond toLoureiro's entity). Waldron,Contr. Bot. Lab. MorrisArbor. Univ. A. asiatica Lour., Fl. cochinch.2: 430. 1790. 4: 312. 1919. cvar.'Valen- " Pennsylvania Lectotype: Habitatculta abundantissime in Cochinchina,& cia' (Krapovickas& Rigoni1960: 225). in China." (BM!). Details of the such A. var. Verh. Bot. " description hypogaea africanaKurtz, as caulis ... pilosissimus, suberectus,ramis VereinsProv. Brandenburg 17: 45. 1875.Latín name diffusisprocumbentibus ... foliolis pilosis ... forthe "Arachide of " d'Afrique" Cordemoy(1866: folliculus3-4-spermus permit the identification of 249). thistaxon. A. hypogaeavar. asiatica (Lour.) Girólaforma A. procumbensBerneaud, in Guerin,Diet. Hist. rosadaGiróla, Cult. Maní Argent., 17. 1921. Nat.1:248. 1834. A. hypogaeaL. var.asiatica (Lour.) Giróla, Cult. 69b(l). A. hypogaea subsp. fastigiata var. ManiArgent. 17. 1922. fastigiata A. hypogaea L. subsp. asiatica (Lour.) Bois, Plantesaliment. 1: 96. 1927. Based on A. asiatica A. hypogaea var. asiatica (Lour.) Giróla Lour. formaoscura Giróla, Cult. Maní Argent. 17, photo. 23. 1921. Erect plant with 3-4-seeded Herbaceous annual, very late-fruiting, fruits. much-branchedand of greatsize. Mainstemis A. hypogaea var. communis A. Chev. erect to prostrate,up to 1 m long, sinuous, subvar. violacea Burkart, Darwiniana 3(2): without inflorescences.Branches are exten- 279. 1939. "Maní negrode Corrientesy Misio-

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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 nes," que "tienegeneralmente 2-4 semillas por SP,US. Placeof origin: Peru, Estación Experimental vaina." Agropecuaria"La Molina,"collection L.D. Tripp (USDA PI 393531). Herbaceous annual,early-fruiting. Mainstem usually little-branched and with some Herbaceous annual. Mainstem with inflorescences.Lateral branchesdecumbent to vegetative branches toward the base and semi-erect, with irregular distribution of reproductivebranches toward the apex. Late- vegetativeand reproductivebranches (sequential ral branches decumbent, with very few or branching).Leaves medium to large, leaflets without vegetative branches. Reproductive with both surfaces glabrous, but may present branches 5-10 cm long, thick,multiflowered, some hairs on the undersidemidvein. Short, sometimes with leaves toward the apex, axillaryinflorescences. Fruits usually withup abundant on the mainstemas well as on the to fourseeds; pericarpmoderately reticulate. lateral branches. Leaves large, somewhat thick. Leaflets glabrous on both sides, with Geographicdistribution. This peanuthas its hairs only on the marginand on the underside mostimportant center of variationin Paraguay midvein.Fruit usually with three seeds, in some and is the most widespread varietyin all of landraces up to four seeds; pericarp strongly South America. Its diffusionoutside South reticulateand with outstandinglongitudinal America must be relatively recent. It only ribs. begins to appear in the literatureat the end of the 19th century.The erect habit and fruits Geographic distribution. The variety with four or more seeds permit its peruviana is grown in almost all of Peru, identification.Handy (1896: 7) mentionsthe especially in the basin of the Marañón River, "Georgia red nut" and the "Tennessee" as and it is commonin Ecuador. Its southernlimit being erect,with 3-4 seeds, and indicatesthat is found in northernBolivia, where a few thereis a peanutgrown in Costa Rica with4 to samples were discovered in Rurrenabaque on 5 seeds. It is illustratedby Dubard (1906, fig. theBeni Riverand in thedepartment of Pando. 6). Beattie (1909: 28, fig. 16, D) illustratesthe A few samples were also obtained in Acre fruitof "Tennessee Red," with 4 seeds, with statein Brazil. an undulatingdorsal surface,with a beak and reticulate pericarp. The name "Valencia" Obs. 1. It is possiblethat it is "thefruit that is appears forthe first time in 19 1 1, whenBeattie grownunderground" of Monardes (1565), that mentionsit as a new introductionfrom Spain, is "all twisted,with a very fine figure ... it similarto, but betterthan, "Tennessee Red." grows underground,on the shores of the Ma- rañón River." 69b(2). A. hypogaea subsp. fastigiata var. peruviana Krapov. & W.C. Gregory nov. Obs. 2. In the var. peruviana there are var. numerouslandraces that vary accordingto the shape of the fruitand the color of the seed. Rami erecti aut decumbentesramißcatione The most frequentcolors are black or violet, sequentiali. Folia subtus glabra, margine and cream, but there are also variegated or nervoquemedio tantum pilos plus minusvelongos streakedseeds, as in the case of "Tingo Ma- gerentia. Fructus 3-4-seminatus.Pericarpium ria," thatare exclusive to var. peruviana. perspicue reticulatumnervis longitudinalibus valde prominentibus. 69b(3). A. hypogaea subsp. fastigiata var. aequatoriana Krapov. & W.C. Gregory Holotype:ARGENTINA. Córdoba.Manfredi, nov. var. EstaciónExperimental Agropecuaria, INTA, crop 86/87:2752, "TingoMaría," granos veteados, 23- Rami erecti aut decumbentesramißcatione III-1987,leg. Krapovickas, Vanni & Williams41725 sequentiali.Folia subtuspilis ca. 2 mm longis (CIES). Isotypes:BAB, CEN, LIL, LPB, MO, NY, SI, vestita. Fructus 3-4-seminatus.Pericarpium

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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. Rrapovickasand W.C. Gregory, Arachis (Leguminosae) perspicue reticulatumnervis longitudinalibus Herbaceous annual. Mainstemerect, highly valde prominentibus. branched, with some inflorescences.Lateral branches decumbent to erect, with short Holotype:ARGENTINA. Cordoba.Manfredi, inflorescences, 1-2 cm long, simple or EstaciónExperimental Agropecuaria, INTA, cult. compound, grouped around the basal nodes. 77/78:1403, "Zanima," 22-111-1978, leg. Krapovickas Medium-sized leaves. Leaflets with both 33756(CIES). Isotypes:BAB, CEN,LIL, LPB,MO, surfaces glabrous, with long hairs on the NY, SI, SP, US. Place oforigin: Ecuador, El Oro, marginand also a few hairs on the underside Zaruma,leg. Polobio A. Romero,1948. midvein. Fructificationclustered around the base of the plant. Fruits small, usually two- Herbaceous annual. Mainstem erect, more seeded; pericarpmoderately reticulate. or less branched,with reproductivebranches usually short, less than 5 cm long. Lateral Geographic distribution. The variety branches decumbent, with reproductive vulgaris, which comprises the "Spanish" branches up to 20 cm long. Large leaves. peanuts, is grown in South America in Uru- Leaflets with upper surface glabrous and guay, in Argentina(Santa Fe, Entre Ríos and underside with hairs 1-2 mm long dispersed Corrientes),in southernBrazil and, to some across entire surface. Fruit 3-4-seeded; extent,in Paraguay. The color of the seed is pericarp very reticulate, with outstanding commonly cream, such as in the typical longitudinalribs. "Spanish" peanuts,but can be black and very rarely red. The peanut with cream-colored Geographic distribution. This variety is seeds passed to southernEurope at the end of almost exclusively fromEcuador where it is the 18th Century (Gilii e Xuarez 1789, Tabares cultivatedprimarily in theprovinces of El Oro de Ulloa 1799). Accordingto McClenny(1935: and Loja. It is cultivated sporadically in 9), "Spanish" peanuts were introducedto Vir- northernPeru. ginia,USA, in 1871 fromMalaga, Spain.

Obs. Two landraces are known in the var. aequatoriana: "Zaruma" with fruitssome 4 Taxa of dubious position cm long, and "Huasquillo" with fruits frequently7 cm long and thinner.The seed color is commonlyviolet, but may be cream or Arachisguaranitica Bertoni, Anales Ci. Parag. more rarelyred. ser.2(5): 331. 1919,nomen nudum non Chodat& Hassler1904. 69b(4). A. hypogaea subsp. fastigiata var. Arachis hypogaea subsp. africana var. vulgaris C. Harz stenocarpaA. Chev.,Rev. Int. Bot. Appl. Agrie. Trop.9(91): 193.1929. "Nzayan, Tigadian (bambara, Harz,Landw. Samenk., 642. 1885.Latin name for ...). Maka diangon(Kassanké)." A runnerpeanut thevar. "comune" of Blanco (1850: 525, figs. 1, 2 & 3). with3-4-seeded fruits that Chevalier identifies with ArachidnaquadrifoliaTrew, PI. rar.,tab. 3. 1763. the"Peruviano" type of Dubardand includesthe Glycinesubterranea auct. non L., Giliie Xuarez, "TenneseeRed" that, for its growth habit, would be Observazionifitologiche. 30-3 1, tab. 4. 1789. a transitionto thesubsp. asiatica. Arachis americana Tenore, Atti Real Ist. Incoragg.Sci. Nat. Napoli 1: 48-50.1811. A. hypogaea var. reticulata Harz, Landw. Acknowledgements Samenk.642. 1885.Latin name for the var. "Rochet" ofBlanco (1850: 525,figs. 6, 7, 8, 9 & 10). A. hypogaea subsp. asiatica var. erecta A. We wish to express our thanks to the Chev.,Rev. Int. Bot. Appl. Agrie. Trop. 9(91): 194. followinginstitutions: ARGENTINA: Instituto 1929. Lectotype:"Guerté volute (de Gandióleau Nacional de Tecnología Agropecuaria Sénégal)."Adam, L'Arachide, 32, fig. 16. 1908. (INTA); Estación ExperimentalAgropecuaria

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(INTA) Manfredi,Cordoba; InstitutoMiguel Nacionaldo Brasil,IBGE - ConselhoNacional de Lillo,Universidad Nacional de Tucumán;Conse- Geografìa. jo Nacional de InvestigacionesCientíficas y Téc- BANKS,D.J. 1976.Peanuts: Germplasm Resources. nicas (CONICET); Facultad de Ciencias Agra- CropScience 16: 499-502. rias,Universidad Nacional del Nordeste, Corrien- . 1984. The Collectionof Arachishypogaea tes; SecretaríaGeneral de Ciencia y Técnica, Germplasm,Technical Report to the IBPGR. Ecua- dor- 1983.MS. 24 UniversidadNacional del Nordeste; Comisión págs. Administradoradel Fondo de Promociónde la . 1985. The Collectionof Arachishypogaea - TecnologíaAgropecuaria, CAFPTA, BuenosAi- Germplasm.Technical Report to the IBPGR. Perù 1985.MS. 33 res.BRAZIL: Anderson-ClaytonCompany Ltd., págs. São Paulo; Centro Nacional de Recursos . 1990.Peanut (Arachis hypogaea L.) collectingin Genéticos e Biotecnología, CENARGEN- coastalPeru. PI. Genet. Resources Newsl. 81/82: 45. EMBRAPA, Brasilia; Empresa Brasileira de BEATTIE, W.R. 1909. Peanuts.USDA Farmer's Pesquisa Agropecuaria,EMBRAPA, Brasilia; Bulletin356, 40 págs.

InstitutoAgronómico, Campinas, São Paulo. . 1911.The peanut. USDA Farmer'sBulletin 431, U.S. A: Plant IntroductionService & Plant 39 págs. ScienceResearch Division, Beltsville, Maryland; BENTHAM,G. 1841.On thestructure and affinities of National Science Foundation,Washington, DC; Arachisand Voandzeia.Trans. Linn. Soc. London 155-lo2. NorthCarolina Agricultural Experiment Station, 18(2):

Raleigh; North Carolina State University, . 1859. Leguminosaeen MartiusC., Flora Raleigh; John Simon GuggenheimMemorial brasiliensis.15(1): 86-87.

Foundation, New York; Texas Agricultural . 1865.Stylosantheae enBentham, G. & J.D.Hooker, ExperimentStation, Texas A&M University. Gen.PI. 1:449. ITALY: InternationalBoard for Plant Genetic BIANCHI-HALL,C M., R.D. KEYS, H.T. STALKER Resources, IBPGR, FAO, Rome. INDIA: & J.P.MURPHY. 1993. Diversity of seed storage in wild Arachis InternationalCrops Research Institutefor the proteinpatterns peanut ( , Fabaceae) PI. Evol.186: 1-15. Semi-Arid Tropics, ICRISAT, Patancheru, species. Syst. AndhraPradesh. BLANCO,Antonio. 1850. Remarques sur la pistachede We also owe our thanksto Drs. Margaret terre.Bull. Acad. Roy. Sci. Belgique 17: 524-528 y unalámina con 10 PflugeGregory and CarmenL. Cristobalfor their figs. support and critical contributions to the BLANCO,Manuel. 1878. Flora de Filipinas, vol.2. developmentof the classificationsystem and to BONDAR, G. 1929. Boletín.do Laboratoriode thecomposition and editingof the work. PathologiaVegetal do Estado da Bahia. We express our thanks to our traveling BRUMMITT,R.K. & C.E. POWELL.1992. Authors of companions, Agr. J.R. Pietrarelli,Dr. C.E. PlantsNames. Royal Botanic Gardens, Kew. Simpson and Dr. J.F M. Vails for their BUNTING,A H. 1955.A classificationof cultivated participationin theplanning and executionof the groundnuts.Empire J.Exp. Agrie. 23(91-92): 158-1 70. expeditions,and especiallyto the last two with BURKART,A. 1939.Estudios sistemáticos sobre las whomwe discussedtaxonomie problems, in the Leguminosas-Hedisareasde la República Argentina Darwiniana 117-302. wild as well as in experimentalplots, for which yregiones adyacentes. 3(2):

they share in the authorshipof some of the . 1942.Los frutosde las especiessilvestres de species. Arachis.Proc. 8th Amer. Sc. Congr.3: 175-178. Thanks to Mr. V. Maruñak who was CABRERA,A L. 1976.Territorios fitogeográficos de la responsiblefor the illustrations and is theauthor RepúblicaArgentina. En[Parodi, L.R.] Enciclopedia de 2o of thedrawings. ArgentinaAgricultura yJardinería, 2(1): 1-84, ed.,ampliada yactualizada bajo la direcciónde W.F. Kugler. Literature Cited CARRANZA,J.M. & J.C. LINDQUIST. 1962. ThecaphoraFrezii n.sp parásitade Arachissp.. Bol.Soc. Argent. Bot. 10(1): 11-18, fig.l. ANDRADE-LIMA,D. de. 1966.Vegetação, en Atlas CASTELLANOS,A. 1945.Las exploracionesbotánicas

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en el época de la Independencia1810-1853. GREGORY,M.P. & W.C. GREGORY.1979. Exotic Holmbergia4(8):3-14. germplasmofArachis L.: interspecifichybrids. J. CHEVALIER,A. 1929 a. L'originebotanique et Hered.70:185-193. l'améliorationdes Arachides cultivées. Essai d'une [Gregory,W.C.]. 1946. Peanutsbreeding program classificationsystématique. Rev. Int.Bot. Appl. underway.Res. & Farming5. Prog.Rep. 4: 41-45. 5-8. Agrie.Trop. 9(90): 97-102, 190-197, pl. , B.W. SMITH & J.A. YARBROUGH.1951. . 1929b. Surune forme ancestrale de l'Arachide Morphology,genetics and breeding, pp. 28-88en cultivée.Compt. Rend. Hebd. Séances Acad. Sci. ThePeanut, the Unpredictable Legume. Nat. Fert. 188,p. 1511. Ass.,Washington. . 1929c. Nouveauxdocuments sur les Arachides. , & M P. GREGORY.1967. Induced mutation and Rev.Int. Bot. Appl. Agrie. Trop. 9(6): 485-496. specieshybridization inthe de-speciation oÍ Arachis . 1933.Monographie de l'Arachide.Rev. Int. Bot. L. Cienciae Cultura19(1): 166-174. Appl.Agrie. Trop. 13(146-147): 689-789. MP. GREGORY,A. KRAPOVICKAS,B.W. CHODATR.&E. HASSLER1898. Plantae Hasslerianae SMITH & JA. YARBROUGH.1973. Structures 1:33. and geneticresources of Peanuts,en C.T.Wilson . 1904.Plantae Hasslerianae 2: 448-450. (ed.) Peanuts- Cultureand Uses. Am. Peanut Res. andEduc. OK. 3. 47- CLUSIUS,C. 1605.Caroli Clusii arrbatis, exoticorum Assoc.,Stillwater, Chapter libridecim: descributur. Ex officinaPlantiniana 134. Lib.ïï:57 Ralphelengii, (cap.29, fig. V). & M P. GREGORY.1976. Groundnut, pp. 151- COBO, B. 1653.Historia del NuevoMundo... Republ., 154.In N.W. Simmonds(Ed.), Evolutionof crop Ed.Jimenez de la Espada,4 vols.1 890-95. V. 1 :359- plants.Longman Group Ltd. London. 12"Del Maní". 360,cap. . 1977.The collection of Peanut Germplasm 1976- CONAGIN,C.H.T.M. 1959. Desenvolvimento dos fru- 1977.Technical Report to theInternational Board tosnas especiesselvagens de amendoim(Arachis forPlant Genetic Resources. Phase I (The Gran spp.).Bragantia 18(5): 51-70. Pantanal,MT, Brazil,December 1976). Phase D and . 1962. Espéciesselvagens do generoArachis. (NorthwesternArgentina Bolivia,March-May PhaseHI and June- Observaçõessobre os da da 1977). (Paraguay Brazil, July exemplarescoleçao seção MS. 118 deCitologia. Bragantia 21 (21): 341-374,est. 1-11. 1977). págs. . 1963.Número de Cromossomiosdas espécies & C.E. SIMPSON.1978. The colletion of Peanut selvagensde Arachis. Bragantia 22(11): 125-129. Germplasm1977-78. Phase IV. Consolidation, . 1964. Númerode cromossomosem Arachis reproductionand distribution ofearlier collections selvagem.Bragantia 23: XXV-XXVn (nota 5). of peanutgermplasm. Technical Report to the InternationalBoard CORDEMOY,C.J. de. 1866.Note sur l'Arachide de forPlant Genetic Resources, (Ule MS. 30 la Réunion,mars 1866). Adansonia 6: 249-253. págs. DONATO,H. 1985. Achegas para a historiade Botucatu, . 1979. IBPGR Technical Report, Peanut Prefeiturade Botucatu, Botucatu. Germplasm.September 1, 1978- August31, 1979. DUBARD,M. 1906.De l'originede l'Arachide.Mus. MS. 25 págs. Nat.d'His. Naturelle, Paris. Bul. 5: 340-344. , A. KRAPOVICKAS& M P. GREGORY.1980. EITEN, G. 1975. The vegetationof the Serrado Structure,variation, evolution and classification in Roncador.Biotropica 7(2): 1 12-1 35. Arachis.en R. J. Summerfield & A.H. Bunting (ed.). FERNÁNDEZ, A. & A. KRAPOVICKAS. 1994. Advancesin legume science, Kew, London. 2: 469- Cromosomas y evolución en Arachis 481. 8: 187-220. (Leguminosae ). Bonplandia HAMMONS,R.O. 1973.Early history and origin of the GARDNER,G. 1842. Tab. 500. Arachismarginata peanut,en C.T. Wilson(ed.) Peanuts- Culture and Gardner.En Hooker, W.J. Icones Plantaram 5. Uses.Am. Peanut Res. and Educ. Assoc. Stillwater, GILD, F.L. & G. XUÁREZ. 1789. Observaciones OK. Chapter2: 17-45. Fitológicas,en G. Furlong,Gaspar Juárez, S.J. y sus HANDRO,O. 1958.Espécies novas de Arachis L. Arq. NoticiasFitológicas. Buenos Aires. 1954. Bot.Estado São Paulo3. 177-181,táb. 42-47. GRAY,A. 1856.The flowersof thepea-nut (Arachis HANDY,RB. 1896,Peanuts: culture and Uses. USDA hypogaeaL ). Amer.Jour. Sci. and Arts, 2nd. Ser. 22: Farmer'sBulletin 25: 3-23. 435-436. HARTLEY,W. 1949. Plant collecting expedition tosub-

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tropicalSouth America, 1947-48. Div. Pl. Ind. Departamentode Agricultura de los EEUU durante Melbourne,Div. Rep. 7, 96 pags. losmeses de mayo yjunio de 1968. Memoria de la m HERMANN,F.J. 1954. A synopsisof the genus ReuniónTécnica Nacional de Maní. Formosa, págs. Arachis.Agrie. Monogr. U.S.D.A. 19, 26 págs. 171-177. HOEHNE,F.C. 1919.Com. lin. telegr. Bot. 8. Legumino- . 1969b.Evolución del géneroArachis. Seminario sas,1-99, táb. 132-159. Avanzadode Genética Agrícola para América Latina. . 1940.Genero: Arachis. Flora Brasílica 25(2) part. Maracay,Venezuela. Resúmenes, SAGA/B(d), 4 122:1-20, táb. 1-15. págs.mimeografiadas, sinn° de página. . 1944.Duas novasespecies de Leguminosasdo . 1973. Evolutionof the genusArachis. En Brasil.Arq. Bot. Estado São Paulon.s. 2(1): 16-18, AgriculturalGenetics. Selected Topics. 1 35-1 5 1 . Ed. táb.5. RomMoav, Jerusalem. & J.G.Kuhlmann. 1951 . Indicebibliográfico e nu- , A. FERNÁNDEZ& P. SEELIGMANN.1974. méricodas plantas colhidas pela Comissão Rondon. Recuperaciónde la fertlidadde un híbrido 400pags. São Paulo. interespecíficoestéril de Arachis(Leguminosae). 129-142. HUTCHINSON,J. 1964.Tribe Stylosantheae en The Bonplandia3(11): generaof floweringplants 1: 484-486.Oxford & W.C.GREGORY. 1974. Arachis Batizocoi nov. UniversityPress. sp. En Krapovickas,A., A. Fernández& P. HUSTED,L. 1931.Chromosome number in speciesof Seeligmann.Recuperación dela fertilidaddeun híbri- peanut,Arachis. Amer. Naturalist 65: 476-477. dointerespecífico estéril de Arachis (Leguminosae). Bonplandia3(1 1): 159,fig.2. . 1936.Cytological studies on the peanut, Arachis. D. Chromosomenumber, morphology and behavior, , C.E. SIMPSON& J.F.M.VALLS. 1985. Arachis andtheir application tothç problem of the origin of villosulicarpa(Leguminosae ), un proceso de domes- thecultivated forms. Cytologia 7: 396-423. ticaciónaislado. Resúmenes. XVI Congreso Argenti- node SAG.Rosario. JOHANSEN,E.L. & B.W. SMITH. 1956. Arachis Genética, hypogaeax A. Diogoi, embryoand seed failure. KRETSCHMER,Jr., A.E. & T.C. WILSON.1988. A Amer.J. Bot. 43(4): 250-258. newseed-producing Arachis sp. withpotential as KAWAKAMI,J. 1930. Chromosomenumbers in foragein Florida. Soil and Crop Sei. Soc.Fla. Proč. Leguminosae.Bot. Mag. (Tokyo) 44: 319-328. 47:229-233. KRAPOVICKAS , A. & V.A. RIGONI. 1949. LAET, Joannesde. 1630. Beschrijvinghevan West- Cromosomasde una especiesilvestre de Arachis. Indien.Elzeviers, Leyden. IDIA2(24): 23-24. LAS CASAS,B. de. 1909.Apologética Historia de las & V.A.RIGONI. 1951 Estudios citológicos en el Indias.Historiadores de Indiaspor D. Serranoy géneroArachis. Revista Invest. Agrie. 5(3): 289-293. Sanz,vol.1, Madrid. 704 págs. & V.A. RIGONI. 1954. Cruzamientos LEÓN,J. 1987. Botánica de los cultivos tropicales. 2oed. interespecíficosenArachis y consideraciones sobre SanJosé, Costa Rica. el origendel maní cultivado. Anais do 2° Congresso LINNAEUS,C. 1738.Hortus Cliffortianus. Panamericanode 29 de a 6 de Agronomía. março . 1748.Hortus Upsaliensis. abril, 266-267. São Brasil. pags. Piracicaba, Paulo, . 1753.Species Plantarum. 2 vols. & V.A.RIGONI. 1957. Nuevas especies de Arachis . 1754.Genera Plantarum, ed.V. vinculadasal problemadel origendel maní. MARCGRAVE,G. 1648.Historiae Rerum Naturalium Darwiniana11(3): 431-458. Brasiliae.En Piso, W. et G. Marcgraf,Historia & V.A. RIGONI.1960. La nomenclaturade las naturalisBrasiliae. Joh. de Laet ed. Lugdun. subespeciesy variedadesde Arachishypogaea L. Batavorum. RevistaInvest. Agrie. 14(2): 197-228. McCLENNY,W.E. 1935.History of thepeanut, The & W.C.GREGORY. 1960. Arachis Rigonii, nueva CommercialPress, Suffolk (Va.). 22 pp. especiesilvestre de maní.Revista Invest. Agrie. 14 MENDES,A.J.T. 1947. Estudos citológicos no genero (2):157-160. Arachis.Bragantia 7(1 1-12): 257-268. . 1969a.Colección de especies silvestres deArachis MÜHLENBROCK,R.H. 1962. TribeHedysareae, y de materialautóctono de maní.Material de espe- subtribeStylosanthinae of CentralAmerica and ciessilvestres recolectado con la participaciónargen- Mexico.Southw. Naturalist 7(1): 29-40. tina,en el viajede exploraciónorganizado por el MONARDES,N. 1565.Historia medicinal de las cosas

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que se traende nuestrasIndias Occidentales que hypogaeae as especiesA villosavar. corren tina, A. sirvenen medicina. Sevilla, (edición de 1574). Diogoie A. villosulicarpa.Ciencia e Cultura29(3): MONTEIROFILHO, H. da C. 1949.As especiesargen- 319-321. tinas... dogenero Sida ( Malvaceae ). Lilloa17: 501- PRINE,G.M. 1964.Forage possibilities in thegenus 522. Arachis.Soil and Crop Sci. Soc.Fla. Proc.24: 187- 4 MURTY, U.R. & M.R. JAHNAVI.1986. The A' 196. Genomeof Arachis hypogaea L. Cytologia51 : 241- , L S. DUNAVIN,R. J. GENNON & R.D. ROUSH. 250. 1986.'Arbrook' rhizoma peanut. A perennialforage OTERO,J.R. de. 1941.Nota de umaviagem de estudos legume.Agricultural Experiment Station. University aos camposdo sul de Mato Grosso.Serviço de ofFlorida, Gainesville. Circular S-332, 16 págs. InformaçãoAgrícola. Ministerio da Agricultura,53 PROUS, A. 1992. Arqueologiabrasileira. Editora págs. UniversidadedeBrasilia. . 1952. Informaçõessobre algumasplantas RESSLAR, P.M. & W.C. GREGORY. 1979. A forrageiras.Serviço de Informaçao Agrícola. Sér. Did. cytologicalstudy of threediploid species of the 11:1-313. genusArachis L. J.Hered. 70: 13-16. OVIEDOy VALDÊS, G.F. de. 1535.La historiageneral . 1980.A reviewof the nomenclature ofthe genus delas índias. Sevilla, 1 vol. ArachisL. Euphytica29: 813-817. . 1547.Coronica de las índias. Salamanca, 1 vol. RUDD,V.E. 1981,Tribe 14. Aeschynomeneae (Benth.) . 185 1 . HistoriaGeneral y Naturalde las índias,ed. Hutch,en Advancesin LegumeSystematics, ed. JA. de los Rios.Madrid. Hay varias reediciones: RM. Polhill& P.H.Raven 1: 347-354.Kew. Asunción,1944, 14 vols. Madrid, 1959, 5 vols. RUMPHIUS,G.E. 1747.Herbarium amboiense 5. PICKETT,T A. 1955.Note on oilsof two wild species RUSBY,H.H. 1901.The nature and uses of the peanut oï Arachis.J. Am. Oil Chem.Soc. 37(10): 521. {Arachishypogaea L ). J.New York Bot. Gard. 2: PIETRARELLI,J.R. 1961 . Informesobre un viaje reali- 114-123. zadopor la provinciade Corrientes (Argentina), Pa- SALAS,AM. 1945.El antigalde Ciénega Grande (Que- raguay,Brasil y Bolivia para coleccionar maníes cul- bradade Purmamarca,Prov. de Jujuy).Pubi. Mus. tivadosy silvestres.Informes Técnicos (INTA) 30, Etnogr.Facultad de Filosofíay Letras.Ser. A. 5: 1- 29 págs. 268.Buenos Aires. . 1968. Nuevo viaje de exploración para coleccionar SCHULZE-MENZ, G.K. 1964. Coronilleae- maníessilvestres y cultivados. Boletín informativo Stylosanthinaeen Melchior, H. 'Engler's Syllabus ManiseroINTA-IADO (Manfredi) 2(12): 6. derPflanzenfamilien' 2: 237. . 1982.Prospecciones enmaní (silvestres ycultiva- SIMPSON, C.E. 1980. The colletionof Peanut dos) conparticipación de técnicosde la Estación Germplasm1980. Argentina,Bolivia, Perú. Agropecuariade Manfredi(Provincia de Córdoba, TechnicalReport to the International Board for Plant Argentina).Oleico 17: 37-46. GeneticResources. MS. 65 págs.. PIRE,S.M.L. 1968.Estudio de los granosde polende . 1981.The collection of Peanut Germplasm 1981. Arachisyde géneros vecinos (Resumen). Memoria II Perú,Brazil. Technical Report to the IBPGR. MS. ReuniónTécnica Nacional de Maní.Cátedra de 47 págs. GenéticaFitotecnia Boletín5: 74. y (Corrientes), . 1982.The collection ofArachis germplasm 1981- . 1974.Estudio palinológico de la tribuHedysareae 82.Technical Report to the IBPGR. MS. 33 págs& (Leguminosae ). Bonplandia 3(12): 1 13-169. 5 figs. PLUKENET,L. 1691.Phytographya. . 1984a.The collection ofArachis germplasm 1983. POITEAU,P.A. 1802.Observations botaniques faites a Argentina,Bolivia, Brazil. Technical Report to the Saint-Domingue.Bull. Soc. Philom.Paris 66: 137- IBPGR.MS. 39 págs. 138. Paris,"Fructidor, an 10 de la Republique". . 1984b.Plant Exploration: Planning, Organization, . 1806.Observations sur Y Arachis hypogaea. Mem. and Implementationwith Special Emphasison Inst.Sci. Divers Savans, Sci. Math. ser. 1.1: 455-462, Arachis, en ConservationofCrop Germplasm - An pl. 7. InternationalPerspective. Crop Science Society of Am. PublicationNo. 8. 1: 1-20. . 1853.Note sur Y Arachis hypogaea. Ann. Sci. Nat. Special Chapter Bot.ser. 3. 19:268-272, lam. 15. & D.L. HIGGINS. 1984. Catalogof Arachis POMPEU, A S. 1977. Cruzamentosentre Arachis germplasmcollections inSouth America, 1976-1 983.

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TAES - IBPGR.79 pp. ofpeanuts with a D genome.Amer. J. Bot. 78(5): . 1986a.The collection ofArachis germplasm 1984. 630-637. Brazil.Technical Report to theIBPGR. MS. 26 TABARESDE ULLOA,F. 1799.Relación sucinta en págs. quese explica sencillamente elnuevo descubrimiento . 1986b.The collection oïArachis germplasm 1985. de sacarel aceytedel cacahuate o maní de América. Brazil.Technical Report to theIBPGR. MS. 28 Valencia. págs. TRINDADE,M.J. 1985. Alegrete do SéculoXVII ao , A. KRAPOVICKAS,JR. PIETRARELLI& R.O. SéculoXX. 1. EditoraMovimento, Porto Alegre. VANNI. 1986c. Evidenceon the Evolutionof VALLS, J.F.M.1983. Reporton the collectionof Arachishypogaea L. APRES,July 1986, Stillwater, GroundnutGermplasm in Brazilcoordinated by Oklahoma,Proc. 18: 27. CENARGEN/EMBRAPA- Expeditions 1 to 5: 1 2 3 SKVORTZOW,B.W. 1920.Notes on theAgriculture, (10-27.VI.1981), (18.VIII-4.IX.81), (2- 4 & 5 Botanyand theZoology of China.J. N. China 21.m.82), (20.IV-12.V.82) (20.HI-15.IV,83). MS. 8 + 2 BranchRoy. Asiat. Soc. 51: 135-158. págs. SLOANE,H. 1696.Catalogus plantaram quae in insula . 1984. Reporton thecollection of Groundnut Jamaicasponte proveniunt. Londini. GermplasminBrazil coordinated byCENARGEN/ SMARTT,J.S. 1964. Cross-compatibility relationships EMBRAPA.Expedition 6 (10-29.V.83).MS. 4 betweenthe cultivated peanut Arachis hypogaea L. págs.& 1 mapa. andother species of the genus Arachis. PhD thesis. , V. RAMANATHARAO, C.E. SIMPSON& A. NorthCarolina State University atRaleigh. KRAPOVICKAS. 1985. Currentstatus of & W.C. GREGORY.1967. Interspecific cross- collectionand conservationof SouthAmerican compatibilitybetween the cultivated peanut Arachis groundnutgermplasm with emphasis on wild hypogaeaL. and othermembers of the genus speciesof Arachis. En Citogeneticasof Arachis, Arachis.Oléagineux 22(7): 455-459. Proceedingsof an InternationalWorkshop, ICRISAT,Patancheru P.O, Andhra Pradesh, India, W.C. GREGORY& M. PFLUGEGREGORY. , págs.15-35 1978 a. The genomesof Arachishypogaea. 1. Citogeneticstudies of putativegenome donors. VERDCOURT,B. 1971.Fl. Trop.E. Air.,Leguminosae Euphytica27: 665-675. 3:442. , W.C. GREGORY& M. PFLUGEGREGORY. WEDDELL,H.A. 1853. Voyagedans le nordde la 1978b. Genomesof Arachis hypogaea L. 2. The Bolivie.Paris. implicationsinthe interspecific breeding. Euphytica WILLIAMS, D.E. 1989. The collectionof crop 27:677-680. germplasmwith emphasis on Arachishypogaea. . 1990.The groundnut, Arachis hypogaea L. en J. Bolivia,1988. Technical Report to the Agricultural Smartt,Grain Legumes, Cambridge University ResearchService, United States Department of Press.Chapter 3: 30-84. Agriculture.MS. 55págs. SMITH,B.W. 1 950. Arachis hypogaea. Aerial flower and . 1991.Peanuts and peanut farmers ofthe rio Beni: subterraneanfinit. Amer. J. Bot. 37(10): 802-815. Traditionalcrop genetic resource management inthe . 1956. Arachis hypogaea. Normal BolivianAmazon. Tesis. The CityUniversity of megasporogenesisandsyngamy with occasional sin- NewYork. MS. 170págs. glefertilization. Amer. J. Bot. 43(2): 81-89. WURDACK,J.J. 1970. Erroneousdata in Glaziou STALKER,H.T. 1991. A newspecies in section Arachis collectionsofMelastomataceae. Taxon 19: 911-913.

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Numerical List of the Taxa

1 - A. guaranitica 42a - A. glabrata var.glabrata 2 - Aņtuberosa 42b - A. glabrata 3 - A. Martii var.Hagenbeckii 4 - A. brevipetiolata 43 - A. glandulifera 5 - A. Oteroi 44 - A. cruziana 6 - A. Hatschbachii 45 - A. montícola 1-A. cryptopotamica 46 - A. magna 8 - A. major 47 - A. ipaënsis 9 - A. Benthamii 48 - A. valida 10 - A. douradiana 49 - A. Williamsii 11 - A. gracilis 50 - A. Batizocoi 12 - A. Hermannii 51 - A. duranensis 13 - A. Archeri 52 - A. Hoehnei 14 - A. stenophylla 53 - A. stenosperma 15a - A. paraguar iensis ssp. 54 - A. praecox paraguar iens is 55 - A. palustris 15b - A. paraguariensisssp. 56 - A. benensis capibarensis 51 - A. trinitensis 16 - A. setinervosa 58 - A. decora 17 - A. Macedoi 59 - A. Herzogii 18 - A. marginata 60 - A. microsperma 19 - A. prostrata 61 - A. villosa 20 - A. lutescens 62 - A. helodes 21 - A. retusa 63 - A. correntina 22 - A. Burchellii 64 - A. Simpsonii 23 - A. Pietrarellii 65 - A. Cardenasii 24 - A. villosulicarpa 66 - A. Kempff-Mercadoi 25 - A. inseminata 61 - A. Diogoi 26 - A. Giacomettii 68 - A. Kuhlmannii 27 - A. sylvestris 69 - A. hypogaea 28 - A. pusilla 69a - A. hypogaeassp. hypogaea 29 - A. Dardani 69al - A. hypogaeassp. 30 - A. repens hypogaeavar. hypogaea - 3 1 A. Pintoi 69a2 - A. hypogaeassp. 32 - A. lignosa hypogaeavar. hirsuta - 33 A. Kretschmeri 69b - A. hypogaeassp. fastigiata - 34 A. Rigonii 69b1 - A. hypogaeassp. - 35 A. chiquitana fastigiatavar. fastigiata 36- A. matiensis 69b2 - A. hypogaeassp. - 37 A. appressipila fastigiatavar. peruviana 38 -A. Vallsii 69b3 - A. hypogaeassp. - 39 A. subcoriacea fastigiatavar. aequatoriana AO-A. Burkart ii 69b4 - A. hypogaeassp. - 41 A. pseudovillosa fastigiatawar. vulgaris

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List of Collectors9

Addor,A. s/n(20). 839(63), 1029(42a), 1057 (42a). AdolfoM. 443 (66). Arbo,M.M. 1088(63). Aguilar,V. 106(63). Arbo,M.M., S.G. Tressens,A. Schinini& M.S. Ahumada,O., L. Ferraro,C. Chiffa& S.G. Tressens Ferrucci1762 (15a), 1764(42a), 1796(42a), 1871 118(63). (42a),1875 (42a). Ahumada,O. 2502(40). Arbo,M.M. & A. Schinini2449 (40). Ahumada,O. & A. Schinini3879 (40). Arbo,M.M., A. Schinini& I. Basualdo2878 (42a). Ahumada,O. 6236(45). Arbo,M.M., R. Monteiro,A. Schinini& A. Furlan Aliem,A.C. 120(68), 644 (37), 646 (37), 661 (67), 685 3485 (19), 3515 (19), 3537 (19), 3555 (19), 3556 (15a), 699 (8), 718 (7), 720 (42a), 721 (42a), 731 (17),3665 (19), 3669 (19). (20), 732 (62), 734 (62), 744 (68), 750 (39), 760 Archer,W.A. & A. Gehrt0017 (42a), 0098 (42a), (37),761 (37). 0137 (42a), 0138 (13), 0151 (5), 3967 (13), 3968 Aliem,A.C. & J.G.A.Vieira 1318(33), 1452(37), (13), 3982 (9) [IAC V. 83], SP 35771 (42a), SP 1455(37), 1663(68). 36469(42a), SP 36470(5), SP 36473(5). Aliem,A.C., J.F.M. Valls, J.G.A. Vieira & J.A. Archer,W.A. 4429 (40), 4439 (40), 4449 (40), 4569 Comastri2 147 (33),2269 (68). (61), 4624 (42a), 4655 (42a), 4655a (42a), 4664 Aliem,A.C., J.G.A.Vieira & W.L. Werneck2796 (42b),4670 (42b), 4758 (42a). (53),2824 (19). Archer,W.A. & T. Rojas4851 (42a). Aliem,A.C., G.L. Webster& W.L. Werneck3070 Archer,W.A. 4904 (42a), 4941 (42a), 4952 (61). (19). Archer,W.A. & T. Rojas 7470(42a). Aliem,A.C. 3485(58). Arechavaleta,J.s.n., herb.Osten 4105 (40), 5368a (61), Allemão,Freire s/n (27). 5369(40), 5369a (40), 5370 (40). Allemão,F. Freire& M. D. Cysneros364 (27). Arenas,P. 416(42a). Anderson,W.R. & al. 6847(19). Arrillagade Maffei,B.R. 644(61), 898 (61). Anderson,W.R. 7021 (22), 11247(42a), 11298(9), Arrillaga de Maffei,B.R., P. Izaguirre & A. 11338 (39), 11339 (20), 11352 (20). Laguardia 1008(40). Anderson,W.R., M. Stieber& J.H.Kirkbride 37175 Arrillagade Maffei,B.R., P. Izaguirre,A. Laguardia (31). & R. Brescia1973 (61). AndradeLima, D. de 3018 (20), 64-4200(25), 65- Báez,J.R., V.A. Rigoni & A. Krapovickas7264 (45), 4317a (25), 67-5006(29), 68-5328(22), 68-5329 [IACV.819 & V.820]. (27). Badcock,W.J. 19 (66). Anzótegui,L.M. 1210(63). Balansa,B. 1526(42a), 1526a (42a). Aranda,D. 69 (42a). BaldwinJr., J.T. 3139 (5), 3140(13), 3141 (37). Arbo,M.M. 21 (42a),449 (63). Bartlett, H.H. 19244(61), 21028 (40), 21201 (61). Arbo,M.M. & A. Schinini548 (63). Basualdo,I., 277(42b). Arbo,M.M., A. Schinini,O. de Coll & R.O. Vanni Baycé,D. & E. Marchesi19970 (40). Baycé,D., Merola,R. Beyhaut & Speroni20763 (61), 20780(61). 9Thislist includes all ofthe specimens studied. In the Berro,M.B. s/n(61), 1117(61), 1501(40), 1897(40), of forwhich there is an abundance descriptionsspecies 5886(61), 7092 (61). of "selectedmaterials" were specimens, chosen,taking Bertoni0558 0695 0775 1905 care not to omitthe localities.Numbers (42a), (42a), (42a), (42a), collecting 2075 3457 5189 appearingin bracketsand in bold indicatethe parent (42a), (42a), (42b). numbersused by M.P. Gregory & W.C.Gregory (1979). Bertoni,G.T. 684-LPS23123 (15a), LPS 23124(42a), Numberspreceeded by V or IAC correspondto the LPS 23125(63), LPS 23126(14), LPS 23127(15a), InstitutoAgronómico de Campinas. LPS 23128(42a).

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Bisby1265 (30). (62),7272 (22). Blanchet,J.S. 2669 (28). Cowan,C.P., S.G. Tressens & L.I. Ferraro4180 (42a). Bondar,G. s/n(27). Cristóbal,C.L., A.Krapovickas & A. Schinini1899 Booke,T. 48 (56). (40). Bordas,E. 1247(42b). Cuezzo,A., De Marco,T. Ruiz1 1 147 (63), 11316 (42a). Bowes,N.D. 7 (61). Davies,P. & E. Marchesi19538 (61). Braun,O. 7 & 8 (56). Davis,P.H. & D.F. Coelho60389 (30). Brooke,T. 108(66). del Puerto,O. 1039(61), 1795(40), 1949(40), 2103 Bruderreck,B. 20 (43). (61), 13039(40), 13078(31). Burche»,W.J. 8328 (22), 8358 (22), 8443 (22). delPuerto, O. & Berreta14787 (61). Burkart,A.E. SI391(61), 0888 (40), 7791 (40), 8166 delPuerto, O. & Borsani2464 (61). (40),8167 (40), 9050 (61), 14111(42a), 15715 (62), del Puerto,O. & E. Marchesi5961 (40), 8388(61), 18159(61), 18222(42a), 21775 (40), 22585 (61). 15363(40). Burkart,A.E. & S.Crespo23004 (61). delPuerto, O. & G. Ziliani15170 (61). Burkart,A.E. & N.S.Troncoso26242 (61). delPuerto, O., G. Ziliani& D. Baycé18171 (40). Burkart,A.E. 27595(63). Descole,H. 3260(42a). CaballeroMármori, G. 1159 (42a). Deslandes,J.A. s/n (40), 69 (29). Cabral,E., L.I.Ferraro& S. Cáceres286 (42a). Diem1584 (42a). Cabrera,A.L. 2585(61). Diogo,J.C. 317 (67). Cáceres,S. & C. Zamudio311 (61). Duarte10313(31). Calcagnini,C. BAB 7952(42a). Duarte,A.P. 13928(42a), 13942 (22). Cárdenas,M. 2988(65), 4741 (50). Duarte,L. 772(42a), 817 (42a). Carnevali,R. 0189 (63), 1643(63), 2271 (63), 2422 Ducke,A. 1970(29). (63), 2978(42a), 3077 (42a), 3131 (40), 3141 (40), Dusen,P.K.H. 13472(53). 4097(63), 4249 (42a), 4547 (42a), 4551 (42a), 4888 Edwall,G. SP 1541(53). (63). Eiten,G. 2549(30), 2746 (30). Carver,W.A. SP 42275(13). Eiten,G. & L.T. Eiten3620 (22), 4005 (27), 9340-B Castellanos,A. s/n(61), LIL 15979(40), LIL 15980 (2), 9904(16), 9425 (42a), 10051 (22). (61),LIL 15981(40). Ekman,E.L. 1718(42a), 1719 (42a). ChebataroffJ.s/n (61). Fernández,A. 363(42a), 364 (40), 385 (42b). Clos, E.C. 1918 (42a), 3225 (61), 5930 (63), 5968 Fernández,J.G.613(51). (42a),5981 (42a), 6031 (42a), 6077 (42a), 6135 (61), FernándezCasas, J. 7371 (42a), 7497 (42a), 7518 6292(61). (42a). Conagin,C.H.T.M. 1 (30) [IAC 18127V.305], 2 (42b) Ferreira,M.B. s/n(9). [IAC 18128V.361], 3 (8) [IAC 18130V.85], 4 (13) Fiebrig,K. 0263(42a), 4195 (42a), 4277 (14). [IAC 18131V.128], 7 (42a) [IAC 18135V.355], 8 Filipovich,R. 404(51), 415 (51). (8) [IAC 18136V.85J, 9 (42a) [IAC 18137V.360], Fonseca,S.G. & E. Onishi325 (42a), 1094 (42a), 1551 10 (45) [IAC 18138V.357J, 12 (13) [IAC 18669 (2). V.128],13 (42a) [IAC 18670V.362], 14 (8) [IAC Fries,R.E. 1465(51). 18671V.85], 15 (8) [IAC 18672V.85], 16 (8) [IAC Gardner,G. 2091(29), 3103 (18), 3104 (19). 18673V.85], 17 (8) [IAC 18674V.85], 18 (24), 19 Gaudichaud,Ch. 53 (53), 864 (53), 883 (61) (leg.Se- (42b) [IAC 18676V.361], 20 (42a) [IAC 18677 llo),1991 (40) (leg.Sello). V.362],21 (42b) [IAC 18678V.361], 22 (8) [IAC Gehrt,A. 4744(53), SP 47535(24), SP 45842(42a). 18129V.85], 23 (42a) [IAC 18134V.362]. Gibbs,P., H.F. Leitão Filho,L. Kinoshita& J.B. Coradin,L., R. Mesquita,L. Valle & J.G.A.Vieira Andrade3506 (53). 1963(29), 2605 (27). Gibbs,P.E., G.J.Shepherd, J.B. de Andrade& G. Coradin,L., R. Schultze-Kraft& G.P. da Silva3224 Buffarah5338 (10), 5394 (42a), 5467 (8). (29). Glaziou,A. 10513(27). Coradin,L. 3466 (20), 3650 (22), 3659a(22), 3660 Götzsche,H. 491(42a), 502 (42a), 514 (42a). (22), 3669 (22), 3715 (22), 3724 (22), 3736 (19), Gregory,W.C. s/nXI-1993 (15a). 3755 (22), 3836 (22), 6859 (20), 6862 (62), 6919 Gregory,W.C., A. Krapovickas & J.R.Pietrarelli LIL

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462183(13), 9530 (63) [22],9531 (63) [22],9548 (19),10538(30) [7]. (63), 9549 (63), 9551 (63), 9553 (42a) [29],9555 Gregory,W.C. & A. Krapovickas10541 (5), [IAC (42a),9557 (63), 9558 (63), 9560 (42a), 9562 (42a), V.784], 10543(5) [53], 10545(5), [IAC V.785], 9563(63), 9564 (42a), 9566 (42a), 9567 (42a) [84], 10546(42a), 10547(9), 10548(9), [IAC V.786], 9568(42a), 9569 (42a), 9570 (42a) [10],9571 (42a), 10549(5), 10550(42a) [30], [IAC V.787],10551 9572(42a), 9573 (42a), 9574 (42a), 9575 (42b), 9576 (11), 10554 (10), 10555 (41), 10556 (10),[IAC (42b), 9577 (42a), 9578 (42a), 9580 (42b), 9587 V.788],10557 (10), 10558(41) [IAC V.789],10559 (42b),9591 (42a) [66],9592 (42a), 9596 (42a), 9606 (41) [60],10560 (41), 10561 (41) [25],[IAC V.790], (42a),9610a & 9610b(42b), 9618 (42b), 9625 (41), 10562(15a), 10563(42a), [IAC V.791],10565 (41) 9626 (41), 9627 (41), 9627a(41), 9628 (41), 9629 [61],[IAC V.792],10566 (41), 10567(42a), 10568 (42a) [24],9630 (41), 9632 (8), 9634(41) [31],9635 (1) [5], 10573(8), [IAC V.795],10574 (42a), [IAC (41), 9636 (10), 9637 (42a), 9638 (8), 9639 (10), V.793],10575 (8), [IAC V.794],10576 (8), 10580 9640(42a), 9641 (8), 9642(42a), 9643 (42a), 9644 (8),[IAC V.796], 10582 (8) [2],[IAC V.797],10585 (42a),9645 (42a) [71],9646 (15a) [3],9647 (42a), (15a) [17],[IAC V.798],10588 (8) [18y 56],10596 9648(14), 9649 (42a), 9650 (41), 9651 (8), 9652 (8), (42a) [27], [IAC V.799],10598 (32) [20], 10602 9664(41), 9665 (1), 9667 (42a) [28],9672 (41), 9674 (67) [37],12787 (31) [96],12881 (25) [97],12922 (42a), 9678 (41), 9679 (41), 9680 (1), 9681 (42a), (25) [98],12939 (29), 12941(29), 12943(29) [99], 9688 (41), 9696 (42a), 9701 (1), 9703 (41), 9706 12945(29), 12946(29) [100], (41), 9735(5), 9736(13), 9747(5), 9753(9), 9761 Grüner,G. 599(42a), 671 (42a). (9), 9763(5), 9764(9) [14],9765 (5), 9766(5), 9769 Hagelund,K. 5482(40), 8317 (40). (9), 9770 (5), 9771 (9), 9772 (11), 9774 (9), 9775 Hagenbeck s/n (42b). (11), 9776 (9), 9784 (9), 9786 (5), 9788 (11) [1], Hammons,R.O. & D.W.Branch Tifton Acc. A28 (63). 9790 (5), 9792 (5), 9793 (5), 9795 (9), 9797 (42a) Hammons,R.O., W.R. Langford & J.R.Pietrarelli 8 [72],9799 (10), 9800 (10), 9801 (42a), 9803 (9), (61), 10 (61), 12 (61), 14 (61), 16 (40) [86],17 (40) 9805(9), 9806(42a), 9810 (5), 9811 (13),9812 (13) [38],23 (40) [87],24 (40),27 (40) [90]. [47],9813 (42a), 9814 (13), 9815(42a), 9816 (13), Hammons,R.O., W.R. Langford& H.R. Ojeda 333 9817(13), 9818 (2), 9819(5), 9820(13), 9821 (13), (42a),334 (42b), 349 (42a) [64]. 9822 (42a), 9823 (42a), 9824 (68), 9825 (8), 9826 Hammons,R.O., W.R. Langford,A. Krapovickas& (8), 9827(42a), 9828 (5), 9829 (1 1),9830 (42a) [23], H.R. Ojeda 354(40) [91],355 (40), 357 (61). 9831(13), 9832 (5), 9834(42a), 9835 (13), 9837 (2) Hammons,R.O., W.R. Langford& A. Krapovickas [4],9839 (12), 9841(12) [16],9843 (12), 9848(6) 362(40) [75],364 (40) [73],365 (30), 408 (53),410 [50],9853 (5), 9855 (5), 9858 (5), 9863 (6), 9865 (6), (53) [42],467 (30). 9869(6), 9875(6), 9880(6), 9882(42a) [77],9883 Hammons,R.O., W.R. Langford,A. Krapovickas& (9), 9889(7), 9893(42a) [78],9894 (42a), 9898 (20), V. Hemsy487 (17), 491 (17), 499 (9), 500(30), 520 9900 (20), 9901 (67) [76], 9903 (20), 9905 (20), (9), 521(5), 523 (9), 524 (5), 525 (3), 526 (3) [85], 9906 (20), 9907 (20), 9909 (20), 9918 (42a),9920 547(13), 548 (5) [69],549 (5), 550(9), 551(8) [65], (20), 9921 (42a), 9922 (42a),9923 (23) [45],9925 552(42a), 553 (42a),554 (9), 555a(8), 555b(42a), (42a), 9926 (62) [33],9927 (20), 9935 (42a) [49], 559 (8) [41], 560 (42a), 562 (8), 563 (42a), 564 9942(2), 9944(42a), 9947 (42a), 9948 (42a), 9949 (42a),565/566 (15b) [63], 567 (42a), 568 (42a), 569 (42a), 9951 (42a), 9953 (42a), 9954 (42a), 9958 (42a) [94],570 (15b),571 (42a),572 (14) [79],573 (42a),9960 (42a), 9964 (42a), 9966 (42a) [57],9974 (15b),574 (1), 575(41), 576 (41) [95],577 (41) [68]. (40), 9975(30), 9977(9) cult.,9978 (9), 9990(37) Handro,O. 682(9), 686(30). [13],9993 (37) [51], 10000(37), 10002(37) [52], Hartley,W. &. T. Rojas SH149(42a), SH155(42a), 10017(65) [34],10026 (65), 10034(34) [21],10038 SH195(42a). (51) [8], 10105(42a) [IAC V.774], 10120(42a) Hassler,E. 0938(42a), 1431(42a), 1706(42a), 3445 [IAC V.775],10127 (17) [15],[IAC V.776],10138 (42a),4261 (42a), 451 1 (41), 4512(42b), 4975 (1), (4), 10139(42a), 10160(20) [43], [IAC V.777], 5069(41), 5863 (42b), 6034 (42a), 6358 (15a), 6513 10174 (20) [44], [IAC V.778], 10176(20), [IAC (42a), 6515 [7115] (42a), 7476 (32), 7542 (15a), V.779],10234 (19), [IAC V.780],10240 (19) [26], 7664(42a), 8439 (42a), 9886 (41). [IAC V.781],10258 (20), 10259(20), [IAC V.782], Hatschbach,G., J.C. Lindeman& J.H. de Haas 10292(19), 10406(18) [12], [IAC V.783],10449 13605(53).

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Hatschbach,G. & A. Krapovickas19240 (53). Irwin,H.S. 2570(20). Hatschbach,G. 20679(53), 21924(42a), 23570(9), Irwin,H.S. & T.R. Soderstrom7281 (2), 7281(20, K 23729 (42a), 25077 (42a), 25104(9), 25149(11), p.p.),7546 (42a), 7605 (20). 25177(42a), 29502 (37), 29566 (42a), 32105 (6). Irwin,H.S., R. Souza,R. Reisdos Santos 10469 (19). Hatschbach,G. & C. Koczicki33179 (9). Irwin,H.S., H. Maxwell& D.C. Wasshausen19055 Hatschbach,G. & R. Kummrow37125 (42a). (19),19196 (19), 21 163 (22), 21448 (22), 21732 (22). Hatschbach,G. 37397(42a), 37425(9), 37492(20), Irwin,H.S., da Fonseca,R. Souza,R. Reisdos Santos, 37496(39), 37674 (42a). Ramos26986 (31). Hatschbach,G. & T.P. Ramamoorthy38197 (19). Irwin,H.S., R.M. Harley& G.L. Smith31651 (27). Hatschbach,G. 38670(41), 39383 (19), 44031 (42a). Irwin,H.S., W.R. Anderson,M. Stieber& E.Y. Lee Hatschbach,G. & R. Kumrow48439 (41), 48441 34701(19). (42a). Isabelle,A. 1833(40). Hatschbach,G. & J.M.Silva 48462 (41). Issouribehere,P. BAB 24067 (42a). Hatschbach,G. & F.J.Zelma 49149 (8). Izaguirrede Artucio,P. & R. Beyhaut19665 (61). Hatschbach,G. 50585(30). Izaguirrede Artucio,P., R. Brescia& M. Marchi Hatschbach,G. & J.M.Silva 56081 (21). 20991(61), 20993 (61), 20995 (61), 21003 (40). Hatschbach,G. & R. Kumrow56351 (19). Izaguirrede Artucio, P., S. Grun& R. Beyhaut19730 Hatschbach,G., A. Schinini& J.M.Silva 58693 (9), (40). 58791(42a), 58796 (42a), 58849 (42a), 58864 (15a), Izaguirrede Artucio,P., Laguarotra& E. Marchesi 58909(41), 58912 (42a), 58913 (68), 58996 (8), 59010 17115(61). (8),59045 (8), 59049 (8), 59053 (42a), 59057 (30). Jorgensen,P. 191(42a), 3637 (42b). Hauman,L. s/n(61). Kirkbride,J.H. & Lieras3042 (64). Heringer,E.P., D. Andrade-Lima,J. de P. Lanna Krapovickas,A 7201(63). Sobrinho& A. CoelhoSarmento 638 (29), 865 Krapovickas,A., V.A. Rigoni & J.R.Pietrarelli], 7830 (29). (63),7847 (42a), 7855 (42b), 7864 (42a), 7870 (42a), Heringer,E.P. 10731(31), 11996 (27). 7890(63), 7897(63), 7904 (42b), 7910 (42a), 791 1 Herter,W.G. 84489 (61). (42a),7934 (42a), 7967 (40). Herzog,Th.K.J. 1110(51). Krapovickas,A. 7988(51) [39],7993 (51), 8010(51), Hoehne,F.C. 0019(52), 2624 (37). 8012(45) [11],8025 (9), 9412(65), 9459 (34), 9484 Hoehne,F.C. & A. GehrtSP 35773(5),[IAC V.82], SP (50) [19],9495 (50), 9496 (50), 9497 (50), 9498 (50), 35775(13), [IAC V.84],SP 36478(13), SP 45842 9503(50), 9504 (50), 9505 (50). (42a). Krapovickas,A. & C.L. Cristóbal11285 (40), 11292 Horovitz,S. s/n(51). (61), 11300(40), 11301(40), 11306(40), 11311 Huidobro,A.M.R. 3697 (61), 3712 (61), 4872 (42a), (61), 11350(63), 11401(42a), 11462(15a) [35], 5182(42a). 11483(42b), 11488 (15a) [36],11609 (42a), 11610 Hunziker,A.T. 1635(51), 5603 (42a), 6683 (42a). (42a),11813 (42a), 11822 (42a), 11851 (42a), 11905 IAC (InstitutoAgronomico de Campinas,SP, Brasil), (63), 11919(63), 11941(63), 11946(42b), 11970 10453(8) [V.128],18127 (30) [V.305],18128 (42b) (42a),12000 (42a). IV.361],18129 (8) [V.85J,18130 (8) [V.85],18131 Krapovickas,A., C.L. Cristóbal& R.A. Palacios (13) [V.128], 18132 (51) [V.356], 18134 (42a) 12142(42b), 12209 (42b), 12290 (42a), 12352 (42a), [V.3621,18135 (42a) [V.355],18136 (8) [V.85], 12363(42a), 12399(42a), 12401(42a), 12404(67), 18137(42a) [V.3601,18138 (45) [V.3571,18139 12450(42a), 12456 (15a), 12460 (42a), 12578 (42a), (63) [V.359],18670 (42a) [V.362],18671 (8) [V.85], 12593(63). 18672 (8) [V.85], 18673 (8) [V.85], 18674 (8) Krapovickas,A. 13663(15a). [V.85], 18675(24) [V.44], 18676(42b) [V.361], Krapovickas,A. & C.L. Cristóbal13735 (42a), 13780 18677(42a) [V.362],18678 (42b) [V.361]. (63), 13782(63). Ibarrola,T. 1104(42a), 1662(40), 2075 (61), 2148 Krapovickas,A., C.L. Cristóbal& L.Z. Ahumada (61), 2391 (61), 2406 (61), 2590 (40), 3610 (42a), 13800(63), 14006(15a), 14055(41), 14168(8), 4229(42a). 14248(32), 14250(42a). Irigoyen,J.M. & A. Schinini137 (42a). Krapovickas,A.& C.L. Cristóbal14318 (63), 14319 (63). Irigoyen,J.M. 370 (63). Krapovickas,A. & V. Hemsy14389 (11), 14424(8),

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(42a),5 162 (42a). Valls,J.F.M., J.P. Moss, M.A.N. Gerin & G.P. Silva Valls,J.F.M. 5786 (30), 5868 (30). 7317 (40), 7320 (40), 7330 (40), 7332 (30), 7333 Valls,J.F.M. & W.L. Werneck5895 (31), 5913 (19), (40),7334 (40). 5916(42a), 5922 (42a), 5924 (42a), 5925 (20). Valls,J.F.M., C.E. Simpson,J.P. Moss, M.A.N. Gerin Valls,J.F.M., R.F.A. Veiga & G.P.Silva6001 (27), & G.P. Silva7335 (40), 7337 (40), 7344 (40), 7345 6110 (27), 6180 (27), 6188 (25), 6215 (29), 6240 (40), 7347 (40), 7349 (40), 7357 (40), 7359 (40), (25),6284(19). 7363(40), 7377 (53), 7379 (53). Valls,J.F.M., C.E. Simpson& A. Gripp6305 (2), Valls,J.F.M., C.E. Simpson& G.P. Silva7382 (53), 6306(42a), 6310 (2), 6324(36). 7384(53). Valls,J.F.M., C.E. Simpson,A. Gripp& C.N. Cunha Valls,J.F.M. & V.R. Rao 7529(31), 7530 (42a), 7531 6325(62), 6326 (62), 6330 (67), 6331 (62). (20),7533 (17), 7536 (17), 7537 (42a), 7539 (19). Valls,J.F.M., C.E. Simpson& A. Gripp6332 (20), Valls,J.F.M., V.R. Rao, M.A.N.Gerin & G.P. Silva 6337 (36), 6338 (20), 6340 (36), 6344 (68), 6345 7540 (42a), 7541 (42a), 7542 (42a), 7543 (42a), (36), 6346 (64), 6351 (68), 6352 (68), 6355 (68), 7545 (2), 7547 (2), 7549 (42a), 7554 (42a), 7555 6356 (36), 6357 (36), 6361 (36), 6366 (36), 6380 (12), 7557(9), 7560(12), 7563(7), 7565(7), 7566 (68), 6389 (68), 6390 (68), 6396 (68), 6404 (68), (7), 7568 (7), 7572 (7), 7574 (7), 7578 (9), 7580 6405(36), 6407(36), 6408a(36), 6408b(68), 6409 (42a),7581 (9), 7585(9), 7586(9), 7588(7), 7590 (36), 6410 (68), 6411 (20), 6412 (20), 6413 (68), (7),7593 (7), 7594 (12), 7596 (7), 7598(5), 7599 (5), 6414 (20), 6415 (20), 6416 (54), 6436 (42a),6441 7600 (5), 7602 (5), 7603 (5), 7606 (13), 7607 (2), (20),6442 (42a), 6443 (42a), 6450 (42a). 7614(13), 7617 (2), 7618(5), 7619(13), 7620 (13), Valls,J.F.M., A. Krapovickas,V.R. Rao & G.P. Silva 7621(5), 7622(9), 7623(42a), 7624 (42a), 7627 (9), 6466 (20), 6475 (19), 6485 (19), 6496 (22), 6517 7628(8), 7630(8), 7631(33), 7632(8), 7633(33), (21), 6522 (19), 6530 (22), 6532 (22), 6536 (55), 7634(42a), 7635 (38), 7637(33), 7638(15a), 7639 6540 (22), 6546 (22), 6547 (27), 6554 (22), 6555 (68), 7641(42a), 7642 (42a), 7644 (8), 7647(42a), (22), 6556 (22), 6558 (22), 6559 (22), 6566 (22), 7648 (42a), 7656 (15a), 7658 (42a), 7659 (42a), 6574 (22), 6575 (27), 6586 (22), 6600 (22), 6602 7669(15a), 7671 (15a), 7672 (8), 7673(42a), 7674 (22), 6604 (22), 6605 (22), 6609 (17), 6610 (17), (15a),7675 (15a), 7676 (8), 7677(15a), 7678 (42a), 6611 (55), 6612 (17), 6626 (17), 6633 (22), 6634 7679(42a), 7680 (42a), 7681 (60), 7683 (15a), 7684 (22), 6635 (22), 6636 (22), 6637 (22), 6638 (19), (15a),7685 (15a), 7686 (42a), 7687 (8), 7688(42a), 6640 (22), 6648 (19), 6649 (18), 6652 (18), 6655 7694(1), 7695 (41), 7698 (42a), 7700 (1), 7701(41), (27),6662 (19), 6667 (19), 6668 (27). 7704(1), 7706(41), 7707 (10), 7709 (10), 7710 (41), Valls,J.F.M., C.E. Simpson& W.L. Werneck6671 7711 (10), 7712 (9), 7713 (42a), 7715 (11), 7716 (30), 6673 (30), 6674 (30), 6676 (27), 6709 (27), (11), 7717 (5), 7718(5), 7719(42a), 7720 (42a). 6727 (31), 6728 (31), 6740 (31), 6741 (31), 6744 Valls,JŠF.M. & C.E. Simpson7725 (19), 7727 (42a), (27), 6759 (18), 6767 (27), 6772 (25), 6773 (27), 7728(42a). 6781(27), 6784 (31), 6785 (27), 6788 (19). Valls,J.F.M., C.E. Simpson.H.T. Stalker,I. Godoy& Valls,J.F.M., C.E. Simpson,W.L. Werneck& J.M. W.L. Werneck7732 (20), 7741 (20), 7742 (20), Santos6791 (31), 6792 (31). 7743(20), 7745 (42a), 7760 (42a), 7761 (42a), 7762 Valls,J.F.M. & J.O.N.Gonçalves 6898 (40). (53), 7764 (53), 7784 (23), 7786 (23), 7793 (23), Valls,J.F.M., A. Krapovickas,R.F.A. Veiga & G.P. 7799 (22), 7805 (22), 7821 (22), 7850 (22), 7861 Silva 7002 (27), 7004 (19), 7037 (27), 7039 (29), (23), 7862 (22), 7863 (22), 7868 (22), 7875 (22), 7055 (29), 7058 (19), 7060 (27), 7062 (29), 7063 7880 (22), 7881 (55), 7883 (55), 7887 (19), 7894 (29), 7065 (27), 7068 (19), 7071 (27), 7076 (27), (19),7895 (19), 7897(19). 7079 (27), 7081 (27), 7086 (29), 7105 (27), 7123 Valls,J.F.M., R.F.A. Veiga, & G.P. Silva8304 (19), (27), 7126 (27), 7130 (27), 7144 (27), 7156 (27), 8308 (19), 8309 (22), 8321 (22), 8335 (22), 8343 7165 (27), 7166 (29), 7172 (27), 7176 (27), 7180 (22), 8345 (27), 8352 (22), 8359 (22), 8373 (27), (27), 7191 (27), 7194 (27), 7197 (29), 7215 (29), 8375 (22), 8377 (55), 8381 (55), 8383 (17), 8386 7232 (25), 7243 (25), 7246 (25), 7287 (27), 7289 (27), 8390 (22), 8392 (27), 8402 (22), 8403 (27), (27),7292 (25), 7294 (27). 8405 (22), 8410 (29), 8413 (22), 8417 (27), 8418 Valls,J.F.M., J.P. Moss & G.P. Silva7300 (42a), 7302 (19), 8422 (27), 8423 (27), 8435 (27), 8437 (27), (42a),7305 (42a). 8440 (29), 8442 (27), 8444 (27), 8451 (27), 8458

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(27),8461 (27), 8462 (29), 8466 (27), 8469 (29), 8471 10837(27), 10891(27), 10892(27), 10912(29), (29),8473 (22), 8475 (27), 8477 (27), 8481 (19), 8482 10913(29), 10921(27), 10922(27), 10932(27), (19),8486 (19), 8491 (27), 8494 (27), 8496 (22), 8501 10933(27), 10936(29), 10939(29), 10940(29), (22),8503 (29), 8505 (29), 8509 (29), 851 1 (27),8516 10945(29), 10946(29), 10947(29), 10953(29), (27),8519 (27), 8520 (27), 8522 (29). 10960(29), 10963(29), 10968(29), 10969(27), Vails,J.F.M., A. Pott,Penteado, L. Jank& M. Araujo 10972(29), 10974(29), 10980(27), 10981(29), 8530 (8), 8555 (15a), 8568 (15a), 8644 (10), 8678 10988(29), 10992(29), 10993(27), 11000(27), (38),871 1 (60),8728 (1), 8729(41). 11001(29), 11006(29), 11008(29), 11016(29), Valls, A. Krapovickas,C.E. Simpson& G.P. 11020 (27), 11022 (27), 11027 (27), 11028 (19). Silva 8736 (36), 8740 (20), 8750 (39), 8753 (68), Valls,J.F.M., C.L. Quarin,M.S. FrançaDantas & 8756(36), 8758 (68), 8763 (68), 8764 (68), 8816 (24), G.P. Silva 11736 (42a), 11746 (5), 11758 (11), 8818(24), 8820 (24), 8887 (68), 8888 (68), 8889 (68), 11768(5), 11778 (13), 11781 (11), 11886 (41), 8890(36), 8891 (68), 8892 (68), 8893 (36), 8896 (64), 11917(41),11922 (42a), 11923 (42a). 8900(64), 8910(36), 8916a(68), 8916b(39), 8918 Vails,J.F.M. & A. Krapovickas12080 (39), 12083 (68), 8920 (39), 8922 (39), 8934 (62), 8935a(68), (62), 12084(36), 12085(23), 12086(23). 8935b(39), 8937 (39), 8941 (39), 8943 (39), 8944 Vails, J.F.M., C.N. Zanin, A.R. Miranda, J.C. (39),8945 (39), 8964 (68), 8965 (54), 8976 (20), 8977 Oliveira& W.L. Werneck12322 (40). (42a),8978 (20), 8979 (68), 8980 (68), 8982 (20), 8986 Valls,J.F.M., M.L. Galgaro,D.M.S. Rocha& G.P. (42a),8988 (42a), 8989 (20), 8995(20), 9000(23), Silva 12488(53), 12516(16), 12523(22), 12567 9002(42a), 9004 (23), 9010 (53), 9012 (53), 9013 (20), (22), 12568(22), 12575(53), 12618(22). 9017(53),9019(20). Valls,J.F.M., M.L. Galgaro& G.P. Silva 12627(22), Valls, A. Pott& L.B. Bianchetti9056 (37), 12646(53). 9060(37), 9061 (37), 9077 (37), 9094 (52), 9095 (52), Valls,J.F.M. 12658 (68). 9129(33), 9130 (37), 9140 (52), 9146 (52), 9147 (67), Valls, J.F.M.,Go, A.R. Miranda& J.C. Oliveira 9148(67), 9153 (48), 9157 (48), 9162 (48), 9214 (68), 12812(61). 9230(68), 9231 (68), 9235 (68), 9236 (68), 9237 (68), Valls,J.F.M., R.N. Pittman& G.P. Silva 12883(21), 9243(68), 9257 (7), 9306(53), 9318 (62), 9348 (20), 12893(58), 12899 (19), 12900 (58), 12901 (27), 12915 9349(20), 9350 (36), 9354 (68), 9355 (68), 9357 (39), (19),12916 (17), 12927 (19), 12939 (21), 12940 (27), 9358(39), 9359 (39), 9375 (68), 9394 (68), 9395 (68), 12941(19), 12952 (22), 12963 (19), 12967 (19), 12976 9397(54), 9401 (39), 9402 (39), 9403 (39), 9455 (7), (19),12978 (19), 13017 (22), 13021 (22), 13022 (27), 9456(9), 9468(8), 9469(8), 9470(68), 9479(68), 13023(55), 13040 (19), 13044 (27). 9481(38), 9482 (38). Valls,J.F.M., L. Faracode Freitas,E.A. Pizarro& Valls,J.F.M. & W.L. Werneck9858 (19). G.P. Silva 13074 (30), 13075(30), 13076 (30), Valls,J.F.M., C.E. Simpson& W.L. Werneck9875 13080(25), 13082(28), 13097(31), 13099(31), (5), 9878 (2), 9882 (5), 9887 (8), 9889 (33), 9894 13102(27), 13104(27), 13105(28), 13107(27), (68), 9896 (68), 9902 (38), 9904 (33), 9906 (15a), 13109(28), 13110(31). 9907(33), 9912 (68), 9913 (68), 9917 (33), 9918 (8), Valls,J.F.M. 13141 (31). 9920 (42a), 9923 (52), 9932 (41), 9937 (5), 9941 Valls,J.F.M., E.A. Pizarro,S.E.S. Valente& W.L. (13),9950 (21), 9952 (17), 9953 (58), 9955 (58). Werneck13142 (19), 13150 (31), 13151 (31), 13153 Valls,J.F.M., S. Miotto& G.P. Silva10229 (53). (31),13154 (31), 13155 (19), 13156 (19), 13157 (19), Valls,J.F.M. & G.P. Silva 10309(53), 10310(53). 13158(19), 13159 (30), 13160 (31), 13161 (31), 13162 Valls,J.F.M., A. Pott,L. Jank& G.P. Silva 10322 (31),13165 (31), 13167 (31), 13172 (31), 13173 (31), (37), 10323(37), 10324(37), 10357(38), 10384(8), 13175(31), 13177 (19), 13182 (31), 13184 (19), 13185 10386(12), 10387(12), 10390(12), 10392(42a), (19),13189 (28), 13193 (28), 13197 (30), 13198 (31), 10396(12), 10397 (8), 10404(33), 10405 (68), 10407 13200(31),13202(26), 13203 (26), 13211 (31). (8), 10415(42a), 10416 (9), 10420(68), 10426(12), Vails,J.F.M. 13250 (66). 10427(12), 10429 (7), 10430(42a), 10433 (7), 10445 Vails, J.F.M.,C.E. Simpson,R.N. Pittman,D.E. (39),10447 (39), 10454 (68), 10467 (54), 10468 (36), Williams& G.P. Silva 13260(53), 13262(53), 10469(36),10470(62), 10471 (62), 10473 (20), 10476 13273(42a). (62),10477 (62), 10506 (68), 10507 (68). Vails, J.F.M.,C.E. Simpson,R.N. Pittman,E.A. Vails,J.F.M., V.R. Rao, & G.P. Silva 10833(27), Pizarro& R.C. dosSantos 13282 (31), 13286(17),

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13288(31), 13290(58), 13294(31), 13296(17), 1273(32), 1291 (32). 13298(31), 13305(17), 13306(27), 13307(58), Vanni,RO., S. Cáceres,G. López& A. Radovancich 13310(31), 13312(31), 13315(31), 13317(17). 1470(63). Valls,J.F.M. & R.C. dosSantos 13322 (29). Veiga,R.F.A. 66 (53),258 (53). Valls,J.F.M. & E.A. Pizarro13326 (30). Venturi,S. 8354(45). Valls,J.F.M., E.A. Pizarro& W.L. Werneck13328 Vieira,J.G.A., A. Amaya& L.A. Cáceres288 (42a), (31), 13330(31). 290(42a), 291 (42a), 292 (42a). Valls,J.F.M. 13335 (31). Vorano,A.E. s/n(51). Valls,J.F.M. & Ek 13341(31). Waechter,J.L. 1021(40). Valls,J.F.M. 13342 (31). Warming,E. 2998(30). Valls,J.F.M., E.A. Pizarro,B. Maass,S.E.S. Valente Weddell,H.A. 2922 (20). & Db 13348(58), 13350(58), 13351(31), 13352 Werneck,W.L. 1 (31),2 (31),3 (19), 17(31), 34 (31), (31), 13354(31), 13355(30), 13356(31), 13357 39 (42a),40 (42a),41 (31),47 (31). (31), 13358(31), 13363(31), 13364(31). Werneck,W.L. & P. Pinheiro178 (26), 179(27), 184 Valls,J.F.M., A.K. Singh& G.P. Silva 13371(58), (28),201 (26). 13382(29), 13383(29), 13391(30), 13392(29), Williams,D.E. 691 (66),697 (66),699 (66),809 (30), 13393(29), 13395(29), 13396(29), 13397(29), 860(56), 861 (56), 866 (57), 867 (49). 13400(29), 13403(27), 13404(28). Williams,D.E. & D. Claure1 1 17 (57), 1 1 18 (49), 1 120 (49). Vanni,RO., M.S. Ferrucci,P. Cowan,R Duré& A. Woolston,A.L. 117(42b). Schinini217 (42b),327 (52), 342 (42a), 345 (52), 350 Zamboni,B. BAB34763 (42a). (15a),359 (15a), 368 (15a), 399 (32), 400 (42a), 438 (15a). Zardini,E.M., N. Deginani,E. Ulibarri& F. Zuloaga Vanni,RO. & S. Cáceres643 (42a). 644(63), 1032(42a). Vanni,RO., L.I. Ferraro& M.S. Ferrucci1152 (42a), Zehntner36-906 (25).

List of Parent Numbers used in Crosses

(Gregory, M.P. & W.C. Gregory, 1979)

Note: Percentage of pollen staining is (*) For number 100 (sp. 29), the very low based on the average of several counts of 500 value of 59.6% was not assigned despite the pollen grains. Whenever possible, 10 counts fact that it was confirmedthree times in the were made foreach parentnumber. same individualwith counts of 500 grainseach.

Colletionno. sp. % pollenstain Colletionno. sp. % pollenstain

1 9788 GKP (11) 93.7-99.1 12 10406 GKP (18) 2 10582 GK (8) 88.4-98.8 13 9990 GKP (37) 97.6-99.0 3 9646 GKP (15b) 92.4-99.8 14 9764 GKP (9) 4 9837 GKP (2) 98.8-100 15 10127 GKP (17) 98.8-99.8 5 10568 GK (1) 89.6-99.8 16 9841 GKP (12) 94.2-99.2 6 PI 210554 (61) 80.0-96.2 17 10585 GK (15a) 89.2-100 7 10538 GKP (30) 96.2-99.8 18 10588 GK (8) 79.8-99.8 8 10038 GKP (51) 90.2-99.4 19 9484 K (50) 86.2-98.4 9 PI 263394 (24) 20 10598 GK (32) 92.8-99.4 10 9570 GKP (42a) 87.4-97.8 21 10034 GKP (34) 96.4-99.8 11 8012 K (45) 85.4-98.0 22 9530-31 GKP (63) 82.2-98.6

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Collettorino. sp. % pollenstain CoUetionno. sp. % pollenstain

23 9830 GKP (42a) 77.4-98.6 66 9591 GKP (42a) 92.0-97.2 24 9629 GKP (42a) 88.0-97.8 67 WAArcher (42a) 67.4-89.8 25 10561 GK (41) 68 577 HLKHe (41) 54.0-77.0 26 10240 GKP (19) 7.2-18.8 69 548 HLKHe (5) 27 10596 GK (42a) 72.0-80.4 70 19455KMoF (47) 97.3-99.0 28 9667 GKP (42a) 90.2-94.2 71 9645 GKP (42a) 29 9553 GKP (42a) 72 9797 GKP (42a) 94.2-98.8 30 10550 GK (42a) 73 364 HLK (40) 60.4-92.0 31 9634 GKP (41) 98.4-99.4 74 30005 KG (67) 99.2-99.8 32 PI 262130 (69al) 97.6-98.0 75 362 HLK (40) 92.8-100 33 9926 GKP (62) 76 9901 GKP (67) 98.8 34 10017 GKP (65) 88.0-98.2 77 9882 GKP (42a) 89.2-94.2 35 11462KC (15a) 91.2-99.0 78 9893 GKP (42a) 97.4-99.0 36 11488KC (15a) 85.0-100 79 572 HLKHe (14) 92.4-99.6 37 10602 GK (67) 91.8-99.8 81 PI 262121 (69b2) 96.4 38 17 HLP (40) 81 PI 262129 (69b2) 39 7988 K (51) 94.0-99.8 82 PI 261942 (69bl) 86.4-87.2 40 "Spantex" (69b4) 90.4 82 PI 261991 (69bl) 41 559 HLKHe (8) 91.4-99.4 82 PI 262075 (69bl) 42 410 HLK (53) 96.2-99.8 83 PI 262092 (69al) 43 10160 GKP (20) 95.6-96.0 84 9567 GKP (42a) 91.2-93.2 44 10174 GKP (20) 96.7-99.6 85 526 HLKHe (3) 97.4-98.0 45 9923 GKP (23) 86 16 HLP (40) 95.2-97.6 47 9812 GKP (13) 98.8-99.0 87 23 HLP (40) 85.0-97.0 49 9935 GKP (42a) 96.0-99.2 88 (NC 2) (69al) 95.8-97.0 50 9848 GKP (6) 84.6-98.2 89 (NC 5) (69al) 97.6 51 9993 GKP (37) 90 27 HLP (40) 49.8-64.6 52 10002 GKP (37) 90.2-100 91 354 HLKOj (40) 75.6-97.0 53 10543 GK (5) 92 (Pearl) (69) 54 10556 GK (10) 93 (TMV2) (69) 56 10588 GK (8) 94 569 HLKHe (42a) 85.4-95.0 57 9966 GKP (42a) 89.0-92.6 95 576 HLKHe (41) 56.8-83.4 60 10559 GK (41) 91.6-97.8 96 12787 GK (31) 75.4-99.8 61 10565 GK (41) 69.4-94.4 97 12881 GK (25) 63 565 HLKHe (15b) 82.4-100 98 12922 GK (25) 91.0-99.4 64 349 HLKOj (42a) 71.2-94.2 99 12943 GK (29) 95.2-97.6 65 551 HLKHe (8) 92.8-97.2 100 12946 GK (29) *90.2-99.8

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Abbreviations of Collectors' Names

A, A.C.Allem,CENARGEN, Brasilia, Brasil. Oj, H.R.Ojeda,Fac. de CienciasAgrarias, Corrientes, Ai, O.Arriola,INIA, Cuzco, Peru. Argentina. Aj, M.Araújo Ok, K.Okada,CIAT, Cali, Colombia. Ar, W.A.Archer,USD A Ov, J.C.Oliveira, EMBRAPA, CNPO, Bagé RS, Bra- B, D.J.Banks,USDA sil. Ba, J.R.Báez,EEA. Manfredi, Córdoba, Argentina. P, J.R.Pietrarelli,EEA, Manfredi,Córdoba, Argenti- Bi, L.B.Bianchetti,CENARGEN, Brasilia, Brasil. na. Bm, B. Maass,CIAT, Cali, Colombia. Pe, M.I.Penteado C, C.L.Cristóbal,IBONE, Corrientes, Argentina. Pm, R.N.Pittman,USDA, Griffin, Georgia, U.S.A. Co, L.Coradin,CENARGEN, Brasilia, Brasil. Po, A.Pott,EMBRAPA, Corumbá, MS, Brasil. Cr, M.Corro,Univ. J.M.Saracho, Tarija,Bolivia. Pz, E.A.Pizarro,CIAT/CPAC-EMBRAPA, Planaltina, F, A.Fernández,IBONE, Corrientes, Argentina. DF, Brasil. Fa, L.Faracode Freitas, CENARGEN, Brasilia, Brasil. Q, C.L.Quarin,IBONE, Corrientes, Argentina. Fd, M.SoterFrança Dantas, CP AC, EMBRAPA, R, V.R.Rao,ICRISAT, India. Planaltina,DF, Brasil. Ri, V.A.Rigoni,EEA Manfredi,Argentina. G, W.C.Gregory,North Carolina State University, Ro, D.M.S.Rocha,CENARGEN, Brasilia, Brasil. Raleigh,NC, USA. Rs, RoseaneC. dos Santos,CNP A, CampinaGrande, Ga, M.L.Galgaro,UNESP, Botucatu, SP, Brasil. Paraiba,Brasil. Gb, R.W.Gibbons, ICRISAT, India. S, C.E.Simpson,Texas A&M University, Stephenville, Gd, I.J.de Godoy,IAC, SP, Brasil. Texas,USA. Ge, M.A.N.Gerin,IAC, SP, Brasil. Sa, H.T. Stalker,North Carolina State University, Gr, A.Gripp,CENARGEN, Brasilia, Brasil. Raleigh,NC, USA. H, R.O.Hammons,USDA Sc, A.Schinini,IBONE, Corrientes, Argentina. He, V.Hemsy,Facultad de Agronomía,Tucumán, Ar- Sg, A.K.Singh, ICRISAT, India. gentina. St, J.L.Stephens,USDA Ht, W.Hartley,SCIRO, Australia. Sv, G.P.Silva,EMBRAPA, Brasilia, Brasil. J, L.Jank,CNPGC, EMBRAPA,Campo Grande, V, J.F.M.Valls, CENARGEN, Brasilia, Brasil. MS, Brasil. Va, S.E.S.Valente,UNESP, Botucatú, SP, Brasil. Jk, L.Janicki,PRODES, La Paz,Bolivia. Ve, R.F.deArruda Veiga, IAC, SP, Brasil. K, A.Krapovickas,IBONE, Corrientes, Argentina. Vn, R.O.Vanni,IBONE, Corrientes, Argentina. L, W.R.Langford,USDA W, W.L.Werneck,CENARGEN, Brasilia, Brasil. M, J.P.Moss,ICRISAT, India. Wi, D.E. Williams,USDA. Mo, L.A.Mroginski,IBONE, Corrientes, Argentina. Z, O.Zurita, Estación Exp. Agrícola,Saavedra, Mr, A.R.Miranda,CENARGEN, Brasilia, Brasil. S.Cruz,Bolivia. O, J.Ramosde Otero,S.Agrostologia, M.Agric., Rio Zi, R.H.Zanini,INTA, Manfredi, Argentina. de Janeiro,BR. Zn, C.N.Zanin,IBONE, Corrientes, Argentina.

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Interspecific Crosses Conducted10

Crosses within each section

Sect.I. TR1ERECTOIDES

A. tuberosax (4) A.guaranitica 4x5 5x4

Sect.II. ERECTOIDES

A. paraguariensissubsp. paraguariensisx (3,17,35,36) A.paraguariensis 3 x 35 24% 35x 3 70.3% A.paraguariensis 3x36 44.3% 36x 3 42.1% A.paraguariensis 35 x36 52.3% 36x35 61.8% A. stenophylla 3x79 16.8% A. Hermannii 3x16 4.3% 16x3 3.5% A. Hermannii 36x 16 0.9% A.gracilis 1x3 1.7% 3x1 0.3% A. major 3x 2 0.65% A. major 3x65 0.1% A. major 2 x 35 0.5% A. Benthamii 3 x 14 0.13% A. Benthamii 35 x 14 A. Oteroi 3 x 53 0.2% 53 x 3 A.Oteroi 35 x53 1.8% A. Oteroi 36 x 53

A. gracilisx 0) A. major 1 x 2 30% 2x1 A. Hermannii 16x 1 20.1% A. Martii 85 x 1 6.2%

A. major x (2,18,41,56,65) A. major 2x41 11.7% A.Ar cher i 47x56 40.4% 56x47 17.6%

A. Hermanniix (16) A. Benthamii 16x14 19.1% A.Oteroi 16x53 17.6%

,0Interspecificcrosses conducted by M.P. Gregory & W.C.Gregory (1979), with the parents' numbers (pp. 192- 193)and the average value (%) ofstained pollen in the hybrids obtained.

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A. Hatschbachiix (50) A.Archeri 50 x 47

A. Mariti x (85) A. stenophylla 85 x 79 0.07% Sect.III. EXTRANERVOSAE

A. villosulicarpax (9) A. marginata 9x 12 A. Macedoi 9x15 A.prostrata 9x26 0.1% A. lutescens 9 x 43 0.2% 43 x 9 0.3% A. lutescens 9 x 44 0.2%

Sect.IV. TRISEMINATAE

A. triseminatax (97) A. triseminata 97 x 98 68.2% 98 x 97 50.7%

Sect.V. HETERANTHAE

A. Dardani x (99) A. Dardani 99x 100 20.6% 100x99 20.5%

Sect.VI. CAULORRHIZAE

A. Pintoi x (96) A. repens 96 x 7 86.8% Sect.VII. PROCUMBENTES

A. appressipilax (13,52) A. lignosa 13x 20 30.9% 20 x 13 26.8% A. lignosa 20 x 52 29.2%

A. Rigonii x (21) A. lignosa 20 x 2 1 54.4% 2 1 x 20 54.4% A. appressipila 21 x 52 40.2% Sect.VIII . RHIZOMATOSAE

A,glabrata v. glabratax (10,23,24,27,77,84) A.glabrata v. glabrata 10x 23 78.1% A. glabratav. glabrata 10x 27 27 x 10 A.glabrata v. glabrata 10x 84 12.9% A.glabrata v. glabrata 24 x 10 91.3% A. glabratav. glabrata 23 x 77 45.2%

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A.glabrata v. glabrata 24 x 27 92.9% 27 x 24 A.glabrata v. glabrata 24 x 77 71.3% A.glabrata v. glabrata 27 x 23 68.6% A.glabrata v. glabrata 27x77 84% 77x27 68.7% pseudovillosa 60 x 10 pseudovillosa 60 x 27

Sect.IX. ARACHIS

Annuals x Annuals

A. duranensisx (8 & 39) A. duranensis 8 x 39 31.3% 39x 8 36.7% A. stenosperma 8x42 17.0% 42x8 18.1% A. stenosperma 39 x 42 21.2% 42 x 39 15.5%

A. Batizocoix (19) A. duranensis 8x 19 0.9 19x 8 0.01% A. duranensis 19x39 4.9 39x 19 17.4% A. stenosperma 19x 42 0.0 42 x 19 0.7% A. ipaënsis 19x70 0.5%

A. ipaënsisx (70) A. duranensis 70 x 8 1.5% A. duranensis 70 x 39 3.7%

Annuals x Perennials

A. duranensisx (8 & 39) A. villosa 8x 6 27.7% 6x8 A. correntina 8x22 41.7% 22x8 A. Cardenasii 8 x 34 52.2% 34 x 8 47.5% A. Diogoi 8 x 37 27.1% 37 x 8 38.8% A. villosa 39 x 6 22.3% 6 x 39 42.3% A. correntina 39 x 22 54.3% 22 x 39 A. Cardenasii 39x34 49.2% 34x39 71.2% A. Diogoi 39x37 33.0% 37x39

A. stenospermax (42) A. villosa 42 x 6 22.5% 6 x 42 27.8% A. correntina 42x22 37.6% 22x42 32.2% A. Cardenasii 42 x 34 50.5% 34 x 42 A. Diogoi 42x37 88.5% 37x42 36.6%

A. Batizocoix (19) A. villosa 19x6 0.2% 6x19 2.9% A. correntina 19x 22 0.97% 22 x 19 A. Cardenasii 19x34 0.07% 34x19 0.07 A. Diogoi 19x37 0.1 %

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Annuals x Tetraploids A. duranensisx (8 & 39) A. montícola 8x11 A. hypogaeav. hypogaea 8x32 18.9% A. hypogaeav. peruviana 8x81 7.8% A. hypogaeav. fastigiata 8x82 A. hypogaeav. hypogaea 8 x 89 3.1% A. hypogaeav. hypogaea 39 x 88 5.2%

A. stenospermax (42) A. montícola 42 x 11 3.3% A. hypogaeav. fastigiata - 82x42 4.8%

A. Batizocoix (19) A. hypogaeav. fastigiata 19x82 0.2% 82x 19 3.5%

A. ipaënsis x (70) A. hypogaeav. hypogaea 70 x 32

Perennialsx Perennials

A. villosax (6) A. correntina 6x22 73.1% 22 x 6 67.3% A. Cardenasii 6 x 34 70.4% 34 x 6 68.4% A. Diogoi 6 x 37 28.9% 37 x 6

A. correntinax (22) A. Diogoi 22x37 32.2% 37x22 35.8%

A. Cardenasiix (34) A. correntina 34 x 22 63.8% A. Diogoi 34x37 48.3% 37x34 44.8%

Perennialsx Tetraploids A. villosax (6) A. montícola 6x 11 11x 6 45.7% A. hypogaeav. fastigiata 6 x 82 A. Cardenasii x (34) A. montícola 11x34 1.0% A. hypogaeav. hypogaea 34 x 32 5.7% A. hypogaeav. hypogaea 83 x 34 28.3% A. hypogaeav. hypogaea 88 x 34 A. hypogaeav. hypogaea 89 x 34 9% A. hypogaeav. fastigiata 81 x34 2%

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A. Diogoi X (37) A. montícola 11 x 37 2.5% A. hypogaea 37 x 82 7.5% A. hypogaeav. hypogaea 37 x 83 18.5% 83 x 37 30.0% A. hypogaeav. hypogaea 88 x 37 3.8% A. hypogaeav. hypogaea 37 x 89 5.3% 89 x 37 3.6% A. hypogaeav. fastigiata 81 x 37 6.6%$

Crosses between sections

Sect.I. TR1ERECT01DESx II. ERECTOIDES

A. tuberosax (4) A.gracilis 4x 1 11.8% 1x4 18.7% A. major 4x 2 2x4 4%

A. guaraniticax (5) A.gracilis 5x 1 10.3% 1x5 3% A. major 5x2 4.8% 2x5 2.6% A.paraguariensis subsp. paraguariensis 5x3 1.1% 3x5 0.4% A.paraguariensis subsp. paraguariensis 5x35 35x5 0.2%

Sect.I. TRIERECTOIDESx VII. PROCUMBENTES

A. guaraniticax (5) A. appressipila 13x5 0.7%

Sect.II. ERECTOIDESx V. HETERANTHAE

A.paraguariensis subsp. paraguariensisx (3,35) A. Dardani 3 x 99 0.6% A. Dardani 35 x 99 2%

Sect.II. ERECTOIDESx VI. CAULORRHIZAE

A. major x (3,18,41) A. repens 3x7 A. repens 18x7 A. Pintoi 96x41

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A. paraguariensissubs p. paraguariensisx (17,36) A. repens 17x 7 6% A. Pintoi 36 x 96 3.5%

Sect.II. ERECTOIDESx VII. PROCUMBENTES

A. paraguariensissubsp. paraguariensisx (3,17,35,36) A. appressipila 3x13 13x3 A. appressipila 3x52 1.8% 52 x 3 4.5% A. appressipila 35x13 1.3% A. appressipila 35 x 52 3.9% A. appressipila 36x13 1.3% A. Rigonii 21 x 3 1.7% A. Rigonii 21 x 17 4.9% A. Rigonii 35x21 21 x35 1.2% A. Rigonii 36x21 1% 21x36 1%

A. major x (18) A. lignosa 18x20 0.1%

A. appressipilax (13,52) A. major 2 x 52 0.7% A. major 13x 2 0.5% A. Benthamii 13x14 A. Hermannii 13x16 4?2% 16x13 4.8% A. Hermannii 16x52 1.9% A. Martii 13x85 0.4%

A. Rigonii x (21) A. Hermannii 16x21 6.9% 21x16 6% A.gracilis 21x1 12.2% A. major 21 x 2 0.6% A. major 21 x 18 0.3% A. major 21x41 1% A. Benthamii 21 x 14 2.6% A.Archeri 21 x47 0.8% A. Oteroi 21 x 53 A. Martii 21 x85 0.3%

Sect.II. ERECTOIDESx VIII.RHIZOMATOSAE

A. major x (2,18,41) A.glabrata v. glabrata 2x10 A. glabratav. glabrata 2 x 24 A.glabrata v. glabrata 2 x 27 A.glabrata v. glabrata 18x 10 A.glabrata v. glabrata 18x 27 A.glabrata v. glabrata 41 x 27 A.glabrata v. glabrata 41 x 94 29.6%

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A. paraguariensissubsp. paraguariensisx (3,17,35,36) A.glabrata v. glabrata 3 x 72 A.glabrata v. glabrata 17x10 A.glabrata v. glabrata 17x 24 A.glabrata v. glabrata 17x 27 A.glabrata v. glabrata 35 x 10 A.glabrata v. glabrata 36 x 10 A.glabrata v. glabrata 36 x 24 A.glabrata v. glabrata 36 x 27 A.glabrata v. glabrata 36 x 66 A.glabrata v. glabrata 36 x 72 A.pseudovillosa 36 x 60

A. Benthamiix (14) A.glabrata v. glabrata 14x10 A.glabrata v. glabrata 14x 24

A. Hermanniix (16) A.glabrata v. glabrata 16x10 A.glabrata v. glabrata 16x 24 A.glabrata v. glabrata 16x 27 zi.glabrata v. glabrata 16x 94 A.pseudovillosa 16x60 46.1% 60x16

Sect.II. ERECTOIDESx IX.ARACHIS

A. duranensisx (8,39) A gracilis 8 x 1 A. Hermannii 16x39 1.2%

Sect.III. EXTRANERVOSAE x V. HETERANTHAE

A. Macedoi x (15) A. Dardani 100x15

Séct.VI. CAULORRHIZAEx VII. PROCUMBENTES

A. Pintoi x (96) A. lignosa 96 x 20 0.2% A. Rigonii 96x21

Sect.VII. PROCUMBENTESx VIII.RHIZOMA TOSAE

A. appressipilax (13) A.glabrata v. glabrata 13x 10 22.2% A.glabrata v. glabrata 13x 27

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A. Rigonii X (21) A.glabrata v. glabrata 21 x 23 14.9% A.glabrata v. glabrata 21 x 27 A.pseudovillosa 21 x 25 A.pseudovillosa 21 x 31

Sect.IX. ARACHISx VIII.RHIZOMATOSAE

A. duranensisx (8,39) A.glabrata v. glabrata 8x10 A.glabrata v. glabrata 8 x 23 A.glabrata v. glabrata 8 x 24 A.glabrata v. glabrata 8 x 27 A.glabrata v. glabrata 8 x 28 A. glabratav. glabrata 8 x 29 A. glabratav. glabrata 8 x 64 A.glabrata v. glabrata 8 x 84 A.glabrata v. glabrata 39 x 27 A glabratav. glabrata 39 x 84 A glabratav. glabrata 39 x 95 pseudovillosa 8x31

A. Batizocoix (19) A. glabratav. glabrata 19x 23 A.glabrata v. glabrata 19x 24 A.glabrata v. glabrata 19x 27 glabratav. glabrata 19x 29 A glabratav. glabrata 19x 30 >4.glabrata v. glabrata 19x 64 A glabratav. glabrata 19x 72 >4.glabrata v. glabrata 19x 94 0.03% A pseudovillosa 19x31 A.pseudovillosa 19x60

/4.stenosperma x (42) A glabratav. glabrata 42 x 10 14□

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List of Scientific Names Cited in Text11'12

Aeschynomeneae,12 sect.Rhizomatosae Krapov. & W.C. Gregorynomen Arachidna[Plumier] Boehmer, 28 nudum,113 Arachidnahypogaea (L.) Moench,167 sect.Rhizomatosae ser. Prorhizomatosae Krapov. & W.C. Arachidnaquadrifolia Trew, 167, 175 Gregorynov. ser., 34, 114 ArachisL., 14,25, 27, 28 sect.Rhizomatosae ser. Prorhizomatosae Krapov. & W.C. sect.Ambinervosae Krap. nomen nudum , 85 Gregoiynomen nudum, 114 sect.Arachis, 24, 28, 34, 123, 197, 201, 202 sect.Rhizomatosae ser. Rhizomatosae, 34, 117 sect.Axonomorphae Krapov. & W.C. Gregorynomen séct.Tetraerectoides Krapov. nomen nudum, 40 nudum, 123 sect.Trierectoides Krapov. & W.C.Gregory nov. sect., 28, sect.Axonomorphae ser. Amphiploides Krapov. & W.C. 33,34, 195, 199 Gregorynomen nudum , 123 sect.Triseminalae Krapov. & W.C. Gregorynomen sect. Axonomorphaeser. AnnuaeKrapov. & W.C. nudum,83 Gregorynomen nudum , 123 sect.Triseminatae Krapov. & W.C.Gregory nov. sect., 28, sect.Axonomorphae ser. PerennesKrapov. & W.C. 33,83, 196 Gregorynomen nudum , 123 A. africanaLour., 172 sect.Caulorrhizae Krapov. & W.C.Gregory nov. sect A. americanaTenore, 175 28,33, 94, 196, 199, 201 A. angustifolia(Chodat & Hassl.)Killip ex Hoehne, 122 sect.Caulorrhizae Krapov. & W.C. Gregorynomen A. angustifoliaauct. non (Chodat & Hassl.)Killip ex nudum, 94 Hoehne,61 sect.Erectoides Krapov. & W.C.Gregory nov. sect., 24, A.appressipila Krapov. & W.C.Gregory nov. sp., 14, 15, 28,33, 40, 195, 199, 200, 201 28,107, 196, 199, 200, 201 sect.Erectoides ser. Procumbensae Krapov. & W.C. A.Archeri Krapov. & W.C.Gregory nov. sp., 13, 16, 59, Gregorynomen nudum , 99 195,200 sect.Erectoides ser. Tetrafoliolatae Krapov. & W.C. A. argentinensisSpeg, nomen nudum, 138 Gregorynomen nudum , 40 A. asiaticaLour., 173 sect. Erectoidesser. TrifoliolataeKrapov. & W.C. A. batizocaeKrapov. & W.C.Gregory nomen nudum, 137 Gregorynomen nudum , 34 A.Batizocoi Krapov. & W.C.Gregoiy, 17, 137, 197, 198, 202 sect.Extranervosae Krapov. & W.C.Gregory nov. sect., A.Xbatizogaea Krapov. & Av.Fernández, 170 24,28, 33, 67, 196, 201 A. benensisKrapov., W.C. Gregory & C.E. Simpsonnov. sect.Extranervosae Krapov. & W.C. Gregorynomen sp.,26, 147 nudum, 67 A.Benthamii Handro, 13, 16, 40, 52, 195, 200, 201 sect.Goniorhiza Resslar nomen nudum , 15 A. brevipetiolataKrapov. & W.C.Gregory nov. sp., 43 secí.Heteranthae Krapov. & W.C.Gregory hov. 28, A.Burchellii Krapov. & W.C.Gregory nov. sp., 78 33,85, 196, 199, 201 A.Burkartii Handro, 13, 14, 15, 16, 28, 114 sect.Procumbentes Krapov. & W.C.Gregory nov. sect., A. CardenasiiKrapov. & W.C.Gregory nov. sp., 14, 15, 26,28, 33, 99, 196, 199, 200, 201 17,158, 197, 198 sect.Pseudoaxonomorphae Krapov. & W.C.Gregory no- A.Cardenasii Krapov. & W.C.Gregoiy nomen nudum, 158 mennudum , 85 A.chacoense Krapov. & W.C.Gregory nomen nudum, 162 sect.Rhizomatosae Krapov. & W.C.Gregory nov. sect., A. chiquitanaKrapov., W.C. Gregoiy& C.E. Simpson 26,28, 34, 113, 196, 200, 201, 202 nov.sp., 105 A. correntina(Burkart) Krapov. & W.C. Gregoiynov. "Page numbersfor descriptions of valid names comb.,13, 16, 155, 197, 198 appearin bold. A. correntina(Burkart) Krapov. & W.C.Gregoiy nomen ,2Theauthors' abbreviations are those recommended nudum,155 byBrummitt & Powell (1992). A. cruzianaKrapov., W.C. Gregory& C.E. Simpson

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nov.sp ., 129 A.hypogaea forma nambyquarae (Hoehne) F.J. Herm., 173 A.cryptopotamica Krapov. & W.C.Gregory nov. sp ., 48 A. hypogaeaforma typica Hoehne, 173 A.Dardani Krapov. & W.C.Gregory nov. sp., 14, 85, 91, A. hypogaeasubsp. africana Bois, 172 196,199, 201 A. hypogaeasubsp. africana var. communis A. Chev.,172 A.decora Krapov., W.C. Gregory & Valls nov. sp ., 149 A hypogaeasubsp. africana var. microcarpa A Chev., 172 A.Diogoi Hoehne, 13, 15, 162, 197, 198, 199 A. hypogaeasubsp. africana var. robustior A. Chev., 172 A. Diogoiauct. non Hoehne, 59, 61 A. hypogaeasubsp. africana var. stenocarpa A. Chev., 175 A. DiogoiHoehne forma minor Hoehne nomen nudum , 59 A. hypogaeasubsp. asiatica (Lour.) Bois, 173 A. DiogoiHoehne forma sericeo-villosa Hoehne nomen A. hypogaeasubsp. asiatica var. erecta A. Chev.,175 nudum,59 A.hypogaea subsp. asiatica var. macrocarpa A.Chev., 172 A. Diogoi Hoehneforma subglabrata Hoehne nomen A.hypogaea subsp. fastigiata Waldron, 173 nudum, 59 A. hypogaeasubsp. fastigiata var. aequatoriana Krapov. A. DiogoiHoehne forma submarginata Hoehne nomen & W.C.Gregory nov. var., 174 nudum, 61, 63 A.hypogaea subsp .fastigiata var. fastigiata, 173, 198, 199 A. DiogoiHoehne subsp. major Hoehne nomen nudum , 49 A. hypogaeasubsp. fastigiata var. peruviana Krapov. & A.douradiana Krapov. & W.C.Gregory nov. sp ., 54 W.C.Gregory nov. var., 174, 198 A. duranensisKrapov. & W.C.Gregory nov. sp ., 13,14, A.hypogaea subsp. fastigiata var. vulgaris Harz, 175 16,17, 24, 25, 26, 28, 138, 197, 198, 201, 202 A.hypogaea subsp. hypogaea, 171 A.duranensis Krapov. & W.C.Gregory nomen nudum , 138 A.hypogaea subsp. hypogaea var. hirsuta Köhler, 27, 173 A. GiacomettiiKrapov., W.C. Gregory,Valls & C.E. A.hypogaea subsp. hypogaea var. hypogaea, 172, 198, 199 Simpsonnov. sp ., 86 A.hypogaea subsp. nambyquarae (Hoehne) A. Chev.,172 A. glabrata,13, 14, 15, 16 A. hypogaeasubsp. oleifera A. Chev.,167 A.glabrata Benth. forma major Hoehne nomen nudum , 119 A. hypogaeasubsp. procumbens Waldron, 172 A.glabrata Benth. forma minor Hoehne nomen nudum , 119 A. hypogaeasubsp. rasteiro (A. Chev.)A. Chev.,172 A.glabrata Benth. subsp. rasteiro (A. Chev.) A. Chev., 173 A. hypogaeasubsp. sylvestris A. Chev., 87 A. glabrataBenth. var. glabrata , 14, 25, 119,196, 200, A. hypogaeavar. aegyptiaca Hassk., 172 201,202 A. hypogaeavar. africana Giróla, 172 A. glabrataBenth. var. Hagenbeckii (Harms ex Kuntze) A. hypogaeavar. africana Kurtz, 173 F.J.Herm., 14, 122 A. hypogaeavar. asiatica (Lour.) Giróla, 173 A. glabrataBenth. var. membranifolia [Benth.?] ex A. A. hypogaeavar. asiatica forma oscura Giróla, 173 Chev.nomen nudum , 1 19 A. hypogaeavar. asiatica forma rosada Giróla, 173 A.glandulifera Stalker, 127 A. hypogaeavar. communis A. Chev.subvar. violacea A.gracilis Krapov. & W.C.Gregory nov. sp ., 55, 195, 199, Burkart,173 200,201 A. hypogaeavar. glabra DC, 172 A. guaranianaBertoni, 172 A. hypogaeavar. indica Kurtz, 172 A. guaraniticaChodat & Hassl.,13, 14, 15,27, 34,35, A. hypogaeavar. nambyquarae (Hoehne) Burkart, 172 195,199 A. hypogaeavar. reticulata Harz, 175 A. guaraniticaBertoni, 175 A.ipaënsis Krapov. & W.C.Gregory nov. sp., 17, 27, 132, A. HagenbeckiiHarms ex Kuntze, 13, 122 197,198 A.Hatschbachii Krapov. & W.C.Gregoiy nov. sp., 47, 196 A. ipaensisW.C. Gregory & M.P. Gregory nomen nudum, A.helodes Mart. ex Krapov. & Rigoni,14, 15, 24, 154 132 A. helodesMart. ex Hoehnenomen nudum , 42 A. KempffMercadoiKrapov., W.C. Gregory& C.E. A. HermanniiKrapov. & W.C.Gregory nov. sp., 58, 195, Simpsonnov. sp., 160 200,201 A.Kretschmeri Krapov. & W.C.Gregory nov. sp., 102 A.Herzogii Krapov., W.C. Gregory & C.E. Simpsonnov. A.Kuhlmannii Krapov. & W.C.Gregory nov. sp., 164 sp.,150 A. lignosa(Chodat & Hassl.)Krapov. & W.C.Gregory A.Hoehnei Krapov. & W.C.Gregory nov. sp., 15, 141 nov.comb., 14, 15, 24, 100, 196, 200, 201 A. hypogaeaL., 13,14, 15, 16, 20, 21, 27, 123, 167 A. lignosa(Chodat & Hassl.)Krapov. & W.C.Gregory A. hypogaeaforma communis (A. Chev.)F.J. Herm., 173 nomennudum, 100 A. hypogaeaforma macrocarpa (A. Chev.) Hoehne, 173 A.lutescens Krapov. & Rigoni,14, 15, 75, 196 A. hypogaeaforma microcarpa (A. Chev.) Hoehne, 173 A.Macedoi Krapov. & W.C.Gregory nov. sp., 69, 196, 201

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A. MacedoiKrapov. & W.C.Gregory nomen nudum , 69 A. prostrataBenth. var. pseudomarginata Chodat & Hassl. A. magnaKrapov., W.C. Gregory & C.E. Simpsonnov. formaangustifolia Chodat & Hassl.ß longicalyx,122 sp., 24,131 A. prostrataBenth. var. pseudovillosa Chodat & Hassl., A. majorKrapov. & W.C.Gregory nov. sp., 13, 16, 49, 117 195,199, 200 A. prostrataBenth. var. villosa (Benth.) A. Chev.,152 A.marginata Gardner, 13, 14, 15, 72, 196 A. pseudovillosa(Chodat & Hassl.)Krapov. & W.C. A. marginataauct. non Gardner, 45, 115 Gregorynov. comb., 14, 15, 117, 197, 201, 202 A.marginata Gardner forma submarginata Hoehne, 53, 1 1 5 A.pusilla Benth., 13, 14, 24, 90 A. marginataGardner subsp. Hagenbeckii (Harms ex A. pusillaauct. non Benth., 42, 83, 129 Kuntze)A. Chev.,122 A. rasteiroA. Chev.,172 A. marginataGardner var. lignosa (Chodat & Hassl.)A. A.repens Handro, 14, 95, 196,199 Chev.,100 A.retusa Krapov., W.C. Gregory & Vails nov. sp., 77 A. MartiiHandro, 16, 42, 195,196, 200 A. RigoniiKrapov. & W.C.Gregory, 17, 26, 28, 99, 104, A.matiensis Krapov., Gregory & C.E. Simpson nov. sp., 106 196,200, 202 A.microsperma Krapov., W.C. Gregory & Vails nov. sp, 151 A.setinervosa Krapov. & W.C.Gregory nov. sp., 68 A.montícola Krapov. & Rigoni,13, 14, 16, 24, 25, 27, 28, A.Simpsonii Krapov. & W.C.Gregory nov. sp., 157 129,198, 199 A. SpegazziniiW.C. Gregory& M.P. Gregorynomen A. nambyquaraeHoehne, 15, 172 nudum,138 A. OteroiKrapov. & W.C.Gregory nov. sp., 13, 14, 16, A.stenophylla Krapov. & W.C.Gregory nov. sp., 61, 195, 45, 195,200 196 A.palustris Krapov., W.C. Gregory & Vails nov. sp., 146 A.stenosperma Krapov. & W.C.Gregory nov. sp., 14, 15, A.par aguar iensis, 13, 15, 26, 63 17,26, 27, 28, 142, 197, 198, 202 A. paraguariensisChodat & Hassl.subsp. capibarensis A. stenospermaW.C. Gregory& M.P.Gregorynomen Krapov.& W.C.Gregory nov. subsp., 66 nudum,142 A. paraguariensis Chodat & Hassl. subsp. A.subcoriacea Krapov. & W.C.Gregory nov. sp., 110 paraguariensis,24,28, 63, 195,199, 200, 201 A.sylvestris (A. Chev.)A. Chev.,13, 14, 26, 87 A.Pietrarellii Krapov. & W.C.Gregory nov. sp., 80 A.trinitensis Krapov. & W.C.Gregory nov. sp., 149 A.Pintoi Krapov. & W.C.Gregory nov. sp., 96, 196, 199, A. inseminataKrapov. & W.C.Gregory nov. sp., 23, 25, 200,201 83,196 A. PintoiKrapov. & W.C.Gregory nomen nudum, 96 A. tuberosaBongard ex Benth., 13, 14, 15, 39, 195,199 A.praecox Krapov., W.C. Gregory & Vails nov. sp., 145 A.valida Krapov. & W.C.Gregory nov. sp., 25, 134 A. procumbensBerneaud, 173 A. VallsiiKrapov. & W.C.Gregory nov. sp., 110 A.prostrata Benth., 13, 14, 15, 67, 72, 196 A. villosaBenth., 13, 15, 16, 151, 198 A. prostrataBenth. forma Hagenbeckii (Harms ex A. villosaBenth. subsp. Diogoi (Hoehne) A. Chev.,162 Kuntze)Hoehne, 122 A. villosaBenth. var. correntina Burkart, 155 A. prostrataBenth. subsp. Hagenbeckii (Harms ex A.villosulicarpa Hoehne, 16, 24, 27, 81, 196 Kuntze)Hoehne, 122 A. WilliamsiiKrapov. & W.C.Gregory nov. sp., 27, 136 A. prostrataBenth. var. angustifolia Chodat & Hassl.ex Arthrocarpum,12 Hoehne,122 Cercosporaarachidicola, 146 A. prostrataBenth. var. genuina Chodat & Hassl.,1 19 Chapmannia,12, 13 A. prostrataBenth. var. genuina forma lignosa Chodat & Glycinesubterránea auct. non. L., 175 Hassl.,100 Hedysareae,12, 13 A. prostrataBenth. var. intermedia Chodat & Hassl.,63 Mandubí[Marcgr.] Adans., 31 A. prostrataBenth. var. lignosa Chodat & Hassl.,119 Pachecoa,12, 13 A. prostrataBenth. var. pseudomarginata Chodat & Poiretinae,12 Hassl.,122 Stylosanthes,12,13, 28 A.prostrata Benth. var. pseudomarginata Chodat & Hassl. S. leiocarpaauct. non Benth., 152 formaangustifolia Chodat & Hassl.,122 Stylosanthinae,12,13 A.prostrata Benth. var. pseudomarginata Chodat & Hassl. Zornia,12, 13 formaangustifolia Chodat & Hassl.a brevicalyx,122

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