David M. Spooner

Reclassification of landrace populations of cultivated potatoes (Solanum sect. Petota)

Cultivated potatoes have been classified as species under the International Code of Botanical Nomenclature (ICBN) and as cultivar-groups under the International Code of Nomenclature of Cultivated Plants (ICNCP); both classifications are still widely used. This study examines morphological support for the classification of landrace populations of cultivated potatoes, using representatives of all seven species and most subspecies as outlined in the latest taxonomic treatment. These taxa are S. ajanhuiri, S. chaucha, S. curtilobum, S. juzepczukii, S. phureja subsp. phureja, S. stenotomum subsp. stenotomum, S. stenotomum subsp. goniocalyx, S. tuberosum subsp. andigenum, and S. tuberosum subsp. tuberosum. The results show some phenetic support for S. ajanhuiri, S. chaucha, S. curtilobum, S. juzepczukii, and S. tuberosum subsp. tuberosum, but little support for the other taxa. Most morphological support is by using a suite of characters, all of which are shared with other taxa (polythetic support). These results, combined with their likely hybrid origins, multiple origins, evolutionary dynamics of continuing hybridization, and our classification philosophy, leads us to recognize all landrace populations of cultivated potatoes as a single species, S. tuberosum, with the eight cultivar-groups: Ajanhuiri Group, Andigenum Group, Chaucha Group, Chilotanum Group, Curtilobum Group, Juzepczukii Group, Phureja Group, and Stenotomum Group. We defer classification of modern cultivars, traditionally classified in Tuberosum Group, to a later study.

Geographic distribution of wild potato species

The geographic distribution of wild potatoes (Solanaceae sect. Petota) was analyzed using a database of 6073 georeferenced observations. Wild potatoes occur in 16 countries, but 88% of the observations are from Argentina, Bolivia, Mexico, and Peru. Most species are rare and narrowly endemic: for 77 species the largest distance between two observations of the same species is <100 km. Peru has the highest number of species (93), followed by Bolivia (39). A grid of 50 × 50 km cells and a circular neighborhood with a radius of 50 km to assign points to grid cells was used to map species richness. High species richness occurs in northern Argentina, central Bolivia, central Ecuador, central Mexico, and south and north-central Peru. The highest number of species in a grid cell (22) occurs in southern Peru. To include all species at least once, 59 grid cells need to be selected (out of 1317 cells with observations). Wild potatoes occur between 38° N and 41° S, with more species in the southern hemisphere. Species richness is highest between 8° and 20° S and around 20° N. Wild potatoes typically occur between 2000 and 4000 m altitude.

Granule-bound starch synthase (GBSSI) gene phylogeny of wild tomatoes (Solanum L. section Lycopersicon [Mill.] Wettst. subsection Lycopersicon)

Eight wild tomato species are native to western South America and one to the Galapagos Islands. Different classifications of tomatoes have been based on morphological or biological criteria. Our primary goal was to examine the phylogenetic relationships of all nine wild tomato species and closely related outgroups, with a concentration on the most widespread and variable tomato species Solanum peruvianum, using DNA sequences of the structural gene granule-bound starch synthase (GBSSI, or waxy). Results show some concordance with previous morphology-based classifications and new relationships. The ingroup comprised a basal polytomy composed of the self-incompatible green-fruited species S. chilense and the central to southern Peruvian populations of S. peruvianum, S. habrochaites, and S. pennellii. A derived clade contains the northern Peruvian populations of S. peruvianum (also self-incompatible, green-fruited), S. chmielewskii, and S. neorickii (self-compatible, green-fruited), and the self-compatible and red- to orange- to yellow-fruited species S. cheesmaniae, S. lycopersicum, and S. pimpinellifolium. Outgroup relationships are largely concordant with prior chloroplast DNA restriction site phylogenies, support S. juglandifolium and S. ochranthum as the closest outgroup to tomatoes with S. lycopersicoides and S. sitiens as basal to these, and support allogamy, self-incompatibility, and green fruits as primitive in the tomato clade.

Selection of highly informative and user-friendly microsatellites (SSRs) for genotyping of cultivated potato

Characterization of nearly 1,000 cultivated potato accessions with simple sequence repeats (SSRs; also referred to as microsatellites) has allowed the identification of a reference set of SSR markers for accurate and efficient genotyping. In addition, 31 SSRs are reported here for a potato genetic map, including new map locations for 24 of them. A first criterion for this proposed reference set was ubiquity of the SSRs in the eight landrace cultivar groups of the potato, Solanum tuberosum. All SSRs tested in the present study displayed the same allele phenotypes and allele size range in the diverse germplasm set as in the advanced potato cultivar germplasm in which they were originally discovered. Thirteen of 13 SSR products from all cultivar groups are shown to cross-hybridize with the corresponding SSR product of the source cultivar to ascertain sequence homology. Other important SSR selection criteria are quality of amplification products, locus complexity, polymorphic index content, and well-dispersed location on a potato genetic map. Screening of 156 SSRs allowed the identification of a highly informative and user-friendly set comprising 18 SSR markers for use in characterization of potato genetic resources. In addition, we have identified true- and pseudo-multiplexing SSRs for even greater efficiency.

Comparison of AFLPs with other markers for phylogenetic inference in wild tomatoes [Solanum L. section Lycopersicon (Mill.) Wettst.]

ac.uk Wild tomatoes (Solanum section Lycopersicon) are native to western South America. The delimitation and relationships of tomato species have differed widely depending upon whether morphological or biological species concepts are considered more important. Molecular data from mitochondrial, nuclear, and chloroplast DNA restriction fragment length polymorphisms (RFLPs), nuclear microsatellites, isozymes, and gene sequences of internal transcribed spacers of nuclear ribosomal DNA (ITS; multiple-copy), the single-copy nuclear encoded Granule-bound Starch Synthase gene (GBSSI or waxy), and morphology, have been used to examine hypotheses of species relationships. This study is a companion to the previous GBSSI gene sequence study and to the morphological study of relationships of all ten wild tomato species (including the recently described S. galapagense), with a concentration on the most widespread and variable species S. peruvianum s.l. These new AFLP data are largely concordant with the GBSSI and morphological data and in general support the species outlined in the latest treatment by C.M. Rick, but demonstrate the distinct nature of northern and southern Peruvian populations of S. peruvianum, and suggest that their taxonomy needs revision. Solanum ochranthum is supported as sister to wild tomatoes, and S. habrochaites and S. pennellii reside in a basal polytomy in the tomato clade.

Extensive simple sequence repeat genotyping of potato landraces supports a major reevaluation of their gene pool structure and classification

Contrasting taxonomic treatments of potato landraces have continued over the last century, with the recognition of anywhere from 1 to 21 distinct Linnean species, or of Cultivar Groups within the single species Solanum tuberosum. We provide one of the largest molecular marker studies of any crop landraces to date, to include an extensive study of 742 landraces of all cultivated species (or Cultivar Groups) and 8 closely related wild species progenitors, with 50 nuclear simple sequence repeat (SSR) (also known as microsatellite) primer pairs and a plastid DNA deletion marker that distinguishes most lowland Chilean from upland Andean landraces. Neighbor-joining results highlight a tendency to separate three groups: (i) putative diploids, (ii) putative tetraploids, and (iii) the hybrid cultivated species S. ajanhuiri (diploid), S. juzepczukii (triploid), and S. curtilobum (pentaploid). However, there are many exceptions to grouping by ploidy. Strong statistical support occurs only for S. ajanhuiri, S. juzepczukii, and S. curtilobum. In combination with recent morphological analyses and an examination of the identification history of these collections, we support the reclassification of the cultivated potatoes into four species: (i) S. tuberosum, with two Cultivar Groups (Andigenum Group of upland Andean genotypes containing diploids, triploids, and tetraploids, and the Chilotanum Group of lowland tetraploid Chilean landraces); (ii) S. ajanhuiri (diploid); (iii) S. juzepczukii (triploid); and (iv) S. curtilobum (pentaploid). For other classifications, consistent and stable identifications are impossible, and their classification as species is artificial and only maintains the confusion of users of the gene banks and literature.

Solanaceae--a model for linking genomics with biodiversity.

Recent progress in understanding the phylogeny of the economically important plant family Solanaceae makes this an ideal time to develop models for linking the new data on plant genomics with the huge diversity of naturally occurring species in the family. Phylogenetics provides the framework with which to investigate these linkages but, critically, good species-level descriptive resources for the Solanaceae community are currently missing. Phylogeny in the family as a whole is briefly reviewed, and the new NSF Planetary Biodiversity Inventories project ‘PBI: Solanum—a worldwide treatment’ is described. The aims of this project are to provide species-level information across the global scope of the genus Solanum and to make this available over the Internet. The project is in its infancy, but will make available nomenclatural information, descriptions, keys and illustrative material for all of the approximately 1500 species of Solanum. With this project, the opportunity of linking valid, up-to-date taxonomic information about wild species of Solanum with the genomic information being generated about the economically important species of the genus (potato, tomato and eggplant) can be realized. The phylogenetic framework in which the PBI project is set is also of enormous potential benefit to other workers on Solanum. The community of biologists working with Solanaceae has a unique opportunity to effectively link genomics and taxonomy for better understanding of this important family, taking plant biology to a new level for the next century.

New Species of Wild Tomatoes (Solanum Section Lycopersicon: Solanaceae) from Northern Peru

Abstract Solanum arcanum and S. huaylasense, two new wild tomato species segregated from Solanum peruvianum sensu lato, are described and illustrated. These two new species are placed in a key with two other segregates of S. peruvianum sensu lato: S. peruvianum sensu stricto and S. corneliomulleri, and the morphologically similar species S. chilense. We also present a list of all 13 species of wild tomatoes we recognize, and their equivalent former names in Lycopersicon.

Potato systematics and germplasm collecting, 1989-2000

This paper reviews the systematics of wild potatoes over the past 11 years, in reference to the latest comprehensive taxonomic treatment by Hawkes (1990. The potato: evolution, biodiversity and genetic resources. Belhaven Press, Washington, D.C.). Included here is information on monographs and floras, new germplasm collections made since 1988 compared to total potato distributional data, new taxonomic changes (including synonymy and new species descriptions), ingroup and outgroup relationships, intraspecific studies, diversity studies, and fingerprinting. In addition, data supporting taxonomic changes and phylogeny from ploidy levels, Endosperm Balance Numbers, and morphological studies of taxonomically important characters are reviewed. A revised list of 206 species is presented (from 232 in Hawkes) that incorporates recent synonymy of names, recognition of new names, and new species descriptions. New germplasm collections of 58 potato taxa were collected that did not occur in genebanks before 1988.ResumenEn este articulo se resume el tratamiento sistematico de las papas silvestres durante los pasados 11 anos, en referencia al mas reciente tratado taxonomico deHawkes (1990. The potato: evolution, biodiversity and genetic resources. Belhaven Press, Washington, D.C.) Aquí se incluye información sobre monografías, nuevas colecciones de germoplasma hechas desde 1988 y comparadas con la distribución geográfica completa de papas silvestres, nuevos cambios taxónomicos (incluidas la descripción de sinónimas y de nuevas especies), parentesco filogenético dentro y fuera del grupo, estudios intraespecíficos, estudios diversos y tipificacion del ADN. Adicionalmente, se revisan los datos que sustentan los cambios taxonómicos y en el parentesco filogenético: niveles de ploidia, Balance Numérico de Endosperma y estudios morfológicos de caracteres taxonómicamente importantes. Se presenta una lista revisada de 206 especies (de 232 de Hawkes) que incorpora la reciente sinonimia de nombres, el reconocimiento de nuevos nombres y la descripción de nuevas especies. Se colectaron 58 taxa de papa que no se encontraban en los bancos genéticos antes de 1988.

Reexamination of series relationships of South American wild potatoes (Solanaceae: Solanum sect. Petota): evidence from chloroplast DNA restriction site variation.

Chloroplast DNA (cpDNA) restriction enzyme site analysis was used to test hypotheses of series and superseries affiliations of 76 taxa, representing 11 of the 13 South American series (material unavailable for two series) of wild potatoes (Solanum sect. Petota) recognized in the latest classification by Hawkes. The cladistic results, combined with those from earlier cpDNA studies of 30 taxa of the Mexican and Central American species (representing eight series; ser. Conicibaccata and ser. Tuberosa have representatives in Mexico and in South America), support four main clades for 17 of the 19 series examined in sect. Petota: (1) the Mexican and Central American diploid species, exclusive of S. bulbocastanum, S. cardiophyllum, and S. verrucosum, (2) S. bulbocastanum and S. cardiophyllum (ser. Bulbocastana, ser. Pinnatisecta), (3) South American diploid species constituting all of ser. Piurana, but also members of ser. Conicibaccata, ser. Megistacroloba, ser. Tuberosa, and ser. Yungasensia, (4) all Mexican and Central American polyploid species (ser. Longipedicellata, ser. Demissa), S. verrucosum (diploid Mexican species in ser. Tuberosa), and South American diploid and polyploid members of ser. Acaulia, ser. Circaeifolia, ser. Commersoniana, ser. Conicibaccata, ser. Cuneoalata, ser. Lignicaulia, ser. Maglia, ser. Megistacroloba, ser. Tuberosa, and ser. Yungasensia. Each of these clades contains morphologically and reproductively very diverse species, and there are no evident morphological features that unite members within a clade to therefore distinguish them. These results strongly suggest a need for a reevaluation of the series and superseries classifications of sect. Petota.