Discovery and characterization of sweetpotato's closest tetraploid relative.

The origin of sweetpotato, a hexaploid species, is poorly understood, partly because the identity of its tetraploid progenitor remains unknown. In this study, we identify, describe and characterize a new species of Ipomoea that is sweetpotato's closest tetraploid relative known to date and probably a direct descendant of its tetraploid progenitor. We integrate morphological, phylogenetic, and genomic analyses of herbarium and germplasm accessions of the hexaploid sweetpotato, its closest known diploid relative Ipomoea trifida, and various tetraploid plants closely related to them from across the American continent. We identify wild autotetraploid plants from Ecuador that are morphologically distinct from Ipomoea batatas and I. trifida, but monophyletic and sister to I. batatas in phylogenetic analysis of nuclear data. We describe this new species as Ipomoea aequatoriensis T. Wells & P. Muñoz sp. nov., distinguish it from hybrid tetraploid material collected in Mexico; and show that it likely played a direct role in the origin of sweetpotato's hexaploid genome. This discovery transforms our understanding of sweetpotato's origin.

Species as a Heuristic: Reconciling Theory and Practice.

Species are crucial to most branches of biological research, yet remain controversial in terms of definition, delimitation, and reality. The difficulty of resolving the "species problem" stems from the tension between their theoretical concept as groups of evolving and highly variable organisms and the practical need for a stable and comparable unit of biology. Here, we suggest that treating species as a heuristic can be consistent with a theoretical definition of what species are and with the practical means by which they are identified and delimited. Specifically, we suggest that theoretically species are heuristic since they comprise clusters of closely related individuals responding in a similar manner to comparable sets of evolutionary and ecological forces, whilst they are practically heuristic because they are identifiable by the congruence of contingent properties indicative of those forces. This reconciliation of the theoretical basis of species with their practical applications in biological research allows for a loose but relatively consistent definition of species based on the strategic analysis and integration of genotypic, phenotypic, and ecotypic data. [Cohesion; heuristic; homeostasis; lineage; species problem.].

A foundation monograph of Ipomoea (Convolvulaceae) in the New World.

A monograph of the 425 New World species of Ipomoea is presented. All 425 species are described and information is provided on their ecology and distribution, with citations from all countries from which they are reported. Notes are provided on salient characteristics and taxonomic issues related to individual species. A full synonymy is provided and 272 names are lectotypified. An extensive introduction discusses the delimitation and history of Ipomoea arguing that a broad generic concept is the only rational solution in the light of recent phylogenetic advances. Although no formal infrageneric classification is proposed, attention is drawn to the major clades of the genus and several morphologically well-defined clades are discussed including those traditionally treated under the names Arborescens, Batatas, Pharbitis, Calonyction and Quamoclit, sometimes as distinct genera, subgenera, sections or series. Identification keys are provided on a regional basis including multi-entry keys for the main continental blocks. Six species are described as new, Ipomoea nivea J.R.I. Wood & Scotland from Peru, I. apodiensis J.R.I. Wood & Scotland from Brazil, I. calcicola J.R.I. Wood & Scotland, I. pochutlensis J.R.I. Wood & Scotland, I. zacatecana J.R.I. Wood & Scotland and I. ramulosa J.R.I. Wood & Scotland from Mexico, while var. australis of I. cordatotriloba is raised to specific status as I. australis (O'Donell) J.R.I. Wood & P. Muñoz. New subspecies for I. nitida (subsp. krapovickasii J.R.I. Wood & Scotland) and for I. chenopodiifolia (subsp. bellator J.R.I. Wood & Scotland) are described. The status of previously recognized species and varieties is changed so the following new subspecies are recognized: I. amnicola subsp. chiliantha (Hallier f.) J.R.I. Wood & Scotland, I. chenopodiifolia subsp. signata (House) J.R.I. Wood & Scotland, I. orizabensis subsp. collina (House) J.R.I. Wood & Scotland, I. orizabensis subsp. austromexicana (J.A. McDonald) J.R.I. Wood & Scotland, I. orizabensis subsp. novogaliciana (J.A. McDonald) J.R.I. Wood & Scotland, I. setosa subsp. pavonii (Hallier f.) J.R.I. Wood & Scotland, I. setosa subsp. melanotricha (Brandegee) J.R.I. Wood & Scotland, I. setosa subsp. sepacuitensis (Donn. Sm.) J.R.I. Wood & Scotland, I. ternifolia subsp. leptotoma (Torr.) J.R.I. Wood & Scotland. Ipomoea angustata and I. subincana are treated as var. angustata (Brandegee) J.R.I. Wood & Scotland and var. subincana (Choisy) J.R.I. Wood & Scotland of I. barbatisepala and I. brasiliana respectively. Attention is drawn to a number of hitherto poorly recognized phenomena in the genus including a very large radiation centred on the Parana region of South America and another on the Caribbean Islands, a strong trend towards an amphitropical distribution in the New World, the existence of a relatively large number of species with a pantropical distribution and of many species in different clades with storage roots, most of which have never been evaluated for economic purposes. The treatment is illustrated with over 200 figures composed of line drawings and photographs.

The temporal dynamics of evolutionary diversification in Ipomoea.

Molecular phylogenies are used as a basis for making inferences about macroevolutionary history. However, a robust phylogeny does not contain the information that is necessary to make many of these inferences. Complex methodologies that incorporate important assumptions about the nature of evolutionary history are therefore required. Here, we explore the implications of these assumptions for making inferences about the macroevolutionary history of Ipomoea - a large pantropical genus of flowering plants that contains the sweet potato (Ipomoea batatas), a crop of global economic importance. We focus on assumptions that underlie inferences of divergence times, and diversification parameters (speciation rates, extinction rates, and net diversification rates). These are among the most fundamental variables in macroevolutionary research. We use a series of novel approaches to explore the implications of these assumptions for inferring the age of Ipomoea, the ages of major clades within Ipomoea, whether there are significant differences in diversification parameters among clades within Ipomoea, and whether the storage root of I. batatas evolved in pre-human times. We show that inferring an age estimate for Ipomoea and major clades within Ipomoea is highly problematic. Inferred divergence times are sensitive to uncertain fossil calibrations and differing assumptions about among-branch-substitution-rate-variation. Despite this uncertainty, we are able to make robust inferences about patterns of variation in diversification parameters within Ipomoea, and that the storage root of I. batatas evolved in pre-human times. Taken together, this study presents novel and generalizable insights into the implications of methodological assumptions for making inferences about macroevolutionary history. Further, by presenting novel findings relating to the temporal dynamics of evolution in Ipomoea, as well as more specifically to I. batatas, this study makes a valuable contribution to our understanding of tropical plant evolution, and the evolutionary context in which economically important crops evolve.

A taxonomic monograph of Ipomoea integrated across phylogenetic scales.

Taxonomic monographs have the potential to make a unique contribution to the understanding of global biodiversity. However, such studies, now rare, are often considered too daunting to undertake within a realistic time frame, especially as the world's collections have doubled in size in recent times. Here, we report a global-scale monographic study of morning glories (Ipomoea) that integrated DNA barcodes and high-throughput sequencing with the morphological study of herbarium specimens. Our approach overhauled the taxonomy of this megadiverse group, described 63 new species and uncovered significant increases in net diversification rates comparable to the most iconic evolutionary radiations in the plant kingdom. Finally, we show that more than 60 species of Ipomoea, including sweet potato, independently evolved storage roots in pre-human times, indicating that the storage root is not solely a product of human domestication but a trait that predisposed the species for cultivation. This study demonstrates how the world's natural history collections can contribute to global challenges in the Anthropocene.

Reconciling Conflicting Phylogenies in the Origin of Sweet Potato and Dispersal to Polynesia.

The sweet potato is one of the world's most widely consumed crops, yet its evolutionary history is poorly understood. In this paper, we present a comprehensive phylogenetic study of all species closely related to the sweet potato and address several questions pertaining to the sweet potato that remained unanswered. Our research combined genome skimming and target DNA capture to sequence whole chloroplasts and 605 single-copy nuclear regions from 199 specimens representing the sweet potato and all of its crop wild relatives (CWRs). We present strongly supported nuclear and chloroplast phylogenies demonstrating that the sweet potato had an autopolyploid origin and that Ipomoea trifida is its closest relative, confirming that no other extant species were involved in its origin. Phylogenetic analysis of nuclear and chloroplast genomes shows conflicting topologies regarding the monophyly of the sweet potato. The process of chloroplast capture explains these conflicting patterns, showing that I. trifida had a dual role in the origin of the sweet potato, first as its progenitor and second as the species with which the sweet potato introgressed so one of its lineages could capture an I. trifida chloroplast. In addition, we provide evidence that the sweet potato was present in Polynesia in pre-human times. This, together with several other examples of long-distance dispersal in Ipomoea, negates the need to invoke ancient human-mediated transport as an explanation for its presence in Polynesia. These results have important implications for understanding the origin and evolution of a major global food crop and question the existence of pre-Columbian contacts between Polynesia and the American continent.

Additional notes on Ipomoea (Convolvulaceae) in Bolivia.

109 species of Ipomoea L. are recorded from Bolivia. This total includes six new country records and two species, I. prolifera J. R. I. Wood & Scotland and I. inaccessa J. R. I. Wood & Scotland, which are described as new, while I. jujuyensis O'Donell is excluded having been included previously in error. The little-known I. subalata Hassl. is described in full and compared with I. chondrosepala Hallier f. and other species with which it has been confused. The paper is illustrated with line drawings, photographs and distribution maps of the main species discussed.

New species of Ipomoea (Convolvulaceae) from South America.

The importance of discovering, describing and cataloguing poorly known species in herbarium collections is discussed. It is a spur to efforts at rediscovery and consequent conservation efforts. The problems faced in describing species from limited material are discussed and our methods and criteria in making a decision are described. Prospects for future novelties are briefly assessed. Fifteen new species are described and illustrated with line drawings and distribution maps: Ipomoea attenuata J.R.I. Wood & Scotland, I. cuscoensis J.R.I. Wood & P. Muñoz, I. dasycarpa J.R.I. Wood & Scotland, I. dolichopoda J.R.I. Wood & R. Degen, I. ensiformis J.R.I.Wood & Scotland, I. fasciculata J.R.I. Wood & Scotland, I. graminifolia J.R.I. Wood & Scotland, I. kraholandica J.R.I. Wood & Scotland, I. longirostra J.R.I. Wood & Scotland, I. revoluta J.R.I. Wood & Scotland, I. scopulina J.R.I. Wood &. Scotland, I. uninervis J.R.I. Wood & Scotland, I. veadeirosii J.R.I. Wood & Scotland, I. velutinifolia J.R.I. Wood & Scotland, I. walteri J.R.I. Wood & Scotland. All species are narrow endemics except I. velutinifolia which is found in Brazil and Peru; of the others, 12 are found in Brazil and one each in Paraguay and Peru.

Remarkable disjunctions in Ipomoea species (Convolvulaceae) from NE Brazil and Central America and their taxonomic implications.

Recent collections of Ipomoea from North East Brazil have revealed a number of unexpected disjunct distributions. The most remarkable is that of I. eremnobrocha D. F. Austin, previously thought to be endemic to Panama but now known from three states in NE Brazil. Revision of Panamanian material named I. eremnobrocha unexpectedly showed that two distinct species had been treated under this one name. Specimens from the Chagres National Park area in Panama are described as a new species under the name I. isthmica J. R. I. Wood & Buril while I. eremnobrocha is retained as the correct name for the plant from the Altos de Campana in Panama and NE Brazil. An amended description of this species is given and a table of differences between the related species is provided. Two recently described species from Bolivia, I. graniticola J. R. I. Wood & Scotland and I. chiquitensis J. R. I. Wood & Scotland are recorded from NE Brazil several thousand km from their type localities. Attention is drawn to the role of granite inselbergs as sites of species with a disjunct distribution. A possible relative of I. chiquitensis is described as a new species from NE Brazil under the name I. melancholica J. R. I. Wood & Buril. The new species are illustrated with line drawings and maps of the unusual distribution patterns are provided.

How the temperate world was colonised by bindweeds: biogeography of the Convolvuleae (Convolvulaceae).

BACKGROUND: At a global scale, the temperate zone is highly fragmented both between and within hemispheres. This paper aims to investigate how the world's disjunct temperate zones have been colonised by the pan-temperate plant group Convolvuleae, sampling 148 of the c. 225 known species. We specifically determine the number and timing of amphitropical and transoceanic disjunctions, investigate the extent to which disjunctions in Convolvuleae are spatio-temporally congruent with those in other temperate plant groups and determine the impact of long-distance dispersal events on diversification rates. RESULTS: Eight major disjunctions are observed in Convolvuleae: two Northern Hemisphere, two Southern Hemisphere and four amphitropical. Diversity in the Southern Hemisphere is largely the result of a single colonisation of Africa 3.1-6.4 Ma, and subsequent dispersals from Africa to both Australasia and South America. Speciation rates within this monophyletic, largely Southern Hemisphere group (1.38 species Myr(-1)) are found to be over twice those of the tribe as a whole (0.64 species Myr(-1)). Increased speciation rates are also observed in Calystegia (1.65 species Myr(-1)). CONCLUSIONS: The Convolvuleae has colonised every continent of the world with a temperate biome in c. 18 Myr and eight major range disjunctions underlie this broad distribution. In keeping with other temperate lineages exhibiting disjunct distributions, long-distance dispersal is inferred as the main process explaining the patterns observed although for one American-Eurasian disjunction we cannot exclude vicariance. The colonisation of the temperate zones of the three southern continents within the last c. 4 Myr is likely to have stimulated high rates of diversification recovered in this group, with lineage accumulation rates comparable to those reported for adaptive radiations.

Widespread mistaken identity in tropical plant collections.

Specimens of plants and animals preserved in museums are the primary source of verifiable data on the geographical and temporal distribution of organisms. Museum datasets are increasingly being uploaded to aggregated regional and global databases (e.g. the Global Biodiversity Information Facility; GBIF) for use in a wide range of analyses. Thus, digitisation of natural history collections is providing unprecedented information to facilitate the study of the natural world on a global scale. The digitisation of this information utilises information provided on specimen labels, and assumes they are correctly identified. Here we evaluate the accuracy of names associated with 4,500 specimens of African gingers from 40 herbaria in 21 countries. Our data show that at least 58% of the specimens had the wrong name prior to a recent taxonomic study. A similar pattern of wrongly named specimens is also shown for Dipterocarps and Ipomoea (morning glory). We also examine the number of available plant specimens worldwide. Our data demonstrate that, while the world's collections have more than doubled since 1970, more than 50% of tropical specimens, on average, are likely to be incorrectly named. This finding has serious implications for the uncritical use of specimen data from natural history collections.

A foundation monograph of Convolvulus L. (Convolvulaceae).

A global revision of Convolvulus L. is presented, Calystegia R.Br. being excluded on pragmatic grounds. One hundred and ninety species are recognised with the greatest diversity in the Irano-Turanian region. All recognised species are described and the majority are illustrated. Distribution details, keys to species identification and taxonomic notes are provided. Four new species, Convolvulusaustroafricanus J.R.I.Wood & R.W.Scotland, sp. nov., Convolvulusiranicus J.R.I.Wood & R.W.Scotland, sp. nov., Convolvuluspeninsularis J.R.I.Wood & R.W.Scotland, sp. nov. and Convolvulusxanthopotamicus J.R.I.Wood & R.W.Scotland, sp. nov., one new subspecies Convolvuluschinensissubsp.triangularis J.R.I.Wood & R.W.Scotland, subsp. nov., and two new varieties Convolvulusequitansvar.lindheimeri J.R.I.Wood & R.W.Scotland, var. nov., Convolvulusglomeratusvar.sachalitarum J.R.I.Wood & R.W.Scotland, var. nov. are described. Convolvulusincisodentatus J.R.I.Wood & R.W.Scotland, nom. nov., is provided as a replacement name for the illegitimate Convolvulusincisus Choisy. Several species treated as synonyms of other species in recent publications are reinstated including Convolvuluschinensis Ker-Gawl., Convolvulusspinifer M.Popov., Convolvulusrandii Rendle and Convolvulusaschersonii Engl. Ten taxa are given new status and recognised at new ranks: Convolvulusnamaquensis (Schltr. ex. A.Meeuse) J.R.I.Wood & R.W.Scotland, stat. nov., Convolvulushermanniaesubsp.erosus (Desr.) J.R.I.Wood & R.W.Scotland, stat. nov., Convolvuluscrenatifoliussubsp.montevidensis (Spreng.) J.R.I.Wood & R.W.Scotland, stat. nov., Convolvulusfruticulosussubsp.glandulosus (Webb) J.R.I.Wood & R.W.Scotland, stat. nov., Convolvuluscapituliferussubsp.foliaceus (Verdc.) J.R.I.Wood & R.W.Scotland, stat. nov., Convolvulushystrixsubsp.ruspolii (Dammer ex Hallier f.) J.R.I.Wood & R.W.Scotland, stat. nov., Convolvulushystrixsubsp.inermis (Chiov.) J.R.I.Wood & R.W.Scotland, stat. nov., Convolvulusrottlerianussubsp.stocksii (Boiss.) J.R.I.Wood & R.W.Scotland, comb. et stat. nov., Convolvuluscalvertiisubsp.ruprechtii (Boiss.) J.R.I.Wood & R.W.Scotland, stat. nov., Convolvuluscephalopodussubsp.bushiricus (Bornm.) J.R.I.Wood & R.W.Scotland, stat. nov. The status of various infraspecific taxa is clarified and numerous taxa are lectotypified. This account represents a new initiative in terms of taxonomic monography, being an attempt to bring together the global approach of the traditional monograph with the more pragmatic and identification-focussed approach of most current floras while at the same time being informed by insights from molecular systematics.

Distributions, ex situ conservation priorities, and genetic resource potential of crop wild relatives of sweetpotato [Ipomoea batatas (L.) Lam., I. series Batatas].

Crop wild relatives of sweetpotato [Ipomoea batatas (L.) Lam., I. series Batatas] have the potential to contribute to breeding objectives for this important root crop. Uncertainty in regard to species boundaries and their phylogenetic relationships, the limited availability of germplasm with which to perform crosses, and the difficulty of introgression of genes from wild species has constrained their utilization. Here, we compile geographic occurrence data on relevant sweetpotato wild relatives and produce potential distribution models for the species. We then assess the comprehensiveness of ex situ germplasm collections, contextualize these results with research and breeding priorities, and use ecogeographic information to identify species with the potential to contribute desirable agronomic traits. The fourteen species that are considered the closest wild relatives of sweetpotato generally occur from the central United States to Argentina, with richness concentrated in Mesoamerica and in the extreme Southeastern United States. Currently designated species differ among themselves and in comparison to the crop in their adaptations to temperature, precipitation, and edaphic characteristics and most species also show considerable intraspecific variation. With 79% of species identified as high priority for further collecting, we find that these crop genetic resources are highly under-represented in ex situ conservation systems and thus their availability to breeders and researchers is inadequate. We prioritize taxa and specific geographic locations for further collecting in order to improve the completeness of germplasm collections. In concert with enhanced conservation of sweetpotato wild relatives, further taxonomic research, characterization and evaluation of germplasm, and improving the techniques to overcome barriers to introgression with wild species are needed in order to mobilize these genetic resources for crop breeding.

Author inflation masks global capacity for species discovery in flowering plants.

Species discovery is a fundamental first step for all of biodiversity science. Recent research has claimed that the increasing number of authors associated with the description of new species represents an expanding workforce discovering the remaining new species from an ever-diminishing pool. Here, we present a comprehensive dataset from The International Plant Names Index (IPNI) of new species of flowering plant published between 1970 and 2011. We show that, on average, 1855 new species have been described annually since 1970. We show that compared to other scientific disciplines the increased number of authors on taxonomic papers is relatively small and may reflect changes in scientific practice rather than an increase in taxonomic capacity. These data, alongside published results demonstrating a lag period of 35 yr between a specimen being collected and published as a new species, strongly suggest that the global taxonomic capacity to describe new species of flowering plant is stagnant at a time of unprecedented concern for conservation and extinction.

Big hitting collectors make massive and disproportionate contribution to the discovery of plant species.

Discovering biological diversity is a fundamental goal--made urgent by the alarmingly high rate of extinction. We have compiled information from more than 100,000 type specimens to quantify the role of collectors in the discovery of plant diversity. Our results show that more than half of all type specimens were collected by less than 2 per cent of collectors. This highly skewed pattern has persisted through time. We demonstrate that a number of attributes are associated with prolific plant collectors: a long career with increasing productivity and experience in several countries and plant families. These results imply that funding a small number of expert plant collectors in the right geographical locations should be an important element in any effective strategy to find undiscovered plant species and complete the inventory of the world flora.

Herbaria are a major frontier for species discovery.

Despite the importance of species discovery, the processes including collecting, recognizing, and describing new species are poorly understood. Data are presented for flowering plants, measuring quantitatively the lag between the date a specimen of a new species was collected for the first time and when it was subsequently described and published. The data from our sample of new species published between 1970 and 2010 show that only 16% were described within five years of being collected for the first time. The description of the remaining 84% involved much older specimens, with nearly one-quarter of new species descriptions involving specimens >50 y old. Extrapolation of these results suggest that, of the estimated 70,000 species still to be described, more than half already have been collected and are stored in herbaria. Effort, funding, and research focus should, therefore, be directed as much to examining extant herbarium material as collecting new material in the field.