David M. Spooner’s research while affiliated with University of Wisconsin–Madison and other places

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Publications (226)


Fig. 1. Maximum likelihood reconstruction and structure of the genetic diversity of 288 accessions of the Daucus carota complex and outgroup, using 29,041 SNPs obtained by genotyping-by-sequencing. Each accession is represented by a horizontal bar, and each color corresponds to a theoretical population (nine in total). Numbers above the branches represent bootstrap values. Daucus syrticus is the outgroup. Accessions previously studied in Arbizu et al. (2016) and Mezghani et al. (2018) are marked in green.
Fig. 2. Distribution of accessions used in our study (possessing sufficient locality data). Red triangles 5 cultivated accessions (Daucus carota subsp. sativus); black triangles 5 wild accessions (all of the other subspecies of D. carota, and D. annuus ([ Tornabenea annua) and D. tenuissimus ([ Tornabenea tenuissima)). See Supplementary Table 1 (Martínez-Flores et al. 2020) for a detailed list of accessions.
Fig. 3. Distribution of the wild accessions (possessing sufficient locality data) from the western Mediterranean Region as well as southern England and adjacent northern France, where all of the gummifer morphotypes occur. Numbers according to Figs. 4-6. See Supplementary Table 1 (Martínez-Flores et al. 2020) for a detailed list of records and corresponding numbers.
Fig. 4. Detail of Clade 1 and the outgroup from Fig. 1. Names on the clade refer to geographic origin of the samples. Numbers before accession names correspond to Fig. 3 map."Coastal morphotypes" are marked in red. See Supplementary Table 1 (Martínez-Flores et al. 2020) for a detailed list of records and corresponding numbers.
Fig. 5. Detail of Clade 2 and Clade 3 part 1 from Fig. 1. Names on the clades refer to geographic origin of the samples. Numbers before accession names correspond to Fig. 3 map. "Coastal morphotypes" are marked in red. See Supplementary Table 1 (Martínez-Flores et al. 2020) for a detailed list of records and corresponding numbers.

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Subspecies Variation of Daucus carota Coastal ("Gummifer") Morphotypes (Apiaceae) Using Genotyping-by-Sequencing
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September 2020

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344 Reads

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8 Citations

Systematic Botany

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Philipp W Simon

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David M Spooner

The genus Daucus is widely distributed worldwide, but with a concentration of diversity in the Mediterranean Region. The D. carota complex presents the greatest taxonomic problems in the genus. We focus on a distinctive phenotypic group of coastal morphotypes of D. carota, strictly confined to the margins to within about 0.5 km of the Mediterranean Sea and the Atlantic Ocean, which we here refer to as coastal morphotypes or D. carota subsp. "gummifer" complex. They are loosely morphologically coherent, sharing a relatively short stature, thick, broad, sometimes highly glossy leaf segments, and usually flat or convex fruiting umbels. We analyzed 288 accessions obtained from genebanks in England, France, and the USA, and an expedition to Spain in 2016, covering the Mediterranean and Atlantic coasts and Balearic Islands, where much of the gummifer complex variation occurs. Our study includes 112 accessions not examined before in this context. Genotyping-by-sequencing identified 29,041 filtered SNPs. Based on high bootstrap support from maximum likelihood and Structure analysis we highlight three main clades. The gummifer morphotypes are intercalated with members of Daucus carota subspecies carota and subspecies maximus in two of these main clades, including a clade containing accessions from Tunisia (also including D. carota subsp. capillifolius) and a clade containing accessions from western Europe (including the British Isles), southern Europe (including the Balearic Islands and the Iberian Peninsula) and Morocco. These results support five independent selections of the gummifer morphotypes in these restricted maritime environments in the Mediterranean and nearby Atlantic coasts. Daucus annuus (5Tornabenea annua) and Daucus tenuissimus (5Tornabenea tenuissima) also fall firmly within D. carota, supporting their classification as morphologically well-defined subspecies of D. carota, which are accepted here under the new combinations Daucus carota subsp. annuus and D. carota subsp. tenuissimus, respectively. Types are indicated for most of treated names, including designation of four lectotypes and three epitypes, which fix their further use.

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What is truth: Consensus and discordance in next‐generation phylogenetic analyses of Daucus

August 2020

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49 Reads

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10 Citations

Journal of Systematics and Evolution

High throughput (next‐generation) DNA sequencing has removed barriers to data quantity and quality and produces phylogenies with high statistical support. Such data are useful to address phylogenetic congruence among individual genes. Concatenated analyses of unlinked genes often produce well‐resolved phylogenetic trees with bootstrap support on major nodes at or approaching 100%, but have been criticized for providing incorrect phylogenies for various reasons to include a history of hybridization, introgression, and incomplete lineage sorting. The present study compares next‐generation sequencing results of the same accessions of Daucus with different genomic regions, three reported before: (1) the entire plastid genome, (2) 47 mitochondrial genes, (3), 94 conserved nuclear orthologs. Here, we report a fourth dataset, (4) 564,895 nuclear SNPs. There are areas of discordance in all four results using the same accessions analyzed with maximum parsimony, maximum likelihood, and with the nuclear data species trees through a coalescent analysis. The nuclear results show significant areas of discordance that were unexpected because these studies used the same DNA samples, the nuclear studies were generated from large and high‐quality data sets with the SNPs distributed on all nine linkage groups of D. carota, and the results were supported by high bootstrap values. These results raise questions of the best data and analytical methods to reconstruct and understand the ‘truth’ of a phylogeny. This article is protected by copyright. All rights reserved.


Mitochondrial DNA Sequence Phylogeny of Daucus

June 2020

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37 Reads

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2 Citations

Systematic Botany

We explored the phylogenetic utility of mitochondrial DNA sequences in Daucus and compared the results with prior phylogenetic results using the same 36 accessions of Daucus (and two additional outgroups) with plastid DNA sequences and with other nuclear results. As in the plastid study we used Illumina HiSeq sequencer to obtain resequencing data of the same accessions of Daucus and outgroups, and analyzed the data with maximum parsimony and maximum likelihood. We obtained data from 47 of 71 total mitochondrial genes but only 17 of these 47 genes recovered major clades that were common in prior plastid and nuclear studies. Our phylogenetic trees of the concatenated data set of 47 genes were moderately resolved, with 100% bootstrap support for most of the external and many of the internal clades, except for the clade of D. carota and its most closely related species D. syrticus. There are areas of hard incongruence with phylogenies using plastid and nuclear data. In agreement with other studies, we conclude that mitochondrial sequences are generally poor phylogenetic markers, at least at the genus level, despite their utility in some other studies.


PTIS Potato Herbarium Transferred to WIS, the Wisconsin State Herbarium

November 2019

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24 Reads

American Journal of Potato Research

The Potato Introduction Station in Sturgeon Bay Wisconsin has maintained herbarium specimens as part of its operations. These specimens mostly were grown from seeds in the field plots or greenhouses, but also include original specimens made during germplasm collecting expeditions. The herbarium of the Station (herbarium acronym PTIS) has been transferred from the potato genebank in Sturgeon Bay Wisconsin to the Wisconsin State Herbarium (acronym WIS) in Birge Hall, University of Wisconsin-Madison where it is now fully integrated and curated and funded by WIS; PTIS is discontinued and is no longer maintained as a separate herbarium. Separately, maps and taxonomic literature that were used by the potato taxonomist in Madison were transferred to the Potato Station in Sturgeon Bay.


Taxonomic richness map for carrot wild relative potential distribution models, (A) across the western Mediterranean, with focus on (B) Tunisia. Darker colors indicate greater numbers of taxa potentially overlapping in the same areas.
Conservation gap analysis results per taxon. Carrot wild relatives are listed by descending priority for further conservation action by priority categories (high priority [HP, red)], medium priority [MP, orange], low priority [LP, yellow], and sufficiently conserved [SC, green]). The red diamond represents the combined final conservation score (FCS‐mean) for the taxon, which is the average of the final ex situ (FCSex) (black circle) and in situ (FCSin) (black triangle) scores. Results of the conservation assessments within each strategy (sampling representativeness score [SRS], geographic representativeness score ex situ [GRSex], and ecological representativeness score ex situ [ERSex] for ex situ; and geographic representativeness score in situ [GRSin] and ecological representativeness score in situ [ERSin] for in situ) are also displayed.
(A) Further collecting priority hotspots map for carrot wild relatives in Tunisia. The map displays richness of areas within the potential distributions of taxa that have not been previously collected for ex situ conservation, with up to seven taxa in need of further collecting potentially found in the same areas. (B) Further in situ protection priorities map for carrot wild relatives in Tunisia. The map displays richness of areas within the potential distributions of taxa that are outside of current protected areas, with up to nine taxa found in the same unprotected areas.
Distributions and Conservation Status of Carrot Wild Relatives in Tunisia: A Case Study in the Western Mediterranean Basin

November 2019

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600 Reads

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18 Citations

Crop wild relatives, the wild progenitors and closely related cousins of cultivated plant species, are sources of valuable genetic resources for crop improvement. Persisting gaps in knowledge of taxonomy, distributions, and characterization for traits of interest constrain their expanded use in plant breeding and likewise negatively affect ex situ (in genebanks) and in situ (in natural habitats) conservation planning. We compile the state of knowledge on the taxonomy and distributions of the wild relatives of carrot (genus Daucus L.) natively occurring within Tunisia—a hotspot of diversity of the genus, containing 13 taxa (27% of species worldwide). We use ecogeographic information to characterize their potential adaptations to abiotic stresses of interest in crop breeding and assess their ex situ and in situ conservation status. We find substantial ecogeographic variation both across taxa and between populations within taxa, with regard to adaptation to high temperatures, low precipitation, and other traits of potential interest. We categorize three of the taxa high priority for further conservation both ex situ and in situ, five medium priority, and five low priority, with none currently considered sufficiently conserved. Geographic hotspots for species diversity, especially in the northern coastal areas, represent particularly high value regions for efficient further collecting for ex situ conservation and for in situ protection in Tunisia.




Figure 3. Specimen of Daucus pusillus (including D. arcanus) collected in Ciudad Real, Spain (ABH78367): A, habit; B, detail of fruiting umbels.
Figure 4. Distribution of Daucus pusillus (including D. arcanus) in the Iberian Peninsula. See Appendix for a detailed list of records.
Chromosome numbers of the species examined here. We found no references for chromosome numbers for Daucus bicolor, D. conchitae and D. gracilis
Extended studies of interspecific relationships in Daucus (Apiaceae) using DNA sequences from ten nuclear orthologues

September 2019

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360 Reads

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7 Citations

Botanical Journal of the Linnean Society

Daucus has traditionally been estimated to comprise 21-25 species, but a recent study expanded the genus to c. 40 species. The present study uses ten nuclear orthologues to examine 125 accessions, including 40 collections of 11 species (D. annuus, D. arcanus, D. decipiens, D. durieua, D. edulis, D. gracilis, D. minusculus, D. montanus, D. pumilus, D. setifolius and D. tenuissimus) newly examined with nuclear orthologues. As in previous nuclear orthologue studies, Daucus resolves into two well-defined clades, and groups different accessions of species together. Maximum likelihood and maximum parsimony analyses provide concordant results, but SVD quartets reveals many areas of disagreement of species within these two major clades. With maximum likelihood and maximum parsimony analyses Daucus montanus (hexaploid) is resolved as an allopolyploid between D. pusillus (diploid) and D. glochidiatus (tetraploid), whereas with SVD quartets it is resolved as an allopolyploid between D. glochidiatus and an unknown Daucus sp. We propose the new combination Daucus junceus (Durieua juncea) for a neglected species endemic to the southwestern Iberian Peninsula often referred to as D. setifolius, and we place D. arcanus in synonymy with D. pusillus. Three lectotypes are also designated.


Daucus: Taxonomy, Phylogeny, Distribution

May 2019

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310 Reads

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15 Citations

Cultivated carrot (Daucus carota subsp. sativus) is the most important member in the Apiaceae family in terms of economy and nutrition and is considered the second most popular vegetable in the world after potato. Despite its global importance, the systematics of Daucus remains under active revision at the species, genus, and subtribal levels. The phylogenetic relationships among the species of Daucus and close relatives in the Apioideae have been clarified recently by a series of molecular studies using DNA sequences of the plastid genes rbcL and matK; plastid introns rpl16, rps16, rpoC1; nuclear ribosomal DNA internal transcribed spacer (ITS) sequences; and plastid DNA restriction sites. Of these DNA markers, the ITS region consisting of ITS1, the intervening spacer, and ITS2 has served as the main marker used. Recently, next-generation DNA sequencing methodologies have been used. We review these techniques and how they are impacting the taxonomy of the genus Daucus.


Carrot Organelle Genomes: Organization, Diversity, and Inheritance

May 2019

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104 Reads

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2 Citations

Cultivated carrot (Daucus carota subsp. sativus) is one of about 25–40 related wild species in the genus Daucus depending on the classification. It is part of a widely distributed and taxonomically complex family Apiaceae (Umbelliferae) containing 466 genera and 3820 species that is one of the largest families of seed plants. Members of the Apiaceae, particularly the genus Daucus, have been the subject of intensive recent molecular studies on the structure and genetics of plastids and mitochondria. This chapter summarizes organellar (plastids and mitochondria) structure, function, mutational rates, and inter-organelle DNA transfer in the Apiaceae and inheritance in the genus Daucus, with a wider focus on the Apiaceae and the sister family Araliaceae, and places these data in the context of other studies in the angiosperms.


Citations (51)


... Models trained on a small diversity of genetically related individuals lack prediction accuracy when tested. Corak et al. (2019) applied five different strategies of sampling from a collection of 433 domesticated carrot (Daucus carota) accessions to develop a smaller, core collection of representative accessions. They found that none of the strategies significantly affected GS model accuracy (r 2 = 0.2-0.3) ...

Reference:

Genomic selection for crop improvement in fruits and vegetables: a systematic scoping review
Comparison of Representative and Custom Methods of Generating Core Subsets of a Carrot Germplasm Collection

... therefore allopolyploidy can only be suspected as it is the most commonly occurring type of hybridization in subtribe Trionum and this would need more study to confirm. Furthermore, the use of the single NRC sequence (despite being an extended region) cannot fully rule out incomplete lineage sorting as more nuclear regions would be needed to provide higher support (Spooner et al. 2020). ...

What is truth: Consensus and discordance in next‐generation phylogenetic analyses of Daucus
  • Citing Article
  • August 2020

Journal of Systematics and Evolution

... After filtering out the raw reads, the total of demultiplexed reads for all 89 genotypes was 579.02 M, with the average reads per accession being 6.51 M. We followed the approach of Arbizu et al. [44] and Martínez-Flores et al. [45] and obtained 338,638 read tags, 36.06% of them uniquely aligned to the L. angustifolius reference genome [30]. A total of 35,760 raw SNPs were obtained, and the filtering approach yielded 5922 high-quality SNPs across the 20 chromosomes of Lupinus with an average marker density of 79.52 kb (Table 1, Figure 1). ...

Subspecies Variation of Daucus carota Coastal ("Gummifer") Morphotypes (Apiaceae) Using Genotyping-by-Sequencing

Systematic Botany

... Several mitogenomes of Apiaceae have been previously reported, which are from Daucus carota subsp. sativus (281,132 bp) and Bupleurum falcatum (463,792 bp) [12,13]. The S. divaricata mitogenome (MZ128146) is a circular molecule of 293,897 bp (Fig. 1B). ...

Recent advances in carrot genomics
  • Citing Article
  • November 2019

Acta Horticulturae

... Climate change is predicted to result in changes in the composition of weed communities, growth dynamics, life cycle, phenology, and degree of infestation (Anwar et al., 2021). Most previous studies on how climate change affects wild carrots have focused on selecting suitable genotypes of carrots, including wild carrots, which suggests that scientists tend to concentrate on the breeding perspective in order to better respond to future climate change Mezghani et al., 2019b;Simon et al., 2021). However, it is crucial to investigate how wild carrots survive and expand under climate change conditions. ...

Distributions and Conservation Status of Carrot Wild Relatives in Tunisia: A Case Study in the Western Mediterranean Basin

... Fay, 2018, and references therein). Following on from papers using 'traditional' and nextgeneration techniques to study species delimitation (Reck-Kortmann et al., 2017;Contreras-Ortiz et al., 2018;Hu et al., 2018;Cortez et al., 2019;Könyves, David & Culham, 2019;Martínez-Flores et al., 2019;Zhong et al., 2019), infraspecific taxonomy (Hardion et al., 2017;Folk et al., 2018;Tarieiev et al., 2019), hybridization (Nakahama et al., 2019) and speciation (Escudero et al., 2019), five papers in this issue focus on species delimitation, using a range of techniques. Zhang et al. (2019) In conservation, a detailed understanding of the genetic structure of remaining populations is of great importance in the development of conservation action plans, particularly at the periphery of the distribution or for populations identified as being at risk due to natural or anthropogenic causes. ...

Extended studies of interspecific relationships in Daucus (Apiaceae) using DNA sequences from ten nuclear orthologues

Botanical Journal of the Linnean Society

... The high coefficients of variation (≥20%) signify a high degree of variability in the collection. This was confirmed by high levels of diversity index H' (0.77 and 0.76) for quantitative and total diversity respectively (Mezghani et al., 2017a) showing Tunisia to contain significant diversity for Daucus in the southern Mediterranean region (Mezghani et al., 2014;Ben Amor et al., 2019). Qualitative characters were evaluated by recording the predominant character states for each trait. ...

ASSESSMENT OF PHENOTYPIC DIVERSITY IN TUNISIAN CARROT (Daucus carota subsp. sativus) LANDRACES

... sativus L.) is a nutrient-rich and delicious root vegetable consumed by people of all age groups [1]. In terms of its economic value and nutritional benefits, carrot is considered as the second most important vegetable crop [2]. Due to its protandrous nature, carrot is a highly outcrossing crop, resulting in broader genetic diversity within the cultivated gene pool. ...

Carrot Organelle Genomes: Organization, Diversity, and Inheritance
  • Citing Chapter
  • May 2019

... carota and D. rouyi Spalik & Reduron. This genus is widely distributed worldwide, but with a concentration of diversity in the Mediterranean biogeographic region (Spooner 2019). The former species has hooked indehiscent fruitsschizocarpswell adapted to external adhesion to fur and feathers of animal dispersers (epizoochory) and is globally distributed in inland environments (Banasiak et al. 2016). ...

Daucus: Taxonomy, Phylogeny, Distribution
  • Citing Chapter
  • May 2019

... illnesses caused by a variety of microbes, inheritable disorders, and contagious factors such as bacteria, fungi, and viruses. Huang et al. (2019) Potato plant leaves have been utilized in this study. The majority of tuber leaf illnesses are caused by fungi and bacteria. ...

Analyses of 202 plastid genomes elucidate the phylogeny of Solanum section Petota