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Korean J. Pl. Taxon > Volume 56(2); 2026 > Article
JORDÃO, KIM, BYUN, SEOL, TAMBOLI, AHMED, CHOI, and MATTHEWS: Plastid genome reveals the genetic diversity and phylogenetic relationships of cultivated taro (Colocasia esculenta) in Korea

Abstract

Taro (Colocasia esculenta) is a culturally and nutritionally important crop with a long history of cultivation in Korea. However, the genetic diversity of Korean taro and its relationship to global taro diversity have not been studied. Here, we report the complete plastid genomes (plastomes) of four cultivated taro samples (CESKR01–CESKR04) from two provinces in South Korea: three from Chungcheongnam-do and one from Jeollanam-do. The plastomes ranged from 162,376 to 162,546 bp and exhibited a GC content, gene composition, and quadripartite structure typical of this species, with a large single-copy region, a small single-copy region, and a pair of large inverted repeats. Phylogenetic analyses of full-length multiple sequence alignments robustly identified two major plastid lineages (CI and CII) in cultivated taro, as previously reported. CESKR01, CESKR02, and CESKR03 grouped with the temperate-associated CII haplogroup, whereas CESKR04 grouped with the tropical CI haplogroup. These maternal lineages of taro in Korea are presumed to reflect independent introduction histories. Through comparative analyses, we identified seven highly variable loci in the CI and CII lineages that can be used in future characterization of Korean taro. A comprehensive taro sampling strategy is required to investigate the phylogeography of Korean taro in relation to local climate conditions and the global context.

INTRODUCTION

Taro, Colocasia esculenta (L.) Schott, is an ancient crop of great agricultural, cultural, and biological significance (Chaïr et al., 2016; Matthews and Ghanem, 2021; Lebot and Ivančič, 2022; Tan et al., 2025). It is cultivated widely across the globe—from humid tropical to warm temperate regions—and is a common component of diets in the Pacific Islands, Asia, Africa, and parts of the Americas. Even at the northern edge of taro’s climatic range, in northern China, South Korea, and Japan (East Asia), taro is commonly and traditionally grown and consumed as a local food (Siegmund, 2010; Matthews and Ghanem, 2021). Despite its wide cultivation and high nutritional value, taro has often been regarded as an “orphan crop” in scientific research (Matthews and Ghanem, 2021). There is a clear need for more comprehensive study of taro, given its importance for food security and cultural heritage in many communities and countries, and potential for further development as a crop in diverse physical environments.
Taro belongs to the arum family (Araceae) and is one of several species in the genus Colocasia Schott, with most being wild species known only to taxonomists. Recent phylogenomic studies have begun to clarify taro’s evolutionary relationships, natural distribution as a wild breeding species, and global phylogeography as a clonally propagated crop (Chaïr et al., 2016; Ahmed et al., 2020; Matthews et al., 2024). Plastid DNA (ptDNA; also referred to as chloroplast DNA) analyses of cultivated and wild taros have revealed a surprisingly complex genetic structure: C. esculenta is comprised of at least three major plastid lineages (haplogroups) that broadly correspond to different environmental adaptations and geographies. The CI lineage is found in wetland cultivars and commensal wild taros in tropical to subtropical Southeast Asia and Oceania, CII has been found mainly in dryland cultivated taros in temperate East Asia and other subtropical-to-temperate regions, and CIII has been found exclusively in wild taro populations from Southeast Asia through New Guinea to northern Australia.
In Korea, taro (toran, or t’oran, meaning “earth eggs”) has been cultivated for centuries (Siegmund, 2010), yet it remains a relatively minor crop compared to staple cereals (Kim et al., 2019). Despite taro’s long presence in Korean agriculture, its phylogenetic position and genetic diversity in a global context remain uncharacterized. Previous authors have suggested that taro may have been introduced to Japan via China, Southeast Asia, or the Korean peninsula. However, their studies relied on limited molecular characterization (Matsuda and Nawata, 2002; Huh and Choi, 2003) or included only a single Korean sample (Motohashi et al., 2024), or no Korean samples (Matsuda and Nawata, 2002), thereby limiting the resolution of introduction history and past dispersal routes into and across East Asia.
To begin addressing this knowledge gap, we sequenced and analyzed the complete plastid genomes (plastomes) of four cultivated taro samples collected in Korea, during a pilot field survey carried out in late 2025. The new plastome data provide a genomic basis for distinguishing Korean taro lineages and provide a foundation for future comparative studies on the origins, introduction routes, and genetic composition of cultivated taros in East Asia.

MATERIALS AND METHODS

Plant materials and next-generation sequencing

Leaves of C. esculenta were obtained from two growers in two localities in Korea in early September 2025 (late-summer/early-autumn), before the main taro harvest season (early winter). Three samples (CESKR01 [field no. CV3], CESKR02 [field no. CV4], and CESKR04 [field no. CV2]) were collected from Seongdong-myeon, Nonsan-si, Chungcheongnam-do (Figs. 13, Online Supplementary Note 1), and one sample [CESKR03 (field no. CV5)] was collected from Gokseonggun, Jeollanam-do (Fig. 4, Online Supplementary Note 1). Grower 1 was a small-scale farming innovator, with a range of subtropical crops being tested alongside traditional Korean crops. Grower 2 was a large-scale commercial grower employing green houses and intensive production facilities to maximize taro yield (Online Supplementary Note 1). The collected leaf samples were immediately dried in silica gel to ensure preservation. Total genomic DNA was later extracted using the NucleoSpin Plant II kit (Macherey-Nagel, GmbH, Düren, Germany), following the sodium dodecyl sulfate method. Sequencing was performed by Novogene (Hong Kong) using the Illumina NovaSeq platform, generating 150 bp paired-end reads.

Plastid genome assembly and annotation

The next-generation sequencing reads were de novo assembled using GetOrganelle v1.8.0.1 with default parameters (Jin et al., 2020). Using Geneious Prime v2025.0.1 (https://www.geneious.com/), the large single-copy (LSC), small single-copy (SSC), and inverted repeat (IR) regions of the plastomes were identified. Genome annotation was performed using PlastidHub (Zhang et al., 2025), followed by manual curation in Geneious Prime using three published C. esculenta plastomes (GenBank accession nos. JN105690, PP817737, and PP817736) as references.

Phylogenetic analyses

To determine the phylogenetic placement of the assembled plastomes from Korean C. esculenta samples, three genomic datasets were generated for phylogenetic analyses: protein coding sequences (CDS), complete plastome, and intergenic regions (IGS) (Table 1). The in-group comprised a total of 18 plastomes, consistent with the sampling used by Matthews et al. (2024). The out group included C. fallax Schott (CFABD01) for the CDS dataset, C. oresbia A. Hay (CORMY01 and CORMY02) for the complete plastome dataset, and C. formosana Hayata (CFOTW03) together with C. spongifolia P. J. Matthews, V. D. Nguyen, Q. Fang & C. L. Long (CSFVN01) for the IGS dataset. In total, 23 plastomes were included in the analysis, including the four newly assembled samples (Table 1, Online Supplementary Table 1). In all datasets, one copy of the inverted repeat (IRa) was removed to avoid redundancy in plastome alignments.
For all three datasets, poorly aligned positions were removed using Gblocks v0.91b with default parameters (Castresana, 2000). Maximum likelihood analyses were conducted in IQ-TREE v3.0.1 with 100,000 bootstrap replicates (Wong et al., 2025). The best-fit nucleotide substitution model was selected following Matthews et al. (2024) (Table 1). Phylogenetic branches with near-zero lengths were collapsed. The resulting trees were visualized using the Interactive Tree Of Life (iTOL) web tool (Letunic and Bork, 2021).

Nucleotide diversity analysis of Colocasia esculenta plastomes

The analysis of nucleotide diversity (Pi) focused on C. esculenta individuals belonging to the CI and CII lineages, which include all four Korean taro individuals analyzed in this study. Accordingly, a total of 11 complete plastome sequences representing these two lineages were included, comprising four newly assembled plastomes and seven previously reported plastomes (Online Supplementary Table 1). Sequence alignment was performed in Geneious Prime v2025.0.1 using the MAFFT algorithm (Katoh and Standley, 2013) with default parameters. The aligned dataset was analyzed in DnaSP v6.12.03 using a sliding-window approach (window length = 600 bp; step size = 200 bp) to estimate nucleotide diversity across the plastome (Rozas et al., 2017). Pi values were visualized in RStudio v2025.05.1 (Posit, PBC) using the ggplot2 package (Wickham, 2016).

RESULTS

Plastid genome characteristics

The plastomes of C. esculenta individuals were successfully assembled and have been submitted to GenBank under the accession numbers PX574384, PX574385, PX574386, and PX574387 (Table 2). In total, 64,110,506–74,710,390 raw reads were generated using the Illumina NovaSeq platform. Of these, the numbers of reads mapped to the plastomes were 2,444,547 (CESKR01; 2,253X), 2,920,662 (CESKR02; 2,692X), 5,296,447 (CESKR03; 4,878X), and 2,242,503 (CESKR04; 2,072X). The plastomes range in length from 162,376 to 162,546 bp and are organized into three major regions: a LSC region of 89,643–89,816 bp, a SSC region of 22,061–22,188 bp, and a pair of IRs of 25,273–25,334 bp each (Fig. 5). The GC content of the plastomes ranges from 36.1 to 36.2%. Each plastome comprises 131 genes, including 86 protein-coding genes (PCGs), 37 transfer RNA (tRNA) genes, and eight ribosomal RNA (rRNA) genes. The LSC region contains 60 PCGs and 22 tRNA genes, the SSC region contains 12 PCGs and one tRNA gene, and the IR regions contain seven PCGs, seven tRNA genes, and four rRNA genes, all of which are duplicated in the IR regions.

Phylogenetic analyses

The resulting phylogenetic trees were well resolved, with most nodes receiving high bootstrap support (BS) (Fig. 6, Online Supplementary Figs. S1, S2). Across all three datasets (CDS, Complete plastome, and IGS), the two major plastid lineages (CI and CII) of C. esculenta were strongly supported. The newly sequenced samples CESKR01, CESKR02, and CESKR03 were consistently placed in the CII lineage across all datasets, whereas CESKR04 was placed in CI. In all analyses, CESKR04 was resolved as sister to CESJP30 (BS = 100), a southern Japanese wetland cultivar known as Taimo (“pond-field taro”). The two plastome sequences were identical, but the two cultivars have very different phenotypes (Ta-imo is a largely green plant forming the lateral corms, not long stolons; see Takaesu et al., 2025). CESKR02 was resolved as sister to CESPK08, a cultivated accession from Pakistan (BS: CDS = 89; Complete = 99; IGS = 98), with no detectable sequence variation between the two plastomes. CESKR01 and CESKR03 formed a well-supported sub-lineage with CESNZ02, a common dryland cultivar in New Zealand (BS: CDS = 100; Complete = 100; IGS = 97). The plastome sequences of CESKR01, CESKR03, and CESNZ02 were identical.

Identification of nucleotide diversity hotspots in Colocasia esculenta

Sliding-window analysis of the complete plastomes from 11 C. esculenta individuals belonging to the CI and CII lineages revealed Pi values ranging from 0 to 0.024, with an overall mean of approximately 0.001. A total of seven highly variable loci (hotspots) were identified (Fig. 7). In the LSC region, three hotspots were identified: trnHpsbA (0.007), trnStrnG (0.006), and accDpsaI (0.005). In the SSC region, the most variable locus was trnNndhF (0.024), with additional hotspots observed in ndhFrpl32 (0.005), ccsAndhD (0.006), and rps15ycf1 (0.006).

DISCUSSION

Comparative analyses of these plastome sequences allowed us to evaluate genetic divergence among Korean taro samples and to assess their maternal lineage relationships. Phylogenetic analyses (Fig. 6, Online Supplementary Figs. S1, S2) based on CDS, IGS, and complete plastome alignments consistently recovered the two major cultivated taro plastid lineages, CI and CII, with strong support, in agreement with previous studies (Ahmed et al., 2012, 2013, 2020; Matthews et al., 2024).
Among the four Korean taro samples, three (CESKR01, CESKR02, and CESKR03) were consistently placed in the CII lineage across all analyses (CDS, IGS, and complete plastome). These appear identical to two closely-related reference CII haplotypes from New Zealand and northern Pakistan (Fig. 6), indicating that the Korean CII taros share maternal ancestry with other temperate-region cultivars that may have originated in the subtropical mountains of Southeast Asia (eastern Himalaya) (Matthews 2014, 2023). In contrast, CESKR04 was placed in the CI lineage together with a reference CI genome from southern Japan that is also identified as the CI Type 1 haplotype shared by tropical taros across Asia and the Pacific (Ahmed et al., 2020; Matthews et al., 2024). The growth of CESKR04 in Korea’s temperate climate is notable, particularly as this recently introduced cultivar was able to reach maturity and flower. During our survey, we also observed ornamental taro varieties flowering at Gungnamji, an artificial pond in Buyeo-si, Chungcheongnamdo. These observations raise the possibility that controlled breeding with winter protection may be possible in Korea. In the context of global climate change, continued warming may further facilitate the cultivation and spread of tropical (CI) taro lineages in Korea, potentially contributing to shifts in the composition of locally adapted taro cultivars. Collectively, these findings show that taro cultivated in Korea does not derive from a single domesticated lineage but instead comprises multiple, genetically divergent maternal origins. This result underscores the utility of plastome-based phylogenomic approaches for revealing infraspecific diversity and for informing strategies in conservation, breeding, and climate-resilient crop development.
The present result arises from just a small pilot field survey, which raises the question of what more might be found in Korea through surveys involving diverse growers with home gardens, experimental farms, small- or large-scale commercial production, and ornamental plant collections, over the entire Korean peninsula and offshore islands. Consulting the popular iNaturalist plant reporting platform (Internet, 15 January 2026) shows that actual distribution of cultivated taro in Korea is much wider than has been stated cumulatively in published reports seen by us. In Korean urban centers (Seoul and elsewhere), potted taro plants are commonly grown alongside other potted vegetables in very small-scale “house gardens” along the sides of buildings and roadsides (authors’ observations). We did not survey such potential sources. Taro is used in Korea not just as a starchy food plant, but also as a source of peeled petioles, fresh or dried. Investigating the varieties used for starch, as green vegetables, or both will also be of historical and practical interest, and can be linked to the food cultures involving taro across East Asia and the world.
The Korean taro samples exhibited pronounced morphological variability (Figs. 14), and different lineages may also differ in chemical composition and taste (Gerrano et al., 2021; Sánchez-Chino et al., 2021; Mitharwal et al., 2022; Tan et al., 2025). Clarifying the evolutionary relationships and geographic origins of these diverse forms requires reliable molecular markers, and Sanger sequencing of short, highly informative target sequences can be cost-effective for wide geographical surveys with many samples. To guide future studies, we identified several plastome regions with elevated nucleotide diversity that may serve as lineage-specific markers (Fig. 7). In particular, variation was concentrated in both LSC and SSC regions, with the trnNndhF intergenic spacer showing the highest variability, indicating its potential as a promising molecular marker.
Several of the highly variable loci identified in this study overlap with previously reported ptDNA markers for taro. Ahmed et al. (2013) described 30 ptDNA markers, five of which—trnS–trnG, accD–psaI, ndhF–rpl32, ccsA–ndhD, and rps15–ycf1—correspond to loci identified here as highly variable. These loci revealed major plastid lineages CI, CII, and CIII (Ahmed et al. 2020) that were subsequently confirmed with whole plastome analysis (Matthews et al., 2024). Two highly variable loci detected in our analysis (trnH–psbA and trnN–ndhF) were not included among previously-identified variable loci. Incorporating these loci—particularly trnNndhF, the most variable region identified—into taro genotyping frameworks could enhance the utility of plastid markers for distinguishing closely-related clonal lineages of taro, including those cultivated in Korea. The trnN–ndhF IGS is located near the IRb–SSC junction, where intraspecific variation can be influenced by IR boundary expansion/contraction and associated gene-conversion processes. Comparable junction-proximal intraspecific polymorphism has been reported in Medicago minima (L.) Bartal., in which small-scale shifts at the IR/SSC boundary were accompanied by gene conversion that homogenizes repeat copies and may overwrite adjacent single-copy sequences, thereby complicating strict positional homology in this region (Choi et al., 2020). For these reasons, variation in IR boundary–associated regions may have limited reliability for phylogenetic inference (Lockhart et al., 2006; Abdullah et al., 2020, 2021). Nevertheless, such highly variable plastome polymorphisms can remain informative for lineage or sub-lineage discrimination.
This study provides the first plastome-level assessment of genetic diversity in Korean taro and contributes to a clearer understanding of intraspecific phylogenetic structure within C. esculenta. Our results establish a foundation for more precise differentiation of taro cultivars in Korea, provide genomic resources relevant to the conservation and utilization of taro crop diversity, and clarify the position of Korea’s germplasm within the wider context of taro’s evolution, domestication and worldwide dispersal.

Online Supplementary Data

Tables S1, Figs. S1, S2, and Note S1 are available at https://doi.org/10.11110/kjpt.2026.56.2.129

NOTES

ACKNOWLEDGMENTS
This research was supported by the Regional Innovation System & Education (RISE) program through the Daejeon RISE Center, funded by the Ministry of Education (MOE) and the Daejeon Metropolitan City, Republic of Korea (2026-RISE-06-013). This research was also supported by the Bio-Health Convergence Research Institute of Hannam University. Fieldwork in Korea by P.J.M. was supported by a Mitsubishi Foundation grant (no. J24M000009) for the project “Food traditions and genetic diversity of taro and giant taro,” with practical assistance from Etsuko Tabuchi. We give special thanks to the local growers who kindly showed us their fields and gardens and granted permission for sampling. We also thank two anonymous reviewers for their helpful comments.
CONFLICTS OF INTEREST
The authors declare that there are no conflicts of interest.

Fig. 1
A. Whole plant: “Purple-green”, CESKR01, field no. CV3; the light-exposed petiole is purple at the base, grading to green toward the upper part; lateral corms are small and numerous. B. The unexposed petiole base and basal ring are white; accessory buds are white (black arrows); the main axillary bud is white (yellow arrow); side corms are globular. C. Cross-section of the corm, showing white interior tissue.
kjpt-56-2-129f1.jpg
Fig. 2
A. Whole plant: “Green”, CESKR02, field no. CV4; the light-exposed petiole is mostly green; the upper petiole is purple at the junction with the blade in some leaves (a commonly variable character in taro); the lateral corms are few (becoming elongate). B. The unexposed petiole base and basal ring are white; accessory buds are white (black arrows); the main axillary bud is white (yellow arrow). C. Cross-section of the corm, showing white interior tissue.
kjpt-56-2-129f2.jpg
Fig. 3
A. Whole plant: “White”, CESKR03, field no. CV5; the light-exposed petiole is mostly green, but the plant is locally referred to as “white”; the upper petiole is purple at the blade junction in some leaves (a commonly variable character in taro); lateral corms are round and sprouting. B. Collection site, a large-scale commercial hothouse, is shown next to rice fields. C. The unexposed petiole base and basal ring are white; accessory buds are white (black arrows); the main axillary bud is white (yellow arrow); cross-sections where the lateral corms were removed are white.
kjpt-56-2-129f3.jpg
Fig. 4
A. Whole plant: “Pink”, CESKR04, field no. CV2; petioles are pink or pinkish when young, becoming green with maturity; stolons, peduncles and spathal tube are also coloured pink to reddish-purple. B. Floral details (spathe and spadix); spadix structure from left to right shows female zone, sterile interstice, male zone, and sterile appendix (6 Sep 2025, early Autumn). C. Accessory buds are small, indistinct (black arrows); main axillary is bud pink and large (yellow arrow); cross-section of the corm, showing white interior tissue.
kjpt-56-2-129f4.jpg
Fig. 5
Plastid genome map of Colocasia esculenta. Genes on the outer and inner circles are transcribed clockwise and counterclockwise, respectively. Gene functional categories are indicated by different colors, and the inner circle represents guanine-cytosine (GC) and adeninethymine (AT) content. Plastome lengths range from 162,376 to 162,546 bp among the four Korean C. esculenta individuals.
kjpt-56-2-129f5.jpg
Fig. 6
Maximum-likelihood phylogenetic tree of Colocasia esculenta based on complete plastome sequences, including Korean individuals. Bootstrap support values are shown at the nodes, and Korean individuals are marked with red asterisks. The CI, CII, and CIII lineages are indicated in yellow, blue, and green, respectively.
kjpt-56-2-129f6.jpg
Fig. 7
Sliding window analysis of nucleotide diversity (Pi) across the complete plastomes of 11 Colocasia esculenta individuals from the CI and CII lineages. The X-axis represents genome position (bp), and the Y-axis indicates Pi values. The red dashed line denotes the hotspot threshold (Pi > 0.00461), and red dots highlight the identified hotspot loci.
kjpt-56-2-129f7.jpg
Table 1
Summary of plastome datasets and alignment statistics.
Genome component No. of species No. of sequences Aligned (bp) Model
CDS 6 23 68,779 TVM + F + I
Complete plastome 5 22 134,054 K3Pu + F + I
IGS 4 20 45,079 K3Pu + F + I

All analyses were performed after removing one copy of the inverted repeat (IRa) and gap regions. Substitution models used were TVM + F + I (Transversion model with unequal base frequencies and a proportion of invariable sites) and K3Pu + F + I (Kimura 3-parameter model with unequal base frequencies and a proportion of invariable sites). CDS, protein-coding sequences; IGS, intergenic spacer regions.

Table 2
Characteristics of four Colocasia esculenta plastomes.
Sample label CESKR01 CESKR02 CESKR03 CESKR04
GenBank accession number PX574384 PX574385 PX574386 PX574387
Size (bp)
 Total 162,546 162,478 162,546 162,376
 LSC 89,816 89,810 89,816 89,643
 SSC 22,063 22,061 22,063 22,188
 IR 25,334 25,304 25,334 25,273
GC content (%) 36.1 36.2 36.1 36.2

IR, inverted repeat; LSC, large single copy; SSC, small single copy; GC, guanine-cytosine.

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