Research Article

Korean Journal of Plant Resources. 30 December 2022. 805-819



  • Introduction

  • Materials and Methods

  •   Plant materials

  •   DNA extraction and PCR amplification

  •   DNA sequencing and alignment

  •   Phylogenetic analysis

  •   Comparison of morphological characters

  • Results

  •   Morphological characters

  •   Phylogenetic analysis

  • Discussion

  •   Morphological characters

  •   Phylogenetic analysis

  •   Naming of a new forma

  •   Taxonomic treatment

  •   Identification Key in O. monacantha complex


At the 1578 Compendium of Materia Medica, Opuntia monacantha f. jejuensis was introduced as a green cactus in the Encyclopedia of Oriental Herbal Medicine and was studied until 1975. Since its introduction in Jeju, it has been used medicinally in the region (Kwon et al., 2017; Koh et al., 2018). Currently, O. humifusa is grown on a large scale in the Chungcheong and Jeolla regions (Kim et al., 2014). In particular, O. stricta is being cultivated on ~320 ㏊ of land (Yang et al., 2020). Cacti of the genus Opuntia, native to or grown in Korea, exhibit superior antibacterial, anti-glycosylation, antioxidant, anticancer, and cholesterol-lowering properties (Chung, 2000; Jung et al., 2012; Park et al., 2013; Choi, 2014; Jung et al., 2014). In addition to kimchi made with Opuntia powder, other foods, such as fermented milk, are also being made (Lee and Bae, 2009; Jung et al., 2016; Kim et al., 2016; Lee, 2017; Lee et al., 2018). Numerous studies have explored the potential of Opuntia as a cosmetic raw material (Lim and Hong, 2016; Kwon et al., 2017).

Cacti is a collective name for succulent plants belonging to the Cactaceae family of Caryophyllaes. In dry areas, during periods of heavy rainfall, water is stored in the plant body and used for plant growth during the dry season. Cactus has a reduced surface area to minimize water loss in arid regions, converting leaves to spines, and shortening stems and roots. Among the 3,000 species of succulent plants found worldwide, ~1,800 are cacti, which originated in South America and evolved in a dry environment (Anderson, 2001). Cactaceae are classified into four subfamilies by the International Cactaceae Systematics Group: Pereskoideae, Maihuenioideae, Opuntioideae, and Cactoideae (Nyffeler and Eggli, 2010). Opuntioideae is classified into seven genera: Brasiliopuntia, Tacinga, Consolea, Miqueliopuntia, Salmiopuntia, Opuntia and Tunilla (Wallace and Dickie, 2002; Griffith and Poter, 2009; Hernandez-Hernandez et al., 2011). In phylogenetic studies, it is classified into ten series: Brasiliopuntia, Tacinga, Elatae, Macbridei, Scheerianae, Humifusa, Macrocentra, Nopalea, Basilares and Microdasys (Table 1), (Majure et al., 2012b).

Table 1.

Classification of Opuntia s.s. based on phylogenetic studies (Majure et al., 2012b)

Sister clade of
Opuntia s.s.
Brasiliopuntia, Tacinga
Opuntia s.s. South American Clades Elatae, Macbridei
North American Clades Scheerianae, Humifusa, Macrocentra, Nopalea, Basilares, Microdasys

Opuntias.s. is a large genus of Cactaceae with ~200 species (Anderson, 2001). In Korea, O. humifusa and O. humifusa f. jeollaensis are native and are cultivated in the regions of Chungcheong, Jeolla, and Gyeongsang (Park et al., 2013; Kim et al., 2014). In the Jeju region, O. stricta Haw. also grows naturally and is cultivated (Yang et al., 2020; Yang and Oh, 2021), and O. monacantha f. jejuensis (this study) grows naturally (Fig. 1), (Yang and Oh, 2021).
Fig. 1.

Distribution map of O. monacantha f. jejuensis (●) and O. stricta (◉) in Jeju Island, Korea.

O. monacantha f. jejuensis is the largest indigenous cactus species in Korea and is found on Jeju Island in the Aewol, Taepyeong, and Bomok regions (Fig. 1). In previous studies, External morphological studies have differentiated O. monacantha f. jejuensis from O. stricta by renaming the misidentified O. stricta, cultivated in the Hallim area of Jeju, and examining the phylogenetic relationship between them (Yang et al., 2020; Yang and Oh, 2021). The taxonomic analysis of O. monacantha sensu lato and its synonymous species is shown in Table 2. O. vulgaris Mill. had been accepted as a synonym of O. monacantha or O. humifusa but was later accepted as O. ficus-indica (L.) Mill. (Leuenberger, 1993). O. archavaletae Speg. was classified as a subspecies based on its different characteristics (flower, fruit) of reproductive organs (O.monacantha subsp. archavaletae (Speg.) Guiggi.) (Guiggi, 2017). A variety of O. monacantha was named O. monacantha var. gracilior Salm-Dyck (Table 2).

Table 2.

The taxonomic history of O. monacantha

Scientific Name Reference
Synonym Accepted Scientific Name:
O. monacantha (Willd.) Haw.
= C. monacanthos Willd.
= O. deflexa Lem.
= O. gracilior Lem.
= O. lemaireana Console ex Bois
= O. lemaireana Console ex F.A.C.Weber
= O. monacantha var. deflexa Salm-Dyck
= O. monacantha var. gracilior Lem.
= O. monacantha var. variegata Anon.
= O. vulgaris var. lemaireana
(Console ex Bois) Backeb.
Haw. 1819. Suppl. pl. succ. 81.
Willd. 1814. Enum. Pl. Hort. Berol. Suppl.:33
Lem. 1839. Cact. Gen. Sp. Nov.:68
Lem. 1839. Cact. Gen. Sp. Nov.:68
Bois. 1898. Dict. Hort. 2:894
Weber, F.A.C.D. 1898. G. J. M. Bois, Dict. hort.2:894.
Salm-Dyck. 1850. Cact. Hort. Dyck.:66
Lem. 1839. Cact. Gen. Sp. Nov.:68
Anon. 1874. Gard. Chron., n.s. 2:49
Backeb. 1958. Cactac.: Handb. Kakteenk. 1:400
Re-evaluation O. vulgaris Mill.

(Accepted Scientific name:
O. ficus-indica (L.) Mill.)
Mill. 1768. Gard. Dict. ed. 8: n 1., nom. rej.

(Leuenberger, 1993; Plant List, 2013)

Subspecies O. arechavaletae Speg.

Accepted Scientific Name:
O. monacantha subsp. arechavaletae (Speg.) Guiggi.
Speg. 1905. Anal. Mus. Buenos Aires, 11: 520.

Guiggi. 2017. Suppl. 5:1.

Variety O. monacantha var. gracilior Salm-Dyck Salm-Dyck. 1850. Cact. Hort. Dyck.(1849). 66, 235.

Generally, phylogenetic analyses are conducted using the maximum likelihood (ML) and Bayesian inference (BI) methods which can be applied to a model of sequence evolution, and phylogenetic trees can easily be constructed using sequence data. Calculating the likelihood, ML maximizes the likelihood of the model for all data, whereas BI maximizes the posterior probability using likelihood. ML typically uses phylogenetic maximum likelihood (PhyML) or randomized accelerated maximum likelihood programs, whereas BI uses Bayesian inference of phylogeny (MrBayes) or Bayesian evolutionary analysis sampling tree (BEAST) programs to construct a phylogenetic tree. The obtained node support value for a phylogenetic tree written in ML is referred to as a bootstrap, and that in BI is called posterior probability. Majure et al. (2012a) used genetic ML and BI methods to conduct a phylogenetic analysis of the genus Opuntia. Kim et al. (2014) conducted genetic ML analysis and morphological classification by floral color for O. humifusa f. jeollaensis. Therefore, it was necessary to identify the position of O. monacantha f. jejuensis in the Opuntia via morphological comparisons and phylogenetic analysis using MI and BI methods.

Comparing the pulp color of O. monacantha f. jejuensis to that of O. monacantha, a distinct difference was observed in the characteristics of the two species (Koh et al., 2018; Yang and Oh, 2021). Owing to the lack of studies confirming the exact taxonomic position of O. monacantha f. jejuensis, which exhibits a distinct difference in pulp color and characteristics from O. monacantha, species identification studies are required before considering it as a raw material for pharmaceuticals, health supplements, and cosmetics. Therefore, to determine the exact phylogenetic position of O. monacantha f. jejuensis within the Elatae series, we compared the morphological characteristics of the O. monacantha complex and comfirmed its taxonomic status using phylogenetic analysis. This study contributes to validating a new forma of the taxon and its potential use as a health-promoting material.

Materials and Methods

Plant materials

The O. monacantha sensu lato used in this study was collected from the Taepyeong-ro habitat in Seogwipo (33°14'40.6" N 126°32'38.0" E). We transferred the collected materials were transferred to the cultivation site of the Baiknyuncho Museum in Hogeun-dong, Seogwipo, Jeju Island, for morphological studies. An immersion specimen was created and stored in Baiknyuncho Museum along with the associated resource management number. For DNA sequencing, one-year-old leaves were collected considering the occurrence of bacteria and contamination levels, and the obtained material was washed with distilled water, frozen in liquid nitrogen, and stored at -80℃ to extract the DNA.

DNA extraction and PCR amplification

Genomic DNA was isolated using the CTAB method (Doyle and Doyle, 1990). The primers and protocol were designed using the plastid intergenic spacers atpB-rbcL, trnL-F, and psbJ-petA, which are suitable for plant taxonomy phylogenetic classification studies, chloroplast gene matK, and nuclear ribosomal gene ITS (Table 3). The polymerase chain reaction (PCR) cycling parameters involved initial denaturation was performed at 94℃ for 5 min. The thermal cycle consisted of denaturation at 94℃ for 30 s, annealing at 55℃ for 30 s, and extension at 72℃ for 1 min, and was repeated 35 times. We performed the final extension at 72℃ for 10 min.

Table 3.

Genomic DNA regions and primer sequences used in this study

Gene sequence or reference Length amplified No. pars. infor. characters Model
861 20 HKY
psbJ-petA psbJ: (Shaw et al., 2007)
1169 72 K81uf+I
trnL-F trnL: (Taberlet et al., 1991)
trnF: (Taberlet et al., 1991)
441 14 K81uf
905 27 F81+I+G
nrITS ITS4: (White et al., 1990)
ITS5: (White et al., 1990)
599 39 TVM+G
cpDNA combined - 3407 115 -
Nuclear combined - 568 57 -
All combined - 3975 172 -

DNA sequencing and alignment

For sequencing of the five genes (atpB-rbcL, psbJ-petA, trnL-F, matK, and nrITS) of the investigated taxon, 25 cycles of sequencing reactions were performed using the Big Dye Terminator Cycle Sequencing Kit v.3.1 and the automated DNA sequencing system, Applied BioSystems DNA 3730XL Analyzer (Applied BioSystems, Foster City, CA, USA). DNA was purified using the Montage SEQ96 PCR clean-up kit (Millipore Corporation, Billerica, MA, USA). The nucleotide sequence was aligned using the ClustalX program (Thompson et al., 1997) and edited and rearranged using BioEdit ver. program (Hall, 1999).

Phylogenetic analysis

After thoroughly confirming the DNA sequence of the target species, the sequences of the species were compared and analyzed from a molecular evolutionary perspective (Majure et al., 2012a,b). Three previously studied (Kim et al., 2014) cacti (O. humifusa, O. humifusa f. jeollaensis, and O. stricta) cultivated in Korea were sampled to obtain evolutionary data on O. monacantha f. jejuensis. Phylogenetic relationships were confirmed using the Tacinga series as outgroups using genetic information registered in the GenBank of the National Center for Biotechnology Information (Clegg, 1993; Soltis et al., 1998; Park et al., 2010), (Appendix 1). Comparative analysis was used to establish the exact position of O. monacantha f. jejuensis within the Elatae series and the form and DNA sequence of O. monacantha complex, including O. monacantha, O. monacantha subsp. arechavaletae, and O. monacantha f. jejuensis species (Appendix 2). BI and ML analyses were used to confirm the phylogenetic relationships between the species, and the programs MrBayes-3.2.5 (Ronquist et al., 2012) and PhyML 3.0 (Guindon et al., 2010) were used. The GTR+I+G (nst = 6, rates = invgamma) model, based on the main parameters affecting the substitution ratio in the DNA sequence, was used for BI analysis. Each of the five genes was analyzed using a random phylogenetic tree of 1×106 generations and 100 generations of Markov chains. In ML analysis, random samples were selected to duplicate the same number of characters as in the existing data set, repeated 10,000 times to form the same tree using PhyML 3.0 program, a phylogenetic tree, and bootstrap.

We constructed a molecular phylogenetic tree to illustrate phylogenetic relationships between species using TRE and PhyML files generated by BI and ML analysis, respectively. The molecular phylogenetic tree was confirmed using the FigTree v1.4.3 program (Rambaut, 2016), which was designed to display the summarized contents and annotated phylogenetic tree produced by BEAST.

Comparison of morphological characters

Six morphological characteristics, i.e., tepal color, stigma lobe color, fruit color, pulp color, and morphological differences between the seed, stem, and fruit, were selected through a comparative morphological study of the investigated taxon, i.e., O. monacantha complex, and evolutionary differentiation patterns within the O. monacantha complex were compared using a phylogenetic tree.


The results of morphological and phylogenetic analyses of the Elatae series, which includes the O. monacantha complex, O. monacantha f. jejuensis, and Korean Opuntia spp., showed different flower colors, cladode shapes, fruit shapes, fruit colors, and pulp colors. O. monacantha f. jejuensis was assigned to the Elatae series in the phylogenetic analysis and was more closely related to O. monacantha subsp. arechavaletae than to O. monacantha at a molecular level.

Morphological characters

The investigated taxon had different morphological characteristics than other Opuntia spp. in Korea (Table 4). The cladodes of mature specimens of O. stricta, O. humifusa, and O. humifusa f. jeollaensis are ovate, but those of O. monacantha f. jejuensis were oblong. Furthermore, plant size, flowering period, fruit shape, and pulp color were significantly different. O. monacantha f. jejuensis is a tree-like plant that grows to a height of > 2 m, while other species are shrub-like plants that grow to a height of GTTCTAGCACCAGAAAGTCG 1 m. O. monacantha f. jejuensis fruits are plum-shaped, whereas those of other species are elongated pear-shaped. The fruit of the investigated taxon is purple-green, whereas those of the others are reddish-purple (Fig. 2). Finally, the pulp of the investigated taxon was yellow-green, whereas that of the other species was purple.

Table 4.

Morphological differences of O. monacantha f. jejuensis and Opuntia spp. in Korea

Taxa O. monacantha
f. jejuensis
O. strictaO. humifusaO. humifusa
f. jeollaensis
Flower color yellow-with
out side purple stripes
yellow yellow yellow-with
red in the red center
Cladode shape Oblong (elongated) Ovate or Obovate Ovate or Obovate Ovate or Obovate
Fruit shape a plums a long western pear a long western pear a long western pear
Fruit color purple-green reddish-purple reddish-purple reddish-purple
Pulp color yellow-green purple purple purple
Plant size 2-5 m
erect, succulent
tree-like plant
less than 0.8 m
erect, succulent
shrubby-like plant
less than 1 m
shrubby-like succulent
less than 1 m
shrubby-like succulent
Fig. 2.

Morpholological characteristics of Cladode, flowers, fruit, and pulp of O. monacantha f. jejuensis, illustrating their morphological characteristics. The scale bar (white) across the bottom of each photograph is 2 ㎝. O. monacantha f. jejuensis (A). Germinated O. monacantha f. jejuensis (B). Oblong shape of a cladode (C). Flower with purple stripes on the outer surface of yellow petals (D,E). Plum-shaped fruit (F-H). Spine (4 ㎝ long) (I). Stigma lobes (×10) (J).

O. monacantha f. jejuensis was observed to be morphologically similar to O. monacantha. However, O. monacantha had yellow tepals with abaxially red midveins, i.e., purple or red stripes on the front and back of tepals (Fig. 3), (Majure and Puente, 2014), whereas these stripes appear only on the back of O. monacantha f. jejuensis tepals. O. monacantha and O. monacantha subsp. arechavaletae had yellow or orange-yellow tepals, whereas O. monacantha f. jejuensis had bright yellow tepals (Fig. 3).
Fig. 3.

Flower and fruits of O. monacantha complex.

Flower of O. monacantha (A-1):

Fruits of O. monacantha (A-2):

Flower of O. monacantha subsp. arechavaletae (B-1)

Fruits of O. monacantha subsp. arechavaletae (B-2)

Stigma lobe color, fruit color, seeds, pulp color, and peculiar phenomenon were the traits used for morphological classification. O. monacantha has creamish stigma lobes, O. monacantha subsp. arechavaletae has greenish stigma lobes (Guiggi, 2017; Maria et al., 2017), and O. monacantha f. jejuensis has creamish stigma lobes (Table 5, Fig. 3). The mature fruit of O. monacantha and O. monacantha f. jejuensis is purple-green, whereas those of O. monacantha subsp. arechavaletae is reddish-purple. The filaments of the stamens of O. monacantha were greenish in color, according to previous studies (Pardo and Alonso, 2017), but in this study they turned out to be yellow. Those of O. monacantha f. jejuensis were yellow, too. Sprouts of O. monacantha f. jejuensis were germinated (Fig. 2). The fruiting pattern on stems in the investigated taxon can be classified into two distinct types. One is that a stem grows from a fruit, with another fruit growing from this stem and the other is that three fruits grow sequentially at the end of a single stem, which is the most characteristic phenomenon that occurs only in O. monacantha f. jejuensis (Fig. 4).

Table 5.

Morphological differences in O. monacantha complex

Taxa O. monacanthaO. monacantha subsp. arechavaletae O. monacantha f. jejuensis
Yellow or orange yellow tepal, Red
stripes on the inside and
outside of tepals
Yellow or orange yellow tepal, Red
stripes on the outside of tepals
Yellow tepal, Red stripes on the
outside of tepals
lobes color
Pale yellow to pale creamish
(Haworth, 1819)
(Guiggi, 2017)
Pale yellow to pale creamish
Fruit color The fruit maturing and reddish purple,
greenish near base or green with
red-purple shades
(Haworth, 1819)
The fruit maturing and reddish-purple,
greenish near base
(club head part)
The fruit maturing and reddish purple,
greenish near base or green
with red-purple shades
Fruit shape Narrowly turbinate to obovoid fruits,
with well-developed loculus
(Maria et al.,2017)
Plum shape
Longer, elongate,
with an apical and smaller loculus
(Guiggi, 2017)
Club shape
Narrowly turbinate to obovoid fruits,
with well-developed loculus
Plum shape
Seeds 3.5×4 mm
(Haworth, 1819)
- Seeds are light tan,
Irregularly elliptic, ca.
2 ea 5×6 mm
More than 50 ea 3×4 mm
Pulp color Greenish
(Maria et al., 2017)
(Maria et al., 2017)
Inside: Greenish
Outside: Green-yellow
- - Fruit+Stem+Fruit,
Fig. 4.

Characteristic phenomenon of O. monacantha f. jejuensis. (A) Consecutively generated fruits. (B) A flower growing from a fruit. (C) a stem growing from a fruit.

Phylogenetic analysis

Bayesian trees based on DNA sequence showed posterior probabilities summarized from a set of post-burn-in trees generated using the GTR+I+G model (Figs. 5 and 6). Opuntia stricta was included in Scheerianae series and O. humifusa and O. humifusa f. jeollaensis were included in Humifusa series (Fig. 5). These three species formed a part of North American clades. However, unlike other Opuntia spp. in Korea, O. monacantha f. jejuensis was included in Elatae series of South American clades (Fig. 6) as it shares numerous morphological characteristics with O. monacantha. The phylogenetic distinctions between O. monacantha f. jejuensis, O. monacantha, and O. monacantha subsp. arechavaletae (Fig. 6) were not distinguished. We conducted ML and BI analyses of the O. monacantha complex and other species in South American clades to identify the detailed molecular evolutionary aspects of O. monacantha f. jejuensis. The results of the analyses were highly relevant, because O. monacantha f. jejuensis branched with O. monacantha subsp. arechavaletae within O. monacantha complex.
Fig. 5.

Bayesian tree showing the phylogenetic relationship within Opuntia s.s., including Korean taxa (trnL-F, matK and nrITS sequences). Bayesian posterior probability values are shown above branch nodes. The BPP values GTTCTAGCACCAGAAAGTCG 0.76 are not provided.
Fig. 6.

Bayesian and ML trees of South American clades including O. monacantha complex taxa (using atpB-rbcL, trnL-F, psbJ-petA, matK and nrITS sequences). BPP (right) and Bootstrap values (left) are shown between branches. South American clades used the names of the series published by Britton and Rose (1920), Engelmann (1856), and Majure et al. (2012a) as a result of their research. The BPP values are > 0.6 and The Bootstrap values are ≥ 60%.


Morphological characters

When compared to O. monacantha (Wagner et al., 1999; Taylor and Zappi, 2004; Navie and Adkins, 2008) and O. monacantha subsp. arechavaletae (Speg, 1905; Guiggi, 2017), O. monacantha f. jejuensis (Yang and Oh, 2021) showed significant differences in flower color, stigma lobe color, fruit color, seed color, pulp color, and peculiar phenomenon (Fig. 4, Table 4). Based on the findings of this study, O. monacantha f. jejuensis has a pulp color that is distinct from that of O. monacantha. Yang and Oh (2021) reported that O. monacantha f. jejuensis, a species native to Jeju Island, was morphologically different from O. monacantha in fruit and stem growth characteristics. As the mature fruits of plants in Opuntia have the same color as pulp, fruit color indicates pulp color (Omweri et al., 2016). However, the pulp of O. monacantha f. jejuensis is greenish-yellow, which differs from the color of its fruit (Table 5). The mature fruit of O. monacantha and O.monacantha f. jejuensis are purple-green, whereas that of and O. monacantha subsp. arechavaletae are reddish-purple. Moreover, O. monacantha f. jejuensis exhibited an unusual flowering phenomenon on the fruit. In other plants, it is common for the fruit to grow on the stem, but O. monacantha f. jejuensis also showed stem growth from in the fruit. Therefore, fruit areole has the same function as stem areole. This study showed that O. monacantha f. jejuensis is morphologically different from O. monacantha and thus can be classified as a new forma.

Phylogenetic analysis

Following Kim et al. (2014), wherein plastid intergenic spacer trnL-F, plastid gene matK, and nrITS, were used, this study included the three taxa, O. stricta, O. humifusa, and O. humifusa f. jeollaensis, in North American clades. O. stricta was included in Scheerianae series and O. humifusa and O. humifusa f. jeollaensis were included in Humifusa series (Majure et al., 2012b). O. monacantha f. jejuensis was included in Elatae series of South American clades, which is consistent with the findings of previous studies (Yang and Oh, 2021) that it is evolutionarily closer to O. monacantha subsp. arechavaletae than to O. monacantha. Furthermore, according to the research findings of Majure et al. (2012a,b), Kim et al. (2014), and Yang and Oh (2021), alloploid species are a radial evolution type in North American clades. When comparing the O. monacantha complex with other species in Elatae series using phylogenetic trees based on plastid and nrITS sequences, the results were consistent with previous findings that O. monacantha f. jejuensis was evolutionarily closer to O. monacantha subsp. arechavaletae than O. monacantha.

Naming of a new forma

Recently, O. monacantha subsp. arechavaletae has been classified as a subspecies of O. monacantha (Guiggi, 2017). The stigma lobes are yellow in O. monacantha but green in O. monacantha subsp. arechavaletae. O. monacantha f. jejuensis differed morphologically from O. monacantha in several areas (Table 5). The fruit shape, size, stem, and spines of the investigated taxon were morphologically similar to those of O. monacantha, making it difficult to regard it as a new species. However, in light of the morphological (flower color, pulp color, and peculiar phenomenon) and phylogenetic (phylogenetic trees) analysis of O. monacantha f. jejuensis, it was deemed reasonable to classify it as a new forma. The species was named O. monacantha (Willd.) Haw. f. jejuensis J. K. Kim ex Y. S. Yang, based on the habitat characteristics, i.e., native or widely cultivated in Jeju, and the first discoverer and scientific name designator was Jekuk Kim, who is a strong advocate for the taxon's protection, cultivation, and breeding.

Taxonomic treatment

Opuntia monacantha (Willd.) Haw.f. jejuensis J. K. Kim. ex Y. S. Yang,for. nov. (Fig. 7)

Korean name: Je-ju-baik-nyun-cho 제주백년초

Origin: Eastern coastal South America (Argentina, Brazil, Paraguay, and Uruguay)
Fig. 7.

Holotype of O. monacantha (Willd.) Haw. f. jejuensis J. K. Kim. ex Y.S. Yang.

A type of succulent tree (thorny shrub); Erect cylinder trunk shrub, trunk diameter 25-30 ㎝; Cladodes shiny green, 2-5 m height, oblong (elongated), 20-35 ㎝ long, 8-12 ㎝ wide, areoles arranged > 2 ㎝ wide; Leaves subulate; Petals always yellow with purple median stripes on the outer perianth segment, 16-25 mm long and 11-16 mm ; Bright yellow inner perianth parts, 22-40 mm long and 16-42 mm wide, with yellow to white staminal filaments (180-200 ea), the style yellow, 10-20 mm long, divided into 5-6 bright yellow stigma lobes and 8-10 mm long; Flowering May-January; Reddish flower buds; Spines, 4-5 ㎝ long, white-grey or yellowish to reddish-brown; Brownish glochids numerous; Seeds, irregularly elliptic, usually 6×5 mm 2 ea and 4×3 mm More than 40 ea, High germination rate (> 90%); Fleshy fruit, purple- green, yellow-green pulp color, plum-shaped, 5-7.5 ㎝ long and 4-5 ㎝ in diameter; Fruiting June to next year April.

Diagnostic characters: Red stripes on the outside of petals, yellow stigma lobes, purple-green fruit, plum-shaped fruit, green-yellow pulp, peculiar phenomenon (securely generated fruit, flower growing from fruit, stem growing from fruit).

Holotype: Taepyeong-ro 200, Seogwipo, Jeju Province, Korea. (33°14'40.6" N 126°32'38.0" E) Oct. 17, 2020. (YANG-202201) Herbarium of Jeju Baiknyuncho Museum. Donation to Herbarium of Jeju Baiknyuncho Museum (JBM).

Isotype: Taepyeong-ro 200, Seogwipo, Jeju Province, Korea. (33°14'40.6" N 126°32'38.0" E) Oct. 17, 2020. (YANG-202202, YANG-202203, YANG-202204) Herbarium of Jeju Baiknyuncho Museum (JBM).

Distribution: Jeju Island, Korea

Etymology: The specific epithet was derived from Jeju Island, where this new taxon is located.

Habitats:O. monacantha f. jejuensis grows on the coast of Seogwipo-si, Jeju-si, Jeju Province, Korea

Identification Key in O. monacantha complex

1a Yellow-orange or orange flowers Elatae series

1b Yellow or Yellow-orange flowers 2(O. monacantha complex)

2a Greenish stigma lobes O. monacantha subsp. arechavaletae

2b Creamish stigma lobes 3(O. monacantha subsp. monacantha)

3a Red stripes inside and outside tepals O. monacantha f. monacantha

3b Red stripes on the outside of tepals O. monacantha f. jejuensis


We thank the Director of the Jeju Baiknyuncho Museum, Jekuk Kim, who first discovered Jeju Baiknyuncho and tried to protect the species.

Conflicts of Interest

The authors declare that they have no conflict of interest.


Anderson, E.F. 2001. The Cactus Family, Timber Press, Portland, OR (USA). pp. 484-525.
Britton, N.L. and J.N. Rose 1920. The Cactaceae. Carnegie Institute of Washington, Washington DC (USA).
Choi, S.H. 2014. Antioxidant effects and melanin synthesis inhibition from Opuntia humifusa extract. Department of Biochemistry, MS. Thesis, Hoseo University, Korea (in Korean).
Chung, H.J. 2000. Antioxidative and antimicrobial activities of Opuntia ficus-indica var. saboten. J. Korean Soc. Food Sci. Nutr. 16:160-166 (in Korean).
Clegg, M.T. 1993. Chloroplast gene sequences and the study of plant evolution. Proc. Natl. Acad. Sci. 90(2):363-367. 10.1073/pnas.90.2.3638421667PMC45662
Doyle, J.J. and J.L. Doyle. 1990. Isolation of plant DNA from fresh tissue. Focus (Madison) 12:13-15.
Engelmann, G. 1856. Synopsis of the Cactaceae of the Territory of the United States and Adjacent Regions. American Academy of Arts and Sciences, Cambridge Mass, Metcalf. vol. III, pp. 259-346. 10.5962/bhl.title.46890
Griffith, M.P. and J.M. Porter. 2009. Phylogeny of Opuntioideae (Cactaceae). Int. J. Plant Sci. 170:107-116. 10.1086/593048
Guiggi, A. 2017. New Combination in the South American Opuntia Ser. Armatae K. Schumann (Cactaceae Opuntioideae). International Cactaceae Research Center, Geneva, Switzerland, Supplementum V to cactology 5:1.
Guindon, S., J.F. Dufayard, V. Lefort, M. Anisimova, W. Hordijk, O. Gascuel. 2010. New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Syst. Biol. 59(3):307-321. 10.1093/sysbio/syq01020525638
Hall, T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/ NT. Nucl. Acids. Symp. Ser. 41:95-98.
Hernandez-Hernandez, T., H. Hernandez, J.A. De-Nova, R. Puente, L.E. Eguiarte, and S. Magallon. 2011. Phylogenetic relationships and revolution of growth from in Cataceae (Caryophyllales, Eudicotyledoneae). Am. J. Bot. 98:44-61. 10.3732/ajb.100012921613084
Jung, B.M. and K.A. Han. 2016. Quality characteristics of Gat Kimchi added with Cheonnyuncho water extract during cold storage. J. Korean Soc. Food Sci. Nutr. 45(12):1808-1815 (in Korean). 10.3746/jkfn.2016.45.12.1808
Jung, B.M., M.O. Shin and H.R. Kim. 2012. The effects of antimicrobial, antioxidant, and anticancer properties of Opuntia humifusa stems. J. Korean Soc. Food Sci. Nutr. 41(1):20-25 (in Korean). 10.3746/jkfn.2012.41.1.020
Jung, E., S.H. Yeon and H.J. Shu. 2014. Hypocholesterol effect of Opuntia humifusa extract on high cholesterol diet-induced hypercholesterolemic Rats. J. Korean Soc. Food Sci. Nutr. 43(4):485-490 (in Korean). 10.3746/jkfn.2014.43.4.485
The Royal Botanic Gardens Kew. Plants of the world online (accessed on 2. June. 2022.)
Kim, E.J., K. Srikanth, E.A. Lee and S.S. Whang. 2014. Opuntia humifusa (Raf.) Raf. f. jeollaensis E. J. Kim & S. S. Whang, a new forma based on three DNA markers. Korean J. Pl. Taxon. 44(3):181-187. 10.11110/kjpt.2014.44.3.181
Kim, M.H., H.J. Kim, M. Jang, T.G. Lim, H.D. Hong, Y.K. Rhee, K.T. Kim and C.W. Cho. 2016. The morphological and chemical composition characteristics of Opuntia ficus- indica and Opuntia humifusa fruits. Korean Soc. Food Preserv. 23(5):711-717 (in Korean). 10.11002/kjfp.2016.23.5.711
Koh, S.C., J.K. Kim and M.H. Kim. 2018. An unrecorded species of Opuntia (Cactaceae) in Korea (Wang-seon-in- jang). Korean J. Plant Res. 31(1):32-36 (in Korean).
Kwon, J.H., T.Y. Kim, J.K. Kim and J.Y. Kim. 2017. Characteristics of Seogwipo Opuntia for the functional raw material production. J. Ind. Eng. Chem. 28(2): 252-256 (in Korean).
Lee, J.A. 2017. Antioxidative capacity and quality characteristics of Yanggaeng added with Beaknyuncho (Opuntia ficus- indica var. saboten) Powder. Culi. Sci. & Hos. Res. 23(4): 33-42 (in Korean). 10.20878/cshr.2017.23.4.004004004
Lee, J.Y. and H.C. Bae. 2009. Preparation of fermented milk added with powder of Opuntia ficus-indica var. saboten and its sensory characteristics. J. East Asian Soc. Diet. Life 19 (6):967-974 (in Korean).
Lee, S.H., H.W. Kim, M.K. Lee, G. Asamenew, Y.J. Kim, S.j. Lee, Y.S. Cha, S.M. Yoo and J.B. Kim. 2018. Characterization of flavonoids from used parts of prickly pear (Opuntia ficus- indica var. saboten) and dragon fruit (Hylocereus undatus) using UPLC-DAD-QToF/MS. J. Korean Soc. Food Sci. Nutr. 47(12):1274-1283 (in Korean). 10.3746/jkfn.2018.47.12.1274
Leuenberger, B.E. 1993. Interpretation and typification of cactus opuntia L., Opuntia vulgaris Mill., and O. humifusa (Rafin.) Rafin. (Cactaceae). Taxon. 42(2):419-429. 10.2307/1223152
Lim, M.H. and S.J. Hong. 2016. Availability of Opuntia ficus- indica var. saboten stem extracts as a natural preservative. Asian J. Beauty Cosmetol. 14(4):449-461 (in Korean). 10.20402/ajbc.2016.0079
Majure, L.C. and R. Puente. 2014. Phylogenetic relationships and morphological evolution in Opuntia s. str. and closely related members of tribe Opuntieae. Succ. Plant Res. 8:9-30.
Majure, L.C., D.E. Soltis, P.S. Soltis and W.S. Judd. 2012a. Cytogeography of the Humifusa clade of Opuntia s.s. Mill. 1754 (Cactaceae, Opuntioideae, Opuntieae): correlations with pleistocene refugia and morphological traits in a polyploid complex. Comp. Cytogenet. 6:53-77. 10.3897/compcytogen.v6i1.252324260652PMC3833768
Majure, L.C., R.P. Majure, M.P. Griffith, S. Walter, S. Pamela, J. Soltis and D.E. Soltis. 2012b. Phylogeny of Opuntia s.s. (CACTACEAE): Clade delineation, geographic origins, and reticulate evolution. Am. J. Bot. 99(5):847-864. 10.3732/ajb.110037522539520
Maria, L.P., O. Luis, C.M. Natalia and B. Gabriel. 2017. Taxonomic and cytogenetic studies in Opuntia ser. Armatae (Cactaceae). Botany 95:1-60. 10.1139/cjb-2016-0048
Navie, S. and S. Adkins. 2008. Environmental weeds of Australia: an interactive identification and information resource for over 1000 invasive plants. CRC for Australian Weed Management Glen Osmond Australia. The University of Queensland, Brisbane, Queensland, Australia. unpaginated.
Nyffeler, R. and U. Eggli. 2010. A farewell to dated ideas and concepts; Molecular phylogenetics and a revised suprageneric classification of the family Cactaceae. Schumannia 6:109- 149.
Omweri, A.H., F.K. Rimberia, S.G. Mwangi and D.N. Sila. 2016. Morphological characterization and distribution of cactus species (Cactaceae) in arid and semi-arid lands of Kenya. Int. J. Agr. Agri. R. 9(1):182-191.
Pardo, F.M.V. and D.G. Alonso. 2017. Aproximación al conocimiento del grupo Opuntia Mill. (s.l.) (CACTACEAE) en Extremadura Folia Bostanica Extremadurensis 11:51-75 (in spanish).
Park, C.M., B.H. Kwak, S.H. Park, H. Kim and D.Y. Rhyu. 2013. Comparison of biologic activities of Opuntia humifusa and Opuntia ficus-indica. Korean J. Plant Res. 26(5):519-525 (in Korean). 10.7732/kjpr.2013.26.5.519
Park, K.R., J.H. Lee, S.T. Kim, S.T. Lee, M.Y. Kim, S.P. Hong and Y.J. Jung. 2010. Plant Systematics (A Phylogenetic Approach), Third Edition. Shinilbooks, Seoul, pp. 112-115 (in Korean).
Rambaut, A. 2016. Tree Figure Drawing Tool Version 1. 4. 3. Institute of Evolutionary Biology, University of Edinburgh, UK.
Ronquist, F., M. Teslenko, P.V.D. Mark, D.L. Ayres, A. Darling, S. Höhna, B. Larget, L. Liu, M.A. Suchard and J.P. Huelsenbeck. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model Space. Syst. Biol. 61(3):539-542. 10.1093/sysbio/sys02922357727PMC3329765
Shaw, J., E.B. Lickey, E.E. Schilling and R.L. Small. 2007. Comparison of whole chloroplast genome sequences to choose noncoding regions for phylogenetic studies in angiosperms: The tortoise and the hare III. Am. J. Bot. 94:275-288. 10.3732/ajb.94.3.27521636401
Soltis, D.E., P.S. Solitis and J.J. dolye. 1998. Molecular systematics of plants II; DNA sequencing. Kluwer Academic Publishers, Boston, MA (USA).
Speg. 1905. Opuntia arechavaletae Speg. Anal. Mus. Buenos Aires. 11:520.
Taberlet, P.G., P. Ludovic, P. Guy and J. Bouvet. 1991. Universal primers for amplication of three non-coding regions of chloroplast DNA. Plant Mol. Biol. 17:1105-1109. 10.1007/BF000371521932684
Taylor, N. and D. Zappi. 2004. Cacti of Eastern Brazil. Cact. Succ. J. Gr. Brit. 77(1):43. 10.2985/0007-9367(2005)77[43:COEB]2.0.CO;2
Thompson, J.D., T.J. Gibson, F. Jeanmougin and D.G. Higgins. 1997. The CLUSTAL-X windows interface: Flexible strategies for multiple sequence alignment adied by quality analysis tools. Nucleic Acids Res. 25:4876-4882. 10.1093/nar/25.24.48769396791PMC147148
Wagner, W.L. Herbst, D.R. Sohmer, S.H. 1999. Manual of the flowering plants of Hawaii. Revised edition. Bernice P. Bishop Museum special publication. University of Hawai'i Press/Bishop Museum Press, Honolulu, HI (USA). p. 420.
Wallace, R.S. and S.L. Dickie. 2002. Systematic implications of chloroplast DNA sequence variation in subfam. Opuntioideae (Cactaceae). Succ. Pl. Res. 6:9-24.
White, T.J., T. Bruns, S. Lee And J.W. Taylor. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In Innis, M.A., D.H. Gelfand, J.J. Sninsky, and T.J. White (eds.), PCR Protocols: A Guide to Methods and Applications. Academic Press, San Diego, CA (USA). pp. 315-322. 10.1016/B978-0-12-372180-8.50042-11696192
Yang, Y.S. and H.S. Oh. 2021. Phylogenetic relationship between two taxa of Opuntia in Jeju Island. Korean J. Plant Res. 34(5):451-461 (in Korean).
Yang, Y.S., B.K. Choi and H.S. Oh. 2020. A taxonomic review and nomenclature of the Opuntia ficus-indica (L). Mill. in Jeju Island. Journal of the Korean Institute of Traditional Landscape Architecture 38(4):68-73 (in Korean).
페이지 상단으로 이동하기