Research Article |
Corresponding author: Hyeok Jae Choi ( skinh@hanmail.net ) Academic editor: Eberhard Fischer
© 2022 Hyun-Do Jang, Kwi-Kwan Jeong, Myoung-Ja Nam, Jun-Ho Song, Hye-Kyoung Moon, Hyeok Jae Choi.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Jang H-D, Jeong K-K, Nam M-J, Song J-H, Moon H-K, Choi HJ (2022) Mosla dadoensis (Lamiaceae), a new species from the southern islands of South Korea. PhytoKeys 208: 185-199. https://doi.org/10.3897/phytokeys.208.89552
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Mosla dadoensis (Lamiaceae), a new species from the southern islands of South Korea, is described and illustrated. The new species is morphologically similar to M. chinensis, but is distinguished from the latter by having two types of hairs on its stems, wider leaf blades, longer corolla length, and ellipsoid nutlets with a narrowly U-shaped extended area of abscission scar. Mosla dadoensis is also distinguished from the Chinese narrow endemic M. hangchouensis by having an included pistil to the corolla, smaller ellipsoid nutlets, and later flowering and fruiting season. Phylogenetic analyses, based on two nuclear ribosomal (ETS, ITS) and three chloroplast (rbcL, matK, trnL-F) DNA regions, confirmed that the new species was constructed as monophyletic, and that M. dadoensis and M. hangchouensis form a sister group with robust support. We hereby provide a detailed morphological description of M. dadoensis with its corresponding geographical distributions, and comparison tables of related taxa.
Elsholtzieae, Korean endemic plant, morphology, phylogeny, taxonomy
Mosla (Benth.) Buch.-Ham. ex Maxim. is a genus within the sixth largest family, Lamiaceae (the mint family). Although Mosla is a small genus of approximately 20 species, it is the second largest genus in the tribe Elsholtzieae (
In Korea, four species of Mosla are recognized, namely M. chinensis Maxim., M. dianthera, M. japonica (Benth. ex Oliv.) Maxim., and M. scabra (Thunb.) C.W.Wu & H.W.Li (
During general floristic study in the southern part of Korea during October 2021, we found an unusual species which is restricted to the southern islands. This species is readily distinguished from previously known Mosla species in Korea by a considerably longer corolla. M. chinensis could be the closest ally, but the leaf shapes and flower features are significantly different. After a thorough literature survey and investigation of the relevant specimens, we designate M. dadoensis K.K.Jeong, M.J.Nam & H.J.Choi as a new species of Mosla from the southern islands of Korea. To clarify the systematic status of M. dadoensis we also conducted barcoding analysis based on nuclear ribosomal (nr) and chloroplast (cp) DNA regions, and observed detailed nutlet morphology, which is well known as a systematically important characteristic in Lamiaceae (
Morphological descriptions were based on specimens from the KB, KH (abbreviations are according to the Index Herbariorum [http://sweetgum.nybg.org/science/ih/]), and the herbarium of Changwon National University. Field surveys were also conducted from October 2021 to February 2022. Materials preserved in 70% ethanol were used for observation and measurement of floral parts. For quantitative characters, measurements were based on at least 50 samples.
For morphological observations and size measurements, the nutlets were first examined using a stereomicroscope (SM; Olympus SZX16, Olympus, Tokyo, Japan). Nutlet sizes were measured using at least 30 randomly chosen individuals from each species. Prior to scanning electron microscopic observations, all the dried nutlets were rehydrated overnight using the wetting agent Agepon (Agfa-Gevaert, Leverkusen, Germany) and distilled water (1:200) at 37–40 °C. The rehydrated materials were dehydrated through an ethanol series (50%, 70%, 90%, 95%, and 100%) at room temperature for 1 h each. The completely dehydrated materials were immersed in liquid carbon dioxide (CO2) for critical point-drying (CPD; SPI-13200J-AB, SPI Supplies, West Chester, PA, USA). For the micromorphological observations, selected nutlets were mounted on aluminum stubs using a double-sided adhesive conductive carbon disk (05073-BA, SPI Supplies, West Chester, PA, USA). Specimens were coated with gold using an ion-sputtering device (208HR, Cressington Scientific Instruments Ltd., Watford, UK), and then observed using a low-voltage field emission scanning electron microscope (FE-SEM; JSM-7600F, JEOL, Tokyo, Japan) at an accelerating voltage of 10 kV and a working distance of 8–10 mm (
To confirm the systematic placement of the putative new species within the genus Mosla, molecular phylogenetic analyses were conducted. The combined cpDNA dataset (rbcL, matK, and trnL-trnF) and nrDNA dataset (ITS, ETS) used in
Fragment | Primer | Sequence 5' → 3’ | Reference |
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ITS | ITS1 | TCCGTAGGTGAACCTGCGG |
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ITS4 | TCCTCCGCTTATTGATATGC | ||
ETS | ETS-B | ATAGAGCGCGTGAGTGGT |
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18S-IGS | GAGACAAGCATATGACTACTG |
|
|
rbcL | rbcL_1F | ATGTCACCACAAACAGAAAC |
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rbcL_724R | TCGCATGTACCTGCAGTAGC | ||
matK | 3F_Kim_F | CGTACAGTACTTTTGTGTTTA | K.J.Kim, pers. comm. |
1R_Kim_R | ACCCAGTCCATCTGGAAATCT | ||
trnL-F | B49317 | CGAAATCGGTAGACGCTACG |
|
A50272 | ATTTGAACTGGTGACACGAG |
Total genomic DNA of M. dadoensis was extracted from silica gel-dried leaf materials using a DNeasy Plant Mini Kit (Qiagen Ltd., Crawley, West Sussex, UK). We conducted PCR with a ProFlex 96-Well PCR System (Applied Biosystems, Foster City, CA, USA). Each reaction mixture contained AccuPower PCR PreMix (Bioneer, Daejeon, South Korea), ca. 10 ng (1 μL) of genomic DNA, and 100 pM of primers in a total volume of 20 µL. Conditions included an initial denaturation at 95 °C for 5 min, followed by 40 amplification cycles comprising 95 °C for 30 sec, 50 °C for 30 sec, and 72 °C for 1 min, with a final extension at 72 °C for 5 min. After the PCR products were visualized on 2% agarose gels, they were treated with a MG PCR Purification kit (MGmed), and sequenced with the ABI 3730xl Analyzer, using the ABI BigDye Terminator v3.1 Cycle Sequencing Kits (Applied Biosystems, Foster City, CA, USA).
List of voucher information and GenBank accessions of species used in this study.
Species | Voucher | ETS | ITS | matK | rbcL | trnL-F |
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Mosla dadoensis 1 | H.J.Choi_210923_001_1 | ON619797 | ON033689 | ON619803 | ON619806 | ON619800 |
Mosla dadoensis 2 | H.J.Choi_210923_001_2 | ON619798 | ON033690 | ON619804 | ON619807 | ON619801 |
Mosla dadoensis 3 | H.J.Choi_210923_001_3 | ON619799 | ON033691 | ON619805 | ON619808 | ON619802 |
Mosla cavaleriei | PNLI20120445 | KY552608 | KY552540 | KY624903 | KY624972 | KY625040 |
Mosla chinensis | PNLI20120245 | KY552609 | KY552541 | KY624904 | KY624973 | KY625041 |
Mosla dianthera | PNLI20120248 | KY552610 | KY552542 | KY624905 | KY624974 | KY625042 |
Mosla hangchouensis | PNLI20120424-1 | KY552611 | KY552543 | KY624906 | KY624975 | KY625043 |
Mosla japonica | PNLI20120416 | KY552612 | KY552544 | KY624907 | KY624976 | KY625044 |
Mosla scabra | PNLI20120427 | KY552613 | KY552545 | KY624908 | KY624977 | KY625045 |
Mosla soochouensis | PNLI20120414 | KY552614 | KY552546 | KY624909 | KY624978 | KY625046 |
Mosla tamdaoensis | C-K-393 | KY552615 | KY552547 | KY624910 | KY624979 | KY625047 |
Keiskea japonica | PNLI20120049-1 | KY552605 | KY552537 | KY624901 | KY624969 | KY625037 |
Phylogenetic analyses were conducted using maximum likelihood (ML). The obtained sequences were aligned using MAFFT with Geneious Prime 2019.2.3 (Biomatters Ltd., Auckland, NZ). To assess the confidence of the phylogenetic relationships, a bootstrap test was conducted with 1,000 replications for the ML analysis. Kimura’s three-parameter model (
This new species is morphologically similar to M. chinensis, but is easily distinguished from the latter by having two types of hairs on its stems, wider leaf blades, longer corolla length, and ellipsoid nutlets with a narrowly U-shaped extended area of abscission scar.
Herbs annual, aromatic. Stems 10–60 cm tall, many branched from base, densely pubescent with white recurved hairs and densely to moderately intermixed with white villous, with impressed glands. Leaves petiolate; petiole 2–5 mm long, pubescent with white villous; blades narrowly lanceolate to lance-ovate, 1–3 cm × 4–10 mm, sparsely pubescent, dotted with impressed glands, adaxially olive green, abaxially gray, base cuneate, margin remotely serrate, apex acute. Racemes terminal, 1–2.5 cm, bracts overlapping, circular-obovate, 5–7 × 4–5 mm, margin ciliate, apex caudate. Pedicel pubescent. Calyx campanulate, ca. 5 × 3 mm, dilated after anthesis, subequally 5-toothed; teeth subulate, ca. 2/3 to 3/4 as long as calyx tube. Corolla slightly 2-labiate, pale purple, ca. 1.5 times longer than bracts, 8–9 mm long, pubescent outside, pubescent with long white villous on lower lip inside; upper lip straight, emarginate; lower lip 3-lobed, middle lobe largest, slightly recurved. Stamens 4, included (non-exserted); filaments shorter than anthers; anthers linear, cells divergent, ca. 2 mm long, connectives distinct. Pistil included; sigma bifid. Nutlets brown to blackish-brown, ellipsoid, 1.2–1.6 × 0.9–1.3 mm, glabrous or sparsely pubescent with gland, pitted with deep depressions, abscission scar basal position, elliptic, extended, extended area narrowly U-shaped at the ventral side, ratio of abscission scar / nutlet diameter 0.51–0.53, primary sculpture outline of cells isodiametric, tetragonal to hexagonal, anticlinal walls straight, raised, thin, periclinal walls concave, secondary sculpture micropapillate.
Flowering and fruiting from August to November.
The specific epithet, “dadoensis”, is based on the name of location, the Dadohae southern coastal region of Korea, where Mosla dadoensis was discovered.
The Korean name of the new species is “Da-do-hae-san-deul-kkae (다도해산들깨)”.
Mosla dadoensis is morphologically similar to M. chinensis, from which it is clearly differentiated by the hairs on its stems [white recurved hairs and intermixed with white villous (Fig.
Comparison of major characters of Mosla dadoensis, M. chinensis, and M. hangchouensis (*: data from
Character | M. dadoensis | M. chinensis | M. hangchouensis* | |
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Habitat | open rocky area along the coast | grassy slope, forest edge, wet land | sunny side of hill peak, forest edge, and under forest along the coast | |
Plant | height (cm) | 10–60 | 10–40 | 20–120 |
Stem | trichome | densely pubescent with white recurved hairs and moderately intermixed with white villous | densely pubescent with white recurved hairs | pubescent, brown glandular sometime intermixed with spreading pilose hairs |
Leaf blade | shape | lanceolate to lance-ovate | linear to linear-lanceolate | lanceolate |
size | 1–3 cm × 4–10 mm | 1–5 cm × 1.3–4 mm | 1.5–4.2 cm × 5–13 mm | |
Corolla | length (mm) | 8–9 | 5–6 | ca. 10 |
length ratio of corolla/calyx | ca. 2.0 | ca. 1.5 | ca. 3.0 | |
Pistil | relative length to corolla | included | included | clearly exserted |
Nutlet | shape | ellipsoid | globose to subglobose | globose to subglobose |
diameter (mm) | 0.9–1.3 | 1.0–1.2 | ca. 2.1 | |
extended area of abscission scar | narrowly U-shaped at the ventral side | widely U-shaped at the ventral side | widely U-shaped at the ventral side | |
ratio of abscission scar/nutlet diameter | 0.51–0.53 | 0.61–0.69 | NA | |
outline of surface cell | tetragonal to hexagonal | rounded | tetragonal to hexagonal | |
anticlinal walls of surface | straight, thin | curved, thick | straight, thin | |
Flowering and fruiting | August to November | June to October | June to September |
The combined dataset has 12 aligned sequences comprising 2,910 bp (609 bp for ITS, 371 bp for ETS, 439 bp for rbcL, 736 bp for matK, and 755 bp for trnL-F), of which 102 occupied variable positions (3.51%). Our phylogenetic tree (Fig.
Phylogenetic tree of Mosla dadoensis and related taxa based on concatenated alignments of two nrDNA (ITS, ETS) and three cpDNA regions (rbcL, matK, trnL-F). The numbers above branches are bootstrap values (BS > 50%) used in the maximum likelihood method. Distribution information was obtained from Plants of the World Online (https://powo.science.kew.org).
Mosla dadoensis (Paratypes): Korea: Jeonnam: Yeosu-si, Geumo-do Isl., 34°30'11.1"N, 127°44'34.2"E, elev. 110 m, 25 Oct 2021 (fr), H.J.Choi 211025-001 (CWNU); Goheung-gun (Naro-do Isl.), Bongrae-myeon, Jangpo-san, 34°25'25.46"N, 127°30'26.90"E, elev. 307 m, 26 Feb 2022, K.K.Jeong s.n. (CWNU); Goheung-gun (Naro-do Isl.), Bongrae-myeon, Bongrae-san, 34°25'45.51"N, 127°30'56.77"E, elev. 150 m, 26 Feb 2022, K.K.Jeong s.n. (CWNU); Jindo-gun, Yeogui-san, 34°23'41.91"N, 126°14'21.00"E, elev. 296 m, 27 Feb 2022, K.K.Jeong s.n. (CWNU); Jindo-gun, Imhoe-myeon, Namdong-ri, Hanbok-san, 34°22'18.1"N, 126°9'42.7"E, elev. 96 m, 9 Oct 2013 (fl, fr), JJP7102 (KB); Goheung-gun, Dohwa-myeon, 34°29'14.01"N, 127°19'26.06"E, elev. 8 m, 3 Mar 2022, K.K.Jeong s.n. (CWNU); Goheung-gun, Dohwa-myeon, 34°26'42.11"N, 127°20'06.30"E, elev. 39 m, 3 Mar 2022, K.K.Jeong s.n. (CWNU); Wando-gun, Bogil-myeon, Jeokja-bong, 3 Oct 2003 (fr), B.Y.Sun et al. s.n. (KB); Wando-gun, Bogil-myeon, Yesong-ri, Geokja-bong, 34°32'38.88"N, 126°55'39.32"E, elev. 303 m, 24 Oct 2013 (fr), kjs 130042 (KB); Wando-gun, Wando-eup, Daeya-ri, 34°22'12.22"N, 126°40'56.03"E, elev. 505 m, 11 Aug 2014, Y.H.Cho & H.J.Na 140811107 (KB); Wando-gun, Bogil-myeon, Buhwang-ri, 34.128717N, 126.535047E, elev. 50 m, 7 Oct 2017, WR-171007-075 (KH); Wando-gun, Bogil-myeon, Yesong-ri, Gyeokjabong, 34°08'15.50"N, 126°33'32.90"E, elev. 148 m, 7 Nov 2009 (fr), HNHM-2010-0355 (KH); Wando-gun, Cheongsan-myeon, Cheonggye-ri, 34.159905N, 126.897922E, elev. 80 m, 8 Sep 2017 (fl, fr), WR-170908-003 (KH); Wando-gun, Saengil-myeon, Bongseon-ri, 7 Sep 2003 (fl), Lee.Y.H. 030062 (KH). Gyeongnam: Namhae-gun (Namhae-do Isl.), Nam-myeon, Eungbong-san, 34°43'40.24"N, 127°53'15.65"E, elev. 268 m, 1 Mar 2022, K.K.Jeong s.n. (CWNU).
Mosla chinensis: Korea: Gyeonggi: Anyang-si, Dongan-gu, Bisan-dong, 37°25'23."N, 126°57'34.4"E, elev. 235 m, (fr), PWK-133 (KH); Suwon-si, Gwonseon-gu, Homaesil-dong, Chilbo-san, 37°15'40.39"N, 126°55'47.4"E, elev. 84 m, 24 Sep 2009 (fr), NIBRVP0000209769 (KB); Incheon-si, Ganghwa-gun, Gilsang-myeon, Donggeom-ri, Donggeom-do Isl., 37°35'25.2"N, 126°31'2.7"E, elev. 63 m, 8 Sep 2012, NIBRVP0000400499 (KB); Paju-si, Tanhyeon-myeon, Bupheung-ri, 37°46'02.4"N, 126°41'19.4"E, elev. 100 m, 30 Aug 2006, VP-NAPI-376034-053 (KB). Chungbuk: Jeungpyeong-gun, Jeungpyeong-eup, Jwagu-san, 36°42'41.8"N, 127°39'39.2"E, elev. 500 m, 25 Aug 2011 (fl), Geumbuk-203 (KH). Chungnam: Seosan-si, Daesan-eup, Ungdo-ri, 36°55'04.4"N, 126°22'24.8"E, elev. 0 m, 15 Aug 2012, DJUIDC20120154 (KH). Jeonbuk: Gimje-si, Dojang-dong, Hwang-san, 35°46'35"N, 126°56'30.1"E, elev. 12 m, 27 Aug 2011, 357014-0420 (KB). Jeonnam: Haenam-gun, Hwangsan-myeon, Wonho-ri, Hakdong village, 34°34'15.14"N, 126°29'2.22"E, elev. 3 m, 17 Sep 2008 (fl), ParkSH81875 (KH); Jindo-gun, Jodo-myeon, Sinyuk-ri, Hajo-do Isl., Sinjeon beach, 34°17'21.5"N, 126°01'88.1"E, elev. 39 m, 6 Sep 2011, HS110899 (KH); Sinan-gun, Docho-myeon, Oryu-ri, Near Simok Sandbeach, 34.725447N, 125.908671E, elev. 30 m, 24 Oct 2007, WR-071024-170 (KH); Sinan-gun, Docho-myeon, Oryu-ri, Near Simok Sandbeach, 34.725447N, 125.908671E, elev. 30 m, 24 Oct 2007, NAM-071024-199 (KH); Yeonggwang-gun, Hongnong-eup, Gyema-ri, Gamami beach, 11 Sep 2012, P126974 (KH); Hwasun-gun, Doam-myeon, Daecho-ri, Cheonbulsan, Unjusa, 34°55'27.3"N, 126°52'14.0"E, elev. 109 m, 6 Sep 2009, SGU 0940 (KH); Gangjin-gun, Byeongyeong-myeon, Jiro-ri, Suin-san, 34°42'56.9'N, 126°50'18.5"E, elev. 174 m, 12 Aug 2014 (fl), HNHM-D-140197 (KH); Sinan-gun, Aphae-myeon, Songgong-ri, Songgong-san wetland, 34.844604N, 126.252203E, elev. 25 m, 19 Sep 2007 (fl), WR-070919-255 (KH); Jindo-gun, Gunnae-myeon, Geumseong-ri, 34°32'54.1"N, 126°17'38.6"E, elev. 30 m, 14 Sep 2005 (fr), ESJeon 52851 (KH); Naju-si, Dado-myeon, Masan-ri, Bulhoe-sa Temple, 34°55'15.7"N, 126°53'35.8"E, elev. 126 m, 14 Sep 2005, ESJeon 52829 (KH); Hampyeong-gun, Hakgyo-myeon, Gokchang-ri, 35.026751°N, 126.570272°E, elev. 100 m, 9 Sep 2012 (fl), WR-20120909-044 (KH); Sinan-gun, Amtae-myeon, Songgok-ri, Amtae-do Isl., 34°50'20.5"N, 126°8'35.9"E, elev. 13 m, 10 Oct 2019, YLJLVP0000006165 (KB); Gangjin-gun, Gundong-myeon, Pungdong-ri, Seongjak-gol, 34°30'48.45"N, 126°40'49.90"E, elev. 294 m, 23 Sep 2010, C201009-0117 (KB); Sinan-gun, Aphae-myeon, Janggam-ri, 34°49'3.9"N, 126°20'58.1"E, elev. 14 m, 4 Oct 2012 (fl), KOSPVP0000256241 (KB); Jindo-gun, Gunnae-myeon, Dunjeon-ri, Geumgol-san, 34°32'24.8"N, 126°17'39.2"E, elev. 81 m, 27 Oct 2013 (fl), KOSPVP0000291190 (KB); Sinan-gun, Jeungdo-myeon, Jeungdong-ri, Gubunpo, Gwakdae-bong to Bunpo reservoir, 28 Sep 1997 (fl), EN97CUB404 (KB). Gyeongbuk: Sangju-si, Jungdong-myeon, Hoesang-ri, Hwanggeum-san, 36°27'55.3"N, 128°16'34.2"E°, elev. 210 m, 9 Sep 2012 (fl), KTPSA-2012076 (KH); Daegu-si, Dong-gu, Jimyo-dong, 35°56'25.09"N, 128°39'49.04"E, elev. 202 m, 21 Aug 2013 (fl), DJUIDC2013-212 (KH); Yecheon-gun, Jibo-myeon, Amcheon-ri, 36°33'06.00"N, 128°27'11.09"E, elev. 11 m, 7 Sep 2011 (fl), Nakdong-1632 (KH); Gyeongbuk, Sangju-si, Jungdong-myeon, Hoesang-ri, 36°27'54.9"N, 128°16'37.0"E, elev. 236 m, 8 Sep 2012, NAPI2012-0153 (KH); Cheongsong-gun, Hyeonseo-myeon, Hwamok-ri, 36°16'24.4"N, 128°52'22.1"E, elev. 387 m, 3 Aug 2018, NIBRVP0000703391 (KB); Gunwi-gun, Bugye-myeon, Changpyeong-ri, San 100, 36°4'56.36"N, 128°41'34.59"E, elev. 225 m, 27 Sep 2019 (fr), NIBRVP0000756907 (KB); Andong-si, Iljik-myeon, Wonho-ri, Jaam-san, 36°29'54.08"N, 128°40'37.14"E, elev. 302 m, 21 Aug 2017 (fl), NIBRVP0000632258 (KB); Gunwi-gun, Bugye-myeon, Changpyeong-ri, 36°4'59.04"N, 128°41'35.97"E, elev. 220m, 29 Aug 2019, NIBRVP0000754852 (KB); Uiseong-gun, Bian-myeon, Jarak-ri, Haemang-san, 36°22'52.31"N, 128°31'3.79"E, elev. 202 m, 3 Oct 2017 (fr), NIBRVP0000643724 (KB); Gimcheon-si, Nam-myeon, Busang-ri, Geumo-san, San 168-7, 36°3'58.8"N, 128°16'34.1"E, elev. 220 m, 20 Sep 2015 (fl), NIBRVP0000585241 (KB); Gumi-si, Namtong-dong, Geumo-san, Peak to Beopseong temple, 36°5'52.5"N, 128°19'46"E, elev. 250 m, 5 Oct 2015 (fr), NIBRVP0000586707 (KB). Gyeongnam: Milyang-si, Muan-myeon, Garye-ri, Yeongchwi-san, 35°29'57.40"N, 128°35'02.20"E, elev. 201 m, 14 Sep 2009 (fr), HNHM-2009-0392 (KH).
This research was funded by the National Institute of Biological Resources, Ministry of Environment of South Korea (Grant Number NIBR202208101).