Corresponding author: Watanabe Yoichi (
Academic editor: P. de Lange
Three new taxa,
Yoichi W, Minamitani T, Oh S-H, Nagano AJ, Abe H, Yukawa T (2019) New taxa of
The genus
The morphological characters were observed and measured based on living materials in the field and herbarium specimens listed in the sections “Additional specimens examined”.
Samples for DNA analyses were collected from three individuals for
Locations of samples used for phylogenetic analyses.
Species | Code | Locality | Latitude / Longitude | Haplotype | Voucher |
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Syk | Mt. Syakagadake, Nara, Japan |
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H1 | Y. Watanabe & K. Yukitoshi s.n. | |
Miu | Mt. Miune, Tokushima, Japan |
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H1 | Y. Watanabe & T. Fukuda s.n. | |
Ttj | Mt. Tsutsujyo, Ehime, Japan |
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H1 | Y. Watanabe & M. Takahashi Ttj02 | |
Ici | Mt. Ichifusa, Kumamoto, Japan |
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H2 | Y. Watanabe Ici01 | |
Mks | Mt. Mukousaka, Kumamoto, Japan |
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H2 | Y. Watanabe Mks04 | |
Sob | Mt. Sobo, Miyazaki, Japan |
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H3 | Y. Watanabe & T. Oi s.n. | |
Gom | Gonam, Jeollabuk-do, South Korea |
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H4 | Y. Watanabe, S. Hwang & N. Yun Gom01 | |
Wol | Mt. Wolbong, Gyeongsangnam-do, South Korea |
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H4 | Y. Watanabe, S. Hwang & N. Yun Wol01 | |
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Snp | Mt. Sanpouiwa, Ishikawa, Japan |
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H5 | Y. Watanabe Snp02 |
Sad | Sado Island, Niigata, Japan |
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H5 | H. Abe s.n. | |
Iid | Mt. Iide, Niigata, Japan |
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H6 | Y. Wataanbe s.n. | |
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Utk | Mt. Utsukushigahara, Nagano, Japan |
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H7 | Y. Watanabe s.n. |
Abe | Abe-touge pass, Shizuoka, Japan |
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H7 | Y. Watanabe Abe01 | |
Kmg | Mt. Kamagatake, Mie, Japan |
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H7 | Y. Watanabe & T. Oi s.n. | |
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Zao | Mt. Zao, Miyagi, Japan |
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H8 | Y. Watanabe Zao01 |
Mus | Mt. Musadake, Hokkaido, Japan |
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H8 | Y. Watanabe s.n. | |
Tor | Mt. Toraidake, Aomori, Japan |
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H9 | Y. Watanabe s.n. | |
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Hkn | Mt. Hakone-komagatake, Kanagawa, Japan |
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H10 | Y. Watanabe Hkn01 |
Kdz | Kouzushima Island, Tokyo, Japan |
|
H11 | H. Abe s.n. | |
Tng | Mt. Tengu, Nagano, Japan |
|
H11 | Y. Watanabe s.n. |
Haplotype, Haplotype codes detected by chloroplast DNA sequences, corresponding to those in Fig.
In addition, genome-wide SNPs were identified from two double digest restriction-site associated DNA (
The phylogenetic relationships were inferred from two data sets, which were obtained from chloroplast DNA sequences and RAD-seq, based on the maximum likelihood method using RAxML v. 8.2.0 (
We found three new entities (Types 1, 2 and 3) that have previously been included within
The three types share the following combination of characters as commonly derived character states, which can be distinguished from the other members of the
Photographs of flowers and leaves for the new taxa described in this study.
Photographs of flowers for the other members of
Diagnostic characters among the
Character |
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Leaf length (mm) | 10–20 | 10–20 | 10–35 | 10–25 | 5–13 | 15–50 | 5–20 |
Leaf width (mm) | 4–7 | 5–10 | 5–15 | 5–12 | 2–5 | 5–20 | 5–10 |
Nerve on abaxial surface of leaf | pinnate 2–3 pairs, obscurely raised | pinnate 2–3 pairs, raised | pinnate 1–3 pairs, raised | nervules reticulate | nervules obscurely reticulate | pinnate 1–2 pairs, prominent, nervules reticulate | nervules obscurely reticulate |
Hair of leaf | densely strigose on adaxial surface, glabrous or sparsely strigose on abaxial surface | densely strigose on adaxial surface, sparsely strigose on abaxial surface | densely strigose on adaxial surface, sparsely strigose on abaxial surface | strigose on both surfaces | densely strigose on adaxial surface, glabrous on abaxial surface | strigose on both surfaces | densely strigose on adaxial surface, glabrous or sparsely strigose on abaxial surface |
Corolla form | tubular-funnelform | tubular-funnelform | tubular-funnelform | tubular-funnelform | tubiform | tubular-funnelform | tubiform |
Number of corolla lobes | 4 | 4 | 4 | 5 | 5 | 4 | 4 |
Corolla tube length (mm) | 2–3 | 3–4 | 2–3 | 2–4 | 5–7 | 2–4 | 3.5–4.5 |
Corolla lobe length (mm) | 3–5 | 2–5 | 3–5 | 4–6 | ca. 2 | 4–6 | 2–3 |
Style length (mm) | 4–10 | 3–4 | 5–6 | 6–13 | 4–5 | 3–7 | 2.5–3.5 |
Style condition | exserted | included | exserted | exserted | included | exserted | included |
Anther | opening by apical pores | opening by apical pores | opening by apical pores | opening by apical pores | opening by longitudinal slits | opening by apical pores | opening by apical pores |
Distribution | Japan: Honshu (Kii Peninsula) and Shikoku | Japan: Kyushu | South Korea: Gyeongsang and Jeolla provinces | Japan: Honshu (Kanto and Tohoku districts) and Hokkaido. Russia: Kunashir Island | Japan: Honshu (Chubu and Kanto districts) | Japan: Honshu (Kinki, Chubu, Kanto and Tohoku districts) | Japan: Honshu (Kinki, Chubu and Kanto districts) |
Phylogenetic relationships of the
Thus the three types can be distinguished from each other and also from the other members of the
Comparative phylogenies for
Neighbor-net for members of the
This species is similar to
Much branched semi-evergreen shrubs 1–1.5 m tall. Branchlets and petioles with dense appressed flattened brownish strigose hairs. Spring leaves scattered or crowded on upper branchlets; petioles 0.5–1 mm long; blade thick chartaceous, oblong, 10-20 mm long (at maximum within each individual), 4–7 mm wide, apex acute and terminating in a gland, base acute, adaxial surface green, abaxial surface pale green, densely strigose on adaxial surface, glabrous or sparsely strigose on abaxial surface without midrib; midrib prominent abaxially; lateral nerves pinnate, 2–3 paired, obscure raised abaxially. Summer leaves oblanceolate, 5–10 mm long, 1–6 mm wide, densely strigose on both surfaces. Flower buds terminal, single, broadly ovoid, acute, ca. 2 mm long, 2 mm wide; scales widely ovate, densely strigose on upper outer surface. Inflorescences umbel-like, 2–4 flowers. Pedicel 2–4 mm long at flowering, densely appressed hirsute. Calyx saucer-shaped, ca. 1.5 mm in diam., densely strigose, shallowly 4-lobed; lobes semiorbiculate, ca. 0.5 mm long. Corolla white, no blotches, openly tubular-funnelform, 8–12 mm long and wide, dissected 1/2 corolla length into 4 lobes; tube 2–3 mm long, ca. 2 mm wide, glabrous outside, pilose on upper inside; lobes elliptic to oblong, rounded, 3–5 mm long, 2–4 mm wide. Stamens 4, subequal, 5–8 mm long, exserted; filaments densely pilose on lower three-quarters; anthers yellow, oblong, ca. 1 mm long. Ovary ovoid, densely soft strigose, ca. 1.5 mm. Style 4–10 mm long, exserted, glabrous. Capsule ovoid, 2–5 mm long, 2–3 mm wide, densely strigose.
JAPAN: Honshu (Kii Peninsula), Shikoku.
The plants inhabit sunny places and grow on mountain ridges and slopes at altitudes over 1000 m above sea level. In such places, there are few trees and established communities of shrubs and dwarf bamboos (
The specific epithet refers to ‘Sohayaki’ a floristic region in Japan that covers Kii Peninsula of Honshu, Shikoku and Kyushu (
Distribution of the
This variety is similar to
Spring leaves scattered or crowded on upper branchlets; petioles 0.5–1 mm long; blade thick chartaceous, oblong-ovate, 10–20 mm long (at maximum within each individual), 5–10 mm wide, apex acute and terminating in a gland, base acute, strigose on both surfaces; midrib prominent abaxially; lateral nerves pinnate, 2–3 paired, raised abaxially. Summer leaves oblanceolate, 3–10 mm long, 1–5 mm wide, densely strigose on both surfaces. Calyx saucer-shaped, ca. 1.5 mm in diam., densely soft strigose, shallowly 4-lobed; lobes semiorbiculate, ca. 0.5 mm long. Corolla white, no blotches, tubular-funnelform, 7–13 mm long and wide, 4 lobes; tube 3–4 mm long, ca. 3 mm wide, glabrous outside, pilose on upper inside; lobes elliptic to oblong, rounded, 2–5 mm long, ca. 2 mm wide. Stamens 4, irregular, 3–5 mm long, as long as or shorter than corolla; filaments densely pilose on lower half; anthers yellow, oblong, ca. 1 mm long. Ovary ovoid, densely soft strigose, ca. 1.5 mm. Style 3–4 mm long, glabrous, shorter than corolla. Capsule ovoid, 3–4 mm long, 2.5 mm wide, densely strigose.
JAPAN: Kyushu.
The plants inhabit sunny and rocky mountain ridges and slopes at altitudes over 1000 m above sea level. Flowering specimens have been collected from July to August; fruiting specimens have been collected from October to November.
The specific epithet refers to ‘Kyushu’ where the new variety is distributed.
Although the style of this variety is included within the corolla, this part is exserted from the corolla in individuals from Mt. Ichifusa.
This variety is similar to
Spring leaves scattered or crowded on upper branchlets; petioles 0.5–1 mm long; blade thick chartaceous, oblong, 10–35 mm long (25–35 mm long at maximum within each individual), 5–15 mm wide, apex acute and terminating in a gland, base acute, strigose on both surfaces; midrib prominent abaxially; lateral nerves pinnate, 1–3 paired, raised abaxially. Summer leaves oblanceolate, 6–20 mm long, 1–8 mm wide, densely strigose on both surfaces. Calyx saucer-shaped, ca. 1.5 mm in diam., densely soft strigose, shallowly 4-lobed; lobes semiorbiculate, ca. 0.5 mm long. Corolla white, no blotches, openly tubular-funnelform, 8–12 mm long and wide, dissected 1/2 corolla length into 4 lobes; tube 2–3 mm long, ca. 2 mm wide, glabrous outside, pilose on upper inside; lobes elliptic to oblong, rounded, 3–5 mm long, 2–4 mm wide. Stamens 4, subequal, 6–9 mm long, exserted; filaments densely pilose on lower three-quarters; anthers yellow, oblong, ca. 1 mm long. Ovary ovoid, densely soft strigose, ca. 2 mm. Style 5–6 mm long, exserted, glabrous. Capsule ovoid, 3–4 mm long, 2.5 mm wide, densely strigose.
SOUTH KOREA: Jeollabuk-do, Gyeongsangnam-do.
The plants inhabit mountain ridges and slopes at altitudes over 800 m above sea level. Flowering specimens have been collected from June to August; fruiting specimens have been collected from October. Honeybees are frequent visitors to the flowers, suggesting that they are pollinators of the variety.
The specific epithet refers to ‘Korea’ where the new variety is distributed.
1 | Corolla 4 lobes |
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– | Corolla 5 lobes |
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2 | Corolla tubiform |
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– | Corolla tubular-funnelform |
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3 | Spring leaves 25–50 mm long at maximum within each individual |
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– | Spring leaves 10–20 mm long at maximum within each individual |
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4 | Lateral nerves of spring leaf 1 paired, prominently raised abaxially, grooved adaxially, nervules reticulate |
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– | Lateral nerves of spring leaf 1–3 paired, raised abaxially, not grooved adaxially |
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5 | Style 4–10 mm long, exserted from corolla; lateral nerves of spring leaf obscurely raised |
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– | Style 3–4 mm long, included within corolla; lateral nerves of spring leaf raised |
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6 | Corolla tubular-funnelform, tube 2–4 mm long; style 6–13 mm long, exserted from corolla |
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– | Corolla tubiform, tube 5–7 mm long; style 4–5 mm long, included within corolla |
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DNA sequences of chloroplast DNA haplotypes reported in this study were deposited in GenBank under accession numbers; LC499847–LC499863 for
We are grateful to Osamu Takahashi for providing a beautiful photograph; and Seung-Hyun Hwang and Narae Yun in South Korea, Takao Oi, Kyohei Yukitoshi, Takumi Fukuda, Minami Takahashi in Japan for their help in collecting plant material. This study was supported by a Grant-Aid for Scientific Research (15K07473, 16K18714) from the Japan Society for the Promotion of Science, JSPS.