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An Overlooked Tree Species, Micromeles Calocarpa (Rehder) M

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An Overlooked Tree Species, Micromeles Calocarpa (Rehder) M ISSN 1346-7565 Acta Phytotax. Geobot. 72 (1): 23–42 (2021) doi: 10.18942/apg.202007 An Overlooked Tree Species, Micromeles calocarpa (Rehder) M. Aizawa (Rosaceae), from Central Japan MINEAKI AIZAWA Department of Forest Science, School of Agriculture, Utsunomiya University, 350 Mine-machi, Utsunomiya, Tochigi 321-8505, Japan. [email protected] In Japan, two simple-leaved species of trees of Rosaceae tribe Maleae have been treated as Sorbus alni- folia and S. japonica or have been assigned to the genus Aria. However, they are morphologically distinct from Aria and Sorbus and should be treated as species of Micromeles. Under S. japonica, variety calo- carpa (hereafter only calocarpa) was described based on specimens collected in Nikko, Japan, where calocarpa is commonly found. Although calocarpa has been neglected, it is hypothesized that it should be treated as a distinct species characterized by larger leaves with a dense white persistently tomentose abaxial surface and round to truncate leaf base. The phylogenetic position of the Japanese simple-leaved species, including calocarpa, in the tribe Maleae was examined using chloroplast (cp) DNA regions. CpDNA analyses demonstrated that the Japanese taxa should be assigned to Micromeles. Second, to test the hypothesis that calocarpa should be recognized as a distinct species, its phylogenetic relationships among Japanese Micromeles using cpDNA and nuclear low-copy-number genes was examined. Leaf morphology of the three taxa was also compared. The phylogenetic and morphological analyses indicated that calocarpa is indeed distinct from M. japonica and M. alnifolia. Therefore, a new status and new combination, Micromeles calocarpa (Rehder) M. Aizawa, is proposed. Key words: Aria, chloroplast DNA, leaf morphology, Micromeles, nuclear low-copy number gene, Sorbus Two simple-leaved species of trees previously ly lobed leaves, flowers in corymbs, a dimerous assigned to Sorbus L. subgen. Micromeles (Dec- to pentamerous gynoecium with styles coalescent ne.) J. B. Phipps, K. R. Robertson & Spongberg at least at the base, inferior ovary, deciduous up- (Phipps et al. 1990) as S. alnifolia (Siebold & per part of the hypanthium and fruit with mem- Zucc.) K. Koch and S. japonica (Decne.) Hedl. branous endocarp and distinct annular cicatrice (Rosaceae tribe Maleae) are common in Japan. at the apex (Kovanda & Challice 1981). The two They were most recently assigned to Aria (Pers.) simple-leaved Japanese species should now be Host (Ohashi & Iketani 1993, Iketani & Ohashi treated as Micromeles alnifolia (Siebold & Zucc.) 2001). Lo & Donoghue (2012) showed that Sor- Koehne and M. japonica (Decne.) Koehne, but bus sensu lato is polyphyletic and that phyloge- their phylogenetic position within Micromeles nies based on chloroplast DNA (cpDNA) and nu- have not been determined. clear ribosomal internal transcribed spacer (ITS) In 1915, Rehder proposed Sorbus japonica supported the recognition of several genera, in- var. calocarpa Rehder based on specimens col- cluding Micromeles from within Sorbus s.l. Ac- lected in Yumoto, Nikko, Tochigi Prefecture, Ja- cordingly, Micromeles is now regarded as distinct pan (Rehder 1915; Fig. 1). According to Rehder’s from Aria and Sorbus sensu stricto (Li et al. 2017, description, var. calocarpa differed from S. ja- Sennikov & Kurtto 2017, Zhang et al. 2017, ponica var. japonica by having larger fruits (ca. Mezhenska et al. 2018). Micromeles is character- 1.5 cm long) ripening orange-yellow or golden- ized by the deciduous, simple, serrate or shallow- yellow and lacking lenticels and larger leaves 24 Acta Phytotax. Geobot. Vol. 72 (about 10 cm long and 8 cm wide) with a dense species using previously used primers (Wang & white, appressed, persistently tomentose abaxial Zhang 2011, Guo et al. 2016) was conducted, but surface. In contrast, Sorbus japonica var. japoni- it either failed PCR amplification or did not ob- ca produces red fruits with white lenticels and tain sufficiently clear sequence chromatograms. has leaves with loose and somewhat floccose to- As an alternative, to elucidate the phylogenetic mentum that sometimes partially wears off to- relationships among the Japanese species of Mi- ward autumn (Rehder 1915). Later studies cromeles and calocarpa, nuclear low-copy-num- (Hayashi 1969, Ohwi 1975, Kitamura & Murata ber or single-copy genes can be used by referring 1979) continued to recognize plants with larger, to the phylogenetic study of Sorbus s.s. (Li et al. orange-yellow fruit without lenticels as S. japon- 2017). ica var. calocarpa, but Yonekura (2005) regarded In the present study, the phylogenetic position it as S. japonica f. calocarpa (Rehder) Yonek. of the Japanese simple-leaved species, including Other treatments did not recognize var. calocar- calocarpa, in the Maleae was examined using cp- pa (hereafter as calocarpa) probably because the DNA regions that can distinguish between Mi- yellow fruit was considered to be merely varia- cromeles, Aria, and Sorbus s.s. (Lo & Donoghue tion within S. japonica, cf. orange in Ohashi 2012, Sun et al. 2018). Then, to test the hypothe- (1989), orange to scarlet in Iketani & Ohashi sis that calocarpa is a separate species, the phylo- (2001), and reddish or sometimes orange-yellow- genetic relationships between calocarpa and M. ish in Aldasoro et al. (2004). My field observa- alnifolia and M. japonica was examined using tions around Yumoto, Nikko, suggested that ca- two granule-bound starch synthase genes, partial locarpa was relatively common in this region, exons (GBSSI-1) and starch-branching enzyme whereas S. japonica var. japonica was absent. I genes (SBEI) of four nuclear genes analyzed by also observed that the large leaves with a dense Li et al. (2017) as well as cpDNA for samples col- white persistently tomentose abaxial surface, lected from populations across their natural range round to truncate leaf base and shallowly lobed of distribution in Japan. Leaf morphology as a di- margins are more reliably diagnostic of calocar- agnostic character for Japanese Micromeles and pa than fruit coloration. Thus, I hypothesized that calocarpa was also evaluated. calocarpa should be treated a distinct species rather than as an infraspecific taxon of Mi- cromeles japonica. Materials and Methods Previous phylogenetic analyses of Sorbus s.l. and the tribe Maleae, Rosaceae, used either a Sample collection and DNA isolation combination of cpDNA and ITS regions (Lo & Samples of calocarpa, Micromeles japonica, Donoghue 2012), exclusively various chloroplast and M. alnifolia were collected from naturally regions (Sun et al. 2018), or exclusively ITS re- growing trees across their typical range of distri- gions (Wang & Zhang 2011, Li et al. 2012, Guo et bution in Japan (Table 1). For calocarpa, a pre- al. 2016). The respective phylogenies were incon- liminary herbarium survey and the description of sistent between studies, perhaps reflecting that Hayashi (1969) indicated that it was present on Sorbus s.l. has undergone complex evolutionary Yatsugatake in Nagano Prefecture. I therefore events as a result of apomixes, polyploidization collected samples on Yatsugatake in addition to and hybridization (Nelson-Jones et al. 2002, Pel- the collections from around Yumoto, Nikko, licer et al. 2012, Dłużewska et al. 2013). More- Tochigi Prefecture, which is the type locality of over, it is assumed that Micromeles may have calocarpa. In total, 13 trees of calocarpa, 25 of originated through hybridization between Sorbus M. japonica, and 32 of M. alnifolia were collected s.s. and Aria (Lo & Donoghue 2012). Since none from 22 sites, including five sites (Omyojin, Mt. of the studies analyzed M. japonica including ca- Takahara, Funyu, Hosoo, and Hiruzen) where M. locarpa, my preliminary analysis of ITS for the alnifolia and M. japonica occur sympatrically February 2021 AIZAWA — An Overlooked Tree Species, Micromeles calocarpa 25 FIG. 1. Holotype of Sorbus japonica (Decne.) Hedl. var. calocarpa Rehder (E. H. Wilson 7643, 16 Oct., 1914, A [00112644]). 26 Acta Phytotax. Geobot. Vol. 72 TABLE 1. Sample list with taxon, sites, number of samples for genetic analyses (Ng) and leaf morphological analyses (Nm), and observed cpDNA haplotypes and nuclear DNA genotypes. cpDNA Nuclear DNA Lat. Long Altitude Haplotype Genotype Taxon Site (°N) (°E) (m) Ng Nm matK+rbcL GBSSI-1 SBEI Micromeles Suganuma, Katashina, Gunma Pref. 36.82 139.36 1,744 1 - CH1 (1) GH1/GH1 (1) SH1/SH1 (1) calocarpa Yumoto, Nikko, Tochigi Pref.‡ 36.81 139.42 1,520–1,667 6 6 CH1 (6) GH1/GH1 (6) SH1/SH1 (6) Mt. Nyoho, Nikko, Tochigi Pref. 36.80 139.53 1,750 1 1 CH1 (1) GH1/GH1 (1) SH1/SH1 (1) Yachiho, Nagano Pref.‡ 36.04 138.39 1,646–1,694 5 5 CH1 (5) GH1/GH1 (5) SH1/SH1 (4); SH1/SH2 (1) 13 12 Micromeles Omyojin, Shizukuishi, Iwate Pref.† 39.66 140.91 274–365 3 3 CH2 (3) GH5/GH8 (2); SH6/SH6 (3) japonica GH5/GH5 (1) Mt. Takahara, Shioya, Tochigi 36.88 139.80 892–913 4 4 CH2 (4) GH5/GH7 (3); SH6/SH6 (4) Pref.† GH5/GH5 (1) Hanazono, Kitaibaraki, Ibaraki 36.86 140.61 675 1 - CH2 (1) GH5/GH7 (1) SH6/SH6 (1) Pref. Funyu, Shioya, Tochigi Pref.† 36.78 139.82 375–381 4 4 CH2 (4) GH5/GH5 (2); SH6/SH6 (3); GH4/GH5 (2) SH3/SH6 (1) Hosoo, Nikko, Tochigi Pref.† 36.71 139.52 1,080–1,145 2 2 CH2 (1); GH5/GH5 (1); SH3/SH6 (1)*; CH3 (1)* GH5/GH11 (1)* SH6/SH6 (1) Shima, Nakanojo, Gunma Pref. 36.70 138.79 803 1 1 CH2 (1) GH5/GH5 (1) SH3/ SH6 (1) Mt. Mitsumine, Chichibu, Saitama 35.92 138.94 1,137 1 1 CH2 (1) GH5/GH5 (1) SH3/SH6 (1) Pref. Hiruzen, Maniwa, Okayama Pref.† 35.31 133.58 650–771 4 3 CH2 (4) GH5/GH7 (2); SH3/SH6 (3); GH5/GH6(1); SH3/SH3 (1) GH5/GH9(1) Ashiu, Minamitanba, Kyoto Pref. 35.30 135.72 371 1 1 CH2 (1) GH5/GH5 (1) SH3/SH6 (1) Kibune, Kyoto Pref.
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