J. Jpn. Bot. 92(1): 1–11 (2017)

Circumscription of Two Phryma Species () in Japan

Yasuhiko Endo* and Tomonari Miyauchi

College of Science, Ibaraki University, 2-1-1, Bunkyo, Mito, 310-8512 JAPAN *Corresponding author: [email protected]

(Accepted on November 4, 2016)

To revise the of Phryma (Phrymaceae) in Japan, we examined the variation in quantitative features of Japanese Phryma corollas and leaves, as well as ratios in various characters, including 1) the ratio between ‘the width of the corolla’s upper lip at its center (ulw)’ and ‘the width of the lobes of the corolla’s upper lip (ul)’, i.e., ul/ulw, and 2) the ratio between ‘the length (ll)’ and ‘the width (lw)’ of the leaf blades with the longest petiole from each specimen, i.e., lw/ll. Based on our analysis, we concluded that Japanese Phryma could be classified into two distinct species, P. nana and P. oblongifolia. At the 80% confidence interval, the ul/ulw ratio of the corolla upper lips ranges from 0.47–0.59 and the lw/ll ratio of the leaf with the longest petiole ranges from 0.81–1.09 in P. nana, while in P. oblongifolia, the ul/ulw ratio ranges from 0.74–1.02 and the lw/ll ratio ranges from 0.43–0.61.

Key words: Corolla, Japan, leaf, lectotypification, morphological variation, Phryma nana, Phryma oblongifolia, taxonomy.

The genus Phryma (Phrymaceae) was 1972, Kitamura et al. 1957, Shimizu 1997, Miwa established as a monotypic genus based on P. and Kigawa 2001, Ohba 2003, Kobayashi 2009, leptostachya L. that is distributed in East Asia Yuzawa 2014). In most cases, the morphotypes and North America (Hara 1948, 1956, 1962, have been treated as separate taxa at the infra- Thieret 1972, Yamazaki 1993, Lee et al. 1996, Li subspecific rank, i.e., form（Ohwi 1953, 1972, 2000, Nie et al. 2006). The populations in East Kitamura et al. 1957, Shimizu 1997, Kobayashi Asia and North America have been distinguished 2009, Yuzawa 2014) or variety (Hara 1948, as different infra-specific taxa, i.e., subspecies Miwa and Kigawa 2001), although they have (Kitamura and Murata 1957, Li 2000) or sometimes been treated as two distinct species varieties (Hara 1948, 1956, Thieret 1972, Lee et (Ohba 2003) or as a single taxon (Yamazaki al. 1996, Nie et al. 2006). Recently, several new 1993), as well. Incidentally, such morphological diagnostic characters were found to recognize variation of Phryma has not been reported in the East Asian and North American populations Korea (Chang et al. 2014) or China (Hong and at the species rank (Endo et al. 2014), i.e., P. Jun 2011). oblongifolia Koidz. and P. leptostachya L., Initially, the two morphotypes of Japanese respectively. Phryma were distinguished from one another Moreover, two morphologically on the basis of differences in leaf shapes distinguishable groups of Phryma have also been (Hara 1948), and Miwa and Kigawa (2001) recognized in Japan (Hara 1948, Ohwi 1953, subsequently described additional diagnostic

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Table 1. List of morphological data, collection sites, and voucher specimens of 58 Japanese individuals and one Chinese individual of Phryma examined llclw ulw lw Collection site a/llclw ul/ulw ll (mm) ar Group Voucher specimen1) (mm) (mm) (mm) Japan Hokkaido Pref., Hakodate-shi, 0.19 1.37 0.69 1.59 39.28 20.04 13.5 OFG Endo 3533 Takizawa-machi 0.54 1.41 0.98 1.60 114.16 43.49 18.0 OFG Endo 3541 0.13 2.74 0.52 2.48 83.31 79.30 18.5 ASG Endo 3543 Cult./Orig.: Hokkaido Pref., 0.21 2.09 0.62 1.99 93.59 66.17 29.5 ASG Endo 3578 Hakodate-shi Cult./Orig.: Aomori Pref., 0.23 1.83 0.75 1.72 147.39 72.62 20.5 OFG Endo 3591 Mt. Iwaki-san Iwate Pref., Hanaizumi-shi 0.19 1.87 0.49 1.97 69.99 66.74 57.0 ASG Miyauchi 41 0.22 2.02 0.60 2.02 70.09 51.38 49.0 ASG Miyauchi 42 0.18 2.02 0.52 2.08 64.24 61.90 56.0 ASG Miyauchi 43 0.15 2.18 0.47 2.29 49.94 50.38 50.5 ASG Miyauchi 44 0.16 1.87 0.50 2.07 71.07 56.49 35.5 ASG Miyauchi 45 Miyagi Pref., 0.28 1.97 1.03 1.77 46.94 34.49 16.5 OFG Miyauchi 46 Higashimatsushima-shi 0.35 1.73 1.06 1.53 55.60 37.17 24.5 OFG Miyauchi 49 0.38 1.79 0.99 1.76 72.88 48.87 19.5 OFG Miyauchi 50 Fukushima Pref., Iwaki-shi 0.34 1.21 0.70 1.78 136.10 62.64 11.5 OFG Miyauchi 11 0.33 1.27 0.64 1.56 69.38 31.56 15.5 OFG Miyauchi 12 0.35 1.48 0.72 1.80 144.87 68.14 12.0 OFG Miyauchi 13 Ibaraki Pref., Hitachiota-shi, 0.30 1.55 0.89 1.79 95.86 43.37 22.0 OFG Endo & Sannohe 3696 Uemiyakawauchi-machi, 0.31 1.42 0.86 1.89 75.00 39.23 17.5 OFG Endo & Sannohe 3697 Mt. Nishikanasa-yama 0.55 1.09 1.14 1.47 55.80 28.37 18.5 OFG Endo & Sannohe 3698 0.34 1.36 0.99 1.73 70.67 29.01 13.5 OFG Endo & Sannohe 3700 Ibaraki Pref., Mito-shi, 0.38 1.54 1.02 1.77 58.00 33.31 25.5 OFG Miyauchi 3 Abokke-cho 0.37 1.58 0.72 1.87 107.39 57.28 21.0 OFG Miyauchi 5 Cult./Orig.: Ibaraki Pref., 0.32 1.58 0.96 1.78 89.68 43.30 29.0 OFG Endo 3589 Mito-shi, Abokke-cho Ibaraki Pref., Mito-shi, Tano 0.24 2.00 0.68 2.20 32.30 40.91 43.5 ASG Miyauchi 6 0.24 1.99 0.60 2.17 54.91 44.19 30.0 ASG Miyauchi 8 0.11 2.20 0.58 2.02 49.98 38.98 27.0 ASG Miyauchi 9 0.11 1.90 0.44 2.15 54.50 62.29 34.5 ASG Miyauchi 10 Cult./Orig.: Ibaraki Pref., 0.22 1.65 0.48 1.96 51.25 58.99 53.0 ASG Endo 3574 Mito-shi, Tano 0.14 1.92 0.55 2.00 60.76 52.77 75.0 ASG Endo 3575 Ibaraki Pref., Higashi-Ibaraki- 0.34 1.58 1.10 1.36 91.72 47.52 15.5 OFG Endo & Sannohe 3676 gun, Shirosato-machi, 0.16 2.08 0.46 2.25 113.94 97.56 38.0 ASG Endo & Sannohe 3677 Gozen-yama 0.15 2.12 0.51 2.37 100.70 103.92 30.5 ASG Endo & Sannohe 3680 Ibaraki Pref., Higashi-ibaraki- 0.25 1.71 0.98 1.63 67.85 30.32 23.5 OFG Endo 3756 gun, Shirosato-machi Ibaraki Pref., Naka-shi 0.30 1.35 0.99 1.41 75.03 48.69 11.5 OFG Endo 3754 0.17 1.83 0.57 1.69 77.28 85.77 48.0 ASG Endo 3766 0.14 1.89 0.58 1.78 58.92 54.59 26.5 ASG Endo 3767 Tochigi Pref., Haga-gun, 0.22 1.70 0.58 1.99 56.15 57.71 32.0 ASG Miyauchi 21 Motegi-machi 0.20 1.94 0.49 2.16 64.44 67.22 40.5 ASG Miyauchi 22 0.23 1.94 0.51 2.23 72.52 65.52 31.5 ASG Miyauchi 23 0.19 2.01 0.54 1.86 60.64 64.77 37.5 ASG Miyauchi 24 0.24 1.41 0.70 1.95 82.57 41.68 21.5 OFG Miyauchi 26 0.19 1.45 0.99 1.56 80.94 43.67 9.0 OFG Miyauchi 29 February 2017 The Journal of Japanese Botany Vol. 92 No. 1 3

Table 1. Continued llclw ulw lw Collection site a/llclw ul/ulw ll (mm) ar Group Voucher specimen1) (mm) (mm) (mm) Cult./Orig.: Tochigi Pref., 0.25 1.72 0.59 1.96 62.18 59.14 51.0 ASG Endo 3580 Haga-gun, Motegi-machi 0.22 1.73 0.51 1.98 64.57 68.97 62.5 ASG Endo 3581 0.16 1.81 0.49 2.01 66.54 63.84 75.5 ASG Endo 3582 0.19 1.93 0.48 2.14 72.49 63.33 68.5 ASG Endo 3583 Saitama Pref., Kumagaya-shi 0.28 1.64 0.78 2.00 77.30 41.97 27.5 OFG Miyauchi 16 0.12 2.21 0.38 2.35 59.64 40.65 23.0 ASG Miyauchi 18 Chiba Pref., Sakura-shi 0.19 2.10 0.47 2.19 59.12 66.94 35.5 ASG Miyauchi 35 Kagawa Pref., Takamatsu-shi, 0.27 1.32 0.93 1.55 98.73 36.05 11.5 OFG Endo 3770 Shionoe-machi 0.26 1.41 0.76 1.60 63.18 30.89 8.0 OFG Endo 3771 0.37 1.39 0.81 1.80 87.28 34.13 8.5 OFG Endo 3772 0.48 1.31 1.05 1.46 82.60 31.70 12.5 OFG Endo 3773 Yamaguchi Pref., 0.36 1.29 0.94 1.47 51.90 33.20 12.5 OFG Endo 3774 Shimonoseki-shi, Takahata Nagasaki Pref., Sasebo-shi, 0.36 1.48 0.87 1.59 68.48 46.20 12.0 OFG Endo 3780 Eboshi-machi 0.33 0.96 0.65 1.17 63.79 33.70 10.0 OFG Endo 3782 0.28 1.65 0.72 1.65 62.63 35.70 12.0 OFG Endo 3783 0.14 2.12 0.82 2.06 99.49 46.38 11.5 OFG Endo 3812 China Sichuan, between Liziping 0.18 1.80 0.58 1.82 61.07 29.98 55.5 Endo & Nemoto and Ximian 593101 Abbreviations: a. The length of the lobes of the corolla’s upper lip. ar. Number of areoles in 5 × 5 mm2 on lower surface of longest leaf in a individual. ll. Length of the leaf with longest petiole in an individual. llclw. Width of the central lobe of the corolla’s lower lip at its center. lw. Width of the leaf with longest petiole in an individual. ul. Width of the lobes of the corolla’s upper lip. ulw. Width of the corolla’s upper lip at its center. ASG. Asiatic-group. OFG. Oblongifolia-group. Cult. Cultivated at the campus of Ibaraki University in Ibaraki Pref., Japan. Orig. Original collection site. 1)Voucher specimens are deposited in INM (Ibaraki Nature Museum, Japan). characters. According to their analysis, one of were examined, i.e., three Japanese, one the Japanese morphotypes has flowers that are Korean, and 10 Chinese. These individuals were 7–10 mm long and 6 mm wide, corollas with phylogenetically divided into the following upper lips that have narrow lobes, and leaves three groups; the first one was composed of two with obvious veinlets on the abaxial surfaces; Japanese individuals, the second one Chinese, whereas the second morphotype has flowers that and the third one Japanese, one Korean, and nine are 6–7 mm long and 4 mm wide, corollas with Chinese (Nie et al. 2006). This result means that upper lips that have lobes that are equal in width the Japanese Phryma could be divided to the base of lips, and leaves without obvious into two phylogenetic groups. veinlets on the abaxial surfaces (Miwa and The present study aimed to re-evaluate the Kigawa 2001). However, Endo et al. (2014) also morphological variation of Japanese Phryma and examined Japanese Phryma and were unable to attempted to propose an infra-generic taxonomic distinguish two morphological groups because treatment of them. of the continuous variations in corolla size and shape. Materials and Methods On the other hand, Nie et al. (2006) did Specimens examined molecular phylogenetic study on the East Asian We examined 58 Japanese and one Chinese Phryma. In this study, 14 E. Asian individuals individuals of Phryma, for which voucher 4 植物研究雑誌 第 92 巻 第 1 号 2017 年 2 月

Fig. 1. Illustrations of dissected corollas from asiatic- (A) and oblongifolia-group (B) Phryma in Japan. a. Length of the lobes of the upper lip. l. Length of the central lobe of the lower lip. llclw. Width of the central lobe of lower lip at its center. t. Width of the corolla. ul. Width of the lobes of the upper lip. ulw. Width of the upper lip at its center. Voucher specimens pictured: A. Endo 3543. B. Endo & Sannohe 3696. Scale bar = 1 mm.

Fig. 2. Scanning images of the abaxial surface of Phryma leaves. A. Areole number = 75.5 in 5 × 5 mm2 (Endo 3582). B. Areole number = 13.5 in 5 × 5 mm2 (Endo 3533). Scale bar = 1 cm. specimens and collection sites are listed in Table were fixed and suspended in FAA (formalin : 1. Most of these materials were shared by our acetic-acid : 50% aqueous ethanol = 0.5:0.5:9) at previous research (Endo et al. 2014). We newly the collection sites. The corollas were dissected examined the Phryma plants distributed in the and opened along the tube (Fig. 1) and were western part of Japan in the present study. photographed under a dissecting microscope. The corolla images were then enlarged by about Corolla shape 20–40 times and were used to measure the To examine the corolla shape, fresh flowers following parts (Fig. 1): length (a) and width (ul) February 2017 The Journal of Japanese Botany Vol. 92 No. 1 5 of the lobes of corolla’s upper lip, width of the Based on the scatter diagram of Fig. 3, upper lip at its center (ulw), length of the central showing the relationships between ul/ulw and lobe of the lower lip (l), width of the central lobe lw/ll, we attempted to separate 58 Japanese of lower lip at its center (llclw), and length (cl) Phryma specimens into two groups (the asiatic- and width (t) of corollas. Each of the lengths and group, ASG, and the oblongifolia-group, OFG). widths were measured using a vernier scale. These two groups were almost clearly separated in the scatter diagrams with ul/ulw and l/llclw Leaf shape (Fig. 4), ll and lw (Fig. 5), lw/ll and ln (ar) (Fig. During our preliminary observations, we 6), a/llclw and ln (ar) (Fig. 7). Therefore, we recognized that, for individual plants, the ratio propose to recognize each of the two groups as between the length (ll) and width (lw) of leaf distinct species. blades (i.e., lw/ll) was largest for the leaf that The comparison of floral and foliar had the longest petiole. Thus, ll and lw were characters between two species, i.e., ASG and measured for the leaf with the longest petiole in OFG, is shown in Table 3. Based on the data, each herbarium specimen, in order to compare there are significant differences in the values leaf shapes. of a, cl, llclw, ln (ar), lw, t, ul, and ulw, and in the ratios of a/uclw, l/llclw, lw/ll, and ul/ulw, Leaf veinlet network between ASG and OFG (Table 3). The range We also measured the density of areoles of the values and ratios were found to overlap on the abaxial surface of leaves, in order to between ASG and OFG. On the other hand, determine whether the leaf veinlet networks the 80% confidence intervals do not overlap were fine or coarse (Fig. 2). We chose the longest between ASG and OFG in the llclw, ln (ar) and, leaf from each herbarium specimen and scanned ulw values and the a/llclw, lw/ll, and ul/ulw its central part, using a scanner with 1600 dpi ratios (Table 3). This indicates that individual resolution (CanoScan 8400f; Canon Inc., Tokyo, characters may be insufficient for discriminating Japan). Then, using the scanned images, we between the two groups, and that using a counted the number of areoles (ar) that occurred combination of characters could be necessary for in an area of 5 × 5 mm2 on each leaf. the discrimination.

Statistic analysis Morphological relationships between Japanese We calculated the means, ranges, and and Chinese Phryma standard deviations of each variable for all The results of the measurements of one specimens (Table 2) and for the two groups Chinese individual in the present study are recognized in this study (Table 3). We conducted shown in Table 1. Based on the results, we t-tests to compare the measurements and ratios recognized the following relationships between of the two groups, using the computer program the two Japanese groups, i.e., ASG and OFG Microsoft Excel for Mac 2011 ver. 14.3.9 groups, and the Chinese individual. (Microsoft Co., Redmond, WA) (Table 3). 1) The ratio a/llclw of the Chinese individual is 0.18, and is put in the range of those of ASG. Results and Discussion 2) The ul/ulw is 0.58, and is in the range of Distinction between the two morphotypes of ASG. Japanese Phryma 3) The lw/ll is 0.48, and is in the range of The results of the measurements of characters OFG. and statistical analyses were shown in Tables 2 4) The ln (ar) is 4.02, and is in the range of and 3. ASG. 6 植物研究雑誌 第 92 巻 第 1 号 2017 年 2 月

Table 2. Floral and foliar characters of Japanese Phryma Range Average Standard deviation Corolla length (cl) (mm) 4.79–9.18 7.05 0.75 Corolla width (t) (mm) 2.31–6.06 4.30 0.68 Length of the lobe of the corolla’s upper lip (a) (mm) 0.20–0.76 0.42 0.11 Width of the lobe of the corolla’s upper lip (ul) (mm) 0.76–1.83 1.31 0.27 Width of the corolla’s upper lip at its center (ulw) (mm) 1.17–2.48 1.84 0.27 Length of the central lobe of the corolla’s lower lip (l) (mm) 1.12–2.43 1.60 0.24 Width of the central lobe of the corolla’s lower lip at its center (llclw) (mm) 0.96–2.74 1.70 0.32 Length of the leaf blade with the longest petiole in a individual (ll) (mm) 32.30–147.39 74.44 22.79 Width of the leaf blade with the longest petiole in a individual (lw) (mm) 20.04–103.92 50.88 16.96 Number of areoles in 5 × 5 mm2 area on the abaxial surface of leaf (ar) 6.5–75.5 26.6 16.3

Table 3. Comparison of floral and foliar characters between two morphological groups in JapanesePhryma Asiatic-group Oblongifolia-group Character p-value Range AVE SD CI (95%) CI (80%) Range AVE SD CI (95%) CI (80%) a (mm) 0.20–0.48 0.35 0.07 0.21–0.49 0.28–0.42 0.26–0.76 0.46 0.12 0.22–0.70 0.34–0.58 *1) a/llclw 0.11–0.25 0.18 0.04 0.10–0.26 0.14–0.22 0.14–0.55 0.32 0.09 0.14–0.50 0.23–0.41 * cl (mm) 6.46–9.18 7.55 0.66 5.59–9.51 6.90–8.20 4.79–7.82 6.70 0.64 5.45–7.95 6.07–7.33 * l (mm) 1.30–2.43 1.63 0.23 1.18–2.08 1.40–1.86 1.14–2.34 1.60 0.29 1.03–2.17 1.32–1.88 0.69 32.30– 34.34– 50.40– 39.28– 30.51– 56.01– ll (mm) 66.48 16.40 81.52 26.03 0.01 113.94 98.62 82.55 147.39 132.53 107.02 llclw (mm) 1.65–2.74 1.98 0.21 1.57–2.39 1.77–2.19 0.96–2.12 1.50 0.24 1.03–1.97 1.26–1.74 * l/llclw 0.67–1.03 0.83 0.09 0.65–1.01 0.74–0.92 0.77–1.66 1.08 0.18 0.73–1.43 0.90–1.26 * ln (ar) 2.92–4.32 3.70 0.36 2.99–4.41 3.35–4.05 2.08–3.37 2.74 0.35 2.05–3.43 2.40–3.08 * 38.98– 32.80– 47.70– 20.04– 18.69– 29.89– lw (mm) 62.61 15.21 41.09 11.43 * 103.92 92.42 77.52 72.62 63.49 52.29 lw/ll 0.68–1.27 0.95 0.14 0.68–1.22 0.81–1.09 0.37–0.73 0.52 0.09 0.34–0.70 0.43–0.61 * t (mm) 4.08–6.06 4.81 0.40 4.03–5.59 4.42–5.20 2.31–5.8 3.92 0.66 2.63–5.21 3.27–4.57 * ul (mm) 0.89–1.51 1.10 0.13 0.85–1.35 0.97–1.23 0.76–1.83 1.46 0.24 0.99–1.93 1.22–1.70 * ulw (mm) 1.69–2.48 2.09 0.18 1.74–2.44 1.91–2.27 1.17–2.06 1.67 0.19 1.30–2.04 1.48–1.86 * ul/ulw 0.38–0.68 0.53 0.06 0.41–0.65 0.47–0.59 0.64–1.14 0.88 0.14 0.61–1.15 0.74–1.02 * Abbreviations: a. Length of the lobe of the corolla’s upper lip. AVE. Average. cl. Corolla length. CI. Confidence interval. l. Length of the central lobe of the corolla’s lower lip. ll. Length of the leaf blade with the longest petiole in an individual. llclw. Width of the central lobe of the corolla’s lower lip at its center. ln (ar). Natural logarithm of the number of areoles (ar) in 5 × 5 mm2 area on the abaxial surface of leaf. lw. Width of the leaf blade with the longest petiole in an individual. SD. Standard deviation. t. Corolla width. ul. Width of the lobe of the corolla’s upper lip. ulw. Width of the corolla’s upper lip at its center. 1)*: p < 0.01.

Therefore, the Chinese individual has a Japanese Phryma plants could be divided into combination of the features of Japanese ASG two monophyletic groups. Similarly, the results group and OFG group. of the present morphological analysis indicates that Japanese Phryma plants could be classified Comparison between the result of the into two morphological groups, i.e., ASG molecular phylogenetic analysis and that of and OFG groups. However, the relationships the morphological analysis on the variation of between the molecular phylogenetic groups Japanese Phryma and the morphological groups are unclear Based on the results of the molecular yet. To know the relationships, we need phylogenetical analysis by Nie et al. (2006), molecular phylogenetic studies on the Japanese February 2017 The Journal of Japanese Botany Vol. 92 No. 1 7

Fig. 3. Scatter plot showing the correlation between the ul/ulw ratios and lw/ll ratios of Japanese Phryma specimens. ll. Length of the leaf blade with the longest petiole in an individual. lw. Width of the leaf blade of the same leaf. ul. Width of the lobes of upper lip. ulw. Width of the upper lip at the center. Open circles indicate the data from 31 oblongifolia-group specimens, and filled circles indicate the data from 27 asiatic-group specimens.

Fig. 4. Scatter plot showing the correlation between the ul/ulw and l/llclw ratios of 58 Japanese Phryma specimens. l. Length of the central lobe of lower lip. llclw. Width of the central lobe of lower lip at its center. ul. Width of the lobes of upper lip. ulw. Width of the upper lip at the center. Open circles indicate the data from oblongifolia-group specimens, and filled circles indicate the data from asiatic-group specimens. 8 植物研究雑誌 第 92 巻 第 1 号 2017 年 2 月

Fig. 5. Scatter plot showing the correlation between the length (ll) and the width (lw) of the leaf blade with the longest petiole from each of Japanese Phryma specimens. Open circles indicate the data from 31 oblongifolia-group specimens, and filled circles indicate the data from 27 asiatic-group specimens. Asterisks indicate the leaves of the lectotypes or original materials for P. oblongifolia (1–3), P. nana (4), P. humilis (5), and P. leptostachya var. asiatica (6).

Fig. 6. Scatter plot showing the correlation between the lw/ll ratio and ln (ar) for Japanese Phryma specimens. ll. Length of the same leaf. ar. Number of areoles in 5 × 5 mm2 of the abaxial surface of leaves. lw. Width of the leaf blade with the longest petiole in an individual. Open circles indicate the data from 31 oblongifolia-group specimens, and filled circles indicate the data from 27 asiatic-group specimens. Asterisks indicate the leaves of the lectotypes or original materials for P. oblongifolia (1–3), P. nana (4), P. humilis (5), and P. leptostachya var. asiatica (6). February 2017 The Journal of Japanese Botany Vol. 92 No. 1 9

Fig. 7. Scatter plot showing the correlation between a/llclw ratios and the natural logarithm of ar for Japanese Phryma specimens. a. Length of the lobes of the upper lip. ar. Number of areoles in 5 × 5 mm2 of the abaxial surface of leaves. llclw. Width of the central lobe of lower lip at its center. Open circles indicate the data from 31 oblongifolia-group specimens, and filled circles indicate the data from 27 asiatic-group specimens. morphological groups. lip at its center (ulw), i.e., ul/ulw, ranges from 0.47 to 0.59 and the ratio of the width (lw) to the Taxonomic treatment length (ll) of the leaf with the longest petiole, i.e., Four Phryma taxa have been previously lw/ll, ranges from 0.81 to 1.09 ...... P. nana described in Japan, i.e., P. oblongifolia Koidz., A. The ul/ulw ratio ranges from 0.74 to 1.02 P. nana Koidz., P. humilis Koidz., and P. and the lw/ll ratio ranges from 0.43 to 0.61 ...... leptostachya L. var. asiatica Hara (Table 4). Any ...... P. oblongifolia type specimens of these four taxa were not cited in their original descriptions (Koidzumi 1929, Phryma nana Koidz. in Acta Phytotax. 1939, Hara 1948). Therefore, we designate here Geobot. 8: 191 (1939). – P. leptostachya L. var. the lectotypes for these four taxa from their nana (Koidz.) H. Hara, Enum. Sperm. Jap. 1: original materials as shown in Table 4. 297 (1948). – P. leptostachya L. subsp. asiatica In the values of ar, ll, ln (ar), lw, and the lw/ll (H. Hara) Kitam. f. nana (Koidz.) Akasawa in ratio, the lectotypes and original materials of P. Bull. Kochi Women’s Univ., Ser. Nat. Sci. 18: 6 oblongifolia and P. humilis belong to OFG, and (1970). Lectotype (designated here): JAPAN. the lectotypes of P. nana and P. leptostachya var. Honshu. Kyoto Pref., Mt. Hiyei-zan, 12 Aug. asiatica belong to ASG (Figs. 5, 6). Therefore, 1936, K. Takeuchi s.n. (KYO). according to the priority rule of nomenclature, L. var. asiatica H. the correct names of OFG and ASG are P. Hara, Enum. Sperm. Jap. 1: 297 (1948). – P. oblongifolia Koidz. and P. nana Koidz., leptostachya L. subsp. asiatica (H. Hara) Kitam. respectively. in Kitam. & Murata in Acta Phytotax. Geobot. Based on the differences of 80% confidence 17: 7 (1957). – P. asiatica (H. Hara) O. Deg. & intervals, the two species are distinguished as I. Deg. in Phytologia 22: 212 (1971). Lectotype follows (Table 3): (designated here): JAPAN. Hokkaido. Hidaka A. The ratio of the width of the lobes of the Subpref., ‘in forests at the foot of Mt. Apoi’, 6 upper corolla lip (ul) to the width of the upper Aug. 1933, H. Hara s.n. (TI). 10 植物研究雑誌 第 92 巻 第 1 号 2017 年 2 月

Table 4. The features of the type specimens of Japanese Phryma plants Scientific name Collection site and collector ll lw lw/ll ar ln (ar) Herb Type P. oblongifolia Koidz. Ganjusan 103.66 42.11 0.41 33.0 3.5 KYO original material U. Faurie 2535 Nuruyu 109.13 40.89 0.37 28.0 3.3 KYO original material U. Faurie 789 Fujiyama 43.46 22.09 0.51 12.5 2.5 KYO lectotype S. Matsuda 31 July 1893 Hakone ------not found1)

Kiusyu: Prov. Higo, Nishize ------not found

P. nana Koidz. Kyoto, Mt. Hiyeizan 17.40 16.19 0.93 35.5 3.6 KYO lectotype T. Takeuchi 12 Aug. 1936 P. humilis Koidz. Rikutiu: Iwategun, Matsuomura 31.15 20.81 0.67 77.5 4.4 KYO lectotype Y. Fukuta 288 P. leptostachya L. In forests at the foot of Mt. Apoi 52.07 46.91 0.90 73.0 4.3 TI lectotype var. asiatica H. Hara H. Hara 6 Aug. 19332) Abbreviations: ar. Number of areoles in 5 × 5 mm2 area on the abaxial surface of leaf. herb. Abbreviations of the name of herbarium, where the lectotype specimens and original materials are deposited. ll. Length of the leaf blade with the longest petiole in an individual. lw. Width of the leaf blade with the longest petiole in an individual. -. No data. 1)Habitats were referred to in the protologues, but the specimens were not referred and not able to be found. 2)Hara (1949) cited his literature, i.e., ‘Hara in Bot. Mag. Tokyo LI: 639 (1937)’, in the description of P. leptostachya var. asiatica. In this literature, Hara (1937) referred only the habitat of the variation, i.e., forests at the foot of Mt. Apoi. On the other hand, a specimen, collected by Hara at Mt. Apoi in 1933, is deposited in TI. Therefore, this specimen is presumed to be an original material of var. asiatica.

Phryma oblongifolia Koidz. in Bot. Mag. The authors thank curators of the herbaria, (Tokyo) 43: 400 (1929); Ohba, Fl. Chiba: 546 KYO, TI, and TNS, for their permission in (2003); Y. Endo & al. in J. Jpn. Bot. 89: 406 examining the herbarium specimens, and (2014). – P. leptostachya L. var. oblongifolia thank Tomoyuki Nemoto (Ishinomaki Senshu (Koidz.) Honda in Bot. Mag. (Tokyo) 50: 608 University, Japan), Yu Iokawa (Joetsu University (1936); H. Hara in J. Fac. Sci. Univ. Tokyo Bot. of Education, Japan), and Xiang-Yun Zhu 6: 377 (1956); & Fl. E. Himal.: 306 (1966); (Academia Sinica, Beijing, China), for their help T. Yamaz. in K. Iwats. & al., Fl. Jap. 3a: 322 in collecting materials in China. (1993); T. Miwa & Kigawa, Fl. Kanagawa: This work was partially supported by 1284 (2001). – P. leptostachya L. var. asiatica H. JSPS KAKENHI Grant numbers 15405013, Hara f. oblongifolia (Koidz.) Ohwi in Bull. Nat. 26440205 (to Y.E.). Sci. Mus. no. 33: 86 (1953); & Fl. Jap.: 1075 (1953). Lectotype (designated here): JAPAN. References Honshu. ‘Fujiyama’, 31 July 1893, S. Matsuda Chang C. S., Kim H. and Chang K. S. 2014. Provisional s.n. (KYO). Checklist of Vascular Plants for the Korea Peninsula Flora (KPF) (Version 1.0). The Designpost, Seoul. Phryma humilis Koidz. in Acta Phytotax. Endo Y., Miyauchi T. and Sannohe T. 2014. Morphological Geobot. 8: 192 (1939). – P. leptostachya L. var. differences in flowers between Eastern Asian and humilis (Koidz.) H. Hara, Enum. Sperm. Jap. Eastern North American Phryma (Phrymaceae). J. Jpn. 1: 297 (1948). Lectotype (designated here): Bot. 89: 394–408. Hara H. 1949. Enumeratio Spermatophytarum Japonicarum JAPAN. Honshu. ‘Rikutiu, Iwate-gun, Matsuo- 1. Iwanami Shoten, Tokyo. mura’, Y. Fukuta 288 (KYO). Hara H. 1956. Contributions to the study of variations in the Japanese plants closely related to those of Europe February 2017 The Journal of Japanese Botany Vol. 92 No. 1 11

or North America. Part 2. J. Fac. Sci. Univ. Tokyo, III Li Z.-Y. 2000. Taxonomic notes on the genus Phryma L. 6: 343–391. from Asia. Acta Phytotax. Sin. 38: 386–391. Hara H. 1962. Racial differencies in widespread species, Miwa T. and Kigawa S. 2001. Phrymaceae. In: The Flora- with special reference to those common to Japan and Kanagawa Association (ed.), Flora of Kanagawa 2001. North America. Amer. J. Bot. 49: 647–652. p. 1284. The Kanagawa Prefectural Museum of Natural Hong D. and Jun W. 2011. Phryamaceae. In: Wu Z., Raven History, Kanagawa Prefecture (in Japanese). P. H. and Hong D. (eds.), Flora of China 19: 493–494. Nie Z.-L., Sun H., Beardsley P. M., Olmstead R. G. and Science Press, Beijing and Missouri Botanical Garden Wen J. 2006. Evolution of biogeographic disjunction Press, St. Louis. between eastern Asia and eastern North America in Kitamura S. and Murata G. 1957. New names and new Phryma (Phrymaceae). Amer. J. Bot. 93: 1343–1356. conceptions adopted in our Coloured Illustrations Ohba T. 2003. Phrymaceae Schauer 1847. In: Ohba T. of Herbaceous Plants of Japan (Sympetalae). Acta (ed.), Flora of Chiba. Chiba Prefectural Government, Phytotax. Geobot. 17 (1): 5–13. Chiba (in Japanese). Kitamura S., Murata G. and Hori M. 1957. Phrymaceae. In: Ohwi J. 1953. Flora of Japan. Shibundo, Tokyo (in Coloured Illustrations of Herbaceous Plants of Japan, 1 Japanese). (Sympetalae). Hoikusha Publishing Co. Ltd., Osaka (in Ohwi J. 1972. Flora of Japan. Revised edition. Shibundo, Japanese). Tokyo (in Japanese). Kobayashi S. 2009. Phrymaceae. Flora of Kochi. p. 453. Shimizu T. 1997. 112. . In: Shimizu T. (ed.), The Kochi Prefectural Makino Botanical Garden, Flora of Nagano Prefecture. pp. 923–928. Shinano Kochi Prefecture (in Japanese). Mainichi Press, Nagano (in Japanese). Koidzumi G. 1929. Contributiones ad cognitionem florae Thieret J. W. 1972. The Phrymaceae in the southeastern Asiae Orientalis. Bot. Mag. (Tokyo) 43: 382–407. United States. J. Arnold Arbon. 53: 226–233. Koidzumi G. 1939. Contributiones ad cognitionem florae Yamazaki T. 1993. Phrymaceae. In: Iwatsuki K., Yamazaki Asiae Orientalis. Acta Phytotax. Geobot. 8: 188–194. T., Boufford D. E. and Ohba H. (eds.), Flora of Japan. Lee N. S., Sang T., Crawford D. J., Yeau S. H. and Kim S.- IIIa: 322. Kodansha Ltd., Tokyo. C. 1996. Molecular divergence between disjunct taxa Yuzawa Y. 2014. Flora of Iwaki. Rekishi Shunju Publishing in Eastern Asia and Eastern North America. Amer. J. Co. Ltd., Fukushima (in Japanese). Bot. 83: 1373–1378.

遠藤泰彦，宮内智成：日本産ハエドクソウ属 2 種の輪郭 日本においてハエドクソウ属は形態の違いから 2 型 0.81–1.09 で あ る 群 と ul/ulw が 0.74–1.02 で， か つ lw/ が知られており，これを品種の階級または変種の階級で ll が 0.43–0.61 である群である．これら 2 群の間では 1) 分類するという考えがある．一方で，種の階級で分類す と 2) の特徴以外に，3) 下唇弁中央裂片の中央部の幅， るべきであるとする考えや同一種とする考えもある．本 ４) 葉裏面の単位面積当たりの小区画数，5) 上唇弁の中 研究では，日本産ハエドクソウ属の形態変異の実態を明 央部の幅，6) 上唇弁の裂片の長さと下唇弁の中央裂片 らかにし，その結果に基づいた，分類学的な取り扱いの の長さの比，についても顕著な違いがあった．以上によ 提案を目的とした． り日本産ハエドクソウ属の 2 群を種の階級で分類するこ 本研究の結果から，ハエドクソウ属の以下に述べる とを提案する．2 群のうち，前者はハエドクソウ（チャ 形質に変異を認めた．花では，1)「上唇弁の裂片部の幅 ボハエドクソウ）Phryma nana Koidz.，後者はナガバハ (ul)」と「上唇弁の中央部の幅 (ulw)」の比 (ul/ulw)，葉 エドクソウ（ヒメハエドクソウ）P. oblongifolia Koidz. では，2) 個体中最長な葉柄を持つ葉の「葉身の幅 (lw)」 に相当する．なお，本研究では同属の日本産分類群の選 と「葉身の長さ (ll)」の比 (lw/ll) である．結果として 1) 定基準標本の指定を併せて行った． と 2) の違いから，調査した植物群を 2 群に分けること （茨城大学理学部） ができた．つまり，ul/ulw が 0.47–0.59 で，かつ lw/ll が