circumscription of two phryma species (phrymaceae) in japan

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Circumscription of Two Phryma Species (Phrymaceae) 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 taxonomy 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.

J. Jpn. Bot. 92(1): 1–11 (2017)

The genus Phryma (Phrymaceae) was established as a monotypic genus based on P. leptostachya L. that is distributed in East Asia and North America (Hara 1948, 1956, 1962, Thieret 1972, Yamazaki 1993, Lee et al. 1996, Li 2000, Nie et al. 2006). The populations in East Asia and North America have been distinguished as different infra-specific taxa, i.e., subspecies (Kitamura and Murata 1957, Li 2000) or varieties (Hara 1948, 1956, Thieret 1972, Lee et al. 1996, Nie et al. 2006). Recently, several new diagnostic characters were found to recognize the East Asian and North American populations at the species rank (Endo et al. 2014), i.e., P. oblongifolia Koidz. and P. leptostachya L., respectively.

M o r e o v e r , t w o m o r p h o l o g i c a l l y distinguishable groups of Phryma have also been recognized in Japan (Hara 1948, Ohwi 1953,

1972, Kitamura et al. 1957, Shimizu 1997, Miwa and Kigawa 2001, Ohba 2003, Kobayashi 2009, Yuzawa 2014). In most cases, the morphotypes have been treated as separate taxa at the infra-subspecific rank, i.e., form（Ohwi 1953, 1972, Kitamura et al. 1957, Shimizu 1997, Kobayashi 2009, Yuzawa 2014) or variety (Hara 1948, Miwa and Kigawa 2001), although they have sometimes been treated as two distinct species (Ohba 2003) or as a single taxon (Yamazaki 1993), as well. Incidentally, such morphological variation of Phryma has not been reported in Korea (Chang et al. 2014) or China (Hong and Jun 2011).

Initially, the two morphotypes of Japanese Phryma were distinguished from one another on the basis of differences in leaf shapes (Hara 1948), and Miwa and Kigawa (2001) subsequently described additional diagnostic

2 植物研究雑誌　第 92巻　第 1号 2017年 2月

Table 1. List of morphological data, collection sites, and voucher specimens of 58 Japanese individuals and one Chinese individual of Phryma examined

Collection site a/llclw llclw (mm) ul/ulw ulw

(mm) ll (mm) lw

(mm) ar Group Voucher specimen1)

JapanHokkaido Pref., Hakodate-shi,

Takizawa-machi0.19 1.37 0.69 1.59 39.28 20.04 13.5 OFG Endo 35330.54 1.41 0.98 1.60 114.16 43.49 18.0 OFG Endo 35410.13 2.74 0.52 2.48 83.31 79.30 18.5 ASG Endo 3543

Cult./Orig.: Hokkaido Pref., Hakodate-shi

0.21 2.09 0.62 1.99 93.59 66.17 29.5 ASG Endo 3578

Cult./Orig.: Aomori Pref., Mt. Iwaki-san

0.23 1.83 0.75 1.72 147.39 72.62 20.5 OFG Endo 3591

Iwate Pref., Hanaizumi-shi 0.19 1.87 0.49 1.97 69.99 66.74 57.0 ASG Miyauchi 410.22 2.02 0.60 2.02 70.09 51.38 49.0 ASG Miyauchi 420.18 2.02 0.52 2.08 64.24 61.90 56.0 ASG Miyauchi 430.15 2.18 0.47 2.29 49.94 50.38 50.5 ASG Miyauchi 440.16 1.87 0.50 2.07 71.07 56.49 35.5 ASG Miyauchi 45

Miyagi Pref., Higashimatsushima-shi

0.28 1.97 1.03 1.77 46.94 34.49 16.5 OFG Miyauchi 460.35 1.73 1.06 1.53 55.60 37.17 24.5 OFG Miyauchi 490.38 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 110.33 1.27 0.64 1.56 69.38 31.56 15.5 OFG Miyauchi 120.35 1.48 0.72 1.80 144.87 68.14 12.0 OFG Miyauchi 13

Ibaraki Pref., Hitachiota-shi, Uemiyakawauchi-machi, Mt. Nishikanasa-yama

0.30 1.55 0.89 1.79 95.86 43.37 22.0 OFG Endo & Sannohe 36960.31 1.42 0.86 1.89 75.00 39.23 17.5 OFG Endo & Sannohe 36970.55 1.09 1.14 1.47 55.80 28.37 18.5 OFG Endo & Sannohe 36980.34 1.36 0.99 1.73 70.67 29.01 13.5 OFG Endo & Sannohe 3700

Ibaraki Pref., Mito-shi, Abokke-cho

0.38 1.54 1.02 1.77 58.00 33.31 25.5 OFG Miyauchi 30.37 1.58 0.72 1.87 107.39 57.28 21.0 OFG Miyauchi 5

Cult./Orig.: Ibaraki Pref., Mito-shi, Abokke-cho

0.32 1.58 0.96 1.78 89.68 43.30 29.0 OFG Endo 3589

Ibaraki Pref., Mito-shi, Tano 0.24 2.00 0.68 2.20 32.30 40.91 43.5 ASG Miyauchi 60.24 1.99 0.60 2.17 54.91 44.19 30.0 ASG Miyauchi 80.11 2.20 0.58 2.02 49.98 38.98 27.0 ASG Miyauchi 90.11 1.90 0.44 2.15 54.50 62.29 34.5 ASG Miyauchi 10

Cult./Orig.: Ibaraki Pref., Mito-shi, Tano

0.22 1.65 0.48 1.96 51.25 58.99 53.0 ASG Endo 35740.14 1.92 0.55 2.00 60.76 52.77 75.0 ASG Endo 3575

Ibaraki Pref., Higashi-Ibaraki-gun, Shirosato-machi, Gozen-yama

0.34 1.58 1.10 1.36 91.72 47.52 15.5 OFG Endo & Sannohe 36760.16 2.08 0.46 2.25 113.94 97.56 38.0 ASG Endo & Sannohe 36770.15 2.12 0.51 2.37 100.70 103.92 30.5 ASG Endo & Sannohe 3680

Ibaraki Pref., Higashi-ibaraki-gun, Shirosato-machi

0.25 1.71 0.98 1.63 67.85 30.32 23.5 OFG Endo 3756

Ibaraki Pref., Naka-shi 0.30 1.35 0.99 1.41 75.03 48.69 11.5 OFG Endo 37540.17 1.83 0.57 1.69 77.28 85.77 48.0 ASG Endo 37660.14 1.89 0.58 1.78 58.92 54.59 26.5 ASG Endo 3767

Tochigi Pref., Haga-gun, Motegi-machi

0.22 1.70 0.58 1.99 56.15 57.71 32.0 ASG Miyauchi 210.20 1.94 0.49 2.16 64.44 67.22 40.5 ASG Miyauchi 220.23 1.94 0.51 2.23 72.52 65.52 31.5 ASG Miyauchi 230.19 2.01 0.54 1.86 60.64 64.77 37.5 ASG Miyauchi 240.24 1.41 0.70 1.95 82.57 41.68 21.5 OFG Miyauchi 260.19 1.45 0.99 1.56 80.94 43.67 9.0 OFG Miyauchi 29

February 2017 TheJournal of Japanese Botany Vol. 92 No. 1 3

characters. According to their analysis, one of the Japanese morphotypes has flowers that are 7–10 mm long and 6 mm wide, corollas with upper lips that have narrow lobes, and leaves with obvious veinlets on the abaxial surfaces; whereas the second morphotype has flowers that are 6–7 mm long and 4 mm wide, corollas with upper lips that have lobes that are equal in width to the base of lips, and leaves without obvious veinlets on the abaxial surfaces (Miwa and Kigawa 2001). However, Endo et al. (2014) also examined Japanese Phryma and were unable to distinguish two morphological groups because of the continuous variations in corolla size and shape.

On the other hand, Nie et al. (2006) did molecular phylogenetic study on the East Asian Phryma. In this study, 14 E. Asian individuals

were examined, i.e., three Japanese, one Korean, and 10 Chinese. These individuals were phylogenetically divided into the following three groups; the first one was composed of two Japanese individuals, the second one Chinese, and the third one Japanese, one Korean, and nine Chinese (Nie et al. 2006). This result means that the Japanese Phryma plants could be divided into two phylogenetic groups.

The present study aimed to re-evaluate the morphological variation of Japanese Phryma and attempted to propose an infra-generic taxonomic treatment of them.

Materials and MethodsSpecimens examined

We examined 58 Japanese and one Chinese individuals of Phryma, for which voucher

Table 1. Continued

Collection site a/llclw llclw (mm) ul/ulw ulw

(mm) ll (mm) lw

(mm) ar Group Voucher specimen1)

Cult./Orig.: Tochigi Pref., Haga-gun, Motegi-machi

0.25 1.72 0.59 1.96 62.18 59.14 51.0 ASG Endo 35800.22 1.73 0.51 1.98 64.57 68.97 62.5 ASG Endo 35810.16 1.81 0.49 2.01 66.54 63.84 75.5 ASG Endo 35820.19 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 160.12 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 35Kagawa Pref., Takamatsu-shi,

Shionoe-machi0.27 1.32 0.93 1.55 98.73 36.05 11.5 OFG Endo 37700.26 1.41 0.76 1.60 63.18 30.89 8.0 OFG Endo 37710.37 1.39 0.81 1.80 87.28 34.13 8.5 OFG Endo 37720.48 1.31 1.05 1.46 82.60 31.70 12.5 OFG Endo 3773

Yamaguchi Pref., Shimonoseki-shi, Takahata

0.36 1.29 0.94 1.47 51.90 33.20 12.5 OFG Endo 3774

Nagasaki Pref., Sasebo-shi, Eboshi-machi

0.36 1.48 0.87 1.59 68.48 46.20 12.0 OFG Endo 37800.33 0.96 0.65 1.17 63.79 33.70 10.0 OFG Endo 37820.28 1.65 0.72 1.65 62.63 35.70 12.0 OFG Endo 37830.14 2.12 0.82 2.06 99.49 46.38 11.5 OFG Endo 3812

ChinaSichuan, between Liziping

and Ximian0.18 1.80 0.58 1.82 61.07 29.98 55.5 Endo & Nemoto

593101Abbreviations: 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).


specimens and collection sites are listed in Table 1. Most of these materials were shared by our previous research (Endo et al. 2014). We newly examined the Phryma plants distributed in the western part of Japan in the present study.

Corolla shapeTo examine the corolla shape, fresh flowers

were fixed and suspended in FAA (formalin : acetic-acid : 50% aqueous ethanol = 0.5:0.5:9) at the collection sites. The corollas were dissected and opened along the tube (Fig. 1) and were photographed under a dissecting microscope. The corolla images were then enlarged by about 20–40 times and were used to measure the following parts (Fig. 1): length (a) and width (ul)

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.


of the lobes of corolla’s upper lip, width of the upper lip at its center (ulw), length of the central lobe of the lower lip (l), width of the central lobe of lower lip at its center (llclw), and length (cl) and width (t) of corollas. Each of the lengths and widths were measured using a vernier scale.

Leaf shapeDuring our preliminary observations, we

recognized that, for individual plants, the ratio between the length (ll) and width (lw) of leaf blades (i.e., lw/ll) was largest for the leaf that had the longest petiole. Thus, ll and lw were measured for the leaf with the longest petiole in each herbarium specimen, in order to compare leaf shapes.

Leaf veinlet networkWe also measured the density of areoles

on the abaxial surface of leaves, in order to determine whether the leaf veinlet networks were fine or coarse (Fig. 2). We chose the longest leaf from each herbarium specimen and scanned its central part, using a scanner with 1600 dpi resolution (CanoScan 8400f; Canon Inc., Tokyo, Japan). Then, using the scanned images, we counted the number of areoles (ar) that occurred in an area of 5 × 5 mm2 on each leaf.

Statistic analysisWe calculated the means, ranges, and

standard deviations of each variable for all specimens (Table 2) and for the two groups recognized in this study (Table 3). We conducted t-tests to compare the measurements and ratios of the two groups, using the computer program Microsoft Excel for Mac 2011 ver. 14.3.9 (Microsoft Co., Redmond, WA) (Table 3).

Results and DiscussionDistinction between the two morphotypes of Japanese Phryma

The results of the measurements of characters and statistical analyses were shown in Tables 2 and 3.

Based on the scatter diagram of Fig. 3, showing the relationships between ul/ulw and lw/ll, we attempted to separate 58 Japanese Phryma specimens into two groups (the asiatic-group, ASG, and the oblongifolia-group, OFG). These two groups were almost clearly separated in the scatter diagrams with ul/ulw and l/llclw (Fig. 4), ll and lw (Fig. 5), lw/ll and ln (ar) (Fig. 6), a/llclw and ln (ar) (Fig. 7). Therefore, we propose to recognize each of the two groups as distinct species.

The comparison of floral and foliar characters between two species, i.e., ASG and OFG, is shown in Table 3. Based on the data, there are significant differences in the values 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, between ASG and OFG (Table 3). The range of the values and ratios were found to overlap between ASG and OFG. On the other hand, the 80% confidence intervals do not overlap between ASG and OFG in the llclw, ln (ar) and, ulw values and the a/llclw, lw/ll, and ul/ulw ratios (Table 3). This indicates that individual characters may be insufficient for discriminating between the two groups, and that using a combination of characters could be necessary for the discrimination.

Morphological relationships between Japanese and Chinese Phryma

The results of the measurements of one Chinese individual in the present study are shown in Table 1. Based on the results, we recognized the following relationships between the two Japanese groups, i.e., ASG and OFG groups, and the Chinese individual.

1) The ratio a/llclw of the Chinese individual is 0.18, and is put in the range of those of ASG.

2) The ul/ulw is 0.58, and is in the range of ASG.

3) The lw/ll is 0.48, and is in the range of OFG.

4) The ln (ar) is 4.02, and is in the range of ASG.


Table 2. Floral and foliar characters of Japanese PhrymaRange 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 Japanese Phryma

CharacterAsiatic-group Oblongifolia-group

p-valueRange 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

ll (mm) 32.30–113.94 66.48 16.40 34.34–

98.6250.40–82.55

39.28–147.39 81.52 26.03 30.51–

132.5356.01–107.02 0.01

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 *

lw (mm) 38.98–103.92 62.61 15.21 32.80–

92.4247.70–77.52

20.04–72.62 41.09 11.43 18.69–

63.4929.89–52.29 *

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 combination of the features of Japanese ASG group and OFG group.

Comparison between the result of the molecular phylogenetic analysis and that of the morphological analysis on the variation of Japanese Phryma

Based on the results of the molecular phylogenetical analysis by Nie et al. (2006),

Japanese Phryma plants could be divided into two monophyletic groups. Similarly, the results of the present morphological analysis indicates that Japanese Phryma plants could be classified into two morphological groups, i.e., ASG and OFG groups. However, the relationships between the molecular phylogenetic groups and the morphological groups are unclear yet. To know the relationships, we need molecular phylogenetic studies on the Japanese


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.


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).


morphological groups.

Taxonomic treatmentFour Phryma taxa have been previously

described in Japan, i.e., P. oblongifolia Koidz., P. nana Koidz., P. humilis Koidz., and P. leptostachya L. var. asiatica Hara (Table 4). Any type specimens of these four taxa were not cited in their original descriptions (Koidzumi 1929, 1939, Hara 1948). Therefore, we designate here the lectotypes for these four taxa from their original materials as shown in Table 4.

In the values of ar, ll, ln (ar), lw, and the lw/ll ratio, the lectotypes and original materials of P. oblongifolia and P. humilis belong to OFG, and the lectotypes of P. nana and P. leptostachya var. asiatica belong to ASG (Figs. 5, 6). Therefore, according to the priority rule of nomenclature, the correct names of OFG and ASG are P. oblongifolia Koidz. and P. nana Koidz., respectively.

Based on the differences of 80% confidence intervals, the two species are distinguished as follows (Table 3):

A. The ratio of the width of the lobes of the upper corolla lip (ul) to the width of the upper

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 length (ll) of the leaf with the longest petiole, i.e., lw/ll, ranges from 0.81 to 1.09 ............... P. nana

A. The ul/ulw ratio ranges from 0.74 to 1.02 and the lw/ll ratio ranges from 0.43 to 0.61 ........ ........................................................ P. oblongifolia

Phryma nana Koidz. in Acta Phytotax. Geobot. 8: 191 (1939). – P. leptostachya L. var. nana (Koidz.) H. Hara, Enum. Sperm. Jap. 1: 297 (1948). – P. leptostachya L. subsp. asiatica (H. Hara) Kitam. f. nana (Koidz.) Akasawa in Bull. Kochi Women’s Univ., Ser. Nat. Sci. 18: 6 (1970). Lectotype (designated here): JAPAN. Honshu. Kyoto Pref., Mt. Hiyei-zan, 12 Aug. 1936, K. Takeuchi s.n. (KYO).

Phryma leptostachya L. var. asiatica H. Hara, Enum. Sperm. Jap. 1: 297 (1948). – P. leptostachya L. subsp. asiatica (H. Hara) Kitam. in Kitam. & Murata in Acta Phytotax. Geobot. 17: 7 (1957). – P. asiatica (H. Hara) O. Deg. & I. Deg. in Phytologia 22: 212 (1971). Lectotype (designated here): JAPAN. Hokkaido. Hidaka Subpref., ‘in forests at the foot of Mt. Apoi’, 6 Aug. 1933, H. Hara s.n. (TI).

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.


Phryma oblongifolia Koidz. in Bot. Mag. (Tokyo) 43: 400 (1929); Ohba, Fl. Chiba: 546 (2003); Y. Endo & al. in J. Jpn. Bot. 89: 406 (2014). – P. leptostachya L. var. oblongifolia (Koidz.) Honda in Bot. Mag. (Tokyo) 50: 608 (1936); H. Hara in J. Fac. Sci. Univ. Tokyo Bot. 6: 377 (1956); & Fl. E. Himal.: 306 (1966); T. Yamaz. in K. Iwats. & al., Fl. Jap. 3a: 322 (1993); T. Miwa & Kigawa, Fl. Kanagawa: 1284 (2001). – P. leptostachya L. var. asiatica H. Hara f. oblongifolia (Koidz.) Ohwi in Bull. Nat. Sci. Mus. no. 33: 86 (1953); & Fl. Jap.: 1075 (1953). Lectotype (designated here): JAPAN. Honshu. ‘Fujiyama’, 31 July 1893, S. Matsuda s.n. (KYO).

Phryma humilis Koidz. in Acta Phytotax. Geobot. 8: 192 (1939). – P. leptostachya L. var. humilis (Koidz.) H. Hara, Enum. Sperm. Jap. 1: 297 (1948). Lectotype (designated here): JAPAN. Honshu. ‘Rikutiu, Iwate-gun, Matsuo-mura’, Y. Fukuta 288 (KYO).

The authors thank curators of the herbaria, KYO, TI, and TNS, for their permission in examining the herbarium specimens, and thank Tomoyuki Nemoto (Ishinomaki Senshu University, Japan), Yu Iokawa (Joetsu University of Education, Japan), and Xiang-Yun Zhu (Academia Sinica, Beijing, China), for their help in collecting materials in China.

This work was partially supported by JSPS KAKENHI Grant numbers 15405013, 26440205 (to Y.E.).

ReferencesChang C. S., Kim H. and Chang K. S. 2014. Provisional

Checklist of Vascular Plants for the Korea Peninsula Flora (KPF) (Version 1.0). The Designpost, Seoul.

Endo Y., Miyauchi T. and Sannohe T. 2014. Morphological differences in flowers between Eastern Asian and Eastern North American Phryma (Phrymaceae). J. Jpn. Bot. 89: 394–408.

Hara H. 1949. Enumeratio Spermatophytarum Japonicarum 1. Iwanami Shoten, Tokyo.

Hara H. 1956. Contributions to the study of variations in the Japanese plants closely related to those of Europe

Table 4. The features of the type specimens of Japanese Phryma plantsScientific name Collection site and collector ll lw lw/ll ar ln (ar) Herb Type

P. oblongifolia Koidz. Ganjusan U. Faurie 2535

103.66 42.11 0.41 33.0 3.5 KYO original material

Nuruyu U. Faurie 789

109.13 40.89 0.37 28.0 3.3 KYO original material

Fujiyama S. Matsuda 31 July 1893

43.46 22.09 0.51 12.5 2.5 KYO lectotype

Hakone - - - - - - not found1)

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

P. nana Koidz. Kyoto, Mt. Hiyeizan T. Takeuchi 12 Aug. 1936

17.40 16.19 0.93 35.5 3.6 KYO lectotype

P. humilis Koidz. Rikutiu: Iwategun, Matsuomura Y. Fukuta 288

31.15 20.81 0.67 77.5 4.4 KYO lectotype

P. leptostachya L. var. asiatica H. Hara

In forests at the foot of Mt. Apoi H. Hara 6 Aug. 19332)

52.07 46.91 0.90 73.0 4.3 TI lectotype

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.


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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型が知られており，これを品種の階級または変種の階級で分類するという考えがある．一方で，種の階級で分類するべきであるとする考えや同一種とする考えもある．本研究では，日本産ハエドクソウ属の形態変異の実態を明らかにし，その結果に基づいた，分類学的な取り扱いの提案を目的とした．　本研究の結果から，ハエドクソウ属の以下に述べる形質に変異を認めた．花では，1) 「上唇弁の裂片部の幅 (ul)」と「上唇弁の中央部の幅 (ulw)」の比 (ul/ulw)，葉では，2) 個体中最長な葉柄を持つ葉の「葉身の幅 (lw)」と「葉身の長さ (ll)」の比 (lw/ll)である．結果として 1) と 2) の違いから，調査した植物群を 2群に分けることができた．つまり，ul/ulwが 0.47–0.59で，かつ lw/llが

0.81–1.09である群と ul/ulwが 0.74–1.02で，かつ lw/llが 0.43–0.61である群である．これら 2群の間では 1) と 2) の特徴以外に，3) 下唇弁中央裂片の中央部の幅，４) 葉裏面の単位面積当たりの小区画数，5) 上唇弁の中央部の幅，6) 上唇弁の裂片の長さと下唇弁の中央裂片の長さの比，についても顕著な違いがあった．以上によ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とを提案する．2群のうち，前者はハエドクソウ（チャボハエドクソウ）Phryma nana Koidz.，後者はナガバハエドクソウ（ヒメハエドクソウ）P. oblongifolia Koidz. に相当する．なお，本研究では同属の日本産分類群の選定基準標本の指定を併せて行った．

（茨城大学理学部）

circumscription of two phryma species (phrymaceae) in japan

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