A Taxonomic Re-Investigation of Papaver Miyabeanum and P. Fauriei (Papaveraceae)
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ISSN 1346-7565 Acta Phytotax. Geobot. 71 (2): 147–155 (2020) doi: 10.18942/apg.201916 A Taxonomic Re-investigation of Papaver miyabeanum and P. fauriei (Papaveraceae) 1,* 2 HIDEKI TAKAHASHI AND MASUMI YAMAGISHI 1The Hokkaido University Museum, Hokkaido University, Kita 10-jo, Nishi 8-chome, Kita-ku, Sapporo, Hokkaido 060-0810, Japan. *[email protected] (author correspondence); 2Research Faculty of Agriculture, Hokkaido University, Kita 9-jo, Nishi 9-chome, Sapporo, Hokkaido 060-8589, Japan Based on the few differences in ITS sequences and outer morphological traits between Papaver miya- beanum of the Kuril Islands and P. fauriei (Papaveraceae) of Rishiri Island, Hokkaido, the two species are treated as conspecific. However, because of the stable differences in ITS sequences, a tendency to- ward smaller dimensions in all morphological features, and the somewhat isolated geographical distribu- tion of P. fauriei, we regard the differences to be recognizable at the subspecific level. We therefore pro- pose the name Papaver fauriei (Fedde) Fedde ex Miyabe et Tatew. subsp. shimshirense (Miyabe et Tatew.) Hideki Takah., comb. & stat. nov. for P. miyabeanum of the Kuril Islands. Key words: ITS, Kuril Islands, Papaver fauriei, Papaver fauriei subsp. shimshirense, Papaver miyabea- num, Rishiri Island Papaver L. (Papaveraceae), with about 70– Tatewaki 1936, Yamagishi et al. 2018), is mor- 80 species, occurs worldwide, but is mainly in the phologically similar to P. miyabeanum, resulting N. Hemisphere (Grey-Wilson 1993, Kadereit in a considerable debate as to whether the two are 1993, Carolan et al. 2006, Mabberley 2008). Pa- conspecific (Shimizu 1983, Eckenwalder 1989, paver is separated into 8 to 11 sections (Kadereit Grey-Wilson 1993, Gardner 1999). 1988, 1993, Grey-Wilson 1993) based on a combi- According to Miyabe and Tatewaki (1935, nation of characters; i.e., habit, leaf vernation, 1936), P. miyabeanum differs from P. fauriei in leaf shape, color of anthers and filaments, mode the mostly pinnatiparted leaves (vs. bipinnati- of capsule dehiscence, and general capsule char- lobed in the latter), broad orbicular leaf lobes acteristics such as size, shape and indumentum. with round, round-orbicular, or corniculate apex Papaver miyabeanum Tatew. and P. fauriei (vs. oblong or oblong-ovate lobes with broadly (Fedde) Fedde ex Miyabe et Tatew. and their re- acute, sometimes mucronulate apex in the latter), lated panarctic-alpine species belong to section larger flowers [(2.5)–3–5 cm vs. 1–2.5 cm in di- Meconella Spach (Kadereit 1988, Carolan et al. ameter when dry], and larger, globose to ovoid 2006, Yamagishi et al. 2010). Papaver miyabea- capsules (1–1.5 cm long and wide vs. 0.7–1 cm num (Fig. 1), an endemic species of the southern long and 0.5–0.8 cm wide). Those traits, especial- to middle Kuril Islands (Miyabe & Tatewaki ly the leaf traits (degree of lobing and shape of the 1936, Barkalov 2009, Takahashi 2015), is a tiny lobes), differ greatly in different habitats within poppy with basal tufts of leaves and solitary sca- the species. pose flowers. The closely related Papaver fauriei Three species of section Meconella have been (Fig. 2), a rare endemic species of Mt. Rishiri (the recognized in the Kuril Islands; P. alboroseum peak 1,721 m above sea level) on Rishiri Island, Hultén, P. microcarpum DC. and P. miyabeanum northwest Hokkaido (Fedde 1909, Miyabe & Tatew. (Barkalov 2009). Papaver miyabeanum is 148 Acta Phytotax. Geobot. Vol. 71 Fig. 1. Papaver miyabeanum [= P. fauriei subsp. shimshi- Fig. 2. Papaver fauriei [= P. fauriei subsp. fauriei] on Mt. rense] on Simushir Island, the Kuril Islands. Rishiri-dake, Rishiri Island. clearly different from Papaver alboroseum (white caule, suggesting a closer or conspecific relation- flowers) and P. microcarpum (long, narrowly ob- ship between P. miyabeanum and P. fauriei. In ovoid capsules) by having yellow flowers and that study, the ITS sequences of 45 individuals of broadly ellipsoidal to globose capsules. P. fauriei, but only 5 individuals of P. miyabea- On Sakhalin, located north of Rishiri Island, num were analyzed. In the present study, we ana- the taxonomy of Papaver has been confused. Pa- lyzed 15 additional specimens of P. miyabeanum, paver on Sakhalin has been treated as comprising and thus 20 individuals from three islands were three species; P. ochotense Tolm. (Miyabe & analyzed in total. Furthermore, with a more in- Tatewaki 1936), P. studenforfii Tolm. (Miyabe & tensive evaluation of the morphological similari- Tatewaki 1936) and P. nudicaule L. (Bezdeleva ties between the two species, we re-investigated 1987, Smirnov 2002), but recent authors recog- the taxonomic relationship between P. miyabea- nize only a single species, P. tolmatschevianum num and P. fauriei. N.S. Pavlova (Barkalov & Taran 2004, Bezdeleva et al. 2006). Papaver tolmatschevianum is char- acterized by having long, narrowly obovoid cap- Materials and Methods sules (Bezdeleva et al. 2006) and is clearly dis- tinct from the P. miyabeanum – P. fauriei asso- ITS sequence ciation. Sequences of the ITS region of 18S–26S nu- Papaver nudicaule L. is a close relative to P. clear ribosomal DNA were used to clarify genetic alboroseum and P. microcarpum in the Kuril Is- closeness between Papaver miyabeanum and P. lands and P. tolmatschevianum in Sakhalin based fauriei. For P. miyabeanum, total DNA was ex- on capsule shape (Takahashi, 2015). Thus, the tracted from dry leaves on herbarium sheets de- opinion of Akiyama (2006), in which P. fauriei is posited in SAPS (20 plants and five native habi- treated as an infraspecific taxon ofP. nudicaule is tats; Table 1, Fig. 3). For P. fauriei, we adopted not accepted here. ITS sequences reported by Yamagishi et al. Yamagishi et al. (2010) showed that nucleo- (2010), in which leaf pieces were taken from 45 tides at three or four positions of the 643 bp (0.5 living plants in three natural habitats on Rishiri or 0.6%) in the internal transcribed spacer (ITS) Island. Methods for total DNA isolation, PCR sequences differed between P. miyabeanum and amplification of ITS regions, and direct sequenc- P. fauriei, but at 15 or 16 positions (2.3 or 2.5%) ing of the PCR products have been described pre- between P. fauriei and P. alboroseum and at 13 viously (Yamagishi et al. 2010). DDBJ accession positions (2.0%) between P. fauriei and P. nudi- numbers of the ITS sequences are shown in the June 2020 TAKAHASHI & YAMAGHISHI — Papaver miyabeanum and P. fauriei 149 legend of Fig. 4. fruiting plants are affixed to each herbarium The ITS sequences of Papaver miyabeanum sheet. All morphological traits were measured on (this study) and those of P. fauriei, P. alborose- each plant. The length / width ratio of the cap- um, P. nudicaule (Yamagishi et al. 2010), P. cro- sules was calculated and compared. The distri- ceum, P. radicatum, P. nudicaule, P. anomalum bution of the measured or calculated values of the and P. alpinum (Carolan et al. 2006) belonging to two species were compared nonparametrically by section Meconella were aligned using the default the Brunner-Munzel test using Bellcurve for Ex- parameters in CLUSTAL X v.2.1 (Larkin et al. cel (Social Survey Research Information Co., 2007). Then, a neighbor-joining (NJ) tree was Ltd.). constructed using PHYLIP 3.6 package (http:// evolution.genetics.washington.edu/phylip.html) and a gene genealogy was estimated using TCS Results and Discussion software v.1.21 (Clement et al. 2000). In some ITS sequences, two nucleotides appeared at sin- ITS sequence gle polymorphic position (nucleotides were The ITS sequences of Papaver fauriei and P. mixed). Because mixed nucleotides are treated as miyabeanum were 643 bp long (Yamagishi et al. missing data in these software packages, genetic 2010). There were four variable sites (Table 2). distance was manually calculated as follows us- One-base substitutions between different ITS re- ing Microsoft Excel: At each polymorphic posi- peats in the same individual were shown as, C + tons, different nucleotide pairs were counted as 1 T, T + G, A + C. Based on the variation in ITS and a pair of mixed nucleotides and single nucle- sequences, we recognized six genotypes in total otide (e.g., T + G versus T) was counted as 0.5. for the two species of Papaver. In P. fauriei, all The sum of the counts per total number of nucleo- 45 individuals showed the same ITS sequence tides was regarded as the distance between the (Yamagishi et al. 2010). We named this genotype two ITS sequences. The matrix of the distances in the ITS region genotype F (Table 2). Among 20 for each pair of ITS sequences was then directly individuals of P. miyabeanum, five different gen- fed into the softwares. otypes were detected (genotypes M1–M5). Yamagishi et al. (2010) reported three se- Measurement of morphological traits quences (genotypes M1, M2 and M5) in P. miya- For Papaver miyabeanum, 22 herbarium sheets beanum. In this study we detected two additional from 16 localities on five of the Kuril Islands sequences in the ITS region (genotypes M3 & were used (Table 1). For P. fauriei, 19 herbarium M4) for P. miyabeanum (Table 2). Among the ITS sheets from 8 collections growing on the gravelly sequences of four species of Papaver; P. fauriei, alpine slopes of Mt. Rishiri, Rishiri Island, were P. miyabeanum, P. alboroseum and P. nudicaule, used (Table 1). Two herbarium sheets of cultivat- the thymine at position 498 was determined to be ed plants of P. fauriei (SAPS 016122, 016121 in unique to P. fauriei in the study by Yamagishi et Table 1) were also compared with the native al. (2010). In the present study, one sample (SAPS plants. To compare the morphology of the two 042355-1: genotype M4) of P. miyabeanum from species taxonomically, only measurements from Iturup showed mixed nucleotides C + T at posi- herbarium sheets of native plants were used.