Variability of the Cones of Picea Jezoensis and P. Koraiensis (Pinaceae) in North Korea

Fragm. Flor. Geobot. 37(1): 241–249, 1992

Variability of the cones of Picea jezoensis and P. koraiensis (Pinaceae) in North Korea

JERZY STASZKIEWICZ

STASZKIEWICZ, J. 1992. Variability of the cones of Picea jezoensis and P. koraiensis (Pinaceae) in North Korea. Fragmenta Floristica et Geobotanica. 37(1): 241–249. Kraków. PL ISSN 0015– 931x.

ABSTRACT: In the present paper variability of the cones of Picea jezoensis (Sieb. & Zucc.) Carr. and P. k o r a i e n s i s Nakai from North Korea on the basis of biometric studies is characterized. It is shown that the differences between the features of the cones of both species are remarkable and this confirms their separate taxonomic status.

KEY WORDS: Picea, cones, variability, biometry, North Korea, East Asia

J. Staszkiewicz, Department of Plant Variability, W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, PL–31–512 Kraków, Poland

INTRODUCTION

The wide distribution of the genus Picea A. Dietr., its marked taxonomic diversity, and considerable historical age provoke the great interest of researchers in particular species which compose the genus. Apart from the classic taxonomic studies, numerous morphological and biometrical studies on different organs are carried out. Much of the research deals with variability of morphological features which often are of taxonomic importance. During his stay in the Korean People’s Republic, the author collected some population samples of the cones of two local species of Picea, namely P. j e z o e n s i s (Sieb. & Zucc.) Carr. and P. koraiensis Nakai. The samples were collected in order to analyse the metric features of the cones and to determine the differences and similarities between the local samples, both within and between the species. It is very important to study variability of the cones because of the geographical position of both species in relation to other taxa of the genus Picea occurring in the borreal and subboreal zones of Eurasia. These species occupy the easternmost, peripheral part of the Asian mainland (Fig. 1). Some interesting relations occur between P. k o r a i e n - sis and P. obovata Ledeb. which, undoubtedly, are closely related. The last problem, however, is going to be presented in a separate work by the present author. Picea koraiensis was described by Nakai relatively lately, i.e, as late as 1919. How- ever, since then its separateness has not been questioned. P. jezoensis described on the

16* 242 Fragm. Flor. Geobot. Ann. 37, Pars 1, 1992

Fig. 1. Geographical range of P. jezoensis (Sieb. & Zucc.) Carr. and P. koraiensis Nakai.

Fig. 2. Distribution of P. jezoensis (Sieb. & Zucc.) Carr. in North Korea. J. Staszkiewicz: Biometric study on Picea jezoensis and P. koraiensis 243

Fig. 3. Distribution of P. koraiensis Nakai in North Korea. basis of a specimen growing in the emperor’s garden in Tokyo is considered by some researchers merely as a garden form (Bobrov 1970) and is also known as P. ajanensis (Lind. & Gord.) Fisch. ex Carr. Some researchers include P. hondonensis Mayr from Honshu in P. j e z o e n s i s in the rank of variety (Rehder 1956; Horikawa 1972). The geographical range of P. koraiensis is not too wide and it is restricted to the northern part of North Korea (Fig. 3) and the Primorye Territory in the Russian Far East (Fig. 1). The distribution of P. jezoensis (without P. hondonensis) is much wider (Figs 1–2). In Russia alone it includes territories situated at the southern shores of the Sea of Okhotsk, basin of Amur River, the Primorye, Sakhalin, central Kamchatka, and the southern part of the Kurile Islands. Additionally, it covers a small patch of north-eastern China, Hokkaido in Japan and the Korean Peninsula, where the southern limit of its distribution reaches slightly further north of lat. 38oN. Picea koraiensis is placed in the type subgenus and section of Picea which includes also P. obovata and P. a b i e s (L.) Karst., while P. jezoensis is included in subgenus and section Casicta Mayr (Bobrov 1970) or in section Sitcha Schmidt (Schmidt 1989). Ac- cording to the latter author P. jezoensis is a more primitive form than P. koraiensis. In spite of the placement in different subgenera, both species very easily hybridize (Bobrov 1970). The specimens recorded from Korea as separate species, namely P. t o - naiensis Nakai, P. pungsanensis Uyeki ex Nakai and P. intercedens Nakai, all originated in the result of hybridization. The same origin was ascribed by Bobrov (1970, 1974) to P. komarovii Vassil. The hybrid between P. koraiensis and P. obovata was described as P. m a n d s h u r i c a Nakai. 244 Fragm. Flor. Geobot. Ann. 37, Pars 1, 1992

MATERIAL AND METHODS

Material for the present study is limited to four samples of cones of P. jezoensis and to three samples of P. koraiensis. The cones were collected at random, from 30 to 50 from each locality, and were examined in respect of the following nine features: A – length of cone; B – width of cone; C – length of scale; D – width of scale; E – length of upper part of scale; F – length/width ratio in cone; G – length/width ratio in scale; H – ratio of length of the upper part of scale to width of scale; I – ratio of length of the upper part of scale to length of scale in per cents. The way of measuring of scale is in Fig. 4. Charac- teristic values for the cone features of both species are given in Tab. 1. In order to show up the differences and similarity between all samples the graphical method of Jentys-Szaferowa (1959) is applied.

Table 1. Arithmetic means of features A–I of Picea jezoensis and P. koraiensis

Picea jezoensis Picea koraiensis

Features General local samples General local samples sample sample 1367 245 A 38.91 41.54 40.81 36.37 36.93 58.15 62.00 61.17 54.53 B 15.12 15.04 16.17 14.13 15.15 19.08 10.07 19.67 18.87 C 11.63 11.56 12.33 10.83 11.80 15.59 16.42 15.79 14.68 D 7.91 7.67 8.29 7.56 8.12 13.62 13.86 14.12 13.17 E 6.27 5.68 6.96 5.90 6.67 5.48 5.55 5.70 5.34 F 2.57 2.74 2.52 2.57 2.44 3.05 3.18 3.13 2.90 G 1.47 1.50 1.49 1.42 1.45 1.14 1.19 1.11 1.11 H 0.80 0.74 0.84 0.78 0.82 0.41 0.40 0.40 0.42 I 54.32 49.40 56.50 54.77 56.50 35.97 34.06 36.17 37.80

The material was collected by the author, unless otherwise stated, from the following stations (Fig. 5): A. PAEKDU-SAN PLATEAU. 1. A small mixed wood at the road between the settlements of Poso-ri and Potae, elev. ca. 800 m; 2. Larch–spruce taiga around the lake, north of Samjiyon, elev. ca. 1200 m; 3. Larch taiga with a slight admixture of spruce, north–east of Taehongdan, elev. ca. 1300 m; 4. Larch taiga with admixture of Abies nephrolepis, coll. W. Wojewoda; 5. Spruce taiga with admixture of Larix olgensis var. koreana, east slopes of Paekdu, elev. ca. 1600 m; 6. Larch taiga on the border of the upper timberline of the forest, east slopes of Paekdu, elev. ca. 2000 m. B. MYOHYANG–SAN MOUNTAINS. 7. Mixed forests on slopes of Mt. Woman, with Pinus koraiensis, Abies holophylla, Betula ermanii and Sorbus commixta, elev. ca. 1400 m.

VARIABILITY OF THE FEATURES OF THE CONES AND DIFFERENTIATION OF THE POPULATION P. JEZOENSIS

Variability of the characters A–E of the cones of P. j e z o e n s i s is presented in Fig. 6. These are small cones, 30 to 50 mm long, rarely longer. The most frequent cones, however, are 33 to 45 mm long. The width of the cones varies from 13 to 17 mm, rarely to 19 mm. Very small are also seed-scales and their length is 8 to 15 mm. The most frequent scales, how- J. Staszkiewicz: Biometric study on Picea jezoensis and P. koraiensis 245

Fig. 4. The way of measuring of the scales. C – the length of a scale, E – the length of the upper part of a scale and D – the width of a scale. ever, are 10.3 to 12.9 mm long. The width of cones does not exceed 6 to 10 mm. Yet, the majority of the scales is 8 mm long. The upper part of the scale is 4 to 8 mm wide. In Fig. 7 the differences between the local samples in relation to the general sample are given. In addition to this, apart from the features of size A–E, the features of the shape were also considered. There occur slight differences between the local samples, which are exemplified by a remarkable similarity of the sample 7 collected in the

Fig. 5. Location of the collectiong stations (Arabic numerals correspond to the list on page 244). 246 Fragm. Flor. Geobot. Ann. 37, Pars 1, 1992

Fig. 6. Frequency diagrams of features A–E of P. jezoens is & Zucc. Carr. (Pj) and P. koraiens is Nakai (Pk). A – length of the cone, B – width of the cone, C – length of scale, D – width of the scale and E – length of upper part of the scale.

Myohyang-san Mountains to the sample 6 collected from the trees occurring at the upper mountain belt limit in the Paekdu-san Plateau, where the habitat conditions are severe.

VARIABILITY OF THE FEATURES OF THE CONES AND DIFFERENCES BETWEEN LOCAL SAMPLES OF P. KORAIENSIS

The cones and seed-scales of P. koraiensis are markedly bigger than those in the case of the former species (Fig. 6). The length of the cones is 40 to 85 mm and the modal length is 55 mm. The width of the cones is also varied, namely from 15 to 25 mm, but 66% of all cones is 17.1 to 21.0 mm wide. The length of the scale does not exceed 11 to 20 mm, most often it is 13.6 to 15.5 mm, whereas the width varies from 10.0 to 18.0 mm. If the standard deviation is considered, the majority of the scales is contained in the J. Staszkiewicz: Biometric study on Picea jezoensis and P. koraiensis 247

Fig. 7. Lines of size and shape of the local samples (angular lines) compared with the general sample of P. jez oensis (Sieb. & Zucc.) Carr. (vertical lines). Features A–I as on page 244.

Fig. 8. Lines of size and shape of the local samples (angular lines) compared with the general sample of P. koraiensis Nakai (vertical lines). Features A–I as in page 244. interval 12.1 to 15.1 mm. The minimum and maximum length of the upper part of the scale is 4 mm and 5.1 mm respectively. The differentiation of local samples is fairly small (Fig. 8). Only in the sample 5, collected in the upper timberline of the forest, the cones and seed-scales were slightly shorter and narrower, however, these differences are contained within the limit of error. 248 Fragm. Flor. Geobot. Ann. 37, Pars 1, 1992

Fig. 9. Line of size and shape of the general sample of P. jezoens is (Sieb. & Zucc.) Carr. (angular line) compared with the general sample of P. koraiensis Nakai (vertical line). Features A–I as in page 244.

DIFFERENCES BETWEEN PICEA JEZOENSIS AND P. KORAIENSIS

The differences between the features of both species are remarkable, which is evidenced by the polygons of the features of variability given in Fig 6. Although variability ranges of their features are partly similar, nevertheless, when the law “2 SIGMA” is applied, they exclude themselves. Erroneous determination of the species based on the cones is im- possible, since their seed-scales are built along a different pattern. The scales of P. j e z o e n - sis have a better developed upper part, frequently delicately serrate, while the scales of P. koraiensis are short and even bordered. Schmidt (1989) includes the cones of P. jezoensis in the morphological type of Casicta, while the cones of P. koraiensis to the type Morinda. The diagram given in Fig. 9 confirms the marked separateness of the cones of P. k o r a i e n - sis from those of P. j e z o e n s i s both in the features of size and shape. The differences are the more convincing if the diagrams in Figs 7 and 8 are considered, which confirm a small differentiation between the local samples of each species.

REFERENCES

BOBROV J. G. 1970. Istoriya i sistematika roda Picea A. Dietr. (Generis Piceo historia et systematica). – Nov. Sist. Vyzsh. Rast. 7: 5–40 (in Russian).

BOBROV J. G. 1974. Introgresivnaya gibridizatsya v rode Picea A. Dietr. (“Introgressive hybridization in the genus Picea A. Dietr.”) – In: Teoreticheskiye osnovy vnutrividovoy izmenchivosti i struktura populyatsyi khvojnykh porod, pp. 60–66. Uralskiy Nauchnyy Centr, Sverdlovsk (in Russian).

HORIKAWA Y. 1972. Atlas of the Japanese flora and introduction to plant sociology of East Asia. 1. 500 + VIII pp. Gakken Co., Tokyo. J. Staszkiewicz: Biometric study on Picea jezoensis and P. koraiensis 249

JENTYS-SZAFEROWA J. 1959. A graphical method of comparing the shapes of plants. – Rev. Pol. Acad. Sc. 4(1): 9–38 REHDER A. 1956. Manual of cultivated trees and shrubs. Ed. 2. 996 pp. Macmillan Co., New York. SCHMIDT P. A. 1989. Beitrag zur Systematik und Evolution der Gattung Pic ea A. Dietr. – Flora 182: 435–461

STRESZCZENIE

W pracy podano charakterystyke˛ biometryczna˛ dwóch korean´skich gatunków s´wierka: P. jezoensis (Sieb. & Zucc.) Carr. i P. koraiensis Nakai. Gatunki te zajmuja˛ najbardziej wschodnia˛, peryferyczna˛ cze˛sć´ kontynentu azjatyckiego. P. koraiensis jest niewa˛tpliwie blisko spokrewniony z P. obovata Ledeb., który charakteryzuje sie˛ bardzo szerokim zasie˛giem. P. koraiensis opisany został stosunkowo pózńo, bo dopiero w 1919 roku. P. j e z o e n s i s opisany na podstawie osobnika rosna˛cego w cesarskim ogrodzie w Tokio i uwaz˙any przez niektórych badaczy tylko za forme˛ ogrodowa˛, znany jest takz˙e pod nazwa˛ P. a j a n e n s i s . Ogólne zasie˛gi, jak równiez˙ ich wyste˛powanie w Korei przedstawiono na Ryc. 1–3. P. koraiensis umieszczany jest w ramach podrodzaju typowego, zas´ P. j e z o e n s i s nalez˙y do podrodzaju Casicta Mayr. Badania przeprowadzono na próbach losowych (Ryc. 3). Analizowano: A – długosć´ szyszki; B – sze- rokosć´ szyszki; C – długosć´ łuski; D – szerokosć´ łuski; E – długosć´ górnej cze˛sći łuski; F – stosunek długosći łuski do jej szerokosći; G – stosunek długosći łuski do jej szerokosći; H – stosunek długosći górnej cze˛sći łuski do jej szerokosći; I – długosć´ górnej cze˛sći łuski w procentach długosći łuski. Po- równanie prób przeprowadzono przy pomocy metody graficznej Jentys-Szaferowej (1959). Zakres zmiennosći cech P. jezoensis jest róz˙ny od zakresu P. koraiensis, a przy uwzgle˛dnieniu prawa „2 SIGMA”, cze˛sto zupełnie sie˛ wyklucza (Ryc. 6 i 9). Wynika to z faktu, z˙e szyszki obu gatunków zbudowane sa˛ według róz˙nego planu. Róz˙nice pomie˛dzy próbami lokalnymi kaz˙dego z gatunków sa˛ bardzo małe (Ryc. 7 i 8), co s´wiadczy o ich duz˙ej jednorodnosći genetycznej.