The Classification (370-285B, C.), (1583)

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A New Proposal for the Classification of the Genus Picea

Tang-shui Liu*

?ti M ag : FVtEOrafiX A

I. Introduction

Before proceeding to the main subject of this paper, it is interesting to mention the history of the name Picea in its use to denote conifer trees, Thc name Piaea is a classical word derived from the Greek word `Wittys", appeared in the works by THEopHRAsTus (370-285 B, C.), "IVbvi In 1536, Brunfels OTTo published a work entitled Nbrbarii". The treatise `CRhapsDdia CLII" was mainly devoted to pines, in it six genera, namely Pinus, Pinaster, Abies, Picea, Lan'x and TZiedo were listed. Thus Picea was mentioned and formally had a share in the conifers. Since then, well-known botanists such as A. CAEsALpiNo (1583), the BAuH!N brothers (Jean BAuHiN, 1650 & Kaspar BAumN, 1623), andJohn RAy (1688) successively foIlowed to cite the name Picea in their monumental works.

{`Institutiones TouRNEFoRT,J, P, in his work entitled Rei Hlarbariae;' (1719) on page 585, used the name Abies in a broad sense to include all spruces, firs and hemlocks. He did

not follow suit to cite Picea to denote only the trees designated by the above authors･

"opecies These works were followed by LiNNAEus, who also listed the name Abies in Plantarum" (1753), volume II, page 1,OOO under the genus Pinus to denote spruces, firs and hemlocks. One year later, in 1754, in the fifth edition of"Genera Plantarum", on page 434, he included spruces, hemlocks, firs, pines, cedars and larches under the name ofPinus, Abies and Larix. However, in l824, A. DiETRicH formally used Picea in place of Abies

to denote all spruces.

Since then, most of the botanists fbllowed the decision of DiETRicH to use Pieea to include spruces in their works, Exceptions were J, C. LouDoN (l838), G. GoRDoN (1858), and K. KocH (1875), who took the names ofAbies for spruces and Picea fbr firs in contrast to DIETRIcH.

II. PreviousWorks

In the family Pinaceae, Picea is a more complex genus than the above-mentioned

* Department of Forestry, College of Agriculture, National Taiwan University, Taipei, Taiwan, Republic of China.

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Abies, Picea is very rich in number of species, being second to Abies or third to Pinus. Morphologically, it shows a great variance among species. The variation ofthe species of Picea is probably caused by the geographical range, ecological habitat and natural

cross-pollinatien, because of their very wide range in the temperate and subarctic regions

ofthe Northern Hemisphere. Many oftheir species are found growing in association with one, two or even more other species of the same genus. Thus, there is a great chance

for two spruce trees to cross, and reproduce intermediates with characteristics which are

botanically not easily identifiable.

In spite ofthe $pecific complexity ofPicea, botanists have successively tried to make subdivisiDns of the genus and to put all the spruces into an orderly arrangement. In 1887, M, WiLLKoMM noticed the shape of needles and first divided the genus inte the

fo11owing two sections : Sect. I. Empicea WiLLK, Needles quadranguIar, with stomata on all fbur surfaces;

with 5 species.

Sect. II. Omorika WiLLK. Needles fiat, with stomata usually on upper surface, lower side greens with 2 species. This subdivision was fbllowed by H. MAyR (l890), who, based on the structure of cones, the shape ofneedle cross-sections, and the presence ofstomata on surfaces ofleaves, proposed two new subdivisions, i.e. Morindd and Casicta, to the section Omorika made by

WiLLKoMM, The three sections set up by MAyR are as follows:

Sect. I. Mon'nda MAyR Needles rhomboicl, stomata on all sides; cone-scales hard

and closed; with7species. . Sect. II. thsicta MAyR Needles flat, stomata on upper surface; cone-scales soft and

loose; with 4 species.

Sect. III. Omorika WiLLK. Needles broad, rhomboid, stomata on lower surface;

cene-scales hard and closed till mature; with 4 species. Based on the shape and structure of needles, buds and cone-scales, M. LAaAssAGNE, in I934, divided Picea into 2 sections as WiLLKoMM did. He classified each section into

phyla according to shape, length of cones and presence or absence of vascular bundles.

These are as follews :

Sect. I. Empicea WiLLK. Phylum A. Cones long, tapering; vascular bundle ciearly present; with 2I species. Phylum B. Cones short, ovate; vascular bundle not clearly present; with 6 species. Sect, II. Omon'ka Wi-K.

Phylum C, Cone-scales orbicular; vascular bundle clearly present; needles slightly

flattened; with 3 species.

Phylum D. Cone-scales awl-shaped; vascular bundle not clear, needles flattened;

with 12 species. Since then, well-known botanists or dendrologists such as R. PiLGER (1960), E. F. DEBAzAG (1964), H. GAussEN (1966), S. G. HARRisoN (l966), C. CoLLEAu (1968) and

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C. KRtissMANN (1972) did a great deal towards the classification of the genus. They fbllowed MAyR to make subdivision of Picea. Instead of using Mon'ncla by MAyR, they

restored WiLLKoMM's Ezipicea as'one ef the 3 sections. Of the above authors, especia}ly,

GAussEN included all spruces in the three sections, Ezipicea, C]Zisicta and Omorika, basing the division as he theught fit on botanical features, as well as geographical regions ofthe species. Based on the size of pollen grains and other morphological and anatomical characters, GAussEN made a phylogenetical arrangement of all the spruce species. In addition to WiLLKoMM's Ezipicea and Omorika, E. F. DEBAzAG, in l964, established Bicolor . as a new sectlon. In 1970, BoBRov made an exce!lent research of the spruces. According to the International Code of Botanical Nomenclature, he adopted Picea in place of Etipicea as a new section to the sections Omorika by WiLLKoMM and Casiota by MAyR; he also added

series to include the species. Following is BoBRov's subdivision ofPicea.

Sect. 1. 0morika WiLLK., inclucles 5 species, Sect. 2. Picea (instead ofEtipicea) Ser. 1. 0bovatae BoBR., includes 5 species. Ser. 2. Asperatae BoBR,, includes 5 species. Ser. 3. Politae BeBR., includes 3 species. Ser. 4. Glehnianae BoBR., includes 4 species. Ser. 5. Glaucae BoBR,, includes 3 species. Ser. 6. Rubentes BoBR., includes 2 species. Sect. 3. thsictaMAyR

Ser. 1. t

III. A New Proposal for the Subdivision of the Genus Picea

Based on the shape and texture of cone-scales, the shape and structure ef leaves, the color and presence or absence of pubescences of shoots with characteristic buds, the

fo11ewing classification ofPicea may be more suitable. As far as the species are concerned,

sorne changes, i.e. combination and reduction ofspecies, have been made.

Picea A, DiETR.

A. DiETR., Fl. Geg. Berl. 2: 794, 1824; LiNK in Abh. Akad. Wiss. Berlin 1827: 18I (Fam. Pinus. 1827) 1830; MAyR, Monogr. Abiet. 43, 1890; PiLG. in ENGL. u. PRANTL,

Nat. Pflanzenfam. 13: 32I, 1926; LAcAss., Trav. Lab. For. Toul. 2: 3, l:l, i934; REHD.,

Bibl. Cult. Tr. Shn 19-28, 1949; KRtissM., Nadelg. 184, 1960; GAuss., Gymn. 8: 568, 1966; DALL. & JAcKs,, Handb. Coni£ 327, 1966; BoBR., Gen. Pieea Hist, Syst. in Nov. Syst. Pl. Vasc. 7: 7-39, Nauka Leningr. 1970; ScHMmT-VoGT,, Fichte I. 1977. Pinus LiNN., Sp. Pl, 1000, l753; Gen. PI. ed. 5, 484, l754, p. p. Abies MiLL., Gard. Dict. abridg. ed. 4,

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1: 1754, p.p.; Gard. Dict ed. 8. 1768, p.p.; Pom., Encycl. Meth. Bot. 6: 509, 1804, p.p. Pinus-Abies WEsT., Bot. Univ. 1: 210, 1770, p.p. Pinus II. Abies MuNcHH., Hausvat. 5: 222, 1770; SwEET, Hort. Brit., ed. 2, 475, 1830. Abies g I, lllaicea DC. in LAM. & DC.,

Fl. Fanc. ed. 3, 3: 275, 1805; Kock, Syn. Fl. Germ. Helv. 667, 1837. Peuce L. C. RiaH. in Ann. Mus. Hist. Nat Paris 16: 298, 1810, p.p. Piius sect. Abies D. DoN in LAMB., Descr. Pinus 3: sub P. pindeow p. (2) 1837, exclud. sp. nonnull. Abies D. DoN ex LouD., Arb. Brit. 4: 2329, l838. Abies sect. I. Picea SpAcH, Hist. Nat. Veg. Phan. 11; 405 1842; K. KocH, Dendr. 2, 2: 232, l873. Pinus sect. IV. Picea (LiNK) KoaH, Syn. Fl. Germ. Helv. ed. 2, 2: 769, l8tft+; ENDL., Syn. Coni£ l12, 1847; PARL. in D(]., Prodr. 16, 2: 413, 1868.

Abies sect. I. Vlara GoRD., Pinet 2, 1858. Vbitchia LiNDL. in Gard. Chron. 1861: 265, 1861, 1874. quoad sp. Piceaj'ezoensis. Picea subgen. Picea BERTR. in Ann. Sci. Nat. ser. 5, 2Q: 83, Lectotypus: Picea abies (L.) KARsT. (Pinus abies L!NN.) Subgen, I. Omorika (wrLLK.) Liu, stat. nov. Gemmae, perulis subulatis saepe extrinsecus obtectae; folia cornplanata vel plus minusve quadrangulata; squamae strobilorum crassae, rigidae, latiusculae, apice rotundatae et saepe integrae, ad fructus rpaturitaterri arcte appressae. Typus: Picea omorika (PANd.) PuRK. Winter buds often with awl-shaped scales outside; leaves flattened or somewhat quadrangular ; cone-scales thick, rigid, rather broad, rounded and often entire at margins, closely appressed before fruit maturity. Based on the shape of leaves and the color and presence or absence of pubescences on shoot bearing characteristic buds, two sections and three series are to be divided.

They are as fbllows: Sect. I. Omorika WiLLK,, emend. Liu WiLLK., Forstl. FI. ed. 2, 66, 1887; MAyR, Monogr. Abiet. Jap. 44, 1890; LAaAss., Trav. Lab. For. Toul. 2: 272, 1934; REHD., Bibl. Cult. Tr. Shr. 28, 1949; GAuss., Gymn. 609, 1966; DALL. & JAaKs., Handb. Conif, Ginkg. 333, 1966; CoLL., Anat. Comp. Feuill. Pieea 1968; BoBR., Gen, Picea Hist. Syst. 27, I970; KRtissM., Handb. Nadelg. 1972. Folia complanata, utrinque carinata, superne non nisi stomatifera, subtus viridia; ramuli juniores pallido- vel luteolo-brunnei, pubescentes vel glabri; gemmae ovoideo-

comcae, ereslnosae.

Typus: P. omorika (PANd.) PuRK. Leaves flattened, keeled on both sides, stomata only on upper surface, lower side green; young shoots pale or yellowish brown, often pubescent; buds ovoid-conical, not reslnous.

The fo11owing species are included in this section. 1. Picea omorika (PAN6.) PuRK, in Oester. Monatschr. Forstw. 27: 446, 1887. PintLs omorika PANd,, Neue Conif. Oestl. Alp. 4, 1876. Abies omorika NyM., Consp. Fl.

Eur. 673, l881.

The Servian or Serbian spruce is a native of Yugoslavia where it grows on limestone rocks at 800-1,600 m altitude on both sides of the river Drina.

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2. Piea breweriana S. WATs. in Proc. Am. Acad, Arts Sci. 20: 378, l885. PiaeaPenduta S. WATs., Bot, Cali£ 2: 122, 1880, nom. subnud. Pinus breweriana (S. WATs.) Voss in Mitt. Deutsch. Dendr, Ges. 1907 (16) : 93, 1908. Brewer's spruce is a rather rare tree in thc wild state, being confined to a few isolated IQcalities at 1,200-2,300m altitude in the Siskiyou Mountains ofN. W. California and

S. W, Oregon in the United States,

3. Picea brachytyla (FRANcH,) PRiTz. in Bot. Jahrb, 29: 216, 1900. Abies braclly4yla FRANcH. in Journ. Bot. 13: 258, 1899, exclud. spec, DELAvAy. Picea ofanensis sensu MAsT. in Journ. Linn. Soc. Lond. Bot. 26 : 553, 1902, non FiscH, ex CARR,, I885. P. alcockiana sensu MAsT. inJourn. Bot, 41 : 269, 1903, non CARR., 1867. P. Pacityctado PATsaHKE in Bot. Jahrb. 47: 630, 1913. P. sa7gentiana ItsHD. et WiLs. in SARG., Pl. Wils. 2: 35, 1914.

3-1. Picea brachytyla (FRANc}i.) PRiTz, var. brachytyla 3-2. Picea brachytyla (FRANcH.) PRiTz. var. cornplanata CMAsT.) CEiENG cx

Hu in Proc. Fifth Pacif. Sci, Congr. 4: 3277, 1934.

Picea eomplanata MAsT. in Gard. Chron. ser. 3, 39: 146, E 57, 1906. 3-3. Picea brachytyla (FRANcH.) PRiTz. var. rhombisquamea STApF in Bot. Mag. 148: t 8969, 1922. Picea ascendens PATscHKE in Bot. Jahr. 47: 630, 1913. This Chinese wceping spruce occurs over an extensive area in W. Hupei and W,

Szechuan, at 2,600-3,100m altitude.

4. Picea spinulosa (GRiFF.) HENRy in Gard, Chron. ser. 3, 39: 219, 1906. Abies spinulosa GRiFF,, Journ. Trav., 259, 1847. Picea alcockiana CARR. var. morindbides MoTT., Conif, Taxac. 273, 1902, nom. nud. P. morindbicles REi{D. in SARG., Tr. Shr. 1:

95, t 48, 1903.

The East Himalayan spruce is a native of E. Himalaya (Sikkim, Bhutan), occurring at 2,400-3,OOO m altitude.

Sect. II. Morinda MAyR, emend. Liu MAyR, Monogr. Abiet. 44, 1890, Picea II. Morindae AscH. et GRAEBN., Syn. Mitt. Fl. 1: l96, 1897. Ramuli juniores pallide brunnei; fblia sectione transversa quadrangulata, undique stomatifera; squamae strobilorum !ate obovatae, basi attenuatae.

Typus: Picea smithiana (WALL.) Boiss. (Picea morindn LiNK). Young branchlets pale brown; leaves quadrangular in cross-section, stomata on all

sides; cone-scales broadly obovate, narrowed at base.

Ser. 1. 0rientales Liu, ser. nov, Folia 6-10 mm longa, plus minusve quadrangulata, utrinque stomatifera; ramuli juniores dense glanduloso-pubescentes; gemmae conicae, eresinosae. Typus: Picea on'entalis CLiNN.) LiNK Leaves 6-1O mm long, more or less quadrangular, with stomata on both sides; young

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branchlets glandular-pubescents buds conical, not resinous. 5. Picea orientalis (LiNN.) LiNK in Linnaea 20: 294, 1847. Pinus erientalis LiNN., Sp. Pl. ed. 2, 1421, 1763. Pinus Abies on'entalis MtiNcHH., Hausvat. 5: 225, 1770. Pinas-Abies orientatis WEsT., Bot. Univ. 1: 21I, 1770. Abies erientalis (LiNN.) Pom., Encycl. Meth. Bot. 6: 518. Picea wittmanniana CARR., Trait6 ConiL 260, 1855; ed. 2. 325, 1867, pro syn. The oriental spruce occurs in mountains ofN. E. Anatolia (Turkey) and Caucasus (USSR), at altitudes from nearly sea-level up to 2,400 m, Ser. 2. Glehnianae BoBR., descr. emend, Liu BoBR., Gen. Picea Hist. Syst. 33, 1970. Folia 6-12 mm longa, quadrangulata, undique stomatifera; ramuli juveniles fiavido- rufescentes vel rufi, dense breviteroque pubescentes; gemmae ovoideo-conicae resinosae. MAsT. Typus: Picea glehnii (Fr. ScHMiDT) Leaves 6-12 mm long, quadrangular, stornata on all sides; young branchlets yellow-

ish-red or red, densely short-pubescent; buds ovoid-conical, resinous.

6. Picea ScHMiDT) ]xuNsT. in Gard. Chron. n. ser. 13: 300, fig. 54, l880. glehnii (Fr. '176, Abies glehnii Fr. ScHMiDT in Mem. Acad. Sci. St. P6tersb. ser. 7, 12, 2: t. 4, fig. Dendr. 8-!2 CReise Amur Saghal.) 1868. Pimas glehnii (MAsT.) Voss in Mitt. Deutsch. Ges. 1907 (16): 93, 1908. Holckaido Saghalin spruce has its distribution in S. Saghalin, the southern Kuriles, and N. Honshu (Pre£ Iwate) ofJapan, with altitudes varying from nearly sea level up to

1 ,600 m. 7. Picea koyamae SHiRAs, in Bot. Mag. Tokyo 27:(128) Pl. II:28-35, 1913. Picea moramomi HoRT. ex REHD. in BAiLEy, Cult. Evergr. 280, I923, pro syn. Found on Mt. Yatsuga-dake at 1,500-2,OOO m altitude in central Honshu ofJapan. 8. Picea maximowiczii REGEL ex MAsT. in Gard. Chron. 13: 363, l880; REGEL,

Ind. Sem. Hort. P6trop. 33, 1865, nom. nud.

Abies obovataj'aponica MAxiM., Ind. Sem. Hort. Petrop, 1, 3, 1866. Picea tschoneskii

MAyR, Fremdl. Parkb. 339, 1906.

8-1. Picea maximowiczii REGEL ex MAsT. var. maximowiczii

8-2. Picea maximowiczii REGEL ex MAsT. var. senanensis HAyAsHJ

ThisJapanese bush spruce is fbund in a restricted area, that is, on Mts. Fuii-san and

Yatsuga-dake in Honshu, ranging between l,100 and 2,OOO m, often situated above the

forest of ITagus crenata BL,

9. Picea koraiensis NAK. in Bot. Mag. Tokyo 33: 195, 1919. Picea obovata sensu KoM. in Acta Hort. P6trop. 20: 195, 1900, p.p. non LEDEB. Picea kqyamai sensu WiLs. in Journ. Arn. Arb. 1(1): 38, l919, non SmRAs. Picea Putagsanetzsis UyEKi in Rep. For. Exp. Stat. Chosen no. 4, Cor. Timp. Tr. 98-!Ol cum t. 1. 1926. Pieea kayamai SHiRAs. var. koraiensis (NAK.) Liou et WANG, 1969. 9-1. Picea koraiensis NAK. var. koraiensis

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9-2. Picea koraiensis NAK. var. intercedens (NAK.) T, LEE, Ill. Woody Pl. Kor. 5, l966, Picea intercedetzs NAK. in Journ. Jap. Bot. 17; 4, pl. 4, 1941. Picea tonaiensis NAK., 1.c., 1. pl. 1. Picea koraiensis NAK. var. tonaiensis (NAK.) T. LEE, 1.c. The range of the Korea spruce extends from N. Kerea, northward crossing Chang- Pai-Shan through Hsiao-Hing-An-Ling of Northeastern China cManchuria), far to the mountains of Ussuri in USSR. In N. E. China, it grows at an altitude of 400-1,800 m. Ser. 3. Smithianae Liu, ser. nov. Folia mm 2040 longa, quadrangulata, undique stomatifera ; ramuli juveniles glabri, cinerascentes, nitentes; gemmae ovoideae, eresinosae. Typus: Picea smithiana (WALL.) Boiss. Leaves 20-40 mm long, quadrangular, with stornata on all sides; young branchlets greyish, shining, without hairs; buds ovoid, without resin. IO. Picea smithiana (WALL,) Boiss., Fl. Or. 5: 700, 1884. PinzLs smithiana WALL., Pl. As. Rar. 3: 24, t. 246, 1832. Pinzas smithiana LAMB., Descr.

Pinus t. 88, 1837, Picea morinda LiNK in Linnaea 15: 522, 1841. Pinus khutrote) RoyLE, Ill. Bot. Himal. 350, 353, t. 84, fig. 1, 1839. Picea khutrotv (RoyLE) CARR,, Trait6 Coni£ 248, 1855. is This a native of W. Himalaya, extending from Afghanistan to central Ncpal, at heights of 2,800-3,600m, occasionally reaching 4,OOO m.

11. PiceaschrenkianaFiscH.etMEyERinBull.Acacl.Sci,St.Petersb.Ie:253,1842. Pieea orientalis l9. Iongijblia LEBEB., Fl. Ili. 671, 1846-1851. Pinzts schrenkiana ANT., Conif. 97, l847. Abies schrenkiana LiNDL. & GoRD. in Journ. Hort, Soc. Lond. 5: 212, 1850; MAxiM. in Bull. Soc. Nat. Mosc. 65: 58, 1879. Picea obovata schrenkiana CARR., Trait6 Coni£ ed. 2, 338, 1867. Pinus obovata 6. schrenkiana PARL. in DC., Prodr. 16, pt. 415, 1868. Abies 2, smithiana HERDER in Bull. Soc. Nat. Mosc. 41, pt. 1, 423 (Enum. Pl. Semenow.) (non FoRBEs), 1868. Picea robertii Vipp., nokn. AH CCCP 61, 2: 373, l948.

PieeaPrDstrata IsAK, {Pn. KH PrCCP 10: 374, 1962,

11-1. Picea schrenkiana FiscH, et MEyER var. schrenkiana 11-2, Picea schrenkiana FiscH. et MEyER var. tianschanica (RupR.) CHENG et Fu in Ill. Fl. China 1: 300, 1980.

Picea tianschanica RupR, in M6m. Acad. Sci, St. P6tersb. ser. 7, 14; 72, l869. Pinus abies £ schrenkiana Voss in Mitt, Deutsch, Dendr. Ges, l6: 93, 1907. Picea schrenkiana

ssp. tianschanica CRupR.) ByK., Spruce fbrest of Tianshan (in Russ.) Izd. AN KazSSR, Alma-Ata, 22, l950. Picea morindd ssp. tianschanica(RupR.) BEREz., Bot. 2urn. 55: 493, 1970. The typical variety ofSchrenk's spruce occurs on Kirgizskiy in Cental Asia, extending eastward along Tien-Shan bordering USSR and Sinkiang of China, far to Pa-Li-Kun-

Shan and northward to the range of Tarbagatai. Tien-Shan spruce is cenfined to the

west ofTien-Shan, ranging between Talass Ala-Tau and Alajskij range. Their altitudinal

range is from I,400 to 3,600m.

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12. Picea wilsonii MAsT. in Gard. Chron. Ser. 3, 33: 133, fig. 55, 56, l903. Picea watsoniana MAsT. inJourn, Linn, Soc. Lond. (Bot.) 37: 419, 1906. Picea mastersii MAyR, Fremdl. Wald-. u. Parkb. 328, fig, 105-107, 1906. Picea shrenkiana REHD, et WiLs.

in SARG., Pl., Wils. 2: 29, 1914, p.p., non FiscH. et MEyER. Wilson spruce is found in Hopei (on Hsiao-Wutai-Shan & U-Lin-Shan), Shansi (Wutai-Shan & Kuan-Lu-Shan), S. Shensi, W. Hupei, S. Kansu ancl N. Szechuan of

China, at heights of 1,400-2,80em.

13. Picea morrisonicola HAy. in Journ, Coll. Sci. Univ. Tokyo 25, Art. 19, 220, ' !908. . This spruce is native to Taiwan, occurring at altitudes ef 1,900-2,80e m in the central ' range. It is the most seuthern spruce in the genus. 14. Picea chihuahuana MART, in Anal. Inst. Biol. Mex. 13: 31, 1942; Pin. Mex.

ed, 3, 9, 1963.

of Chihuahua and Chihuahua spruce is a southern species occurring in the provinces Durango of Mexico. In the western Sierra Madre it grows at heights of 2,300-3,200 m.

Subgen. II. Picea Picea subgen. Pioea BERT'R. in Ann, Sci. Nat. ser. 5, 20: 83, 1874. Gemmae perulis latiusculis, interdum subulatis extrinsecus obtectae; folia sectione flexibilesvel transversa quadrato-rhombica vel compressa; squamae strobilorum tenues et subcoriaceae et laxe appressae vel patentes, ebovatae vel rotundae apice saepe truncatae

et denticu!atae vel erosae, interdum rotundatae et emarginatae. Typus: Picea abies (LiNN.) KARs'r. (Pinus abies LiNN.) Buds with rather broad, sometimes with awl-shaped scales outsicle; leaves quadrate-

rhombic or compressed in cross-section; cone-scales thin and flexible or subcoriaceous and

and den- loosely appressed or spreading, obovate or rounded in outline, often truncate

ticulate or erose, sometimes rounded and emarginate at apex.

･ Based on the shape and texture of cone-scales as well as leaves, and the color and 'on buds, the following presence or absence of pubescences shoot bearing characteristic two sections and eight series are to be subdivided.

Sect. I. Picea Fl. BoBR., Gen. Picea Hist. Syst. 29, 1970. Picea sect. Etipicea WiLLK., Forstl. ed. 2, 66, 1887. Picea sect. 1. Morinda MAyR, Monogr. Abiet.Jap. 44, 1890, p,p. min. Picea II. MerindaeAsaH, et GRAEBN,, Syn. Mitt. Fl. 1: 196, l897, p.p. min. Picea sect. Alcockiana PARDE in Bull. Soc. Dendr. France 1911: 9, 19!1, p.p. quoad P. alcockiana. Picea sect. Bicotor FiTzp. in Sci. Prec. Roy. Irish Acad. n. ser. 19: 198, 1929, p.p. quoad P. bicolor. Piclaa sect. Bicolor DEB., Man, Conif 1964, p.p. quoad P. bicolor. . characteristics ofbud, fiveseries . Based on the shape and texture of cone-scales and ' are to be subdivisions under this section. Ser. 1. Politae BoBR., descr. emend. Liu BoBR., Gen. Picea Hist. Syst, in Nov. Syst. Pl. Vasc. 7: 7-32, 1970.

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Ramuli juveniles flavido-brunnei, glabri, nitentes; gemmae globosae, vix resinosae; folia 12-20 mm longa, rigida, apice spinoso-acuta.

Typus: PiceaPolita CSmB, et Zucc.) CARR. Young branchlets yellowish-brown, glabrous, shining; buds globose, scarcely re- ' sinous; leaves 12-20 mm long, rigid, spiny-pointed at apex.

15. Picea polita (SiEB. et Zuca.) CARR., Trait6 Coni£ 256, 1855. Abies Polita SiEB. et Zucc., Fl. Jap. 2: 20, 1842. Picea torano KoEHNE, Deutsche Dendr. 22, 1893.

The tigertail spruce is found in the cool-temperate region of Honshu, Shikoku and Kyushu ofJapan, at altitudes varing from 400 to 1,850m. 16. Picea bicolor (MAxiM.) MAyR, Monogr. Abiet 49, t. 3, £ 8, 1890. Abies bicolor MAxiM. in Bull. Acad. Sci, P6tersb. 10: 488, 1866. Picea alcockiana CARR,, Trait6 Conif. ed. 2, 343, 1867. P. j'`iponica MAxiM, in Ind. Sem. Hort. Petrop. 1865: 33,

l866, pro syn. P. bicolor REGEL. ex SENEcL,, Coni £ 24, i868, nom. Pinzasj`aponica PARL.

in DC., Prodr. 16, 2: 431, 1868, sp. dub. P, alcoguiana PARL., 1.c., 431. 16-1, Picea bicolor (MAxiM.) MAyR var. bicolor 16-2. Picea bicolor (MAxiM.) MAyR var. reflexa SHiRAs. et KoyAMA in Bot. Mag. Tokyo 27: l29, t, 2, f. 9-17, I913. The Aicock's spruce isindigenous to the subalpine fbrest ofcentral Honshu ofJapan, with altitudes ranging from 700 to 2,180 m. 17. Picea shirasawae HAyAsm, Nitzl. Geh. Erl. Abb. 43, l969. Picea bicolor MAyR (MAxiM.) var. acicularis SmRAs. et KoyAMA in Bot. Mag. Tokyo 27: l29, 1913. Picea alcockiana acicutaris FiTscH, This spruce has a narrow distribution in Honshu ofJapan. It grows on the west slope of Mt. Yatsuga-dake at heights of 1,200-1,950m.

Ser. 2. Asperatae BoBR,, descr, emend. Liu BoBR,, Gen. Picea Hist. Syst. 31, l970, Ramuli rubescenti-brunnei juveniles vel aurantiaci, subinde pubescentes; gemmae conicae, resinosae; folia 12-25 mm longa, rigida, apice spinoso-acuta,

Typus: Picea asperata MAsT. Branchlets reddish-brown or orange, occasionally pubescent; buds conical, resinous; leaves l2-25 mm long, rigid, spiny-pointed at apex, 18, Picea asperata MAsT. in Journ. Linn. Soc. Lond. Bot. 37:419, 1906. Piceaaurantiaca MAsT. 1.c. 421. Picea retr(V7exa MAsT., 1.c. Picea asperata MAsT. var. notabilis REHD. et WiLs, in SARG., Pl. Wils. 2: 23, 1914. Picea notabilis (REHD. & W-s,) LAcAss. in Trav, Lab. For. Toul. II, 3, 1: 180, fig. (Et. Anat. Syst. Picea), 1934. Picea heterolopis REHD. et WrLs., l.c. 24. Picea asperata MAsT. var. Pondkerosa REHD. & WiLs., Lc.,1914. Piceaponderosa (REHD. & WiLs.) LAaAss.,l.c., 203, fig,, 1934, Picea mperata MAsT. var. heterolopis (REHD. & WiLs.) CHENG ex REHD., Man, Cult, Trees Shrubs ed. 2, 24, 1940. The dragon spruce covers a wide area in W. China, reachlng an altitude of 3,900 m.

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236 Acta Phytotax. Geobot. Vol. XXXIII

19. Picea crassifblia KoM., Bot. Mat. 4: !77, 1923. This spruce has its distribution in Tsinghai (east of Lake Kuku-Nor, southern slepe ofTsi-Lien-Shan), northern part ofKansu and Holan-Shan in China, at altitudes of2,600-

3,100 m. 20. Picea meyeri REHD. et WiLs. in SARG., Pl. Wils. 4: 28, 1914. Found in Shansi (on Wutai-Shan & Kuan-Lu-Shan), Hopei (en U-Lin-Shan & Hsiao-Wutai-Shan), Kansu (Tao-Chan Ting) and Inner Mongolia in N. China, at

altitude 1,600-2,IOO m, 21. Picea gemmata REHD. et WiLs. in SARG., Pl. Wils. 4: 28, 1914. Occurring on Tapao-Shan, northeast of Ta-Chien-Lu, in W. Szechuan, at altitude . 2,200-3,600 m. 22. Picea neoveitchii MAsT. in Gard. Chron, ser. e, 33:1l6, fig. 50, 51, 1903. This tree is found in W. Hupei, S. Shensi and S. Kansu, extending northeastward to Chahar and Jeho! in China, at altitudes of 1,500-2,OOO m. Ser. 3. 0bovatae BoBR., descr. emend. Liu BoBR., Gen. Picea Hist Syst. 29, 1970. Ramuli juveniles rubescenti-brunnei vel aurantiaco-rubri, minute pubescentes; !O-20 mm gemmae conicae, perulis subulatis extrlnsecus obtectae, eresinosae; folia longa, rigida, apice obtusa. Typus: Picea obovata LEDEB. Branchlets reddish-brewn or orange-red, with minute pubescence; buds conical, with awl-shaped scales outside, without resin; leaves 1O-20mm long, rigid, obtuse at apex. 23. Picea abies (LiNN.) KARsT., Deutsch. Fl. Pharm, Med. Bot. 324, 1881. Pinus abies LiNN., Sp. Pl. 10e2, l753. Abies Picea MiLL., Gard. Dict. ed. 8, A. no. 2, 1768. Pinas-AbiesPicea WEsT., Bot, Univ. 1: 211, l770. Pinuspieea Du Roi, Obs. Bot.

"excelstts." 37, 1771. Pintts exeelsa LAM., Fl. Frang. 2: 202, 1778, Abies Pectinata GiuB., Exerc. Phytol. 2: 411, 1792, non Pinus Peetinata LAM., 1778. Pintts Mrarnidaiis SALisB., Prodr. Stirp. Chap. Allect. 399, 1796, Abies excelsa Pom., Encycl. Meth. Bot 6: 518, 1805. Peuce rubra L. C. RicH. in Ann. Mus. Hist. Nat. Paris 16: 298, 1810, Picea rtibra A. DiETR., Fl. Geg. Berlin 2: 795, 1824. Pieea vuigaris LiNK in Abh. Akad. Wiss. Berlin Man. 365, l836. 1827: 180 (Fam. Pinus, 1827) I830, Abies aommunis LAws., Agric. Abies exeelsa var. 1. commtinis LouD., Arb. Brit. 4: 2294, 1838. Picea excelsa (LAM.) LiNK in Linnaea 15: 517, 1841. Pinus Picea var. a vu(garis LouD, ex ENDL., Syn. Conif 117, 1847. Abies vuigaris WEND,, Pflanz. Bot. Gart. I. Conif. I5, 1851, non PoiR. 1805. Abies abies (L.) BoRBAs in Magyar Bot. Lap. 1: 28, 1902, Pinzaf abies L. (subsp.) vuigaris Voss in PuTuTz & MEyER, Landlex. 4: 770, 1913, 23-1. Picea abies (LiNN.) KARsT. var. abies 23-2. Picea abies (LiNN.) KARsT. var. acuminata (BEcK) DALL. et JAcKs. in HARR. & DAL.L., Handb. Conif Ginkg. 335, 1966.

Pieea excetsa acuminata BEcK 23-3. Picea abies (LiNN,) KARsT. var. europaea (TEpL.) JuRK. et PARF., Ilju!1.

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glavn, bot, sada ANSSSR 64, 42, 1967, Picea vuigaris var. ezaropaea TEpL. in Bull. Soc. Nat. Moscou 41, 2: 249, fig. I, 1868.

Abies excetsa a. vuigaris WiLLK., Forstl. Fl, 66, 1875. Picea vuigaris a. vuigaris BEcK in Blatt.

Ver. Landesk. Niederoestr. 1890: 73, 1890, Picea excelsa var. 7. europaea (TEpL.) SaHROT･ in Viertelal'ahrschr, Naturf Ges, ZUrich 43: 142, 282, figs., 1898, Piceaabies£ europaea

LiNDM., Svensk, Fanerog. 39, fig., 1918.

The common spruce is widely distributed in N. and C. Europe, reaching an altitude

of 2,300 m, often in large forests.

24. Picea ebovata LEDEB., Fl. Alt. 4: 201, 1833. Pinus abies sensu PALL., Fl. Ross. 1, 1: 6, f, 1, fig. G, 1788, non LiNN,, }753. Abies obovata (LEDEB,) D. DeN ex LouD., Arb. Brit. 4: 2329, 1838. Pinusobovata (LEDEB.) ANT･, Conif. 69, t. 37, fig. 2, 1847, Pinus abies obovata AND. ex A. MuRR. in (Ravenscroft))

Pinet. Brit. 2: 162, l868. Picea vuigaris var. altaica TEpL. in Bull. Soc. Nat. Moscou 41, 2: 250, fig. 4, 1868. Abies excetsa var. obovata (LEDEB.) K. KocH, Dendr. 2, 2: 238, 1872. Picea excelsa 6. obevata (LEDEB.) BLyTT, Norges Fl. 2: 391, 1874. Abies excelsa 7. attaiea (TEpL.) WiLLK., Forstl. Fl. 67, 1875. Piceaaipestris BRuEGG, inJahresber, Naturfl Ges. GraubUnd. n. ser. 29: 167, 1886. Picea excelsa aipestris (BRuEaG.) BEissN., Handb, Nadelh･ 357, l891. Pinus abies f. obovata (LEDEB.) Voss in Mitt. Deutsch. Dendr. Ges, 1907 (16) : 93, l908; in PuTLiTz & MEyER, Landlex. 4: 770, 1913. Picea obovata var. aipestris CBRuEGG.) HENRy in ELwEs & HENRy, Trees Gr, Brit. Irel. 6: 1360, 1912. Picea excelsa B- 1: 1913. P. aipestris (BRuEGG.) AscHERs. & GRAEBN., Syn. Mitteleur. Fl. ed. 2, 306, Picea exce"fa D. P. obovata (LEDEB.) AscH. & GRAEBN., Syn. Mitteleur, Fl. ed. 2, 1: Be8, 1913, Picea abies f, obovata (LEDEB.) LiNDM., Svensk Fanerog. 39, fig. 49, 5, 1918. Picea Petschorica GovoR.

24-1, Picea obovata LEDEB. var. obovata

2C2. Picea obovata LEDEB. var. fennica (REG.) HENRy in ELwEs & HENRy, Trees Gr. Brit Irel. 6: IS60, 1912. Pinus abies var, fonnica REGEL in Gartenfi. 12: 95, fig. n, 1863, Pinus abies L, var. medioxirna NyL. in Bull. Soc. Bot. France 10: 501, l863. Picea vuigaris var. zaze;arewii N,

KAuFFM., Moskovsk. Fl. 605, 1866. Abies excetsa medioxima A. MuRR. in (Ravenscroft), Pinet. Brit. 2: 138, 1867; 3: 827, 1884. Picea vuigaris var. ntratensis TEpL. in Bull. Soc-

Nat. Moscou 41, 2: 249, fig, 2, 3, 1868. Picea excelsafotinica REGEL, Russ. D.endr. 18, 1870, nom. Picea excelsa 6. medioxima (NyL,) WiLLK,, Forstl. Fl. ed. 2, 75, 1887; KoEHNE,

``subsp.";LipA, Deutsche Dendr. 23, 1893, Dendrofl. U.R,S.S. 1: 69, !939. Picea vuigaris e. fcnnica (REGEL) BEcK in Blatt. Ver. Landesk, Niederoster. 1890: 74, 1890, Picea excelsa var, fonnica subvar. medioxima (NyL.) ScHRbTER in Vierteljahrschr. Naturf, Ges. ZUrich 43: 140 (Vielgest. Fichte, 20), 1898. Picea excelsa c, P.fonnica (REG.) AscHERs. & GRAEBN., Syn. Mitteleur, Fl. ed. 2, 1: 307, 19I3. Picea abies f.fonnica (REG,) LiNDM., Svensk Fanerog. 39, fig. 49, 3, 1918, Pieea fonnica (REG.) KoM., Fl. U,R.S.S. 1: 145, 1934, Picea excelsa soptentrionalis HoRT, ex KoM., I.c., 1934, pro syn.

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238 Acta Phytotax. Geobot. Vol. XXXIll

Siberian spruce is the most widely distributed species in the genus, occurring from

N. E. Europe through N. and E. Russia, Siberia, to Kamtschatka and N. E. China.

Ser. 4. Glaucae BoBR., descr. emend. Liu

BoBR., Gen. Picea Hist. Syst. 34, 1870. Ramuli juveniles plerumque glabri, pallido-viridi, demum pallide brunnescentes vel atroflavidobrunnescentes; gemmae ovato-ovoideae, leviter resinosaej folia 8-l8mrn longa, apice calloso-acuta.

Typus: Picea glauca (MoENoH) Voss Young branchlets rnostly glabrous, glaucous, becoming pale brown or dark yellow-

ish-brown afterward; buds ovoid-ovate, slightly resinous; leaves 8-18 mm Iong, callose-

acute at apex.

25. Picea glauca (MoENaH) Voss in Mitt. Deutsch. Dendr, Ges. 1907(l6) : 93, l908. Pinus glauca MoENcH, Verz. Ausl. Baume Weissenst. 73, 1785. Abies canadensis

MiLL., Gard. Dict. ed. 8, A. no, 4, 1768. Pinus abies alba MtiNaHH,, Hausvat. 5: 222,

1770. Pinus-Abies canadensis WEsT., Bot. Univ. 1: 211, 1770. Pinus alba AiT., Hort･

Kew. 3: 37!, 1789. Pinus canadensis a. alba CAsT., Viagg. Stati Uniti 2: 314, 1790. Abies curwij?]lia SALisB, in Trans. Linn. Soc. Lond. 8: .315, l807. Abies alba MicHx., Hist. Arb.

Amm. Sept. 1: 133, t. I2, 18IO, non MiLL., 1768. Picea atba LiNK, Handb. Erkenn. Gew.

2: 478, 1831. Abies acutissima KNiGHT ex PERRy, Syn. Cloni£ 37, 1850, p.p. Picea canadensis (MiLL.) BRiTT., STERNs & PoGG., Prelim. Cat. Anthoph. Pterid. N. York 71, 1888. Picea laxa (EHRH,) SARG. in Gard. & For. 2: 496, l888, non Pinus laxa EHRH. Picea gtauca Voss var. porsidi RAup in Sargentia 6: 102, 1947. 25-l. Picea glauca (MoENcH) Voss var. g!auca 25-2, Picea glauca (MoENcH) Voss var. albertiana (S. BR.) SARG. in Bot. Gaz. 67: 208, 1919.

Picea agbertiana S. BR., Torreya 7: l26, 1907. Picea columbiana sensu RyDB. in Mem･

New York Bot. Gard. 1: 11, 1900, non LEMM., 1897. Picea alba sensuREHD. in MoLLER's

Deutsch. Gartn.-Zeit. 20: l16, fig. 11, l905, non LiNK, I831. Picea alba albertiana CS. BR.) BEissN,, Handb. Nadelh. ed. 2, 273, 1909, Picea canadensis var. albertiana (S, BR.) REHD. in Mitt. Deutsch. Dendr. Ges. 1915(24) : 213, 1916.

The white spruce has a transcontinental distribution. It extends from Alaska, east across Canada to Hudson Bay, LabradQr, and Newfbundland, south to New York, Minne-

sota, Montana, and British Columbia, occurring from near sea level up to the tree line at

305-1,607 m,

Ser, 5. Rubentes BoBR., descr. emend. Liu BoBR., Gen, Picea Hist. Syst. 35, 1970. Ramuli juveniles brunnei-rubescentes, dense glandulosi vel atro-pilosi; gemmae ovoideo-conicae, eresinosae, perulis subulatis extrinsecus obtectae; folia 6-18mm longa,

apice obtusa vel brevi-acuta.

Typus: Picea rubens SARG. Young branchlets reddish brown, with dense glandular or black pilose hairs; buds

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ovoid-conical, with awl-shaped scales outside, without resin; Ieaves 6-l8 mm long, ebtuse

or short-pointed at apex.

26. Picea rubens SARG,, SIiva N. Am. 12: 33, t, 597, I898.

Pintts-Abies eanadensis 2. rubescens WEsT., Bot. Univ. 1: 2ll, l770. ]Pinus abies actttissima

MVNcHH., Hausvat. 5: 225, ]770. Pinus mariana rubra Du Roi, Obs. Bot. 39, 1771.

Pinus amen'cana rubra WANG., Beytr. Deutsch, Holzger, Forstwiss. Nordam. Holz. 75, t. 16,

fig. 54, l787. Pinus rabra LAMB., Descr. Pinus 1: 43, t. 28, 1803, nen Mim., 1768, nec MicHx. f., 1810. Abies rubra Pom., Encycl. M6th. Bot. 6: 520, 1805. Picea rubra (Du Roi) LiNK, Handb. Erkenn. Gew. 2: 478, 1831, non A. DiETR., 1824. Abies nigra 6.

SpAcH, Hist. Nat, Veg, Phan, 11: 377, 1842, Abies americana K. KoaH, Dendt 2, 2:241,

1873, non MmL., 1768, nec PRovENaHER, 1862. Abies arctica CuNNiNGHAM ex GoRD･,

Pinet. ed. 2, l7, 1875, pro syn,, nonA. MuRR., 1867. Abies rubra arctica HoRT, ex GoRD., I,c-, 1875, pro syn. Picea nigra var. rubra ENGELM. in Gard. Chron. ser. 2, 11: 334, 1879,

Picea acutissimaJ. G.JAcKin Gard. & For. 10: 63, 1897, nom, tentat. PiceaatLstralis SMALL,

Fl. Southeast. U, S. 30, 1903.

The red spruce is a native of N. E. North America. Its range extends frorn Cape Breton Island, Nova Scotia, to Maine and S. Qyebec and Ontario, and southward to W. North Carolina and E. Tennessee, reaching from sea level te about 1,500 m altitude

in the Adirondacks.

27. Piceamariana (MmL.) BRiTT,, STERNs & PoGG., Prelim. Cat. Pl, New York City 7I, 1888.

Abies mariana MiLL,, Gard. Dict, ed. 8, A. no. 5, 1 768. Pinus Abies mariana MttNcHH.,

Hausvat. 5: 224, 1770. Pinus-Abies canadensis 3. nigricans WEsT., Bot, Univ. 1: 211, l770. Pinus mariana (Mim.) Du Roi, Obs, Bot. 38, l771. Pinus nigra AiT., Hort. Kew. 3: 370, l789. Pinus canaclensis fi, nigra CAsTiGLioNi, Viagg. Stati Uniti 2: 315, 1790. Pinus americana GAERTN., Fruct. Sem, 2: 60, t, 91, fig. a-e, 1791,nonMmL,, 1768. Abies denticulata MicHx., Fl. Bor,-Am, 2: 206, l803. Abies nigra (AiT,) Pom, Encycl, M6th, Bot.

6: 520, 1804, Pinus denticulata MiaHx. ex MuHL., Cat. PI. Am. Sept. 89, 18}3. Picea

nigra (L.) LiNK, Handb. Erkenn. Gew, 2: 478, l831. Picea brevijblia PEcK, Spruces Adirond, 13, 1897. Picea nigra var. brevijblia (PEcK) REHD. in BAiLEy, Cycl, Am, Hort. 3: l334, l910. Picea mariana var. brevijblia (PEcK) REHD, in Rhodora 9: 109, 1907. Abies matylandica HoRT. ex BEissN., Handb. Nadelh. ed. 2, 257, 1909, pro syn, Black spruce spans the continent from Newfoundland and the northeastern United States west and northward to northwestern Alaska. It grows usually at hei.crhts between 150 and 8eO m, occasionally grows at i,800 m in the Rocky Mountains, Sect. II. Casicta MAyR, descr. emend. Liu

MAyR, Monogr. Abiet. 44, 1890. I'icea sect. Alcockiana PARDE in Bull. Soc, Dendr. France 1911: 9, 1911, p,p. Picea sect. Bicolor FiTzpATR in Sci. Proc. Roy, Irish Acad. n, ser, 19: 198, 1929, p.p. Vkitchia

LiNDL, in Gard. Chron. 1861: 265, 1861.

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Gemmae plerumque resinosae ; folia compressa, carinata vel inaequalitcr quadrangu- Iata, superne interdum utrinque superficiebus ambabus folii stomatifera; squamae

strobilorum ovato-oblongae, tenues et flexibiles, ad frutus maturitatem laxe imbricatae

vel patentes, apice eroso-denticulatae. Typus: PiceaJ'ezoensis (SmB. et Zucc.) CARR. (Picea cijanensis FiscH.) Buds very frequently resinous; leaves compressed, keeled, stomatiferous above, sometimes on both surfaces ; cone-scaies ovate-oblong, thin and flexible, loosely overlapped

or spreading before fruit maturity, erose-denticulate at apex.

Ser. 6. Ajanenses BoBR., descr. ernend. Liu Sect. thsicta, ser. 4ianenses BoBR., Gen. Picea Hist, Syst. 37, l970. Ramuli juveniles flavido-brunnei, glabri, nitiduli; gemmae ovoideo-comcae, resmo- sae; folia I5-25 mm longa, apice acute acuta, sectione transversa lineari-rhomboidea,

superne stomatifera; squamae strobilerum ovato-oblongae.

Typus: Picea J'ezoensis CSmB. et Zucc.) CARR. (Picea njanensis FiscH.) Young branchlets pale yel!owish brown, glabrous, shining; leaves 15-25 mm long,

sharply acute at apex, linear-rhomboid in cross-section, stomatiferous on upper side;

cone-scales ovate-oblong.

28. Picea jezoensis (SiEB. et Zuac.) CARR., Trait6 Conif. 255, 1855. AbiesJ'eaDensis SiEB. et Zucc., Fl. Jap. 2: 19, t. 110, 1842. Pinusj'ezoensis ANToiNE, Coni£ 97, t. 37, fig. 1, I847. Abies ofanensis LiNDL. & GoRD. inJourn. Hort. Sec. Lond. C`A. `iPicea 5: 212, 1850, nom., Sphalm. njonensis"; cum syn, njonensis." Picea ay'anensis FiscH. ex CARR., Trait6 Coni£ 259, !855, descr. manca. Pintis.tirma var. fi. j'ezoensis ENDL., Syn, Coni£ 99, 1847. Abiesj7rma 6. j'ezoensis CARR., Trait6 Conif. 212, 1855. Abies microsperma LiNDL. in Gard. Chron. 1861: 22, 186I. Vlaitchia j'ciponica LiNDL., 1.c.

265, 1861. Picea microsperma aARR., Trait6 Cloni£ ed. 2, 339, 1867, Abies meneiesii

$ensu FRANcH. et SAv., Enum. Pl. Jap. 1: 467, 1875, non LiNDL., l833. Picea ay'anensis FisaH. var. microsperma (LiNDL.) MAsT. in Gard. Chron. ser 2, 13; 115, !880. Abies ay'anensis microsperma VEiTcH, Man. Conif, 66, 1881. Tszrga ay'anensis REGEL, Russ. Dendr. ed. 2, 39, 1883. Pinusj'ezoezasis f microsperma (MAsT.) Voss in PuTLiTz & MEyER, Landlex.

4: 772, 1913. Picea kamtschatkensis LAcAss. in Bull. Soc. Hist. Nat. Toul. 58: 637, 1929;

Trav. Lab. For. Toul. 3(1):138, t. I41, 1934. Picea mantishurica NAK. inJourn.Jap. Bot.

19: 138, i943. Picea komarovii V. VAssiL., Bot. >KYPH. 35(2): 504, 1950.

2g-l. Picea jezoensis (SiEB. et Zucc.) CARR. var. jezoensis 28-2. Piceajezoensis (SiEB. et Zucc.) CARR. var. hondoensis (MAyR) REHD. jn Mitt. Deutsch. Dendr. Ges. 1915 (24): 314, 1916. Abies atcegzaiana J. G. VEiTH ex LiNDL. in Gard. Chron 1861: 23, 1861, p.p. Picea hondbensis MAyR, Monog. Abiet, Jap. 51, t, 4, fig. 9, 1890. Pinus hondbensis (MAyR) Voss in Mitt. Deutsch. Dendr. Ges. 1907 (16): 93, 1908. Picea ay'anensis FisaH. var. miero- sperma sensu BEissN., Handb. Nadelh. ed. 2, 291, 1909, non MAsT., 1880, The Yezo spruce has a very wide range in N. E. Asia and Japan, extending from

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Ayan on the sea of Okhotsk southward through Siberia to N. E. China (Manchuria) and N. Korea-the southern-most limit, east to Kamchatka and southward through S. Kuriles,

Saghalin to Hokkaido; occurring at heights of l,OOO-l,300 in. The variety, Hondo spruce, often forms subalpine forests in central Honshu ofJapan, with higher distribution at 1,1OO-2,650 m as compared with the typical one at 1,OOO-l,300 m.

29. Picea sitchensis (BoNG.) CARR., Trait6 Coni £ 260, l855. Pinus sitchensis BoNG. in Mem. Acad. Sci. St. P6tersb. 2: I63, I833. Abies falcata RAFiN., Med. Fl. 2: 182, I830, nom. Pintts menziesii DouGL. ex D. DoN in LAMB., Descr.

Pinus ed. 3, 2:･ p. sine num. inter 144 et 145 inserta, 1832. Abies fneneiesii LiNDL. in Penny

Cycl. 1: 32, 1833. Picea menziesii ENGELM. in Trans. Acad. St. Louis2:214, l863. Picea

sirkaenst's MAyR, Wald. Nordam. 338, 1890. Pinus sitkaensis BoNG. ex Voss in Mitt. Deutsch. Dendr. Ges. 1907 (16): 93, I908. Pinus sitkaensis £ menziesii Voss in PuTuTz & MEyER, Landlex. 4: 772, l913. Picea.fblcata (RAF.) VoLcK,-SuRiNG. in Mitt. Deutsch. Dendr. Ges. 1927 (38): 321, 1927. The sitka spruce grows in a narrew strip along the Pacific coast from northwestern

California to southern Alaska, over an area abeut 2,500 km long. It extends from sea

Ievel to the timberline, about 1,l89 m in the coastal mountains, but occurs mainly at

altitudes below 457 m.

Ser. 7. Likiangenses BoBR., descr. emend Liv BoBR., Gen. Pieea Hist. Syst. 39, 1970. 'rufobrunnei, Ramuli juveniles flavido-aurantiaci vel pallide flavo-cinerei, interdum

setulosi; fblia 8-15 mm longa, sectione transvera compresso-quadrata utrinque stomati-

fera; squamae strobilorum rhomboido-ovatae.

Typus: Picea likiaugensis (FRANcH,) PRiTz. Branchlets yellowish orange or pale brown, sometimes reddish brown, minutely setose; leaves 8-15 rnm long, compressed-quadrate in transverse section, stomatiferous

on both sides; cone-scales rhomboid-ovate. 30. Picea likiarigensis (FRANaH.) PRiTz. in Bot. Jahrb. 29: 217, 1900. Abies likiangensis FRANcH. inJourn. Bot. 13: 257, 1899. Picea alcockiana sensin MAsT. in Journ. Linn. Soc. Lond. Bot. 37: 418, 1906, non CARR. 30-1. Picea likiangensis (FRANaH.) PRiTz. var. Iikiangensis The Likiang spruce is fbund in N. W. Yunnan and S. W. Szechuan, China, at altitudes of 2,500-3,900m. 30-2. Picea likiangensis (FRANaH.) PRiTz, var. balfouriana (REHD. et WiLs.) HrLL. (1932), Man. Tr. Shr. rev. ed. 502, 1975. Piceabaijbntriana REHD. et WiLs. in SARG., Pl. Wils. 2: 30, 19I4. Picea sikangensis (]HENG in Contr. Biol. Lab. Sci. Soc. China (Bot.), ser. 6, 33, 19Bl. Picea likiairgensis 'Cin var. baijberriana (REBD. & WiLs.) CHENG lit. May 15, 1939) ex REHD., Man Cult. Trees Shrubs, ed. 2, 29, 1940. Picealunnanensds ViLM. ex LAcAss. in Trav. Lab. For. Toul.

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242 Acta Phytotax. Geobot. Vol. XXXIII

II, 3, 1: 246, fig. i934. Pieeayecnnanezasis WiLs. inJourn. Arn. Arb. 7; 47, 1926, pro syn. Found in S. Tsinghai, N. W. & W. Szechuan, and area of Charndo (Sikang) and

westward to E. Hsitsang, at altitude 3,OO04,100m. 30-3. Picea 1ikiangensis (FRANcH.) PRiTz. var. hirtella (REHD. et WiLs.) WANG Picea hirtella REHD. et WiLs. in SARG., Pl. Wils. 2: 32, 1914. Western Szechuan (west of Kuan-Hsien, northeast of Kangting), altitude 3,OOO-

4,100 m. 3on. Picea likiangensis (FRANcH.) PRiTz. var. montigena (MAsT.) Liu, comb. nov.

Picea montigena MAsT. in Gard. Chron. ser. 3, 39: i46, fig. 56, 1906. Mountains of N. W. Yunnan, (Likiang Snow Range) northward to those around

Kanting in W. Szechuan, alt. 3,60an,OOOm. 30-5. Picea 1ikiangensis (FRANaH.) PRiTz. var. rubescens REHD. et WiLs. in

SARG., PL Wils. 3: 3I, 1916. In forests areund Ta-chien-lu, W. Szechuan, alt. 3,OOO-4,IOO. 31. Picea purpurea MAsT. inJournL Linn, Soc. Lond. Bot 37: 418, 1906. Picea likiangensis var.PuiPurea (MAsT.) DALL. etJAaK., Handb. Coni£ 334, fig. 74, 1923. This is a native of southwestern China. Its range covers N. W. Szechuan, S. W. & Range), often C. Kansu, and E, Tsinghai, at altitudes of 2,600-3,800 m (in Min-Shan

forming pure forest. Ser. 8. Pungentes Bo]R., descr. emend, Liu

BoBR., Gen. Pieea Hist. Syst. 38, l970. Ramuli juveniles flavido-brunnei; gemmae ovoideae, eresinosae; folia 20-43 mm longa, apice acuminata, pungentia, sectione transversa inaequaliter quadrangulata,

utrinque stomatifera; squamae strobilorum rhomboideo-oblongae.

Typus: Picea Pungens ENGELM. 32. Picea pungens ENGELM. in Gard. Chron. n. ser. 11: 334, l879. Abies menziesii sensu ENGELM. in Amer. Journ. Sci. ser. 2, 34: 330, 1862, nQn LiNDL.,

menziesii Sci. St. Louis 2: 214 non CARR., 1833. Picea ENGELM. in Trans.Acad. (1863), ' l855. Picea mengiesii part?ana ANDRE in Ill. Hort. 23: 198, I876. PiceaParrp,ana BARR. ex FREuD., Bekannt. Cult. Nadelh. 6, l886, nom. subnud. PinusPartyana (EHRH., 1788) ex Voss in Mitt. Deutsch, Dendr. Ges. 1907 (16) : 93 (1908), non GoRD., 1858, nec ENGELM., 1862. Pinus armata Voss in op: cit 95, l908, nom. event. The range of blue spruce extends from Colorado to New Mexico, Utah, and

Wyoming ofU.S.A. at altitudes from 1,800 to 3,300m. 33. Piceaengelmanni(I'ARRy)ENGELM.inTrans.Acad.Sci.St.Louis2:212,1863. Abies angelmanni PARRy in Trans. Acad. Sci. St. Louis 2: I22, 1863, nom. nud. Pinus commutata PARL. in DC., Prodr. 16, 2: 417, 1868. Pieea colzambiana LEMM. in Gard. & Forest 10: 183, 1897. Picea engelmanni var. colambiana (LEMM.) BEissN. in Mitt. Deutsch. Dendr.

Ge$. 1897(6): 52, 1897,

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Engelmann spruce has a very wide distribution. Its range extends from British Columbia and Alberta, south to New Mexico and Arizona. It grows as low as 800m

in the Coastal Range, and as high as 4,OOO m in the S. Rocky Mountains.

34. Picea mexicana MART. in Anal. Inst. Biol. Mex. 32 (1-2): I37, 1961. As the specific name shows, it is a native of Mexico, occurring in Nuevo Leon and Coahuila (Saltillo), at an altitude of about 2,700 m.

IV. DistributionoftheGenus

The genus Picea consists of 34 species and fifteen varieties all confined to the temperate and subarctic regions ofthe Northern Hemisphere. Geographically it extends from Lat. 72025' N. (far exceeding the Arctic' Circle) southward to Lat, 230N, (a !ittle over the Tropic ofCancer, Taiwan, ROC), and from Long. 5e27' W, (Massifde Vercors lying eastofRh6ne,France) to Long. I55eW. (Kamchatka,USSR). Its rangeincludesregions

of northcrn Eurasia, northeastern Turkey and Caucasus, Central Asia and Tianshan-

Himalaya, central and southwestern China, eastern Asia and many islands bordering the Pacific Ocean in the northeast, and North America. It grows at heights starting at sea level in the boreal region to a height of 4,lOO m (occasionally attaining 4,800 m) on the

higher mountains in southwestern China, however, it is found most abundant from sea

level to 3,600m.

V. Acknowledgements

I am mest gratefu1 to Dr. H. HARA of the University of Tokyo, Dr. R. ORNDuFF

of the University of California at Berkeley, Dr, D. H. NicoLsoN of the Smithsonian

Institution in Washington, D. C. and Dr. T. KoyAMA of the New York Botanical Garden

who gave rne kind permission to freely study the spruce specimens in their outstanding herbaria. r am also extremely gratefu1 to Pro£ de FERRE of Universite P. Sabatier at Toulouse, Dr. J. E. VmAL of the Mus6um National d'HistQire Naturelle in ?aris, Dr. T. YA"fAzAKi of the Koishikawa Botanical Garden in Tokyo and Dr. T. I. CHuANG of the

Illinois State University for their kind assistance in providing many important references regarding the spruces. Finally I wish to express my gratitude to JCRR now CAPD (Council fbr Agricultural Planning and Development, the Republic of China) fbr partly financing this study.

VI. ImportantReferences

BEissNER, L.: Handbuch der Nadelholzkunde, rev, by FiTsaHEN,J, (1930). BoBRov, E. G.: Generis Picea Historia etSystematica, Nov. Syst. Plant. Vasc. 7. 7-39, Nauk, Leningrad (1970). CHEN, Y.: Illustrated Manual of Chinese Trees and Shrubs (1937), CHENG, W. C.; Les Forest du Se-tchouan et du Si-Kang eriental (1939). CoLLEAu, C.: Anatomie compar6e des feuilles du Picea (1968). DA-iMoRE, W. & A, B. JAcKsoN; A Handbook ef Coniferae and Ginkgoaceae, rev. by HARRisoN, S, G, <1966).

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244 Acta Phytotax， Geobot． Vo 至． xxxm

DEBAzAG E F ： Manuel des 1964 ，．． Conifer6s， Nancy （）．

DIETRIcH， A ．： Florader Gegend um BerlinI1， 794 （1824）． Fow 肌 Ls ， H ． A ．： Sil崛 cs of 正brest trees Qf the United States（1965）， − GAussEN ， H ．； Les Gymnospermes，563 628 （1966）． indigenous to 1952 ． HAYAsHI， Y ．： The natur 盆 distribution of important trees Japa旦（）： NUtzliche Geb61ze mit Er1註uterungen und Abbildungen （1969）．， HILLIERs Manual of trees ＆ shrubs （1975）．

1． D ．＆ V 。 L PARFENov ： Zur Fragc der Systematik von Picea abies L KARsT ．i皿 Russ． JuRKEvr6，（）（） Bjull sada 1967 ．glavn．bot， AN SSSR 64 （）． KRUssMANN ， G ．： Die Nadelgeh61ze（1960）．

KuRATA S ： Illustrated fbrest trees ofJapan ，． important （1964）． ’ ’ LA 鯔 sAGNE M ： Unc nouvelle csp ξce d Epicca ： Pioea s ，． kamtchatkensi（1929）．： Etude morphologique ， anatomiquc et systematique du genre Picea （1934）． L 匸NNE ， H ．： Species Plantarum （1753＞．．： Genera Plantarum （1754）． LITTLE ， E ． L Jr．： Check list ofnative and natura 玉ized trccs ofthc Unjted States（including Alaska）（1953），， MAEKAwA ， F． et al ．； MAK 【No s New Illustrated Flora ofJapan （1970）．

MARTINEz M ： Las Pinaceas Mexicanas ed 3 1963 ，．，．（）． MAsTERs ， M ・T ．： On the conifers of China （Journ． Linn ， SQc ． Bot ．37，1906）・ MAYR H ．： Monographle der Abietineen des R．eiches 1890 ．， Japonischen （）， MELcHIoR H ＆ E WERDERMANN ： A ENGLER s Syllabus PHanze 曲 milien 1954 ．，，，， der （）

Mr 跚 E K ＆ Y KuDo ： Icon （s of the essential forest trees ef Hokkaido 1932 ，．．（）．

NAKAI T ．： Taxaceae et coniferae in flora coreano −Manshuricae novae ．皿 n ． Bot ．2 1919 ．，（Jap JQ ，） ● ad asiae orientalis c Notulae plantas （1．．17， 1941 ＆ 19， 1943）。 OHWI ．： Flora ofJapan engL ed ．1965 ．， J ，（＞ PAN616 ．： Eine neue cenifere in den Alpen 1876 、， J 6stlichen （） REHDER ， A ．： Manual of Gultivated Trees and Shrubs， ed ．2 （1961）．一．．．： Bibliography of Cultivated Trees and Shrubs Hardy in the Cooler Temperate Regions of the Northern Hemisphere （1949b ）．

REHDER ＆ E H WILsoN ： Pinaccae in SARG Plantae Wilsonianae 1914 ，A ．．．，， 2 （）．

SARGENT G S ： Manual of trees of North America 1961 ，．．（）．

SGHMIDT −VoGT H ： Die Fichte 1977 ，（）．

sHIR 胡 AwA H ＆ M KoY 蝋 A ： some ncw species inJapan Bot Ma Tokyo 27 1917 ，．． ofP 嫐 and 朋 β5 （．呂・，）・

TsENG W ．： DendrQlogy of Ch …na 1961 ．，（）．．： Far −eastern in For Abstr 13 1950 ． VAsrLEv， V N spruces of the section Omerika（Russ ．）．．，27 （） V 凪 REcK L A ．＆ E ． L LITTLE ，： Alaska Trees and Shrubs 1972 ．，，Jr （）

WILLKoMM M ： Forstliche fiora von DeutSchland und eesterrcich cd 2 1887 ，，．（）．

ヒビートウ属 Picea の分類の歴史をレュするとともに，全世界の種を対象とした新しい分類

体系を提唱した。本属は北半球の温帯および亜寒帯に分布する34種からなるが，それらは以下

の 2 亜属， 4 節， 11系にまとめられる。

Subgen ．1，0morika （WILLK ．）Llu Ser．2． Asperatae BoBR ． Sect．1． Omorika WiLLK． Ser．3． Obovatae BoBR ，

Sect． II． Morinda MAyR Ser．4． Glaucae BOBR ．

Ser．1，0rientales LIu Ser．5． Rubentes BoBR ．

Ser．2。 Glehnianae BoBR ． Sect． II． Casicta MAYR

Ser．3． Smithianae Llu Ser．6． Ajanensis BOBR ． Subgen ． II・Picea Ser．7．1．ikiangenses BoBR ． Sect．1． Picea Ser・8・Pungentes BOBR ． Ser。 L Politae BoBR ．

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April 1982 ， Acta Phytotax ． Geobot． 245

これらの分は，のののいののよび類群球果鱗片形状，葉形と構造，若枝毛有無，冬芽おその

の形などのにづいつ鱗片状形質基て定義された。また，各種にいて異名が整理された。

日のについてアエマツ P ．ヤツガヒヒバ本産種は，カゾ glehnii，タケトウ P．koyamae ，メ

モ maximowiczii ラミ P ・が上記の Glehnianae にハモ P ．マツハ P ． − 系，リミ polita，ダ bi

ヒマハ shirasawae color ，メツダ P，が Politae にエゾマツ P ．系， jezoensisが Ajanenses

系に所属する。

小山博滋：オオキダチハマグルマ（新称）

キチハマルマ Wcdelia bifloraL ． DC ，はミクロシアメラシアアダグ（）ネ，ネからフリカに

かけての主として熱にする KosTER in Blume ：帯広く分布（ 25 268， 1979）キク科植物で，九ー州南部が分の限であるにはタイやマレシアンシ布北。京大，イドネアで採集された標本が 30 枚ほどある。それのにら茎は上向き長く伸長し，よく分枝する。葉は有柄で，葉身が長卵形な

いでいし卵形，先端が鋭頭なし鋭尖頭でやや尾状となる傾向がある。 1 頭花あたりの小花数は

30〜50 8 〜12 〜35 のの個（舌状花が個，筒状花が 20 個）で，開花時頭花直径は 1．5cm 内外であ

る。総苞は卵形でさ〜片狭長，等長，畏 5 7mm ，上半分が草質でしばしば反転する。ところ

が，やでの 1 〜〜台湾琉球採集された標本中には，頭花あたりの小花数が 60 85個（舌状花が 14

15個筒状花が 45 〜 70 と〜つ，個）多く，開花直径も 2．5 3．Ocm と大きい頭花をけるものがあ

る。これらの標本では葉は卵形で先端が尾とならないがやや，状。質厚くなり，茎や花柄が太

く，全体としてすらっとした感じがない。

琉球や台にはが生るがアジアでののにことい湾両者育す，東南は後者ようなも出合うはな。

これをキダチハマルマののハマグ分布域北方寄りで分化した変種と考え，オオキダチグルマ

W ． biflora var ．． ryukyucnsis H KoYAMA と命名し，記載する。

つハマ葉がやや厚いため，小さい葉をけた個体はオオグルマにてるがそれ似来，とは葉脚がやや心形となるここと，総苞片が狭長卵形であるとなどで区別出来る。そのどちらとも判断し

かねる屋久島・琉球と小笠原産の標それぞれ 10 と 2 についてべこ本枚枚，花粉を調たとろ，そ

のい二 “ ” れら花粉はずれも倍体が示す正常なものではなかった。これはオオキダチハマグル

マとオオハマグルマとのこ間に雑種が生じているとを示唆するものである。

Wedelia biflora L DC ． var ． ryukyuensis H ． KoYAMA var ． nov ．（），

Wedeliα 6 o γα ： KITAM ・ Comp ．． III： 260 1942 ．．哲， Jap （）， p p

Caulis elongatus scandens ． Folia caulina longe i．5− ．5 cm petiolata， petiolis 2 longis laminis − − ・， ovatis 6 12 cm longis 3 8 cm latisJ ，， textura sub crasse chartacea ．

Gapitula ．− cm − − 2 5 3 diametro sub anthesi ， 4 8 cm longis． Flosculus60 85 pedunculis ，

flores radii uniseriales f（）eminei 14− 45 − ， 15， floresdiscihermaphroditi， 70．

Type cQllection ：， RYuKyu ： IsL Iriomote Ko ama et aL 177 Holotype in KYO Isotype in TNS ， Japan （？，）

Other specimens examiled ． KYusHu ： IsL Yakushima FtdZmoka Japan，（＆ Okamoto I190 KYO ）J IsL

Tokara 26 V 1953 Hori KYO （）． RYuKYu ； Okinawa （13V1923 Koidzumi KYO ； S． Sakaguchi KYO ）； IsL

Iriomote Kayama et al ． 276 KYO 「（， TNS ； Okada 磁 L eda 2103 KYO ）． Ghina ， TAIwAN ：剣潭山（4 V l904 Nagasawa KYO ・）（H ． KOYAMA ）

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