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ISSN OOOI-6799 Acta Phytotax. Geobot, 45 (2): 111-118 (1994)

Phytogeographic Notes on Some Aquatic Plants

in Yakutia, Eastern Siberia

HIDEKI TAKAHASHIi, TOSHIYUKI SAT02,

NIKITA G, SOLOMONOV3 and BORIS I. IVANOV3

iBotanic 2institute Garden, fuculty ofAgricutture, Hokkaido U}ziversity, Sapporo 060; ofLow 3Ytikutian Tlenrperature Science, Hbkkaido Uhiversity, Sturporo 060; Institute of Biotogy, Ytikutsk, Ybukutia, Russia

Abstract. In 1992- 1993, botanical surveys of Yakutia in eastern Siberia were carried out.

Aquatic plants were collected and new phytogeographic information was added. Utricularia macrorhiza Le Conte was collected from four sites in Yakutia, suggesting that this species is more widely distributed in northeastern Asia. Sparganium natans L. was recorded from east- cm Siberia. This eyidencc fi11s up thc gap in its circumpolar distribution. Thc prcscnt collec- tion of spirodela polyrhiza (L.) Schleid, is the second formal record from Yakutia, probably thc northcrn limit of this species in Asia.

Key words: aquatic plants, Lemnaceae, phytogeography, Siberia, spargani"m, Utricutaria, Yakutia.

Received September 5, J994; accapted October 22, 1994

Specimens of aquatic plants have been scarcely recorded from Siberia, and therefore their taxonomy, ecology and phytogeography are still insuW- ciently known. Our botanical field surveys were carried out in Yakutia, which is located in eastern Siberia, including the taiga to tundra regions in 1992 and 1993. Here we note the phytogeography of representatives of the Yakutian aquatic plants; Utricutaria, Elparlganium, and Lemnaceae.

Materials and Methods

Yakutia is situated in eastern Siberia (Fig. 1) and has a severe con- tinental climate and low diversity of vascular plants (Takahashi, 1994). In Yakutia the six main rivers from west to east are the Anabar, Olenek, Lena, Yana, Indigirka and Kolyma, all of which flow northwards into the Arctic Ocean. Our collection sites in 1992-1993 botanical surveys were the following five regions along the Lena and Yana river systems; from south to north Olekminsk, Yakutsk, Zhigansk, Verkhoyansk and Tiksi (Fig. 1). The posi- tion of the main city or town in each region is as follows: Olekminsk, 120030'E, 60030'N; Yakutsk, 129045'E, 62e05'N; Zhigansk, 1230E, 660N;

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FIG. 1. Map showing research sites in Yakutia.

Verkhoyansk, 133023'E, 67033'N; Tiksi, 128052'E, 71e38'N. We collected aquatic plants from various water habitats, and detailed locality and other data relevant to each collection are given below for the specimens ex- amined. Brief water condition at every site was indicated by the electric conductivity (EC; TOA CM-11P EC meter). EC values at 25eC were noted in each habitat when measured. All specimens are deposited in the Her- barium, Faculty of Agriculture, Hokkaido University (SAPT), and dupli- cates of some specimens are in SASY, TUS and H.

Results and Discussion Utricularia All plants of Yakutian Utric"taria collected in our surveys belong to U. vuigaris L. or U. macrorhiza Le Conte. Taylor (1989) recognized U. macrorhiza as a definite species distinguished from U. vuigaris in having markedly upwards curved distal part of spur of corolla with acute apex and internal glands on both dorsal and ventral surfaces. Based on these fea- tures all Yakutian specimens (Fig. 2) should be attributable to U. mac- rorhiza. Although Hulten (1968) regarded U. macrorhiza as a subspecies of U. vuigaris, he treated U. macroi:hiza as a North American taxon and Ul vul- garis as a Eurasian taxon. Later Hult6n and Fries (1986) demonstrated that

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FIG. 2. A dried inflorescence of Utric'ularia mac- rorhiza frem Olekminsk, 15662. The arrow in- dicates the long spur.

`American' U. macrorhiza occurs also in Kamtschatka and the Amur re- gion in eastern Asia. In addition to these localities, Taylor (1989) noted its occurrence in Sakhalin, the Ussuri region and westwards to the Altai in the USSR. But, until now specimens identified as U. macrorhiza have not been recorded from eastern Siberia. In our expeditions, U. macrorhiza was collected from the Olekminsk and Yakutsk regions along the Lena River and the Verkhoyansk region along the Yana River. This means that U. macrorhiza occurs more widely in eastern Siberia than expected.

Utriclllaria macrorhiza Le Conte U. vuigaris auct. non L., Tolmachev, Man. High. Pl. Yakut.: 440 (1974). Specimens examined, OLEKMINSK: Neryuktyai village, Biryuk set- tlement, a somewhat eutrophic, long and narrow pond (695-842 ptS cm'i, more than 40 cm in depth) by the settlement, July 2, 1993, fl., H. Taka- Bgs.h,gei,s,i･-g,5f,62.(,SA,I}IIIs[U,,,S)hg,A,K,U.Ifitt.Y.a,k,u,t,sli,git,ys.ftgtso,p,h,ig `,agi.1i90g,',fi2",P,'.3i.fl>8,o.",t }s",a.Rilxti'r6e"slx,ii,k,`9,I,gi,g･b.lllaTf,k,a-hgsBj£ Forest Station of Yakutsk Institute of Biology, July 8, 1993, fl., H. Taka- hashi et al. 15787 (SAPT). VERKHOYANSK: Verkhoyansk tewn, Appyt settlement, an oxbow lake (154-162 ptS cm-i) of the Yana River by the pasture, July 12, 1993, fl., H. Takahashi et al. 15938 (SAPT, TUS).

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F](}. 3. Dncd specimens et Yakutiun sparganium emersum. (a) from thc Yakutsk region. 12967; (b) the Zhigansk region. 13438. F indicates fernate and M ind)cates male heads. Spai:ganium Spat:ganium emers"m is the most common in Yakutia. It was found in somewhat disturbed or unstable places in more or less eutrophic standing water habitats. All specimens collected were erect and emergent, and some could tolerate periods of drought in summer. Cook and Nicholls (1986) re- ported that there is a clinal variation of stigma length and anther length for S. emersum from east to west in Eurasia. But plants of S. emersum even from eastern Siberia show variation in those characters. Specimens from the Yakutsk region (Fig. 3a) possess relatively small anthers (0.8-1.1 mm long) and stigmas (1.0-1.6 mm long) in comparison with the value by Cook and Nicholls (1986), and those from the Zhigansk region (Fig. 3b) possess larger anthers (1.1-1.3 mm long) and stigmas (1.5-2.0 mm long). Thus more specimens of S. emersum should be examined for ascertaining the geographical cline in Eurasia. SParlganium natans was first formally confirmed from eastern Siberia in this study. Plants which produced aerial leaves and grew in clear waters of an oligotrophic lake in northern taiga (Fig. 4) from the Zhigansk region, are determined as S. natans by the following features-all female heads are axillary, the male head is solitary and remote from the uppermost female head with 15-25 mm long internodes, and the lowest bract is scarcely longer than the inflorescence (Fig. 5). The occurrence of S. natans from eastern Siberia fi11s the gap in its circumpolar distribution. Although Tolmachev (1974) noted S. natans (as S. minimum) in his Yakutian manu- al, Hulten and Fries (1986) wrote a question mark on eastern Siberia in the distribution map of this species and Cook and Nicholls (1986) did not

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Fi(}. 4. A clear pond in which Sparganium natans grows, in the northern taiga at thc Zhigansk region,

record it from this region, Typical North American plants of S. natans has the lowermost female heads which are usually sessile or have less than 10 mm long peduncles (Harms, 1973). Yakutian plants of S. natans sometimes show up to 20 mm long peduncles in the lowermost female heads. This trait of a tendency to longer peduncles in the lowermost female heads is also found in Alaskan and northwestern Canadian populations (Harms, 1973),

Sparganium emersum Rehm. S. simplex Huds., Yuzepchuk in Komarov, Fl, USSR 1: 223 (1934); Tolmachev, Man. High. Pl. Yakut.: 37 (1974), Specimens examined. YAKUTSK: Pokrovsk-Lena Pillars, on a big island in the Lena River, a somewhat eutrophic, shallow pond (up to 40 cm in depth) by the pasture, August 12, 1992, fr., H. Takahashi et al. 13701 (SAPT); About 40 km E of Yakutsk city, a lake at the Alas Station of Yakutsk Institute of Biology, July 23, 1992, fir., H. Takahashi et al. 12967 (SAPT, SASY), ZHIGANSK: About 50 km S of Zhigansk town, the confluence of the Undyulyung River and thc Lena River, a dried up pond, August 5, 1992, flr., H. Takahashi et al. 13438 (SAPT, SASY, TUS, H), fr., H, Takahashi et al. 13441 (SAPT, SASy).

Sparganium natans L. S. minimum Hill, Yuzepchuk in Komarov, Fl. USSR 1: 225 (1934); Tolmachev, Man. High. Pl. Yakut.: 36 (1974). Specimens examined. ZHIGANSK: About 50 km S of Zhigansk town, the confluence of the Undyulyung River and the Lena River, a clear oli- gotrophic pond, August 5, 1992, emergent, flr., H. Takahashi et a}. 13409 (SAPT, SASY, TUS, H),

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Fie. 5. A dried specimen of Yakutian Sparganium natans from the Zhigansk rcgion, 13409. F, femalc heads; M, male heads.

Lemnaceae Tolmachev (1974) reported three species of Lemnaceae from Yakutia; SPirodela polyrhiza (L.) Schleid., Lemna trisulca L. and L. minor L. `L. Landolt (1986) regarded all the northern Asian minor' as L. turionijlera Landolt, but it is difficult to distinguish the two using dried specimens. In the present study, we use L. minor following the conservative treatment. In any case, we confirmed all the three species from Yakutia. Collection records of these species are quite rare from Yakutia (Land- olt, 1986), and those of Elpirodela are especially rare. Only one collection of Spirodela was observed for Yakutia in the distribution map of Hulten , and Fries (1986). The present collection of SPirodela is the second formal record from Yakutia and is near the northern limit of it in Asia (Hulten and Fries, 1986; Landolt, 1986). Similarly, the Lemna minor population of Verkhoyansk may be near the northern limit of this species in Asia.

Lemna minor L. ? L. turionijl7ra Landott, Aquat. Bot, 1: 355 (1975); Fam, Lemn. 1: 443 (1986). 'nfti,'hY.:;5dU,T.S.K,i'.Xain".t,Zkhiifi:'tY6..e,"grgPfh,'hC.P.O,O.'g (242oS-P8giiM,e".Sse,XiilMtY River, July 18, 1993, st., H. Takahashi et al. 16090 (SAPT); About 30 km NWForest of Yakutsk city, a pond (198-229 ptS cm-i) at the Spaskayapad Station of Yakutsk Institute of Biology, July 21, 1992, st., H. Taka- hashi et al. 12793B (SAPT), July 8, 1993, st., H. Takahashi 15789 (SAPT). VERKHOYANSK: Verkhoyansk town, a somewhat eutrophic, long and

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narrow oxbow pond within the town, July 10, 1993, st., H. Takahashi et al. 15829 (SAPT). Lemna trisulca L.

,,..g.P8CIM,e.".',e.x,a.I?ig.eg,6,O,L,,?kl.l#gN.i",:..N,,esy."k,`.y.ak?gi6gg-e,･,gi7gk,,\{g}; more than 40 cm in depth) by the settlement, July 2, 1993, st., H. Taka- hashi et al. 15663 (SAPT, TUS). YAKUTSK: Yakutsk city, eutrophic pools (2420-3310 ptS cm-i) with a bad smell on marshlike banks of the g,eri,iiiGVSi>r6;"iCX,ii,k,'29,,3,',S.t'p.IilaT,a,k,a-h,a8Bi2t,aJi.IigOg,2,kS.",P,T.3i.",,b,O.",` Forest Station of Yakutsk Institute of Biology, July 21, 1992, st., H. Taka- hashi et al. 12792 (SAPT), July 8, l993, st., H. Takahashi et al. 15788 (SAPT, TUS); About 40 km NE of Yakutsk city, a pond at Ogrodtakh, July 27, 1992, st., H. Takahashi et al. 13113 (SAPT). Spirodela polyrhiza (L.) Schleid. Specimens examined. YAKUTSK: About 30 km NW of Yakutsk city, a pond (198-229 ptS cm-i) at the Spaskayapad Forest Station of Yakutsk Institute of Biology, July 21, 1992, st., H. Takahashi et al. 12791 (SAPT).

The ficld surveys of 1992-1993 in Yakutia w ¢ re assisted by the following scientists: Gaku Kudo, Gen Takaku, Hideaki Watabe from Japan, and Rornan V. Desyatkin, Trophim K. Maximov, Danil P. Yefremov, Kostya Volotovsky, Izabella P. Matveeva, N. Danilova, Pyotr A. Timofeyev, N. N. Vinokurov, A, I, Avcrensky, Alexandor Kononov

and Ayal Maximov from Russia, and Dr. Shigeru We are gratefu1 to Dr. Hugh Tan, National University of Singapore Ogawa, Joctsu University of Education for their critical reading and valuable comments on

the manuscript. This study was financially supported by a Grant-in-Aid for International Scientific Re- `tCLimatic search, Project influences to cryosphere-biosphcre in Siberian permafrost" (No. 04041014) to Dr. Masami Fukuda, ILTS Qf Hokkaido University, from the Ministry of

Education, Science and Culture, Japan.

References

Cook, C. D. K, and M. S. Nicholls, 1986. A monographic study ef the genus Sparganium (Sparganiaceae). Part 1, Subgenus Xanthosparganium Helmberg. Bot, Helv. 96: 213-267. Harms V, L, 1973. Taxonomic studies of North American Spai:ganium, I, S, hyperboreum and S. minimum. Can, J. Bot, 51: 1629-1641. Hulten, E. 1968. Rora of Alaska and neighboring territories. Stanford University Press, Stanford. and M. Fries. 1986. Atlas of north European vascular plants, vols, 1-3. Koeltz Scientific Books, K6nigstcin. Landolt, E, 1986. The family of Lemnaccae-a monographic study, vol, 1. VerOff, Geobot. Inst. ETH, Stiftung, Rilbcl, ZUrich, 71 Hcft. pp. 1-566, Takahashi, H, 1994. Phytogcography of vascular plants in Yakutia (Sakha), Proc. Jap. Soc. Pl. Tax. 10: 21-33. (In Japanesc with English summary), Taylor, P. 1989. Thc genus Utricutaria-a taxonomic monograph, Kew Bulletin Additional Scrics XIV, Her Majesty's Stationery Office, London,

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． ． Tolmachcv ， A 1 （Editor 1974． Manual of higher of Yakutja ． Nauka Publ． Co ． ） plants ， Siberian Division Novosibirsk ， ．（ln Russian）． Yuzepchuk S． V ．1934． Sparganiaceae． ln V ． ed 正ora ， L Komarov （ ．）， F of the USSR 1； − ’ ， 216 229．lzdate1stvo Akademil Nauk SSSR ， Lcningrad．

摘 要

1 ・ 2 ・ ・ 1 ー 高橋 英 樹 佐 藤 利 幸 N ．G ． Solomonov3 B ．1． Ivanov3 ： ベ ア 東 シ 丿 ， ヤ ク チ ア産 水 生 植物 の 植物地 理 ー − 東 シ ベ リ ア ヤ ク チ ア の 植 物 調査 （1992 1993）で 採 集 した 水 生 植 物 の うち，植 物 地 理学

つ い 〃 ー 的 に 興 味 の あ る種 に て 報 告 し た 。 Utriculariamacrorhiza をヤ ク チ ア の 4 地点 か ら採 し た い れ に い で 集 。 ず も人家 近 富 栄養化 した池 沼 あ っ た。本種 は Hult6n（1968）に よ り， タ

ヌ キ モ ひ vulgaris の 北 ア メ リ カ 産 亜 と もみ ら れ て い た が の カ ム カ ム ー 種 ， そ 後 チ ャ ッ ， ア

ル 地方 な どで も見 つ か っ て い た 。 の 査 に よ u ． macrorhiza は こ で 今 回 調 り， ， れ ま 考 え られ

て い た よ に い りも さ ら 広 く，北 東 ア ジ ア に分 布 し て る 事 が 明 ら か と な っ た 。 Spargani“m

natans ジ ガ ン ス の の ベ を ク 付近 貧栄 養 自然湖 沼 か ら採集 し た 。 こ れ に よ り，東 シ リ ア で 欠

し て い た が つ Spirodeta ー 落 周 北 極 分布 な が っ た 。 ウ キ クサ potyrhizaをヤ ク ッ ク の 人家 近 1 くの や や 富栄 養化 した 湖沼 か ら採 集 した 。 ア ジ ア に お る北 限 に 近 い で る 〔〕 け 集 団 あ 。 （〒 60 2 札 幌市 中央 区北 3 条 西 8 丁 目 北 海道 大 学農 学部 附属植 物 園 ； 〒 060 札幌 市北 区北 19 条 3 ー ー 西 8 丁 目 北 海道 大学 低 温 究 ロ シ ア ヤ ク チ ア ヤ ク ツ ー 科学研 所 ； （ ） ， ク 市 ， ヤ ク ト 生物学 研究 所 ）

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