Phytogeographic Notes on Some Aquatic Plants

The Japanese Society for Plant Systematics

The JapaneseSociety forPlant Systematics

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 Garden, fuculty

ofAgricutture,

^Hokkaido U}ziversity,Sapporo 060;^2institute

ofLow

TlenrperatureScience,Hbkkaido Uhiversity,

Sturporo

060; ^3Ytikutian Institute

of

^Biotogy,

Ytikutsk, Ybukutia, Russia

Abstract. In 1992-1993,botanicalsurveys of Yakutiain eastern Siberiawere carried out.

Aquatic plants^{were collected and new} phytogeographic informationwas 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.Thiseyidencc fi11sup thcgap initscircumpolar distribution.Thc prcscnt^collec- tion of

spirodela

polyrhiza

(L.) ^{Schleid, is}

^{the second}

^formal

^record

^{from Yakutia,}

probably thc northcrn limitof thisspecies inAsia.

Key words: aquatic plants,Lemnaceae, phytogeography, Siberia,

spargani"m,

Utricutaria,

Yakutia.

ReceivedSeptember5,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;

112 Acta Phytotax. Geobot.

f

"

Vol. 45

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

The Japanese Society for Plant Systematics

The JapaneseSociety for Plant Systematics

December

1994 TAKAHASHI et al.: Aquatic

plants in Siberia 113

FIG.

2. A dried inflorescence

of Utric'ulariamac- ^rorhiza frem Olekminsk, 15662.The arrow

in-

dicatesthe longspur.

`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

}s",a.Rilxti'r6e"slx,ii,k,`9,I,gi,g･b.lllaTf,k,a-hgsBj £ `,agi.1i90g,',fi2",P,'.3i.fl>8,o.",t

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).

114 Acta Phytotax. GeobQt. Vol. 45

tlle//･tL･

l･i,l:11111'1i

i.111.1

.11

F](}.3. Dncd specimens et Yakutiun

sparganium

^emersum.

(a)

^{from thc Yakutsk region.}

^the Zhigansk region. 13438.F indicates

fernate

and M ind)catesmale heads.

12967; (b)

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

The Japanese Society for Plant Systematics

The JapaneseSociety for Plant Systematics

December 1994 TAKAHASHI

etaL :

Aquatic plants in Siberia 115

Fi(}.4. A clear

pond

^inwhich

Sparganium

natans grows,inthe northern taigaat 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),}

116 Acta Phytotax. Geobot, Vol. 45

^{,,,, ,,･ 1..1}

1･,i/,1}//,l,l,l,,

'

t/t///ttttttttt

Fie. 5.

A driedspecimen of Yakutian

Sparganium

^natans

from

the

Zhigansk

rcgion, 13409.

F,

femalcheads;M, male

heads.

Lemnaceae

^Tolmachev (1974)

^reported

^three

^{species of}

^{Lemnaceae from Yakutia;}

SPirodela polyrhiza (L.) ^{Schleid., Lemna trisulca L. and L.}

^minor

^L.

Landolt (1986)

^{regarded all}

^the

^northern

^Asian

^{`L. 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).

(242oS-P8giiM,e".Sse,XiilMtY

'nfti,'hY.:;5dU,T.S.K,i'.Xain".t,Zkhiifi: 'tY6..e,"grgPfh,'hC.P.O,O.'g

River, July 18, 1993,

st.,

H. Takahashi

et al.

16090 (SAPT); ^{About 30 km}

^of

^Yakutsk

^{city, a}

^pond (198-229 ^ptS

^{cm-i) at}

^the Spaskayapad NWForest

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

,

The Japanese Society for Plant Systematics

The JapaneseSociety forPlant Systematics

Decembcr1994 TAKAHASHI

^{et aL :}

Aquatic plants in Siberia 117

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

^{ficldsurveys 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,

We

^are

gratefu1

^to

Dr. Hugh

^Tan,

National University

of

Singapore

and

Dr. Shigeru 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-

search,

Project

^`tCLimatic

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,}

118

Acta 　Phytotax

．

　

Geobot ． Vo1 ，45 Tolmachcv

_，　

A ．1．

_（

Editor）1974．



Manual

　of　

higher

　

plants

^of　Yakutja

．

　Nauka 　

Publ．



Co ．

_，

　　Siberian



Division

_，　

Novosibirsk．（ln



Russian）．

Yuzepchuk

_，　

S．

　

V ．1934．

　

Sparganiaceae．

　

ln

　V

．

　

L

　

Komarov

_（ed

．

_），　

F

正ora　of

the



USSR

_，

1

；

　　216 − 229．lzdate1’

^stvo　

Akademil

　

Nauk

　

SSSR ，



Lcningrad．

摘 　 要

高 橋 英 樹¹

・

_{佐 藤 利 幸}²

・ N ． G ．



Solomonov3 ・B ． 1．



Ivanov3

：東 シ^{ベ 1}丿^ア， ヤ ク

ー

_チ

_ア

_{産 水 生} 植 物

^の

植 物 地 理

　 ^{東 シ}

^ベ

^リ

^ア

^ヤ

^ク

^{ー 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 的

に 興

味のあ

る 種

^{につ い て}報 告 ^し

^た

。

Utricularia

^〃macrorhiza

を ヤ

ク

ー

チ

ア の

4

_{地 点}

か

ら採

集 ^{し た}

。 い ず

^れ

も人 家

^に

近^い富 栄 養 化

し

た 池 沼^で 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

種

^{と も}

^み

^ら

^{れ てい た が}，

そ

^の

後

^カム

^チ

^{ャ ッ カ}

，

^アム

ー

ル地 方 な ど^で

も

見^つ

か

っ て^い

た 。

今 回^の調

^査

^に

^よ

り

，u ．

^{macrorhiza は}， こ

れ ま

^で

考

え

ら れ

て ^い

た

よ り も

さ ら

に広

く ， ^北

東^ア

^ジ

^ア

^に

分 布

^し

^{てい る}事

^が

明^ら

か と な

っ

た

。

Spargani

“m natans を

^{ジ ガ}

^{ン ス}

ク

付 近^の貧 栄 養^の 自然

湖 沼 か ら 採

集

^{し た}

。

こ れ に よ

り

，

東^シベ

リ

ア で欠 落

^し

^てい

^た

周 北 極 分 布^{が つ}

な が

っ

た

。

ウ キ ク サ Spirodeta

　

potyrhiza

^{を ヤ}

^ク ー

_ッ

_ク

_{の人 家 近} く^{の や や}富 栄 養 化 し

^た

湖 沼 か ら採 集

し た 。

ア

ジ

ア

に お

け

^{る 北 限 に 近}

^い 集 団^であ

^る

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