Nat Hist Res.. Vol. 9 NO.2: 45 -62. March 2007

T. F. Allen's C ollection of Characeae in Herbarium of AIgology, Natural History Museum and Institute, Chiba (CBM) , Japan

Masahiko Miyata 1) and Jenny Bryant Moore2)

。 Department of Plant Sciences, Natural History Museum and Institute, Chiba

955-2 Aoba-cho, Chucはu， Chiba 26()..8682, ]apan

E-mail: mi砂yata@chiba-ml凹II山us詑E印.OIぽr.j2剖) Department of Botany, the Natural History Museum

Cromwell Road, London SW7 5BD, UK

Abstract Characeae ]aponicae Exsiccatae by T. F. Allen (1895) preserved in the Herbarium of Algology,

Natural History Museum and Institute, Chiba (CBM) , ]apan, which consist of 1 species and 3 varieties in

Chara and 9 species and 1 form in Nitella. Especially, of those, Ni，信lla japonica Allen, N. orie1ltalis Allen, N.

plllchella Allen, N, pSlIcicostata Allen and N. slIblllcens Allen are important specimens following the type mateｭ

rial of species which is described as new species in ]apan by T.F.Allen. These authentic specimens are essenｭ

tial for taxonomy of Characeae in ]apan, as standard specimens.

Key words: Characeae ]aponicae Exsiccatae, T. F. Allen, taxonomy, Characeae, Nitel/a, Chara, ]apan, CBM.

Characeae J aponicae Exsiccatae

Characeae ]aponicae Exsiccatae (Allen, 1895) deｭ

posited in the Herbarium, Natural History Museum

and Institute, Chiba (CBM) , ]apan, which consist of

Chara fragilis Desv.. 2 specimens of Chara gymllopi，砂S

A.Braun var. "alJフha"， ABraun, 2 specimens of Nitella

coronaωZiz. ， 2 specimens of Nitella japonica AlIen,

Nitella orientalis AlIen, Nitella Plllchellan AlIen, Nitella

pallcicostata AlIen, Nitella Plllchella Allen, Nitella

sllblllcens Allen and Nitella oligospira A.Braun. Espeｭ

cially, of those, Nitella japonica Allen, N. orientalis Allen,

N. Plllchella Allen, N. pSllcicostata Allen and N. slIblllcells

AlIen are important specimens following the type maｭ

terial of species which is described as new species in

]apan by T. F. AlIen (1894, 1895). This exsiccata is on eternalloan 仕om the University Museum, the Univerｭ

sity of Tokyo (TI). These specimens are essential for

taxonomy of Characeae in ] apan, as standard speciｭ

mens. Each taxon identified by T. F. AlIen in the list

are enumerated (Table 1).

Taxonomy of Charophytes

Charophytes (Charophyceae, Charales) are among

the largest and most structurally complex of the green

algae. The main axis have, at intervale, whorls of short

lateral branchlets so that the plants bear a superficial

resemblance to Eqllisetllm.

In 77 A.D. Pliny the Elder (Plinius 23-79 A.D.) in

“ Historie Naturalis' assinged the green alga Chara

to the land plants as an aquatic horse-tail (Plinius,

1741). Since then systematic position of Chara and reｭ

lated algae within the plants has been controversial.

Linnaeus (1753) includ巴d four species of

Charophyta in a single genus, Chara, in his “ Species

Plantarum' , namely, C. tomentosa, C. vlllgaris, C. hispida

and C. flexilis , and these names are unchanged today,

except that C. flexilis is now in Nitella. After that there

are new genera, species, varieties and forms, at preｭ

sentωtalling over 300 t拡a 泊 the world, on the taxoｭ

nomic limits of the Linnean species (Wood, 1962; Wood

and Imahori, 1964, 1965; ]ohn and Moore, 1987).

During the 19th century, five additional genera

were recognized Nitella (Agard, 1824) , Lychnothamllnlls

(Ruprecht, 1剖5) ， Tolypella (A. Braun, 1849) ,

Lamprothamllnlls (A. Braun, 1882) and Nitellopsis (Hy,

1889). In 1916, ]. Groves substituted 也.e name

Lamprothamnillm for Lamprothamnlls as the latter had

been used for a genus of Rubiaceae. Representatives of

all these genera are found in ]apan without

Lychllothamlllllls and Toか'pella.

20 years later, Karsten et al. (1936) recognized a

common ancestry of the green plants (i.e.,

Chlorophyceae, Bryophyta and Pteridophyta) , but

Characeae and Conjugatophyceae were still excluded

from the green algae (Volker et al.. 1997).

It is now generally accepted that these genera make

up a highly specialized family, Characeae, within the

Chlorophyta, although their position in relation to

other green algae, this group have treated as a

ロ-u

A告

Masahiko Miyata and Jenny B. Moore

Table 1. List of specimens of Characeae Japonicae Exsiccatae (T. F. Allen, 1895) , (1) the number of specimens in Characeae Japonicae Exsiccatae (2) the scientific name (3) the collection site (4) the plate number.

Allen's No. Scientific name

Chara fragilis 0ε:SV.

2 Nitella coronata Ziz.

3 Nitella corollata Ziz.

4 Nitella Japollic，αAllen

7 Nitella orie1l/alis Allen

8 Nitella Plllchellan Allen

9 Nitella Japollica Allen

10 Chω'a gylllllopitys A.Braun var. "alp/lll" A目Braun

11 Chara gymllopityδ A.Braun var. "alp/w" A.Braun

12 Nitella paucicostata Allen

14 Nitella Plllchella Allen

16 Nitella sublucells Allen

17 Nitella oligospira A.Braun

separate division as Charophyta. But Mattox and

Stewart (1984) suggests that the Chlorophyta should

be divided into two classes, the Chlorophyceae and the

Charophyceae, the second class organizing the

restwhile members of the Chlorophyceae such as

Charales, Chlorokybales, Coleochaetales, Klebsorｭ

midales and Zygnematales. The term Charophyte or

stonewort is used as a common-name for the memｭ

bers of the family Characeae, which is divided into

two tribes, Charaeae (Chαra， LamprothαmnÍwn，

Lychnothamunlls, Nitellopsis) and Nitelleae (Nitella,

Tolypella)

1n history of taxonomy of ]apanese Characeae,

Kotaro Saida (1887) firstly described 2 sp巴cies of

Chara (Chara brallnii Gmelin，仁 foetida Braun) and 5

species of Nitella (N. flexilis Agardh, N. gracilis (=

Tolypella graci!is) Agardh, N. opaca Agardh, N.

tenllissima Kuetzing, N. translllcens Agardh) and T. F

Allen (1894, 1895, 1898) described 3 species of Chara

(仁 coronata Ziz. (=仁 braunii) ， C. fragilis Desv. (= C.

globularis Thuiller) , C. gymnopiか's A. Br, var, 'alpha' A.

Br.) and 17 species, 2 varieties and 4 forms of Nitella

(N. expansa Allen, N. gracillima Allen, N. japollica Allen.

N. oliogospira A. Br., N. orientalis Allen, N.目 mucronata

Miq. var. tenllior A. Br., N. II1l1ltipartita Allen, N.

mllltipartita Allen form. intermedia Allen, N. mllltipartita

Allen form. sllbsrecta Allen, N. mllltipartita Allen form.

transil仲間a Allen, N. pseudoflabellata A. Br.. N.

pseudoflabellata A. Br. form. testaglabra Allen, N.

Collection site Plate number

Province Tokio No.l

Province Mikawa No.2

Province Tokio NO.3

Province Tokio No.4

District of Toyohashi, and Province Ise NO.5

Province Ise, District Yamakami, pond Kaida-ike No.6

Ise,Yamagami NO.7

Province Mikawa Tennu pond, Toyohashi NO.8

Province Tokio, Shinbashi pond No目9

Province Tokio No.lO

Province Mikawa, Tennu poid, Toyohashi and No.ll Province Tokio Shinbashi pond

Province Sagami, Localty Koduz No.l2

Province Mikawa, Seishin pond No.l3

46

pseudoflabel!ata A. Br. var. imeprialis Allen , N

pselldoflabellata A. Br. var. ramuscllla Allen, N.

pallcicostata Allen, N. Plllchella Allen, N. rigida Allen, N.

saitoiana Allen, N. sllbgloll1erata A. Br. var. japonica

Allen, N. sllblllcens Allen, N. tanakaiana Allen) from

]apan, on the basis of specimens collected by Kenzo

Saito and Y oshio Tanaka. Of those, 12 species, 2 varieｭ

ties and 4 forms were new to science, and 4 species

and 1 varieties were new to ]apan. As a results of his

study. Characeae ]aponicae Exsiccatae. which consist

of 17 specimens, 1 species and 2 varieties of Chara and

7 species and 2 varieties of Nitella. has distributed to

several Europe herbarium such as the British Muｭ

seum (Natural History) and Tokyo Imperial Univerｭ

sity (J apan).

Tomitaro Makino (1929) had called attention to

the study of Characeae of ]apan on T. F. Allen's reｭ

ports. (1894, 1895, 1898) Then Walter Migula (1935)

described 3 species of Chara and 8 species and 1 form

of Nitella on specimens coll巴cted by Shigeru Miki. Of

those , N. dill1oqJ!w Migula and N. polyglochill A. Br.

form. japollica ν1igula were new to science. and N.

aCllmlllat，α A. Br. , N. dispersa A. Br. , N. flaccida A. Br. , N.

hyalina A. Br. and N. polyglochin A. Br. form. japonica

Migula were new to Japan. After that Hideo Morioka

(1940, 1941a, 1941 b) reported 5 species of Chara and

11 species of Nitella. Of those, 7 species (爪 coreana

Morioka collected in Korea, N. fallosa Morioka. N.

gracilens Morioka, N. lIloniliforll1is Morioka, N.

II1l1sashiensis Morioka, N. spicifol7nis Morioka, N.

T. F. AJJen's C oJJection of ]apanese Characeae

sllblllcens Morioka) were new to science and 3 species

(c. corallina Willdenow, C. benthamii A. Br., c. zevlanica

Willdenow) were new to ]apan. Afterwards, Kozo

Imahori (1964) described 8 species of Chara and 47

species of Nitella containing 14 species as new to scト

ence (N. allenii Imahori, N. axilliformis Imahori, N. crispa Imahori, N. erecta Imahori, N. horikawae Imahori, N. inversαImahori， N. pusilla Imahori. N. robllsta

1mahori, N. sejuncta 1mahori, N. shinii 1mahori, N. spillosa 1amahori, N. stabilis 1mahori, N. stricta 1mahori, N. tllyamae 1mahori). Then Hideo Kasaki (1964) reｭ

ported 5 species, 7 varieties and 1 subspecies of Chara,

21 species and 35 varieties of Nitella , 1 species of

Nitelopsis and 1 species of Lamprotamnillm. Of those, 4

varieties and 1 form are new to science (N. flexilis var.

flexilis Kasaki, N.目 shinii var. graciliformis Kasaki, N. pselldoflabellata form. macrophylla Kasaki，仁 bellthalllii

var. longicorollαta Kasaki, C. benthalllii var.

brevibr，αcteata Kasaki and c. globlllaris var. hakonellsis

Kasaki). Kozo 1mahori and Hideo Kasaki (1977) reｭ

viewed ]apanese species of Charophyte with 4 genera

(Chara , Lalllprothamnilllll, Nitella , Nitellopsis) , 26 speｭ

cies, 42 varieties and 1 subspecies.

With the development and broader application of

molecular genetic techniques, the number of investigaｭ

tions increased especially in the late 1990s. McCourt et

al. (1999) investigated systematic relations between

Charophytes by analyzing the chlororplast-encode

large subunits of the Rubisco均gene (rbcL) in recent as

well as extinct species, whereas Meiers et al. (1999)

studied the relationships between the subsections of

the genus Chara by means of small subunit (18S) of

the ribosomal DNA gene. McCourt et al.. (1999)

could prove the monophyletic origin of the tribe

Characeae. After that rbcL-based data segeregated

Lamprotanmillm from Chara (McCourt et (11.. 1999).

whereas an 18S-rDNA analysis placed Lamprotanmillm

within from Chara (Meiers et al. , 1999). Furthermore

investigation on the intra-species level, either by

AFLP (Mannschreck et al., 2002) or by microsatellite

analysis (Schaible, 1999) , a tool especially valuable for

discrimination between ecotypes and morphological

acclimination, have become a field of increasing interｭ

est. Recently taxonomic reexamination has been done

by comparative morphology and multiple DNA

marker analysis (Sakayama et al. , 2005).

Under these circumstances, it is still necessary to

study morphologically species of Charophyte on type

methods (McNeil et a l., 2006). The specimens of

“ Characeae ]aponicae Exsiccatae (Allen, 1895)" is

essential voucher specimens in taxonomy of

Charophyte in ]apan

47

Acknowledgements

Authors give great thanks to the late Professor

Kozo 1mahori for giving informations, where is the

type materials described by 1mahori, K. and distribuｭtion of authentic specimens in ]apan. This contribution

is a part of results of The Japan-United Kingdom

Charophyta Dαta Base Project by financial support from

the Daiwa Foundation Small Grants in the Daiawaｭ

Anglo Foundation (2004-2006) and The Study 01 Natural History 01 Boso Peninsula, Flora 01 AIgae. Then authors extend our thanks to Professor Hideaki Ohba, Univerｭ

sity of Tokyo, for the eternal loan of Characeae

]aponica巴 Exsiccatae (Allen, 1895) to Natural History

Museum and 1nstitute, Chiba. Then to the memory of

Dr.Hid巴o Kasaki (Morioka) , a pioneer of taxonomy of

Charophyte in ]apan, who was dead on March 27th ,

2007.

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Allen, T. F. 1894. ]apanese Characeae 1. Bull. Torrey

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lassenen Manuskripten A. Brauns herausgegeben

von Dr. O. Nordstedt. Abh. Kgl. Akad. Wiss. Berlin

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(Accepted 10 March 2007)

千葉県立中央博物館・藻類標本室 (CBM)

が保管する日本産シャジクモ科・

T. F. Allen (1895) コレクション

宮田昌彦 1) ・ジ‘ェニー・ブライアント・ムーア 2 )

1) 千葉県立中央博物館・植物学研究科

干 260・8682 千葉市中央区青葉町 955-2

E-mail: miyata@chiba-muse.or.jp

2 )英国自然史博物館・植物研究部門

クロムウエルロード，ロンドン SW75BD. 英国

日本産シャジクモ科エキシカータ (T. F. Allen, 1895) を千葉県立中央博物館・ i築類標本室 (CBM) が保管し

ている.当標本群は，シャジクモ属 1 種 3 変種，フラス

コモ属 9 稜 I 品種からなる.特に ， Nitella japonica

Allen, N. orientalis Allen, N. pulchellαAllen， N,

psucicostata Allen そして N. sublucens Alle日の標本は，

日本を基準産地として T. F. Allen が原記載した種の

基準標本に準じる証拠標本であり，日本におけるシヤ

ジクモ科の分類学的な研究のための重要参照標本であ

る.

T. F. Allen's Collection of ]apanese Characeae

Explanation of Plates (1 -13)

(habitus with an original label)

Plat巴 1 No.1 Chara fragilis Desv.

Plate 2 No.2 Nitella coronata Ziz.

Plate 3 No.3 Nitella coronata Ziz.

Plate 4 No.4 Nitella japonica Allen

Plate 5 No.7 Nitella orientalis Allen

Plate 6 No.8 Nitella pulchellan Allen

Plate 7 No.9 Nitella japonica Allen

Plate 8 No.10 Chara gymnopitys A.Braun var. "alpha" A.Braun

Plate 9 No.11 Chara gymnopitys A.Braun var. "alpha" A.Braun

Plate 10: No. 12 Nitella paucicostata Allen

Plate 11: No. 14 Nitella pulchella Allen

Plate 12: No. 16 Nitella sublucens Allen

Plate 13: No. 17 Nitella oligospira A.Braun

-49 一

ル'lasa hi ko Miyata and Jenny B. Moore

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CHARACEAE JAPONICAE EXSICCATAE. -NO.2.

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CHARACEAE JAPON1CAE EXSICCATAE.

No.7

Ni色ella orien也li.守 Allen.

Distriot of 百oyohashん

and 舟ovìnce Ise.

Science College, Irnperial University, Japan. 貴重 科大 祭ーーーーー・--=-O~唱・ーーー一一

Nirella orienralis Allen

anｭ-hd

No.7

1'. F. Allen's C oLlection of ]apanese Characeae

Plate 6

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Province IB~. District Yamakami. pond Kaida.ike.

Science Oollege, Imperial University, Japan・

理科大 事

ー--.恥~，~ペ‘・ー一一一一ー

Nitella fJulchellan Allen

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No.8

Masahiko Miyata and ]enny B. Moore

Plate 7

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Science College, 1 m卵rial Univ回i帆 Japan.

理科大皐

H

O. 9 Nitella japonicC/ Allen

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FHU

T. F. A II巴 11's C ollection of J apanese Characeae

Plate 8

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N Chara gy田Dopity.， .A. Br.,

町，. "alpha". A. Br.

Province Mikawa.

Ttnnu pond, Toyohsahi.

Science College, ImperiaJ Unive悶ity， J，晶伊n.理科大串

NO.10 Chara gymnopirys A.Braun var. "alpha" A.Braun

ウJFD

1-,1!asahiko .Miyata and ]enny B. Moore

j4 61W 身

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Plate 9

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了 F. AlLEN

CHARACEAE JAPON'CAE EXSICCATAE.

No.11

Chara gy四D.opitys， A. Br. , "はr. "alpha'二 .d .Br

Shinbashi pond.

Science College, 1 mperial U niversiも'y， Japan.

理科大穆一一一ー骨-=e~叫・--

PrOl,ince Tok;o.

Chw司a gym170pitys A.Braul1 var. "alpha." A.Braul1

。《U

Fhd

NO.ll

T. F. A II巴n's Collection of japanese Characeae

Plate 10

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Scicnce College, Imperinl University, Japan. 理科大串

一一一一軒~OC>-<.・c一一一ー

2

τ F' . ALLEN

CHARACEAE JAPON1CAE EXSICCATAε. -No. 12

Nit・Ua. paucicoetata., All~ lI

Nitella paucicostata Allen

Qd

F「U

舟ovince Tok;o.

O. 12

色、王、、ー-.

可

Masahiko Miyata and Jenny B. Moore

Plate 11

T, f". AlLEN

CHAAACEAE JAPONICAE EXS1CCA.TAE.

Nn. I ・ e

NiteUa pu1eh・11la.， .. tIl.帽

Pro~ince Mikowa. T~nnu ρond，

Toyoh�M.

.nd 舟ov;noe T04;0. ShinbB~hi pond.

Science College, Imperial 可恥n凶町問i討同附ve附自町削r理科大'

No. 14 Nilella pulchella Allen

-60 -

T. F. AlIen's C ollection of J ap an巴se Characeae

Plate 12

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CIiARAOEAE JAPON/CAE g)(S/CCATAE. 一ーーーーーーー

No. 16 ~

Nitella ・ublu~1l・， ~IIer1

Province 8a.9aml, Locality Koduz.

Sci邸ce College, Imperial University, Japan. 理科大事ーーーー唄ー・ÞO-‘00--<"・ーーーーーー

NO.16 Nitella sublucens AUen

l

phv

Masahiko Miyata and ]enny B. Moore

Plate 13

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Provinoe Milcawa.

Seishin ρond.

Science Oollege, Imperial University, Japan・

理科大皐ー一ーー一司惨ト~OC-<.. ・ーーー一ーー

No. 17 Nitella oligospira A. Braun

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Phu