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Edaphologia, No. 91:21-30, December 28, 2012 21

Cubarisiriomotensis, a junior synonym of the species Cubaris murina (Crustacea: Isopoda:pantropicalOniscidea)

Shigenori Karasawa

Fkeculty ofEducation, Fleikuokti Uhiversity ofEducation, Fleikuoha 811-4192, Jtu)an

Received: 1 7 December 20 1 1 ; Accepted: 6 August 201 2

Abstract This study was conducted to reassess the taxonomic status of Cubaris iriomotensis (Nunomura, 1990).

IYpe series from Iriomotejima Island, southern Ryukyu in Japan, and new specimens collected from Iriomotejima

and Ishigakijima Islands were examined. Taxonomic assessrnent of the specimens included detailed morphological

observations using a scanning electronic microscope (SEM). Specimens from the two islands closely correspond with the morphology of C. murina Brandt, 1833, a species widely distributed in the tropics, Thus, C iriomotensis can be

regarded as a synonym of C. murina.

Key vvords: Armadillidae, Cubaris iriomotensis,Cubaris murina, Ryukyu Islands

1998). No morphological comparisons have been made Introduction between C iriomotensis and other species within the genus

Terrestrial isopods are one ofthe most abundant Cubaris since Nunomura (1999a) transferred S. iriomotensis

macrofaunal groups fbund with in the litter layer of the to the genus Cubaris, however. The morphologies of male

subtropical forests of Japan, and play an important role pleopod 1 and pereiopod 1 ofC. iriomotensis, as described by in the decomposition of leaf litter (Karasawa, 2009). The Nunomura (1990), are similar to those of C. murina; hence family Armadillidae Brandt and Ratzeburg, 1831 (Isopoda: these similarities suggest that C iriomotensis may be ajunior

Oniscidea) is diverse and includes about 600 species, synonym of C. murina.

distributed throughout tropical and temperate zones (Taiti To investigate this further I collected new Cubaris

et aL, 1998). However, taxonomy within the Armadillidae specimens from the type-Iocality, Oraha in Iriomotejima

remains unclear. Reassessment of the taxonornic status Island, and Ishigakijima Island, a neighbor of Iriomotejima

of several specific taxa in the family is important fbr Island. Morphological approaches using a scanning electronic

understanding terrestrial isopod diversity and also for microscope (SEM) were used to identify the species collected

understanding soil ecosystems. from Ishigakijima Island and to elucidate the taxonomic status

Nunomura (1990) described one isopod species, ofthe C iriomotensis specimens from type-locality, Moreover, sphaerillo iriomotensis Nunomura, 1990, from Iriomotejima the morphology of the holotype of C. iriomotensis and the

Island in the southern part of the Ryukyu Archipelago, new specimens collected from Iriomotejima and Ishigakijima

southwestern Japan. This species was later transferred to the Islands were examined and compared.

genus Cubaris Brandt, 1833 by Nunomura (1999a). In the Materials and Methods world catalogue of terrestrial isopods (Schmalfuss. 2004), C

iriomotensis is considered endemic to Iriomotejima Island. Samplecollection

Although most species of Cubaris have limited distributions Specirnens were collectecl from leaf litter and/or under

because oftheir low migration ability, one species, C, murina stones and decaying logs, using aspirators. Specimens were

Brandt, 1833, is widespread throughout the tropics (Taiti et preserved in 99.5% ethanol at room ternperature. Some

al,, 1998; Schmalfuss, 2004). In the vicinity of Iriomotejima of the specimens were deposited in the collection of the

Island (i.e., south-western China, Hong Kong, and Taiwan), University Museum, University of the Ryukyu (RUMF-ZC).

C murina is the only Cubaris species that has been reported I also examined the holotype and allotype of C. iriomotensis

(Kwon and Jeon, 1993; Kwon and Taiti, 1993; Dai and Cai, deposited in the Ibyama Science Museum (TOYA-Cr).

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22 Shigenori Karasawa

120. Synonymized with Cubaris murinus (sic) Brandt, Morphologicalobservation 1833 by Barnard (1932: 379). Transferred to Cubaris by

Specimens were observed under a stereomicroscope Jackson(1941:16).

(Olympus SZH, at a magnification of 7-64 × ) and under Armadillo borellii Dollfus, 1894: 1-2; Budde-Lund 1904: a biological light microscope (Nikon Eclipse E400, 120. Synonymized with thbaris murina Brandt, 1833 by magnification of 40-400 x). High resolution irnages were Van Name (1936: 387).

taken with Collinge, 1914: 209-21O. Synonymized with using SEM (JCM-5100,JEOL).Specimensused Cubaris,fi'agilis the SEM were immersedin absolute ethanol and driecl at room Cubaris murina Brandt, 1833 by Ferrara and Taiti (1981 :

temperature or freeze-dried using t-butanol, then mounted on 485).

brass stubs with double-sided tape, and coated with platinum, Cubaris murinus (sic): Barnard 1932: 379-380. Synonymized

with Cubaris murina Brandt, 1833 by Ferrara and Taiti [BixonomicAccount (1979: 174). Cubaris murina Brandt, 1833 Cubaris y'avanensis: Jackson 1941: 16. Synonymized with [Japanesename:Nettai-koshibiro-dangomushi] Cubaris murina Brandt, 1833 by Green (1961: 331). (Figs 1-8) ?Vesodilla (sic) murinus: Verhoeff 1946: 4-5. Synonymized

with Cubaris tnurina Brandt, 1833 by Ferrara and faiti

Cubaris brunnea Brandt, 1833: 190. Transferred to Armadillo (1979: 174).

by Milne-Edwards (1840: 179). Synonymized with SPhaerillo iriomotensis Nunomura. 1990: 2426, Transferred Cevbaris murina Brandt, 1833 by Gerstaecker (1873: to Cubaris by Nunomura (1999a: 90).

527). Cubaris iriomotensis: Nunomura 1999a: 90; Nunornura

Cubaris morrina Brandt, 1833: 190; Gerstaecker 1873: 527; 1999b: 623; Saito et aL 2000: 94; Schmalfuss 2004: 79.

Richardson 1905: 645-647; Budde-Lund 1913: 377-378; Nunomura 2011: 71. Syn. nov. Van Name 1936: 387-389; Green 1961: 33l; Lemos de Castro 1971: 12-13; Schultz 1972: 2-4; Ferrara and Tlaiti Diagnosis. Body color violet-brown rnottled with yellowish

1979: 174; Ferrara and Taiti 1982: 485-486; Schmalfuss color; pereonite 1 with a semicircular lobe on ventral surface;

and Ferrara 1983: 122; AraLijo et al. 1996: 132; Dai and pereonite 2 with a quadrangular lobe on ventral surface; male

Cai 1998: 141; Schmalfuss 2004: 81. Transferred to pereiopods 1-3 each with brush of long setae on carpus and

Armadilto by Milne-Edwarcls (1840: 179). merus.

Armadillo branneus: Milne-Edwards 1840: 179; Budde-Lund

1904: 120, Synonymized with Cubaris murina Brandt, Materia] examined. 3 males, dissected, Ohara, Iriomotejima

1833 by Richardson(1905:645). Island, Japan, N24 16.271, E123 52.883 (dd mm.mmm), 27

Armadillo murinus: Milne-Edwards 1840: 179; Budde-Lund November 20", S Karasawa leg.; 14 males and 8 females,

1885: 27-29; Budde-Lund 1904: 119-120; Budde-Lund dissected. Shinei Park, Ishigakijima Island, Japan, N24

1908: 273. Synonymized with Cubaris murina Branclt, 20.433, E124 09.189, 29 September 2008, S Karasawa leg.;

1833 by Richardson (1905: 645) and Budde-Lund (1913: 6 males, dissected, Manta Park, Ishigakijima Island, Japan, 377). Transferred to IVbsodilla (sic) by Verhoeff (1946: N24 20.054, E124 09.761, 29 September 2008, S Karasawa

45), leg.; 17 males, dissected, Hamazaki Green Park, Ishigakijima

Armadillo cubensis Saussure, 1858: 481-482. Transferred to lsland, Japan, 29 September 2008, N24 20.320, El24 08.960,

Cubaris by Miers (1877: 666). S Karasawa leg.; 1 males, dissected, Hamazaki Green Park, Cubaris qMnis Miers, 1877: 666. [Not Cubaris qfi7nis (Dana, Ishigakijima Island, Japan, N24 20.320, E124 08.960. 30 1854)] Synonymized with Armadillo murinus (Brandt, November 2011, S Karasawa leg. Two males (RUMF- 1833) by Buclcle-Lund (1885: 28). ZC-Ol701-2) and 2 females (RUMF-ZC-O1703-4) from Cubaris cubensis: Miers 1877: 666. Synonymized with Shinei Park, 2 males (RUMF-ZC-O1705-6) from Manta Park, Armadillo murinus (Brandt, 1833) by Budde-Lund (1885: and 2 males (RUMF-ZC-O1707-8) from Hamazaki Green

28). Park are deposited in the University Museum, University

ArmadilloJ'avanensis Dollfus, 1889: 91; Buclde-Lund 1904: of the Ryukyus. Three males (SK-ve-1969-1971) from

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Taxonomy ofJapanese Cztbaris species 23

' utpt

Fig. 1. Cubaris murina Brandt, 1833, cotlected from Ishigakijima island (RUMF-ZC-Ol7e7, non-type, male). A, Lateral view; B, dorsal view. Seale bar: 2 mm.

Ohara, 12 males (SK-ve-17-24, 27-30) and 6 females (SK- ve-1771-1776) from Shinei Park, 4 rnales (SK-ve-55-57, 60) from Manta Park, and 16 males (SK-ve-80-84, 86, 88-96,

1980) from Hamazaki Green Park are kept in a personal

collection (SK-ve) of S. Kan'asawa.

Comparative material. Cubaris iriomotensis; TOYA-

Cr-8884, holotype, maLe; TOYA-Cr-8885, allotype, female.

Description of specimens from Japan. Body length O,65-

1.04 cm in adult male (n =[ 4l), O.60-1,08 cm in adult female (n = 8) (Fig. 1). Body color violet-brown mottled with yellowish co]or in ethanol. Eyes each with about 20 ommatjdia.

Frontal larn{na protruding above the vertex and margina] line 1833, collected continues on both sides within the frontal rnargin (Fig. 2), Fig. 2. Cubaris inurina Brandt, frorn Ishigakijima Island CSK-ve-19, non- Upper edge ofcephalothorax convex in the middte and frontal type, male), cephalothorax, lateral view. Scale shield trapezoid in frontal view (Fig, 3A). bar: 300 pm.

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24 Shigenori Karasawa

Fig. 3. Cubaris murina Brandt, 1833, co]]ected from Ishigakijima Island (non-type, male). A, Cephalothorax, frontat view, RUMF-ZC-O1702; B, second antenna, SK-ve-28i C, inner endite of secopd maxi]]a, SK-ve-28; D, right mandibte, SK-ve-28; E, teft mandible, SK-ve-89; F, maxiitiped, SK-ve-28; G, outer endite ofsecond rnaxi]la, SK- ve-28. Scale bars: 500 prn forA and B; 50 ,um for C-G.

First antenna consisting of three articles; third article pleonite 3 with distinet horjzontat lobe on posterior part (Figs.

with 1O-12 aesthetascs, Second antenna wjth five peduncular 4C-H),

articles and two fiagelluin articles (Fig. 3B). Male pereiopods 1-3 each with brush of long setae on Mandible with at least five plurnose setae between hairy carpus and merus (Figs. 5A-C, left); male pereiopod 7 with

lobe and pars molaris; lacinia mobilis of left mandibie three- more than 15 strong spines on carpus (Fig. 5G, left); male

headed; right lacinia rnobilis square (Figs. 3D, E). pereiopods 4-6 (Figs. 5D-E left) and a]1 female pereiopods First maxilla with outer endite bearing 1O tooth-}ike setae (Figs. 5A-G, right) with fewer than IS spines on carpus and at the apex (Fig. 3G); inner endite with two penicits (Fig. merus. 3C). Second maxilla apically bilobate, covered with short fine Male pleopod 1 with straight endopodite and apicai part

setae. bent outwards bearing a row ofsmall short setae, with a small

Maxilliped with reetangular endite bearing about five swelling on inner margin (Figs. 6A, B); exopodite triangular setae; palp consisting of two articles tapering toward apex (Fig. with several small spines on distal margin. Male pleopod 2

3F). with narrow endopodite; exopodite triangular with distal half

Pereonite 1 with a sernicircular lobe on ventral surface elongated, exopodite as long as endopodite (Fjg. 6C). Female (Fig. 4A); pereonite 2 with rectangular epimera, bearing a pleopod 1 with minute exopodite (Fig. 7A). Perispiracular quadrangular lobe on ventral surface (Fig. 4B); pereonites area located on the outer part of exopodjtes, a rnonospiracular 3-7 each with an oblique lobe just visible on ventral surface; covered lung (Figs. 7B, C).

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Tbxonomy ofJapancse Cztbans spec]es 25

ag

es be,.･:t,1.vt･f,gy･,.1･,1･l･,r-･t eetz x

Fig. 4. Cubaris murina Brandt, l833, collected from Ishigakljima Island (SK-ye-24, non-type, male). A, Pe]eonite 1; B, peleonite 2; C, peleonite 3; D, pe]eonite 4; E, peleenite 5; F, peleonite 6; G, petcon]te 7; H, pleenite 3. ScaLe bars: 30e pm.

Pleotelson hour-glass-shaped, with ctistal part shorter Discussion than basal part (Fig. 8A). Uropod with protopod enlarged;

exopodite inserted on dorsal surface near the middle margin The new specimens collected from Iriomotejima and

ofprotopod; endopodite about twice as long as the exopodite Ishigakljima Isiands are rnorpho]ogically identical and show

the morphological characteristics regarded as (Fig. 8B). 5 diagnostic features of S. iriomotensis by Nunomura (] 990): 1) a flatter

body shape than other arrnadiltid species m Japall; 2) a

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26 Shigenori Karasawa

Fig,5. Cletbaris murina Brandt, 1833, collected from Ishigakijima Island (non-type). Left row: male (A-D, SK-ve-28; E-9, SK-ve-24), right row/ female (A-E, SK-ve-l771; F-G, SK-ve-1773). A, Pereiopod 1; B, pereiopod 2; C, pereiopod 3; D, pereioped 4; E, pereiopod 5; F, pereiopod 6; G, pereiopod 7. Scale bars/ 300 Pm for male A-D and female A-G; SOO pm for male E-G.

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Ibxonomy efJapanese Cttbaris species 27

A

K{'s'e1,Es1i

B

Fig. 6. Cubaris murina Brandt, 1833, collected frem Ishigakijima Island (non-type, maie). A, Pleoped 1, exepodite and endopodite, frontal view, RUMF-ZC-O1708; B, pleepod 1, posterior view, SK-ve-l980; C, pleopod 2, exopodite and endopeciite, frontal view, RUMF-ZC-O1708. Scale bars/ 30e pm forA and C; 20 pm for B.

pleotelson with a straight distal margin; 3) a stouter processus to the new specimens collected in this study, although some

molaris of the mandibles; 4) a triangular exopodite of the morphological characteristics of the hoiotype were not able to

male first pleopod; and 5) many setae on the merus and carpus be examined because ofpoor condition.

of the male first pereiopod. Moreover, examination of the C. The new specimens and holotype have morpholegical

iriomotensis holotype clearly showed that it was identical characteristics consistent with the generic diagnosis of

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28 Shigenori Karasawa

ww ww

asss"fzaxagt.ngptag ' s,,.. ge k-

&･

Fig. 7. Cubaris murina Brandt, 1833, collected from Ishigakijima Island (non-type, femaie). A, Pleoll, ventral view, SK-ve-1776; B, per{spiracular area of pleopod 2 exopodite, dorsal view, SK-ve-1775; C, perispiraeu]ar area ofpleopod 2 exopodite, SK-ve-1776, Dor: dorsal side, Ven/ ventral side. Sca]e bars: 1 mm tbrA; 30 pm fbr B and C.

Fig. 8. Cubaris murina Brandt, 1833, collected from lshigakijima fsland (non-type, male). A, Pleotelson, RUMF-ZC-O1701; B, uropod, SK-ye-24. Scale bars: 300 pm.

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Taxonomy of 亅apanese Cubaris species 29

Cubaris described by Green （1961 ）， e ．g ． the lobe on the Refe「ences ventral surface of pereonite l does not connect with the − epimeral border；the ventral lobe of pereonite 2 does not Araujo， R ， Buckup ， L 旧 nd Bond Buckup， G ，，1996， Is6podos

Rio pr〔）ject beyond the epimcral border；the exopodite is inserted terrestres（Crustacea，Onisoidea）de Santa Catarinae − on the dorsal su face of the protopod and is removed frem the Grande do Sul， BrasiL Iheringia Stirie Zoologia，81： lll 138．

inner border of the protopod． These observatbns suggest that Barnard ， K ．，1932． Contribution to the fauna of South Afhca ． l l．

the species belongs to the genus Cubaris． Moreover， the many Terrestrial isopoda ，！4nnats げ ’he South African ルfuseu〃 1，30 ： − setae of the male pereiopod l （Fig．5A ）and the morphology 179 388 ．

of the male 1 Figs 6A B are congruent with C Brandt 1 1833 Conspectus Monographiae Crustaceerum pleopod （ ． ， ） ． ，．， ，

Fig 12 in Dai 1998 Figs 35−37 in Oniscodorum Latrei且liL ulleten moskovskogo Obshchestva murina （c£ ． and Chai， ， ． βソ − SchmalfUs and Ferrara，1983）． Thus， the speclmens collected IspStateteiPritり dfi，6： 171 193． − − 丘om type 10cality and Ishigakijima Island， and the hobtype Budde Lund ， G 、，1885．Crustacea lsopoda terrestria per familias et

of C ． iriomotensis itself， suggests that on morphological genera et species descripta． Sumtibus Auctoris， Copenhagen，

grounds the specimens belong within the C ， murina group． 319PP ，

in summary C ．’尸’omotensis should be regarded as a Budde −Lund G ，1904 ． A Revision ofCrustacea 置sopoda ， junior ， ，

synonym of C mu 厂ina， and that C 加 躍 加 α are distributed on Terrestria， with Additions and Illustrations，2， Spher 田 oninae ． − Iriomot句ima and Ishigakijima Islands in southwestcm Japan． 3．．4rmad〃 o ． H ． Hagemp Publisher， Copenhagen， pp 33 144． − Budde Lund， G ，，1908」 sopoda von Madagaskar und Osta飼 ka Acknow 且edgements − mit Diagnosen verwandter Arten． pp．265 308，ノn ： Reise

− Iwish to thank Mr ． N ． Nunomura （Toyama Science in Ostafrika inden Jahren l903 1905．Wissenschaftliche

Museum ）＆）r providing the holotype and allotype of Cubar ’∫ Ergebnisse ， voL 2 ．（ed ． VDehzkow ， A ．）， E ． Schweizerbartsche

iriomotens ’∫ and Dr Taiti Italian National Research Verlagsbuchhandlung Stu賃 ， ．S． （ ， gart． − Council）fbr critical reading of the manuscript and usefUl Budde Lund， G ．，1913． Terrestrial Isopoda， particularly considered − advice ． This research was partially supported by the Ministry in relation to the distribution of the southern lndo Pacific − − of Education， Science， Spoれ s and Culture， Grant in Aid fbr species ・Transactions （ゾ the Linnean ∫oo ’θ り20J ヂLondon ，2n ｛ノ − Scientific Research （no ．21310025 ）， a Research Fellowship Ser ’es ， Zoology ，15 ： 367 394

丘om the Japanese Society負）r the Promotion of Science fbr Collinge， W ． E ．，1914 ． On some new terrestrial isopods 行 om the

Young Scientistsno 19−6360 and a Scientific Research Andaman Islands and southem India． Record of the Indi硼 （ ． ）， − Grant from Leave a Nest Co，， Ltd， Japan． Museum ，10：207 210．

Dai， A 、 and Cai， Y ，1998．］℃rrestrial Isopoda（C 田 stacea ： Isopoda： 摘 要 Oniscidea）from the Xishuangbanna Region of Yunnan

唐 沢 重 考 （〒 811−4192 福 岡 教 育 大 学 ： 福 岡 県 宗 像 市 赤 間 Province， south −westem China． Acta Zootaxonomica Sinica，

文教 町 1−1）： Cubaris iriomotensisは ，汎 熱帯re Cubaris murina 23 ： 128−155．

（Crustacea：頁sopoda ； Oniscidea）の 新 参 異 名 ． Dana， J．，1854． Catalogue and description of Crustacea collected ・ Cubaris iriomotensis の タ イ プ 標 本 ．お よ び ，西 表 島 石 垣 in California by Dr．亅ohn L． Le Conte． Proceedings qズthe ’ づ ノ − 島 か ら採集 し た Cubaris属 の 標本 に 基 き Ciriomotensisの 分 ！4cadenリ ofnatura ’Sciences（〜〆Philadellワhia，7； 175 177． 一 類 学 的 検 討 を 行 っ た ．形態観察 の 結 果 ， こ れ ら の 標本 は 同 Doll釦 s ， A ，，1889． Sur quelques isopodes du Mus6e de Leyde， − 種 で あ り， そ の 形 態 的 特 徴 は ，汎 熱 帯種 の Cmurina の そ れ Atotesfrom the 乙eiden ルtuseum，11：91 94， 一 と 致 す る こ とが 分 か っ た ．こ の 結 果 ，Ciriomotensisは ， C Doll 負ls ， A ，，1894 ． Viaggio del dott． Alfredo BORELLI nella

murina の 新参異 名 で あ る と 結 論 づ け た ． Republica Argentina e nel Paragua》へ VI， Isopodes terrestres．

80 〃ett ゴno 6／ei Musei di ZOO109’α ed ／ゴnato 〃lia comparata

ー ー ： コ ロ ゴ コ ロ − キ ワ ド イ リ オ モ テ シ ビ ダ ン ム シ ， シ ビ ダ ン de”a 「ea 〜e こ”ziレ e 「sitd diTorino，9： 1 3． − ゴ ム シ 科 ，ネ ッ タ イ コ シ ビ ロ ダ ン ゴ ム ．琉球列 島 Ferrara， F． and Taiti， S．，1979． A check list of terrestrial isopods

from Afhca （south of the Sahara）．ルfonitore Zoologico − lta’iano，ノVuova Serie， SUPP’e 〃 lento ，12：89 215．

Ferrara， F． and Taiti， S，，1982 ． Ricerche zoologiche della REEF 78

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