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J-'l# VENUS Jour. Malac.) (Jap. '- Vol. 53. No. 2(1994): 119 127

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NewClassification of the Squid Family Mastigoteuthidae

(Cephalopoda: Oegopsida)

Mario Alejandro SALcEDo-VARGAs and Takashi OKuTANI

(Tokye University of Fisheries, 4-S-7, Konan, Minato-ku, Tokye 108)

Abstract: The two genera with two subgenera each are recognized for the squid family Mastigoteuthidae. The genus imstigoteuthis contains two subgenera, Mastigoteuthis s. str. and Ebhinoteuthis. The genus ldioteuthis is here resurected, and it is divisible into two sub- genera, ldioteuthis s. str. and Mbgnoteuthis nov.

The family Mastigoteuthidae was first established by Verrill (1881) and subsequently Chun (1910) placed it as a subfamily of the Chiroteuthidae including eight species. Roper et al. (1969) raised it again to the family by incorporating with the genus Ebhinoteuthis Joubin, 1993, which was based on three paralarvae of questionable taxenomical position. Nesis (1977) attempted a preliminary revision of the Mastigoteuthis and prepared a key to the species. In a recent work, he (l987) proposed two subgenera for this genus, namely, Mtzstigoteuthis s. str. and Mastigopsis Grimpe, 1922, but none for the genus Echinoteuthis that contained two species. Besides Mastigoteuthis s. str. and Mastigopsis he (1977, 1987) recognized the grimaldii-species group based on the structure and position of tentacular suckers. During an extensive and critical examination of the morphology of the 310 specimens collected from the Northwest Pacific, it became clear that the family Mastigoteuthidae is

reasonably divisible into two genera, each contains two subgenera. For designating the genera five major characters are selected among 18 characters used for the more extensiye revision on Northwest Pacific fauna (Salcedo-Vargas, 1993) (Fig. 1). Five more eharacters lead to define the subgenera (Fig. 2). The discussion on detailed sucker morphology is referable to Salcedo-Vargas (in press). In Figs. 3-6 are shown the four sub- genera recognized for the family Mastigoteuthidae.

1. Genus Mastigoteuthis Verrill, 1881 See the definition of Mastigoteuthis s. str. below

1.1. Subgenus Mastigoteuthis s. str. 7)lpe species: Mastigoteuthis agassizii Verrill, 1881

* Presentaddress :Unidad Cuitlahuac edificio95-B-101, Mexico, D.F, CP025oo,AZCAPOTZALCO, MEXICO

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Ditrgnosis: Body small to moderately large. The dorsal margin of the antero-dorsal lobe of the mantle is slightly produced to the funnel. Body covered by numerous white small photophores embedded in the integument, covered each one by small, dark chromatophores in the center. They are mainly concentrated in the ventral side of the body, especially apparent in the anterior end of the tail and posterior extremities of the fins in ventral and dorsal sides. Funnel cartilage ovate in outline with weakly developed tragus and low antitragus, forming a deeper cavity in posterior margin rather than the real antitTagus. Tentacle long and narrow, longer than mantle length, whip-shaped, not expanded in the center of the club, with suckers covering 60Celb of the club. In the proximal part of the

club, at the beginning of the sucker rows, a group of suckers are separated from the rest "circular forming a pad"-like structure. Middle part suckers develop small cushions from the proxirnal margin of the inner ring. Photophore on the eye sinus very small. Included species: 1. Mastigoteuthis agassizii (Verrill, 1881) syn. ML grimaldii Joubin, 1895; M. dentata Hoyle, 1904; M. flammea Chun, 1910; Ml schmidti Degner, 192S.

MssnGOITUTMS MIOITU7Hrs

Bodyphobophores

Ptaent Abeeat

a --.= Sinusphotombore S.c: 9...-.HixN 1 "

Pregent Ab-ent

J-ny= Conusofgladius EilESit====:= = e

emm Itrul '''' ' fF/ -pa) Olfactorypapilla

Sttmandla-g Iboedandthet

Armsudkers

PentxbedTinge Smeothtu

Fig. 1. Comparative table of the characters selected to separate two genera, Mitstigoteuthis and ldioteuthis. tsa)S>ma.

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Salcedo-Vargas . Okutani :NewClassificationof Mastigoteuthidae 121

Mastigoteuthis Idtoteuthis

Medigateuthis Echineteuthis ldiateuthis Magnoteuthis

o t 'N'tY " Tbnta(mlar

t suclcer ' -

innarpegB innerpegs and rim alpmdyenhrged enlarged Ovabe pegu Palygonalpegs

im sucker s Si(IilO

Regulardentition Iimgu1ardmtitim SoftTing imthring

..l,

Sldnt,iberdes

'" i " A thnus 2liilg (ervss sectioni ?}s # O @

//r1,r?Lx1- JttNNi Furme- mantle ,tjilS (II,)s cartilages

Fig. 2.Comparative table of the characters used in the definition of the four subgenera of the family Mastigoteuthidae. gua05}taflvadittwt.

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Remarks: The weak development of the tragus, the relative small size of the eyes, the distinctive small size of the arm suckers from Arm IV are important characters to separate

this subgenera from other taxa. Rancurel (1971) in the redescription of M, grimaldii emphasized that the presence of luminous organs in the body and the teethed suckers are not enough significant to separate

species. He added other external and internal characters to his description in order to improve the unsufficient description by Joubin (1895). He suggested for the first time a possible synonymy of grimaldii with dentata and the two closely related species flammea and schmidti. In spite of this, Young (1972) decided that ML grimatdii can be a species dubia. Nesis (1977, 1987) defined the grimaldii-group in which he included the grimaldii. schmidti and dentata, and rvrodes being less related to them, based on the number and size of

tentacular suckers. After the close examination of the type specimen of ML schmidti and the original descrip- tions and redescriptions of the members of the grimaldii-group, Salcedo-Vargas (1993) con- cluded that grimaldii, dentata, .fTammea and schmidti, are synonymous with agassizii.

1.2. Subgenus Ebhinoteuthis Joubin, 1933 II}{pe species: Echinoteuthis donae Joubin, 1933 (=fometica Berry, 1909) Diagnosis: Body medium to large. Funnel cartilage ovate to subtriangular in outline, tragus strongly developed and oriented into the cavity, antitragus moderately to strongly developed inside. Tentacle suckers become from two rows to gradually more than 12 up to the tip of the club. Tentacular suckers in middle part slightly enlarged, but not visible to naked-eye. The inner ring (==sucker disc) develop two or three elongated pegs oriented toward the cavity. Photophore on the eye sinus sometimes enlarged.

Included species: 1. Mastigotheuthis .fametica Berry, 1909 syn. Echinoteuthis clanae Joubin, 1933; Mastigoteuthis psychrophita Nesis, 1977.

2. Mastigoteuthis gtaukopis Chun, 1910 syn. Mastigoteuthis iseiini MacDonald and Clench, 1934. 3. Mastigoteuthis atlantica Joubin, 1933 4. Mastigoteuthis nvrodes Young, 1972 5. Mastigoteuthis n. sp. Salcedo-Vargas, 1993

Remarks: This subgenu$ is separable from the Mastigoteuthis s. str. by the well excavated

mid-line of the dorsal margin of the mantle, by the relatively bigger size of the eyes, and the presenee of enlarged, spherical photophores in surface of the mantle, head and arms. Though the fin size is variable, this species have leaf-like to rhomboid-like fins. The Echinoteuthis was established by Joubin (1933) as the second genus of this family. Joubin separated it from the genus Mastigoteuthis in tentacle morphology and possession of the so-called "spiny tubercles" that cover all the body. It has been considered as the valid taxon by Young (1972), Nesis (1977, 1987) and Sweeney et al. (1992). But, later, Young (1991) in the examination of some paralarvae from Hawaii synonymized it with Mastigoteuthis.

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6

Fig.3-6.Subgenera of the Mastigoteuthidae. 3. Ventral view of the subgenus Mastigoteuthis.

135 mm DML, 4. Ventral view of the subgenus Ethinoteuthis. 255 mm DML. 5. Ventral

view of the subgenus ldioteuthis, 318 mm DML. 6. Ventral view of the subgenus Mtrgnoteuthis. 105 mm DML. 4]iti.Kcr)t

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Taking Youngs' (1991) statements into consideration, it is concluded that the genus Eehinoteuthis is a subgenus of Mastigeteuthis, even though one of the three specimens

of Joubin is of doubtful taxonomic position. The name danae is considered to be a synenym of ML .famelica. Joubin himself indicat- ed the ctose relationship to that species, Young also (1991) mentioned the close relationship with .fametica in his redefinition of the species from Hawaii. The morphology of the paralarval

specimens from the Northwest Pacific agrees with their views.

2. Genus ldioteuthis Sasaki, 1916

See the definition of ldioteuthis s. str. below

2.1 Subgenus ldioteuthis s. str. 7)lpe species: ldioteuthis latipinna Sasaki, 1916 Diagnosis: Body moderate to very large. The dersal margin of the mantle clearly produced to the funnel. Body covered by hundreds of grey and dark, very small chromatophores. Funnel cartilage oyate in outline, with moderately developed tragus and antitragus. Tentacle

leng, whip-shaped, slightly widened in the central part of the club. Central suckers enlarged, develop cushions from the teeth of the infundibulum or from its internal wall through the infundibulum or swollen distal margin. Photophores embedded in each division of the

trabeculae. Protective membrane covers all the club.

Included species: 1. ldioteuthis cordijbrmis (Chun, 1908) syn. Mastigote"this latipinna Sasaki, 1916

2. ldieteuthis bjorti (Chun, 1913).

Remarks: Chun (1910) described Mastigoteuthis cordijbrmis and few years later M. bjorti (1913). Later, Sasaki (1916) erected a new subfamily Idioteuthinae based on his new genus and species, ldieteuthis latipinna. However, Sasaki (1929) resumed this genuslspecies into the Mastigoteuthis. Sasaki (1916) originally gave the naine Idioteuthis, however, he (1929)

misspeled fridioteuthis in the subsequent synonym list. Grimpe (1922) established the genus Mastigopsis, typifying M. bjorti Chun (1913), and was revived by Nesis (1977). But Sasaki's name was proposed in 1916 although it has

long been ignored by the subsequent authors.

By close examinations on the large-sized specimen in the present study (Salcedo-Vargas, 1993), and by the re-evaluation of suckers morphology (Salcedo-Vargas, in press), its taxo- nomic position and relationship with the ether mastigoteuthid species became clear. The large eyes, the mantlelfin index, the soft conical elastic tubercles over the skin and the morphology ef the gladius were re-evaluated which separated this from the Mastigoteuthis. This is also separable from the genus Mtzstigoteuthis by the absence of luminous organs in the body, the nearly smooth rings in arm suckers, morphology of the funnel cartilage

and the olfactory papillae.

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2.2 Subgenus Magnoteuthis nov. 7)Tpe species: Mastigoteuthis magna Joubin, 1913

Diagnosis: Body medium to very large. Body covered by hundreds of dark chromatophores.

Funnel cartilage ovate in outline. Tragus moderately developed straight, opposite margin

of the tragus also widened into the cayity giying the bottle-shaped contour. Antitragus ab- sent. The central part of the mantle cartilage has a straight crest. Tentacle long, whip-

shaped, club never expanded, neither evidence of cushions, nor teeth in the infundibulum

of the suckers. Suckers very small, with more than 25 rows, invisible to naked eye. Two

white rounded photophores, one in central and other one in distal part of the club. Included species: 1. ldioteuthis magna (Joubin, 1913) syn. Mastigoteuthis magna Joubin, 1913; Mtzstigoteuthis inermis Rancurel, 1972.

Remarks: Features that make separable from the another subgenus are the moderate size

of eyes, the mantlelfin index, the different structures of skin tubercles which are triangularly

shaped, and the morphology of the tentacle.

Table 1. Comparison of the taxonomic systems of Nesis (1977, 1987) and the present study, Nesisa)fimei+ckta)bkpt.

Nesis

(1977, 1987} Present study

Echinoteuthis danae fontelica

Echinoteuthis g

astigoteuthis

Mastigoteuthis Mastigoteuthis

' M(rgnoteuthis

ldioteuthis

ldioteuthis

Mastigopsis

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Conc 置皿 sive Remarks

The morphological relationships among members of the family Mastigoteuthidae have been discussed by Young （1972） and Nesis （1977）． The present paper is the result of an

examination of 310 specilnens from Northwest Pacific， besidessome reference specimens

museums from overseas ， includingthe type specimens of 砿 atlant ’ca and M ．schmi ゴti．

Studies on familial relationships were first done by Chun （1910）． Young and Roper

considering morphology − （1968 ， 1969） the gladius ， funne1 mantle connectives and the tentacle

morphology ， stressed the close relationship among the Chiroteuthidae Grimalditeuthidae ， ， − Joubiniteuthidae， Mast 旭oteuthidae and Batoteuthidae． Clark （1988）added the Promachoteuthi

to this ． on the Hawaiian waters dae families group Young （1991）， based paralarvae from ， ‘‘ ” difined the following members of these families as chiroteuthid lineage ： Ch ’roteuthis ，

Gr’〃 ialdi ’euthis ， Plt〜n た’o ごeuthis ， Mastigo ’eu ’his and ・Lep ’do ’euthis ． However these families all need more extensive and precise revision and proper estima − tion on characterstates based on common standards among these members ， because most

of them not to the standards and currently used cephalopod do fit （Roper Voss ， 1983） for

morphological taxonomy ．

In this sense ， the firstapproach to establish standardization of some of these characters − （Salcedo Vargas ，1993 ）， may help in near future to solve the taxonomic problems of these related families leading to a better approximation for phylogenic studies （Table 1），

要 約

外 洋 中 層 性 の ム チ イ カ 科 Ma8ligoteuthidae を 2 属 4 亜 属 に 分 け る 新 分 類 法 を 提 唱 す る 、，

1．MastigOte” thi，s 属

は さ ． に を つ ． つ 鰭 小 く，体 表 ．発 光 器 も 。 眼 瞼 に も発 光 器 が あ る 。嗅 突 起 は 球 状 ，腕 吸 盤 に は 鋭 歯 を も 。

］ 軟 甲 の i1錐 は 細 く尖 る 。 ］aMaStigoteuthis 亜 属

い 皮 膚 は 平滑 ，触 腕 の 吸 盤 角質環内 側 の 楔状 突起 は 肥 大 し な 。 漏 牛軟 骨 器 の 溝 は 広 い 。 ’ 1b Echinvteuthih亜 属 皮 膚 に 突 起 が あ る 。触 腕 の 吸 盤 角質 環 内 側 の 楔 状 突．起 は 著 る し く 肥 大 ，、漏 斗 軟 骨 器 は 狭 く 「く」

の 字 型 。

2． Idioteuthis es

・ ． い 鰭 は 比 較 的大 き く，体 表 眼 瞼 に も発光 器 は な 、，嗅 突 起 は 棍 棒 状 r ，腕 吸 盤 に は 鈍 歯 が あ る ，，軟 甲 の 円 錐 は や 々 幅 広 い 、，

2a Idiote“ this mp．属

皮 膚 に は 丸 み の あ る 疣 状 突 起 が あ る 。 触 腕 吸 盤 の 楔 状 突 起 は 丸 み が あ る 。軟 甲 は 中 部 か ら 管 状 と な る ， 2b Magnoteuthis 亜 属 （新 ）

皮膚 に は 円 錐状 の 疣状突 起 が あ る 。 触 腕吸盤 の 楔状 突 起 は 多角形 。 軟 甲 は 後 端 に な っ て か ら 管 と 状 な る 。

− ’ Acknowledgments ： We extend our gratitude to Capitan and crews of the R ／V s Kt亦 o −Maru and Kotaka 一物 彫 ， as well as scientists on board for their great effort of samp 且ing oceanic specimens

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Salcedo-Vargas.Okutani: New Classification of Mastigoteuthidae 127

on which this study was based, To Mr. M. J. Sweeney (NMNH-Smithsonian Institution, Washington) for the permission to examine the reference cellection, and to Dr. T. Schiotte (Zoological Museum, Copenhagen) for the loan of the type specimens under his care.

References

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``Michael Chun, C. 1913. Cephalopoda from the Sars" North Atlantic Deep-Sea Expedition, 1910. R(lp. Scient. Results Micheat Sars N, Atlant. deep Sea Iixped. 3: 1-21. Clarke, M. R. 1988. Evolution ef the Recent Cephalopda-A Brief Review. 331-340 pp. in: M. R. Clarke and E. R. Trueman (eds.) Paleontology and neontology of cephalopods. The Mollusca, Vol, 12. Academic Press, New York. Grimpe, G. 1922. Systematiche Ubersicht der europaischen Cephalopoden. Sben natuctl Ges. Lpz, 1918-1921: 36-52.

Jeubin. L. 1985, Cephalopedes. Result. Camp, scient. Prince Albert L 9: 3-63. Joubin, L. 1913. Etudes preliminaires sur les cephalopodes recueilis au cours de croisieres de S.A.S. Ie Prince de Monaco, 3. Mastigoteuthis magna. nov. sp. Butl. Inst. oce'anogr. Monaco, No. 275: 1-IL Joubin, L. 1933. Notes preliminaires sur les c6phalopodes des croisieres du Dana (1921 - 1922) 40 Partie. Annts. Inst, ocdnogr. Monaco. 13: 1-49.

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"Larval'' Sweeney, M. J., Roper, C, F. E,, Mangold, K., Clarke, M. R. and Boletzky, S. 1992. and juvenile cephalopeds: A manual for their identification. Smiths. Contrib. ZooL S13: 1-277. Verrill, A. E. 1881. Report on the cephalopods and on some additional species dredged by the U.S, "Fish-hawk'', Fish Commission Steamer during the season of 1880. Butl Mus. comp. ZooL H7arv. 8: 99-116.

Young, R. E. and Roper, C. F. E. 1968, The Batoteuthidae, a new family of squid from Antartic

waters. Antartic Res. Ser. 11: 185-202, Young, R. E. and Roper, C. F. E. 1969. A Monograph of the Cephalopoda of the North Atlantic: The Family Joubiniteuthidae. Smiths. Contrib. ZooL 15: 1-10. Young, R. E. 1972. The systematics and aereal distribution of pelagic cephalopods from the seas off Southern California. Smiths, Contrib, ZooL 97: 1-159. Young, R. E. 1991. Chiroteuthid and related paralarvae from Hawaiian waters. BulL Mar. Sci, 49(1 & 2): 162 -- 185.

[Received: November 30, 1993]

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