ARTICLE

Memoir of the Fukui Prefectural Dinosaur Museum 21: 1–58 (2022)© by the Fukui Prefectural Dinosaur Museum

EARLY PERMIAN (CISURALIAN) BRACHIOPODS FROMNAGAIWA‒SAKAMOTOZAWA, SOUTH KITAKAMI BELT, JAPANJun-ichi TAZAWA and Tomohiko SHINTANI
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Hamaura-cho 1-260-1, Chuo-ku, Niigata 951-8151, Japan
Dia Consultants Co. Ltd, Toyotsu-cho 5-9, Suita, Osaka 564-1051, Japan

ABSTRACTThis study describes 44 species of early Permian (Cisuralian) brachiopods in 37 genera (including two new
species, Jakutoproductus japonicus sp. nov. and Anemonaria kitakamiense sp. nov.) from the Sakamotozawa andKanokura Formations in the Nagaiwa–Sakamotozawa area, eastern part of the South Kitakami Belt (southern
Kitakami Mountains), northeastern Japan. Based mainly on brachiopod biostratigraphy, the lower part (SK1 Unit) of
the Sakamotozawa Formation is correlated with the Sakmarian, the upper part (SK4 Unit) of the Sakamotozawa
Formation with the lower Kungurian, and the lower part (KN1 Unit) of the Kanokura Formation with the upper
Kungurian. In terms of palaeobiogeography, the Sakmarian fauna (the SK1 assemblage from the SK1 Unit) is amixed Boreal–Tethyan fauna with a predominance of Boreal elements, and has a close affinity with the faunas ofcentral Russia (southern Urals), northern Russia (Timan and northern Urals) and northern China (Inner Mongolia).
The Kungurian fauna (the SK4 assemblage from the SK4 Unit and the KN1 assemblage from the KN1 Unit) is amixed Boreal–Tethyan–Panthalassan fauna, and is similar to the faunas of northwestern (Xinjiang) to northeastern(Jilin) China. Thus, the South Kitakami region belonged to a transitional zone (the Northern Transitional Zone)
between the Boreal and Tethyan realms in the Northern Hemisphere. This region was probably a shallow sea
bordering a microcontinent (the Proto-Japan Block) that was located near and to the east of the North China Block at
the eastern end of the Central Asian Orogenic Belt during the Sakmarian–Kungurian.
Key words: Brachiopoda, early Permian, Japan, Nagaiwa–Sakamotozawa, South Kitakami Belt
田沢純一・新谷友彦 (2022) 南部北上帯長岩‒坂本沢地域から産出する前期ペルム紀（南ウラル世）腕足類．
福井県立恐竜博物館紀要 21：1‒58.
南部北上帯長岩‒坂本沢地域に分布する坂本沢層の下部（SK1 ユニット）と上部（SK4 ユニット）および
叶 倉 層 下 部（KN1 ユ ニ ッ ト）か ら 産 出 す る 37 属 44 種（2 新 種，Jakutoproductus
Jakutoproductus japonicus sp. nov.,
Anemonaria kitakamiense sp. nov. を含む）の前期ペルム紀（南ウラル世）腕足類を記載した．主にこれらの腕足類により，坂本沢層は Sakmarian 〜 Kungurian 下部に，また，叶倉層下部は Kungurian 上部に対比される．古生物地理学的に，Sakmarian のフォーナ（SK1 ユニットから産出する SK1 群集）は ボレアル型
優勢のボレアル型‒テチス型混合フォーナで，ロシア中央部（ウラル山脈南部），ロシア北部（チマン，ウ
ラル山脈北部），中国北部（内蒙古）のフォーナに類似する．Kungurian のフォーナ（SK4 ユニットから産出する SK4 群集および KN1 ユニットから産出する KN1 群集）はボレアル型‒テチス型‒パンサラッサ型混合フォーナで，中国西北部（新彊）〜中国東北部（吉林）のフォーナに類似する．南部北上地域は，前期ペルム紀（Sakmarian‒Kungurian）に北半球のボレアル区とテチス区境界付近にあった北漸移帯に属し，おそらく中央アジア造山帯の東端，北中国地塊の東方にマイクロコンチネントとして存在した原日本地塊（新称）の縁辺浅海域であったと推定される．
かの くら

シンジャン

ジーリン

INTRODUCTION

Hikoroichi-cho, Ofunato City, Iwate Prefecture; Figs. 1–4) inthe eastern part of the belt is the type locality of the lower
Permian (Cisuralian) Sakamotozawa Formation (named by
Onuki, 1937). In this area, the stratigraphy of the Sakamotozawa
Formation has been studied by Onuki (1937, 1956, 1969),
Yamada (1959), Mikami (1965), Kanmera and Mikami (1965a,
1965b), Minato et al. (1979b) and Shintani (2009, 2011).
Recently, Ueno et al. (2007, 2009) described fusulines from thelower and upper parts of the Sakamotozawa Formation, andTazawa and Shintani (2010, 2015) and Shintani (2011)

Permian marine sedimentary rocks are widely distributedin the South Kitakami Belt, northeastern Japan. The
Nagaiwa–Sakamotozawa area (i.e., Nagaiwa and Sakamotozawa,
Received October 9, 2021. Accepted June 6, 2022.
Corresponding author―Jun-ichi Tazawa
E-mail: j1025-tazawa*memoad.jp
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JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIFIGURE 1. Maps showing the location and geology of the Nagaiwa–Sakamotozawa area, South Kitakami Belt. A, Geotectonic map of the JapaneseIslands, showing the distribution of the South Kitakami Belt (based on Tazawa, 2018); B, geotectonic map of the northeastern Honshu, Japan, showing the
distribution of the Permian rocks in the South Kitakami Belt (based on Kawamura et al., 2013); C, topographical map of the Nagaiwa–Sakamotozawa area
and the surroundings. Enclosures are corresponding to Figs. 7 and 10 (using the electronic topographical map of GSI).

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

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FIGURE 2. A view of the central to northern Nagaiwa–Sakamotozawa area, showing two Permian limestone quarries, the Sakamotozawa Quarry and the
Nagaiwa Quarry.

described brachiopods from the same horizons. However, little
is known about the stratigraphy, fossil content and age of theoverlying Kanokura Formation (named by Onuki, 1937).
Mikami (1965) and Kanmera and Mikami (1965a) studied thestratigraphy of the Kanokura Formation but did not report any
fossils; therefore, the age of the Kanokura Formatrion in the
Nagaiwa–Sakamotozawa area has remained uncertain.
In the present study, we describe brachiopods from the lower
a n d u p p e r p arts o f th e S ak amo to z a w a F o rma tion a ndf r o m the lower part of the Kanokura Formation in the
Nagaiwa–Sakamotozawa area and discuss the age andpalaeobiogeography of the brachiopod faunas. Most of thebrachiopod specimens described herein were collected by T.
Shintani in 2006–2009 during the course of undergraduate andgraduate studies at the Niigata University under the supervisionof J. Tazawa. The specimens are now registered and housed in
the Faculty of Science, Niigata University, Niigata, Japan(prefix NU-B, numbers 1212–1286, 1826–1830, 1867–1900.
2280–2286, 2301 and 2302).

PREVIOUS WORK
Stratigraphy
Previous studies on the stratigraphy of the lower Permianrocks corresponding to the Sakamotozawa Formation in the
Nagaiwa–Sakamotozawa area are summarized in Fig. 5. In theearly studies, Endo (1924) and Onuki (1937) named thelimestone-dominated formation the “Fusulina Limestone” andthe Sakamotozawa Stage, respectively. Modern stratigraphicalstudies on the Permian of the Nagaiwa–Sakamotozawa area
began in 1950s after Minato et al. (1954) establishedthe Permian stratigraphy in Setamai, 6 km west ofNagaiwa–Sakamotozawa. Onuki (1956) proposed threemembers in the Sakamotozawa Formation, namely, theYubanosawa Slate Member, the Shiratorizawa Limestone
Member and the Motoiwazawa Sandstone Member in ascending
stratigraphic order. However, the ages of the members wereuncertain due to a paucity of palaeontological data. Mikami(1965) and Kanmera and Mikami (1965a, 1965b) established an

4JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIFIGURE 3. A distant view (A) and a close view (B) of the fossil localities SSK27 and SSK28 in the upper stream of the Imahorazawa Valley, northern
Nagaiwa–Sakamotozawa area.

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

5FIGURE 4. A view of outcrop of sandstone of the KN1 Unit, lower Kanokura Formation in the fossil locality SSK17, 630 m SE of Nagaiwa, central partof the Nagaiwa–Sakamotozawa area.

excellent fusulinid biostratigraphy of the Sakamotozawa
Formation in the Nagaiwa–Sakamotozawa area. According tothose studies, the Sakamotozawa Formation is upper
Wolfcampian–lower Leonardian in age, and is divided into two
subformations and four members: the Lower Subformation(containing the Sa and Sb members) and the Upper
Subformation (containing the Sc and Sd members). The
boundary between the two subformations is an angular
unconformity. Recently, Shintani (2009, 2011) proposed four
members (the Yubanosawa Sandstone Member, theTashiroyama Limestone Member, the Shiratorizawa Limestone
Member and the Shiraishi Sandstone-Limestone Member inascending stratigraphic order) in the Sakamotozawa Formation.
However, the names of the members are nomina nuda, because
the proposal lacked designations and descriptions of the type
localities of the members.
Palaeontology

Many taxonomic studies on the early Permian biota of the
Sakamotozawa Formation in the Nagaiwa–Sakamotozawa area
have been published: fusulines (Hanzawa, 1938, 1939; Igo,
1964; Kanmera and Mikami, 1965b; Ueno et al., 2007, 2009),
corals (Minato, 1955; Minato and Kato, 1965a, 1965b),brachiopods (Tazawa and Shintani, 2010, 2015; Shintani, 2011)
and calcareous algae (Endo, 1951, 1952). Fusulines andbrachiopods are abundant in the studied area. Tazawa andShintani (2010, 2015) and Shintani (2011) described 17
brachiopod species (in 14 genera) from the lower part of the
Sakamotozawa Formation: Echinauris opuntia (Waagen),
Reticulatia cf. donetziana (Licharew), Echinaria sp., Juresaniasp., Waagenoconcha humboldti (d’Orbigny), Edriosteges cf.
multispinosus Muir-Wood and Cooper, Scacchinella sp.,
Linoproductus simensis (Tschernyschew), Auriculispina
kanmerai Tazawa and Shintani, Terrakea sp., Cyclacanthariasp., Derbyia crassa (Meek and Hayden), D. dorsosulcata Liu
and Waterhouse, D. sakamotozawensis Shintani, Meekellastriaticostata (Cox) and M. nagaiwensis Shintani and6JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIFIGURE 5. Comparison of stratigraphic schemes proposed for the lower Permian (Cisuralian) Sakamotozawa Formation in the Nagaiwa–Sakamotozawa
area, South Kitakami Belt.

Rhynchopora sp.
In terms of biostratigraphy, Kanmera and Mikami (1965a,
1965b) proposed five fusuline zones in the Sakamotozawa
Formation: the Zellia nunosei Zone and the Nipponitella
explicata–Monodiexodina langsonensis Zone in the Lower
Subformation, and the Pseudofusulina vulgaris, Pseudofusulina
fusiformis and the Pseudofusulina ambigua zones in the Upper
Subformation. They concluded that the Sakamotozawa
Formation can be correlated with the upper Wolfcampian–lower
Leonardian of California and Texas, the USA. Subsequently,
Ueno et al. (2007, 2009) concluded that the Sakamotozawa
Formation can be assigned to the Sakmarian–Bolorian based on
fusulines from both the basal and the uppermost parts of theformation in the type area.
In terms of palaeobiogeography, Tazawa and Shintani (2010,2015) proposed that the Sakamotozawa brachiopod fauna from thelower part of the Sakamotozawa Formation was a mixed

Boreal–Tethyan fauna with a predominance of Boreal elements.
This faunal affinity suggested that the South Kitakami region wasprobably part of the continental shelf bordering the eastern marginof the North China Block during the early Permian (Sakmarian).
STRATIGRAPHY
Permian rocks are exposed in the Nagaiwa–Sakamotozawa
area, and they form the core and eastern wing of a syncline withan axis trending N–S to NNW–SSE and plunging gently to thesouth (Fig. 6). The Permian strata are divided into threeformations: the lower limestone-dominated Sakamotozawa
Formation; the middle sandstone-dominated Kanokura
Formation; and the upper shale and conglomerate-dominatedToyoma Formation (named by Mabuti and Noda, 1934). In the
Nagaiwa–Sakamotozawa area, brachiopod fossils occur in boththe Sakamotozawa and Kanokura Formations. In this paper weCISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

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FIGURE 6. Geologic map of the Nagaiwa–Sakamotozawa area, Legend 1, 2, upper Permian Toyoma Formation (1, Usuginu-type conglomerate; 2, shale),
3, lower Permian Kanokura Formation (KN1 Unit); 4–7, lower Permian Sakamotozawa Formation (4, SK4 Unit; 5, SK3 Unit; 6, SK2 Unit; 7, SK1 Unit),
8, upper Carboniferous Nagaiwa Formation; 9, lower Carboniferous Onimaru Formation; 10, lower Carboniferous Hikoroichi Formation; 11, middle
Devonian Nakazato Formation; 12, lower Devonian Ono Formation; 13, conglomerate; 14, limestone; 15, Cretaceous granite; 16, Hikami Granite; 17,dyke rock.

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JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIFIGURE 7. Map showing the fossil localities SSK5, SSK14, SSK17, SSK24, SSK27 and SSK28 (using the electronic topographical map of GSI).

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

FIGURE 8. Generalized columnar section of the Sakamotozawa
Formation in the Nagaiwa–Sakamotozawa area, South Kitakami Belt,
showing the fossil horizons SSK27, SSK14, SSK28, SSK5 and SSK24.

divide the Sakamotozawa Formation into three parts (lower,
middle and upper) and four lithostratigraphic units (the SK1,
SK2, SK3 and SK4 units) and erect one unit (the KN1 Unit) in
the Kanokura Formation.
Sakamotozawa Formation

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FIGURE 9. Generalized columnar section of the Kanokura Formation(Lower part. KN1 Unit) in the Nagaiwa–Sakamotozawa area, South
Kitakami Belt, showing the fossil horizon SSK17.

The Sakamotozawa Formation in the Nagaiwa–Sakamotozawa
area has a total thickness of 685 m and is subdivided into alower part (the SK1 Unit: sandstone with thin layers ofconglomerate and shale, 45 m thick), a middle part (the SK2
Unit: limestone with thin layers of shale, 280 m thick; and the
SK3 Unit: limestone with a conglomerate bed at the base, 240 m

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JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIFIGURE 11. Stratigraphic distributions of brachiopod species of theSK1 assemblage in the Sakamotozawa fauna. Broken line shows rangeof the genus.

FIGURE 10. Occurrence of brachiopod species from the Sakamotozawa
and Kanokura Formations in the Nagaiwa–Sakamotozawa area, South
Kitakami Belt. A: abundant, C: common, R: rare.

thick) and an upper part (the SK4 Unit: sandstone and limestone
with subordinate shale, 120 m thick; Fig. 8). Brachiopod fossils
were collected from four localities (SSK5, SSK14, SSK27 and
SSK 28) in the SK1 Unit, and one locality (SSK24) in the SK4
Unit (Fig. 7). The Sakamotozawa Formation unconformably
overlies the upper Carboniferous (Pennsylvanian) Nagaiwa
Formation (Minato, 1942; Yamada, 1959; Kanmera andMikami, 1965a; Mikami, 1965). The boundary between the SK2

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

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FIGURE 12. Stratigraphic distributions of brachiopod species of theSK4 assemblage in the Sakamotozawa fauna. Broken line shows rangeof the genus.

Unit and the SK3 Unit is a disconformity as noted by Mikami(1965) and Kanmera and Mikami (1965a).
Kanokura Formation
The Kanokura Formation is represented in the
Nagaiwa–Sakamotozawa area only by the lower part (KN1
Unit); the middle and upper parts (KN2 and KN3 Units) areabsent, although they are developed in the Setamai area. The
KN1 Unit (460 m thick) consists mostly of sandstone and shale,with a thin (1–5 m thick) basal conglomerate (Fig. 9).
Brachiopod fossils were collected from greenish-greyfine-grained sandstone 30 m above the base of the Kanokura
Formation at locality SSK17 (Fig. 7). The boundary
between t h e K a n o k u r a F o r m a t i o n a n d t h e u n d e r l y i n g
Sakamotozawa Formation is an angular unconformity in the
Nagaiwa–Sakamotozawa area (Choi et al., 1979; Shintani, 2009)
as well as in the Setamai area (Minato et al., 1954).
MATERIALS AND FOSSIL LOCALITIESThe brachiopods described herein were collected from six
localities (stations): four localities (SSK5, SSK14, SSK27 andSSK28) in the SK1 Unit, one locality (SSK24) in the SK4 Unit,
and one locality (SSK17) in the KN1 Unit. The topographic andstratigraphic locations, lithologies and brachiopod species of theseven fossil localities are indicated in Figs. 7‒10 andsummarized below.
SSK5: Roadcut at 30 m west of junction of the Sakamotozawa
and Shiratorizawa valleys (39°08’04” N, 141°38’52” E;

FIGURE 13. Stratigraphic distributions of brachiopod species of the
KN1 assemblage in the Kanokura fauna. Broken line shows range of thegenus.

Figs. 7, 8), grey calcareous medium-grained sandstone,
top of the SK1 Unit, with Rhynchopora sp.
SSK14: Roadcut at 500 m west of Tashiroyashiki (39°08’17” N,
141°39’12” E; Figs. 7, 8), grey medium to coarse-grainedsandstone, middle part of the SK1 Unit, with Terrakeasp., Cyclacantharia sp., Derbyia crassa, D. dorsosulcata,
D. s a k a m o t o z a w e n s i s , M e e k e l l a s t r i a t o c o s t a t a ,M. nagaiwensis, Streptorhynchus sibiricus, Pinegathyris
royssiana, Choristites sp. and Spiriferellina cristata.
SSK17: Roadcut at northern slope of Mt. Tashiroyama, 630 m SE
of Nagaiwa (39°08’36” N, 141°39’04” E; Figs. 4, 7,9), dark grey to light greenish-brown fine-grainedsandstone, 30 m above the base of the Kanokura Formation,
lower part of the KN1 Unit, with Transennatia insculpta,
Echinauris sp., Anidanthus sp., Pseudoleptodus sp.,Dicystoconcha lapparenti, Stenoscisma sp., Hustedia
ratburiensis, Martinia lata, Jilinmartinia sp., Martiniopsis
sp. and Crenispirifer sagus.
SSK24: Roadcut at 880 m south of Nagaiwa (39°08’25” N,
141°38’33” E; Figs. 7. 8), light grey fine-grained sandstone,
lower part of the SK4 Unit, with Anemonaria kitakamiense
sp. nov., Xenosteges adherens, Costatumulus pseudotruncata,12

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIFIGURE 14. Geographic distributions of brachiopod species of the SK1 assemblage in the Sakamotozawa fauna.

Auriculispina kanmerai, Trasgu confinensis, Hustedia
ratburiensis and Crenispirifer sagus.
SSK27: Upper stream of the Imahorazawa Valley (39°09’09” N,
141°38’57” E; Figs. 3A, 7, 8), dark grey fine-grained sandstone,
lower part of the SK1 Unit, with Rugaria semicircularis,
Jakutoproductus japonicus sp. nov., Anemonaria kitakamiense
sp. nov., Echinauris opuntia, Reticulatia cf. donetziana,
Echinaria sp., Juresania sp., Waagenoconcha humboldti,
Scacchinella sp., Linoproductus simensis, Auriculispina
kanmerai, Meekella striatocostata, M. depressa, M.
nagaiwensis, Schuchertella cooperi, Schuchertella sp.,
Streptorhynchus sp., Hustedia ratburiensis, Choristitessp., Phricodothyris asiatica, Callispirina ornata andSpiriferellina cristata.
SSK28: Roadcut at the upper Imahorazawa Valley (39°09’10” N,
141°38’56” E; Figs. 3A, 3B, 7, 8), grey medium tocoarse-grained sandstone, middle part of the SK1 Unit,
with Edriosteges cf. multispinosus, Terrakea sp., Derbyia
crassa, D. dorsosulcata, Meekella striatocostata, M.
nagaiwensis, Rhynchopora sp., Hustedia ratburiensis,
Callispirina ornata and Spiriferellina cristata.
BRACHIOPOD FAUNAS
The brachiopods described herein are the following 44
species in 37 genera, including two new species(Jakutoproductus japonicus sp. nov. and Anemonaria
kitakamiense sp. nov.): Rugaria semicircularis Afanasjeva,
Tazawa and Shintani, 2015, Jakutoproductus japonicus sp. nov.,
Transennatia insculpta (Grant, 1976), Anemonaria kitakamiense
sp. nov., Echinauris opuntia (Waagen, 1884), Echinauris sp.,
Reticulatia cf. donetziana (Licharew, 1938), Echinaria sp.,

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

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FIGURE 15. Sakmarian reconstruction map of the world (adapted from Ziegler et al., 1997), showing the geographic distributions of brachiopod speciesof the SK1 assemblage in the Sakamotozawa fauna. Solid circles indicate numbers of brachiopod species listed in the SK1 assemblage. Station numbers
are same in Fig. 14.

Juresania sp., Waagenoconcha humboldti (d’Orbigny, 1842),
Edriosteges cf. multispinosus Muir-Wood and Cooper, 1960,
Xenosteges adherens Muir-Wood and Cooper, 1960,
Scacchinella sp., Linoproductus simensis (Tschernyschew,1902), Costatumulus pseudotruncata (Ustritsky, 1960),
Anidanthus sp., Auriculispina kanmerai Tazawa and Shintani,
2015, Terrakea sp., Cyclacantharia sp., Pseudoleptodus sp.,Dicystoconcha lapparenti Termier and Termier in Termier et al.,
1 9 7 4 , D e r b y i a c r a s s a ( M e e k a n d H a y d e n , 1 8 5 8 ) , D.
dorsosulcata Liu and Waterhouse, 1985, D. sakamotozawensis
Shintani, 2011, Meekella striatocostata (Cox, 1857), M.
depressa Schellwien, 1900b, M. nagaiwensis Shintani, 2011,
Schuchertella cooperi Grant, 1976, Schuchertella sp.,
Streptorhynchus sibiricus Zavodowsky, 1968, Streptorhynchussp., Stenoscisma sp., Rhynchopora sp., Trasgu confinensis(Schellwien, 1892), Pinegathyris royssiana (von Keyserling,1846), Hustedia ratburiensis Waterhouse and Piyasin, 1970,
Martinia lata Grabau, 1936, Jilinmartinia sp., Martiniopsis sp.,
Choristites sp., Phricodothyris asiatica Chao, 1929, Callispirina
ornata (Waagen, 1883), Spiriferellina cristata (von Schlotheim,
1816) and Crenispirifer sagus Cooper and Grant, 1976b.
The brachiopods of Nagaiwa–Sakamotozawa is classified into
two faunas and three assemblages: the Sakamotozawa fauna (theSK1 and SK4 assemblages) and the Kanokura fauna (the KN1
assemblage).

Sakamotozawa fauna
SK1 assemblage
The SK1 assemblage, from the SK1 Unit of the
Sakamotozawa Formation (localities SSK5, SSK14, SSK27 andSSK28), includes 31 species in 25 genera, with two new species(Jakutoproductus japonicus sp. nov. and Anemonaria
kitakamiense sp. nov.). The species are Rugaria semicircularis,
Jakutoproductus japonicus sp. nov., Anemonaria kitakamiense
sp. nov., Echinauris opuntia, Reticulatia cf. donetziana,
Echinaria sp., Juresania sp., Waagenoconcha humboldti,
Edriosteges cf. multispinosus, Scacchinella sp., Linoproductus
simensis, Auriculispina kanmerai, Terrakea sp., Cyclacanthariasp., Derbyia crassa, D. dorsosulcata, D. sakamotozawensis,
Meekella striatocostata, M. depressa, M. nagaiwensis,
Schuchertella cooperi, Schuchertella sp., Streptorhynchus
sibiricus, Streptorhynchus sp., Rhynchopora sp., Pinegathyris
royssiana, Hustedia ratburiensis, Choristites sp., Phricodothyris
asiatica, Callispirina ornata and Spiriferellina cristata. Of these
species, Rugaria semicircularis, Jakutoproductus japonicus sp.
nov., Waagenoconcha humboldti, Linoproductus simensis,
Auriculispina kanmerai, Derbyia crassa, D. dorsosulcata, D.
sakamotozawensis, Hustedia ratburiensis, Choristites sp.,Phricodothyris asiatica and Spiriferellina cristata are abundant;
Terrakea sp., Meekella striatocostata, M. depressa and M.
nagaiwensis are common; and the other species are rare (Fig.

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JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

Sakamotozawa Formation

FIGURE 16. Geographic distributions of brachiopod species of the SK4
assemblage in the Sakamotozawa fauna and the KN1 assemblage in the
Kanokura fauna.

10).
SK4 assemblage
The SK4 assemblage, from the SK4 Unit of the Sakamotozawa
Formation (locality SSK24), includes seven species in seven
genera: Anemonaria kitakamiense sp. nov., Xenosteges
adherens, Costatumulus pseudotruncata, Auriculispina
kanmerai, Trasgu confinensis, Hustedia ratburiensis andCrenispirifer sagus. Of these species, Anemonaria kitakamiense
sp. nov. and Crenispirifer sagus are abundant; Trasgu confinensis iscommon; and the other species are rare (Fig. 10).
Kanokura fauna
KN1 assemblage
The KN1 assemblage, from the KN1 Unit of the Kanokura
Formation (locality SSK17), includes 11 species in 11 genera:
Transennatia insculpta, Echinauris sp., Anidanthus sp.,
Pseudoleptodus sp., Dicystoconcha lapparenti, Stenoscisma sp.,
Hustedia ratburiensis, Martinia lata, Jilinmartinia sp.,
Martiniopsis sp. and Crenispirifer sagus. Of these species,
Transennatia insculpta and Echinauris sp. are abundant;
Martinia lata is common; and the other species are rare (Fig.
10).
AGE AND CORRELATION

Lower part of the Sakamotozawa Formation
The stratigraphic distributions of the brachiopod species of
the SK1 assemblage are described in the “Systematic
descriptions” section of the present paper, and summarized inFig. 11. Of the brachiopods listed above, four species (Rugaria
semicircularis, Derbyia sakamotozawensis, Meekella depressa
and M. nagaiwensis) are known only from the Sakmarian;
Streptorhynchus sibiricus is known from the Asselian–Sakmarian;
Derbyia crassa from the Moscovian–Kungurian; Linoproductus
simensis from the Kasimovian–Roadian; Meekella striatocostatafrom the Bashkirian–Wordian; and Waagenoconcha humboldti
from the Gzhelian–Capitanian. In contrast, two species(Auriculispina kanmerai and Schuchertella cooperi) are known
from the Sakmarian–Kungurian; another two species (Derbyia
dorsosulcata and Pinegathyris royssiana) from the
Sakmarian–Wordian; and Hustedia ratburiensis from the
Sakmarian–Wuchiapingian. The other species are long-ranging:
Echinauris opuntia has a range of Sakmarian–Changhsingian;
Phricodothyris asiatica of Moscovian–Wuchiapingian; and two
species (Callispirina ornata and Spiriferellina cristata) of
Kasimovian–Changhsingian. At the generic level, Jakutoproductus
is known from the Asselian–Kungurian (Brunton et al., 2000);
Reticulatia from the Bashkirian–Artinskian (Brunton et al.,2000); Echinaria from the Gzhelian–Kungurian (Brunton et al.,2000); Juresania from the Kasimovian–Sakmarian (Brunton et al.,
2000; this study); Edriosteges from the Asselian–Changhsingian(Shen et al., 1992; Brunton et al., 2000); Scacchinalla from the
Sakmarian–Wordian (Brunton et al, 2000; Tazawa and Araki,1999); Terrakea from the Asselian–Wuchiapingian (Brunton etal., 2000); and Cyclacantharia from the Sakmarian–Wordian(Brunton et al., 2000; this study). The other genera are
long-ranging: Schuchertella has a range of Late Devonian–early
Permian (Williams and Brunton, 2000); Streptorhynchus of
Carboniferous–Permian (Williams and Brunton, 2000);
Rhynchopora of Tournaisian–Tatarian (Savage, 2002); andChoristites of Mississsippian?–Cisuralian? (Carter, 2006). In
summary, the age of the SK1 assemblage is identified as
Sakmarian.
As discussed above, the SK1 assemblage is identified as
Sakmarian; thus, the lower part of the Sakamotozawa Formation(SK1 Unit) is correlated with the Sakmarian. This conclusion isconsistent with the previous study (Ueno et al., 2007); in which
the age of the lowest part of the Sakamotozawa Formation is
assigned to the Sakmarian based on some fusulines,
Schubertella sp., Quasifusulina? sp., Rugosofusuina sp.,
Eoparafusulina aff. perplexa (Grozdilova and Lebedeva) andNipponitella explicata Hanzawa.
Upper part of the Sakamotozawa Formation
The stratigraphic distributions of the brachiopod species of
the SK4 assemblage are described in the “Systematic
descriptions” section of the present paper, and summarized inFig. 12. Of the brachiopod species listed above, four species(Xenosteges adherens, Costatumulus pseudotruncata,CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

15

FIGURE 17. Kungurian reconstruction map of the world (adapted from Ziegler et al., 1997), showing the geographic distributions of brachiopod speciesof the SK4 assemblage in the Sakamotozawa fauna and the KN1 assemblage in the Kanokura fauna. Solid circles indicate numbers of brachiopod species
listed in the SK4 and KN1 assemblages. Station numbers are same in Fig. 16.

Auriculispina kanmerai and Trasgu confinensis) are known
from the Sakmarian–Kungurian; Hustedia ratburiensis from the
Sakmarian–Wuchiapingian; and Crenispirifer sagus from theArtinskian–Kungurian. At the generic level, Anemonaria is
known from the Sakmarian–Wuchiapingian (Brunton et al.,
2000; Tazawa, 2011). In summary, the SK4 assemblage isassigned to the Artinskian–Kungurian in age.
As discussed above, the SK4 assemblage is identified asArtinskian–Kungurian; thus, the upper part of the
Sakamotozawa Formation (SK4 Unit) is correlated with theArtinskian‒Kungurian. However, Ueno et al. (2009) concluded
that the age of the upper part (SK4 Unit) of the Sakamotozawa
Formation is Bolorian (= early Kungurian) on the basis offusulines, Pseudofusulina dzamantalensis (Leven), Darvasites
minatoi (Kanmera and Mikami), Kubergandella? sp. andMisellina sp. Therefore, the upper part of the Sakamotozawa
Formation (SK4 Unit) is correlated with the lower Kungurian.
Kanokura Formation
Lower part of the Kanokura Formation
The stratigraphic distributions of the brachiopod species of
the KN1 assemblage are described in the “Systematic
descriptions” section of the present paper, and summarized inFig. 13. Of the brachiopods listed above, Transennatia insculpta
is known from the Artinskian–Wordian; Dicystoconcha

lapparenti from the Kungurian–Wuchiapingian; Hustedia
ratburiensis from the Artinskian–Wuchiapingian; Martinia lata
from the Asselian–Wuchiapingian; and Crenispirifer sagus fromthe Artinskian–Kungurian. At the generic level, Echinauris is
known from the Gzhelian–Wuchiapingian (Cooper and Grant,
1975; Brunton et al., 2000; Shen and Zhang, 2008); Anidanthusfrom the Asselian–Wuchiapingian (Brunton et al., 2000; Klets,2005); Pseudoleptodus from the Asselian–Capitanian (Cooper
and Grant, 1974; Grant, 1976; this study); Stenoscisma from thelower Carboniferous to the Changhsingian (Shen et al., 2000;
Carlson and Grant, 2002); Jilinmartinia from the
Moscovian–Kungurian (Pavlova, 1991; Carter and Gourvennec,
2006); and Martiniopsis from the upper Carboniferous to the
Lopingian (Carter and Gourvennec, 2006). In summary, theKN1 assemblage is identified as Kungurian in age. The lower
part of the Kanokura Formation (KN1 Unit) in the
Nagaiwa–Sakamotozawa area is probably correlated with theupper Kungurian, considering that the base of the Kanokura
Formation (KN1 Unit) covers the upper part of the
Sakamotozawa Formation (SK4 Unit) with an angular
unconformity in broad area of the South Kitakami Belt (e.g.,
Minato et al., 1954; Choi et al., 1979).
PALAEOBIOGEOGRAPHY
Sakmarian

16

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIThe geographic distributions of the Sakmarian brachiopod
species of the SK1 assemblage are described in the “Systematic
descriptions” section of the present paper, and are summarized
in Figs. 14 and 15. Of the 31 species of the SK1 assemblage, six
species also occur in central Russia (southern Urals); fivespecies have been reported from northern China (Inner
Mongolia); four species are found in northern Russia (northern
Urals and Timan); and three species also occur incentral-southern China (Guangxi). To summarize, the SK1
assemblage has a close affinity with the Sakmarian brachiopod
faunas of central Russia (southern Urals), northern Russia(northern Urals and Timan) and northern China (Inner
Mongolia). Furthermore, the SK1 assemblage includes both
antitropical genera (Jakutoproductus, Anemonaria, Juresania,
Waagenoconcha, Auriculispina, Terrakea, Rhynchopora,
Pinegathyris and Choristites) and tropical genera (Echinauris,
Scacchinella, Cyclacantharia and Meekella). Thus, theSakmarian brachiopod fauna (SK1 assemblage) is a mixed
Boreal–Tethyan fauna with a predominance of the Boreal
elements, and exhibits affinities with those of central Russia(southern Urals), northern Russia (northern Urals and Timan)
and northern China (Inner Mongolia).
This conclusion is consistent with the previous works on the
Sakmarian brachiopod faunas of the Nagaiwa–Sakamotozawa
area (Tazawa and Shintani, 2010, 2015) and from the Kamiyasse
area (Tazawa and Shintani, 2014).
Kungurian
The geographic distributions of the Kungurian brachiopod
species of the SK4 and KN1 assemblages are described in the“Systematic descriptions” section, and are summarized in Figs.
16 and 17. Among the 16 species of the SK4 and KN1
assemblages, two species also occur in the USA (Texas) andsouthern Thailand; and one species has been reported from
central Japan (Hida Gaien Belt), southwestern Japan(Mizukoshi), Slovenia, Afghanistan, northwestern China(Xinjiang), northern China (Inner Mongolia), eastern China(Zhejiang), central-southern China (Hubei, Guangdong andGuangxi), north-central Thailand and Malaysia. To summarize,
the Kungurian fauna (SK4 and KN1 assemblages) somewhat
resembles those of the USA (Texas) and southern Thailand. In
addition, the Kungurian brachiopod fauna (SK4 and KN1
assemblages) includes both antitropical genera (Anemonaria,
Costatumulus, Anidanthus, Auriculispina and Jilinmartinia) andtropical genera (Transennatia, Echinauris, Xenosteges,
Pseudoleptodus and Dicystoconcha). It is noteworthy that
Jilinmartinia occurs mostly from northwestern China tonortheastern China (Shen et al., 2017). Thus, the Kungurian
brachiopod fauna (SK4 and KN1 assemblages) is a mixed
Boreal–Tethyan–Panthalassan fauna, and exhibits affinities withthose of northwestern–northeastern China.
This conclusion is consisted with previous work (Tazawa andNakamura, 2015), which described a Kungurian brachiopod

fauna consisting of 15 species in 15 genera from the basal partof the Hosoo Formation in the Nakadaira area in the South
Kitakami Belt. The Nakadaira fauna is also a mixed
Boreal–Tethyan fauna, and exhibits a close affinity with theearly Permian brachiopod faunas of northwestern China(Xinjiang) and northern China (Inner Mongolia).
DISCUSSION: CISURALIAN GEOGRAPHY AND
BIOGEOGRAPHY OF THE SOUTH KITAKAMI BELT
From the information provided above, we can conclude thatearly Permian (Sakmarian–Kungurian) brachiopod faunas of theSouth Kitakami Belt are characterized by a mixture of both
Boreal and Tethyan elements, and belong to a transitional zone
between the Boreal and Tethyan realms in the Northern
Hemisphere, i.e., the Northern Transitional Zone (Shi et al.,1995) [= the Inner Mongolian–Japanese Transisional Zone
(Tazawa, 1991) and the Sino–Mongolian–Japanese Province(Shi and Tazawa, 2001)]. Shen et al. (2013) noted that theNorthern Transitional Zone between the Boreal and the Tethyan
realms was not recognized before Kungurian based on an
analysis of global database; however, it is clear that the zone
was present in the Sakmarian–Kungurian in terms of the faunas
described herein. The South Kitakami region belonged to theNorthern Transitional Zone in the Northern Hemisphere, andwas probably a shallow sea bordering a microcontinent (theProto-Japan Block) at the eastern end of the Central Asian
Orogenic Belt and near and to the east of the North China Blockduring the Sakmarian–Kungurian.
The co-occurrence of the Permian coral groups Durhaminidae
and Waagenophyllidae in the lower Permian strata of the South
Kitakami Belt (Minato and Kato, 1965a, 1965b. 1970), as notedby Tazawa (2018), indicates that the South Kitakami region was
located in a transitional zone between the Boreal and Tethyan
realms. The area probably lay near the North China Block,which was located at midlatitudes of the Northern Hemisphere
during the early Permian.
Among early Permian land plants, the Maiya flora (Asama,
1967, 1981) from the Nishigori Formation of Maiya closely
resembles that of northern Korea (Sadon), and the Setamai flora(Asama and Murata, 1974) from the Sakamotozawa Formationof Setamai is similar to that of northern China (Shanxi). Thus,
the early Permian Maiya and Setamai floras of the South
Kitakami Belt have affinities with those of the North China
Block, which belonged to the Cathaysia Northern Subprovince
during the Permian (Huang and Chen, 1987).
Some objections to the placement of the South Ktakami
region near the North China Block were listed by Ehiro (2001),
Okawa et al. (2013) and Isozaki et al. (2017) who placed theSouth Kitakami region near the South China Block or as part of
the Greater South China during the Permian. However, the early
Permian brachiopod faunas of southern China (Jin et al., 1974;
Yang et al., 1977; Liu et al., 1982) completely lack the Boreal(antitropical) genera such as Jakutoproductus and Rhynchopora
in the Sakmarian faunas, and Anemonaria, Costatumulus andCISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

Jilinmartinia in the Kungurian faunas.
CONCLUSIONS

17

Rugaria semicircularis Afanasjeva, Tazawa and Shintani, 2015,
p. 23, pl. 3, figs. 1–6.

In this study, early Permian (Cisuralian) brachiopods aredescribed from the lower part (SK1 Unit) and the upper part(SK4 Unit) of the Sakamotozawa Formation and the lower part
(KN1 Unit) of the Kanokura Formation in the
Nagaiwa–Sakamotozawa area, South Kitakami Belt,
northeastern Japan. A total of 44 species in 37 genera aredescribed, of which two (Jakutoproductus japonicus sp. nov.
and Anemonaria kitakamiense sp. nov.) are new. In terms ofbiostratigraphy, the lower part of the Sakamotozawa Formationis correlated with the Sakmarian; the upper part of the
Sakamotozawa Formation is corelated with the lower
Kungurian; and the lower part of the Kanokura Formation iscorrelated with the upper Kungurian. Palaeobiogeographically,
the Sakmarian fauna (the SK1 assemblage from the SK1 Unit) isa mixed Boreal–Tethyan fauna with a predominance of the
Boreal elements. The assemblage exhibits a close affinity withthose of central Russia (southern Urals), northern Russia(northern Urals and Timan) and northern China (Inner
Mongolia). The Kungurian fauna (the SK4 assemblage from theSK4 Unit and the KN1 assemblage from the KN1 Unit) is amixed Boreal–Tethyan–Panthalassan fauna, and exhibits
affinities with those of northwestern–northeastern China. We
conclude that the South Kitakami region, including the
Nagaiwa–Sakamotozawa area, belonged to the Northern
Transitional Zone of Shi et al. (1995), and was probably a
shallow sea bordering the newly proposed Proto-Japan Block,which was located at near and to the east of the North China
Block at the eastern end of the Central Asian Orogenic Belt
during the Sakmarian–Kungurian.

Material.—Six specimens from locality SSK27: (1) ventral
internal mould and dorsal external mould of a conjoined shell,
NU-B1830; and (2) external and internal moulds of four ventral
valves, NU-B1826, 1827 (holotype), 1828, 1829.
Remarks.—These specimens were described by Afanasjeva
et al. (2015, p. 3, pl. 3, figs. 1–6) as Rugaria semicircularis
Afanasjeva, Tazawa and Shintani, 2015 from the SK1 Unit of
the Sakamotozawa Formation in the Nagaiwa–Sakamotozawa
area, South Kitakami Belt, northeastern Japan. Rugaria
semicircularis resembles the type species, Rugaria hessensis(King, 1931), redescribed by Cooper and Grant (1975, p. 1296,
pl. 496, figs. 18–25; pl. 498, figs. 1–12; pl. 499, figs. 70, 71)
from the Hess Formation (Taylor Ranch Member) of Texas insize and shape of the shell, but differs in having thinner andmore numerous capillae on both ventral and dorsal valves.
Occurrence.—SK1 Unit.
D i s t r i b u t i o n . —S a k m a r i a n : n o r t h e a s t e r n J a p a n(Nagaiwa–Sakamotozawa in the South Kitakami Belt).

SYSTEMATIC DESCRIPTIONS(by J. Tazawa)

Etymology.—Named after the country, Japan, including thefossil locality.
Material.—Six specimens from locality SSK27: (1) external
and internal moulds of a ventral valve, NU-B2363; (2) internalmould of a ventral valve, NU-B2364; and (3) external andinternal moulds of four dorsal valves, NU-B2365 (holotype),
2366–2368.
Diagnosis.—Small, transverse Jakutoproducus, with sparcespine bases on ventral valve.
Description.—Shell small in size for genus, transversely
subrectangular in outline; hinge shorter than greatest width atabout midlength; length 8 mm, width 11 mm in the best
preserved ventral valve specimen (NU-B2363); length 7 mm,
width 11 mm in the best preserved dorsal vale specimen(holotype, NU-B2365). Ventral valve moderately and unevenly
convex in lateral profile, most convex at umbonal region; umbo
small; ears small, triangular, not clearly differentiated fromvisceral portion; sulcus very shallow or absent. Dorsal valve
flattened on visceral disc, geniculated and followed by short
trail; ears small, distinct; fold narrow and low. External surface
of ventral valve ornamented with numerous irregular concentric

The suprageneric classification given herein mainly follows
that of “Treatise on Invertebrate Paleontology, Part H
Brachiopoda Revised, Volumes 2–6”, edited by Kaesler (2000a,
2000b, 2002, 2006) and Selden (2007), with exception thatclassification of the suborder Productidina follows Waterhouse(2002).
Order PRODUCTIDA Sarytcheva and Sokolskaya, 1959
Suborder CHONETIDINA Muir-Wood, 1955
Superfamily CHONETOIDEA Bronn, 1862
Family RUGOSOCHONETIDAE Muir-Wood, 1962
Subfamily RUGOSOCHONETINAE Muir-Wood, 1962
Genus RUGARIA Cooper and Grant, 1969
Type species.—Chonetes hessensis King, 1931.
Rugaria semicircularis Afanasjeva, Tazawa and Shintani, 2015
(Fig. 18F–H)

Suborder PRODUCTIDINA Waagen, 1883
Superfamily PRODUCTELLOIDEA Schuchert, 1929
Family AVONIIDAE Sarytcheva, 1960
Subfamily TUBERSULCULINAE Waterhouse, 1971
Genus JAKUTOPRODUCTUS Kaschirtzev, 1959aType species.—Marginifera verchoyanica Fredericks, 1931.
Jakutoproductus japonicus sp. nov.
(Fig. 18A–E)

18

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIFIGURE 18. Brachiopods of the SK1 assemblage (1). A–E, Jakutoproductus japonicus sp. nov.; A, external latex cast (A1, A2) and internal mould (A3) ofventral valve, NUB2363; B, external latex cast (B1, B2) of ventral valve, NU-B2364; C, external latex cast (C1, C2), external mould (C3), internal latex cast(C4) and internal mould (C5) of dorsal valve, NU-B2365 (holotype); D, external latex cast (D 1, D2), external mould (D3), internal latex cast (D4) andinternal mould (D5) of dorsal valve, NU-B2367; E, external latex cast (E1, E2), external mould (E3), internal latex cast (E4) and internal mould (E5) of
dorsal valve, NU-B2366; F–H, Rugaria semicircularis Afanasjeva, Tazawa and Shintani; F, internal mould of ventral valve, NU-B1827 (holotype); G,
external mould of ventral valve, NU-B1828; H, external mould (H1) and internal mould (H2) of ventral valve, NU-B1829. Scale bars are 1 cm, except for
those of F–H.

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

19

FIGURE 19. Brachiopods of the SK1 assemblage (2). A, Anemonaria kitakamiense sp. nov., ventral external latex cast (A1, A2), ventral external mould(A3) and dorsal internal mould (A4) of conjoined shell, NU-B2370; B, C, Reticulatia cf. donetziana (Licharew); B, external latex cast (B1) and internalmould (B2, B3) of dorsal valve, NU-B1873; C, external latex cast of dorsal valve, NU-B1874; D, Echinauris opuntia (Waagen), external latex cast (D1,
D2), external mould (D3) and internal mould (D4) of ventral valve, NU-B1867. Scale bars are 1 cm.

20

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

rugae and sparse elongate spine bases. External ornament of
dorsal valve consisting of numerous irregular lamellae andsparce dimples. Interior of ventral valve not well preserved.
Dorsal interior with a bilobate cardinal process, a pair ofelongate adductor scars, and a thin, long median septum
extending to half valve length.
Remarks.—The present new species resembles the typespecies, Jakutoproductus verchoyanicus (Fredericks, 1931, p.
211, pl. 1, figs. 11–13) from the lower Permian of the Dulgalakh
River, western Verkhoyansk, northern Russia, in size andoutline of the shell, but differs in having sparce spine bases on
the ventral valve. Jakutoproductus crassus Kaschirtzev, 1959b,
redescribed by Abramov and Grigorjeva (1988, p. 115, pl. 5,
figs. 15–19, 22–26; pl. 6, figs. 1, 2) from the lower Permian(Asselian–Artinskian) of Verkhoyansk, has also sparce spine
bases on the ventral valve, but differs from Jakutoproductus
japonicus sp. nov. in the larger size and in having distinct sulcus
on the ventral valve. Jakutoproductus tatjanae Abramov andGrigorjeva (1983, p. 67, pl. 1, figs. 19–26), from the upper
Carboniferous of Verkhoyansk, similar to the present new
species in size and outline of the shell, but differs in having
numerous spine bases on the ventral valve. The shells describedby Shi (1994, p. 115, fig. 4) as Jakutoproductus verchoyanicus(Fredericks, 1931), from the lower Permian of northern Yukon
Territory, Canada, has also sparce spine bases on the ventral
valve, but differs from the present new species in its larger sizeand less transverse outline.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).

insculpta (Grant, 1976), originally described by Grant (1976, p.
135, pl. 32, figs. 1–37; pl. 33, figs. 1–16) as Gratiosina
insculpta Grant, 1976, from the Ratburi Formation of Ko Muk,
southern Thailand, in the small and transverse shell (length 12mm, width 19 mm in the best preserved dorsal valve specimen,
NU-B2283) and the coarse reticulate ornament on the visceraldiscs of both ventral and dorsal valves. Transennatia cf.
insculpta (Grant, 1976), described by Sone (in Sone et al., 2003,
p. 528, figs. 7, 8c) from the middle Permian (Roadian–Wordian)of Johore, Peninsular Malaysia may be a synonym of T.
insculpta. The type species, Transennatia gratiosa (Waagen,
1884, p. 691, pl. 72, figs. 3–7), from the Wargal and Chhidru
Formations of the Salt Range, Pakistan, differs from T. insculpta
in the larger size and in having finer reticulate ornament on thediscs of both valves.
Occurrence.—KN1 Unit.
Distribution.—Artinskian–Wordian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt),
northwestern China (Xinjiang), southern Thailand (Ko Muk)and Malaysia.

Superfamily MARGINIFEROIDEA Stehli, 1954
Family MARGINIFERIDAE Stehli, 1954
Subfamily MARGINIFERINAE Stehli, 1954
Genus TRANSENNATIA Waterhouse, 1975

Etymology.—Named after the fossil locality, Kitakami
Mountains.
Material.—Six specimens from localities SSK24 and SSK27:(1) external and internal moulds of four ventral valves,
NU-B2325 (holotype), 2326, 2327, 2370; and (2) internalmoulds of two ventral valves, NU-B2328, 2329.
Diagnosis.—Small, equidimentional Anemonaria, with smallears and low sulcus on ventral valve.
Description.—Shell small in size for genus, rounded
subquadrate in outline, hinge slightly shorter than greatest widthat about midlength; length 10 mm, width 11 mm in the holotype(NU-B2325). Ventral valve strongly and unevenly convex inlateral profile, most convex at umbonal region; umbo small, ears
small, blunt; sulcus low on anterior half; lateral slopes steep.
External surface of ventral valve nearly smooth except for a few
faint costae on trail; a row of large spine bases, numbering 3 atbase of each ears. Ventral interior with widely flabellate
diductor scars and narrow elongate adductor scars
Remarks.—Anemonaria kitakamiense sp. nov. mostresembles Anemonaria sulankherensis Manankov (1998, p. 52,
pl. 8, figs. 8, 9), from the Tsaganul Horizon (Ufimian) of Mt.
Dzhirem-Ula, southeastern Mongolia, in the small, rounded
quadrate ventral valve with a low sulcus. But the Mongolian
species differs from the present new species in being moreType species.—Productus gratiosus Waagen, 1884.
Transennatia insculpta (Grant, 1976)(Fig. 32A–C)
Gratiosina insculpta Grant, 1976, p. 135, pl. 32, figs. 1–37; pl.
33, figs. 1–16.
Transennatia cf. insculpta (Grant). Sone in Sone et al., 2003, p.
528, figs. 7, 8c.
Transennatia insculpta (Grant). Chen, 2004, p. 14, pl. 2, figs.
5–12.
Material.—Six specimens from locality SSK17: (1) external
and internal moulds of a ventral valve, NU-B2281; (2) externalmould of a ventral valve, NU-B2282; (3) external and internalmoulds of two dorsal valves, NU-B2283, 2284; (4) externalmould of a dorsal valve, NU-B2285; and (5) internal mould of aconjoined shell, NU-B2286.
Remarks.—These specimens are referred to Transennatia

Family PAUCISPINIFERIDAE Muir-Wood and Cooper, 1960
Subfamily PAUCISPINIFERINAE Muir-Wood and Cooper,
1960
Genus ANEMONARIA Cooper and Grant, 1969
Type species.—Marginifera sublaevis King, 1931.
Anemonaria kitakamiense sp. nov.
(Figs. 19A, 30A–C)

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

21

FIGURE 20. Brachiopods of the SK1 assemblage (3). A, Echinaria sp., external latex cast (A1, A2) and internal mould (A3) of ventral valve, NU-B1899;
B, Juresania sp., internal latex cast (B1, B2) and internal mould (B3) of dorsal valve, NU-B1900; C, D, Waagenoconcha humboldti (d’Orbigny); C,
external latex cast (C1, C2) of ventral valve, NU-B1282; D, external latex cast (D1) and internal latex cast (D2) of dorsal valve, NU-B1283; E, Edriosteges
cf. multispinosus Muir-Wood and Cooper, external latex cast (E1) and internal latex cast (E2) of ventral valve, NU-B1868. Scale bars are 1 cm.

22

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANItransverse outline. Anemonaria sublaevis (King, 1931),
redescribed by Cooper and Grant (1975, p. 1103, pl. 408, figs.
1–26), from the Bone Spring, Cathedral Mountain and Road
Canyon Formations of Texas in the USA, differs from A.
kitakamiense sp. nov. in the transverse ventral valve with deeper
sulcus. Anemonaria auriculata Shi and Waterhouse (1996, p.
68, pl. 6, figs. 10–28, text-figs. 22–24), from the upper Jungle
Creek Formation of northern Yukon Territory, Canada, is also asmall-sized Anemonaria, but differs from the Kitakami species
in having alate ears and deep sulcus on the ventral valve.
Occurrence.—SK1 and SK4 units.
Distribution. —Sakmarian–Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Family COSTISPINIFERIDAE Muir-Wood and Cooper, 1960
Subfamily COSTISPINIFERINAE Muir-Wood and Cooper,
1960
Genus ECHINAURIS Muir-Wood and Cooper, 1960
Type species.—Echinauris lateralis Muir-Wood and Cooper,
1960.
Echinauris opuntia (Waagen, 1884)(Fig. 19D)
Productus opuntia Waagen, 1884, p. 707, pl. 79, figs. 1, 2;
Broili, 1916, p. 17, pl. 117, figs. 9, 10.
Echinauris opuntia (Waagen). Grant, 1968, p. 27, pl. 8, figs.
1–8; pl. 9, figs. 1–8; Licharew and Kotlyar, 1978, pl. 20, fig.
14; Shen et al., 2000, p. 742, figs. 10.24–10.32; Shen and Shi,
2009, p. 158, figs. 3Y–3CC; Tazawa and Shintani, 2015, p.
41, fig. 3.5.
Material.—One specimen from locality SSK27, external andinternal moulds of a ventral valve, NU-B1867.
Remarks.—This specimen was described by Tazawa andShintani (2015, p. 41, fig. 3.5) as Echinauris opuntia (Waagen,1884) on account of its small, slightly transverse and rounded
subtriangular ventral valve (length 11 mm, width 14 mm), withnumerous long, thin spines on the lateral slopes. The typespecies, Echinauris lateralis Muir-Wood and Cooper (1960, p.
222, pl. 68, figs. 1–13), from the Word Formation of the Glass
Mountains, Texas, differs from the present species in the larger
size and in having stronger and thicker spines on the ventral
valve.
Occurrence.—SK1 Unit.
Distribution. —Sakmarian–Changhsingian: northeasternJapan (Nagaiwa–Sakamotozawa in the South Kitakami Belt),
eastern Russia (South Primorye), central-southern China(Guangxi), southwestern China (Xizang), Pakistan (Salt Range)
and Timor.
Echinauris sp.
(Figs. 20A, 32D–F)

Material.—Six specimens from locality SSK17: (1) external
and internal moulds of two ventral valves, NU-B2287, 2288; (2)external and internal moulds of two dorsal valves, NU-B2289,
2290; and (3) external moulds of two dorsal valves, NU-B2291,
2292.
Remarks.—These specimens are safely assigned to the genus
Echinauris MuirWood and Cooper, 1960 by the small, elongate
shell (length 11 mm, width 15 mm in the best preserved dorsal
valve specimen, NU-B2289), and the external ornaments ofnumerous stout spine bases on the ventral valve and numerous
dimples and fine growth lines on the dorsal valve. The
Nagaiwa–Sakamotozawa species is most like Echinauris crassa
Cooper and Grant (1975, p. 1006, pl. 327, figs. 1–36), from the
Leonardian of Texas, the USA, in the transverse outline and inhaving large ears, but differs from the latter in being wider
configuration and larger size. Echinauris interrupta Cooper and
Grant (1975, p. 1006, pl. 328, figs. 1–33), from theWolfcampian of Texas, differs from the present species in thelarger size and in having finer spine bases on the ventral valve.
The type species, Echinauris lateralis Muir-Wood and Cooper(1960, p. 222, pl. 68, figs. 1–13), from the Word Formation of
Texas, is readily distinguished from the present species in theless transverse outline and in having smaller ears. The
Tashiroyama species may be a new species of Echinauris,
although the specimens are poorly preserved.
Occurrence.—KN1 Unit.
Distribution.—Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakai Belt).
Superfamily PRODUCTOIDEA Gray, 1840
Family DICTYOCLOSTIDAE Stehli, 1954
Subfamily DICTYOCLOSTINAE Stehli, 1954
Genus RETICULATIA Muir-Wood and Cooper, 1960
Type species.—Productus huecoensis King, 1931.
Reticulatia cf. donetziana (Licharew, 1938)(Fig. 19B, C)
Reticulatia cf. donetziana (Licharew, 1938). Tazawa andShintani, 2015, p. 43, fig. 3.3, 3.4.
Material.—Two specimens from locality SSK27: (1) external
and internal moulds of a dorsal valve, NU-B1873; and (2)external mould of a dorsal valve, NU-B1874.
Remarks.—These specimens were described by Tazawa andShintani (2015, p. 43, fig. 3.3, 3.4) as Reticulatia cf. donetziana(Licharew, 1938). The Nagaiwa–Sakamotozawa species
resembles Reticulatia donetziana (Licharew, 1938), from theUpper Carboniferous (Kasimovian) of the Donetz Basin,
western Russia, in having a peculiar bordering flange atantero-lateral margins of the dorsal valve. The same ringlike
structure has been described and figured by Sutton (1942, p.
464, pl. 71, figs. 12, 13) in the specimens of Reticulatia
americana (Dunbar and Condra, 1932) from the Pennsylvanian

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

23

FIGURE 21. Brachiopods of the SK1 assemblage (4). A, Scacchinella sp., ventral (A1), anterior (A2), posterior (A3) and lateral (A4) views of internalmould of ventral valve, NU-B1286; B–D, Linoproductus simensis (Tschernyschew); B, external latex cast (B 1, B2) of ventral valve, NU-B1875; C,
external latex cast (C1) and internal latex cast (C2) of dorsal valve, NU-B1879; D, internal latex cast of dorsal valve, NU-B1881; E–G, Auriculispina
kanmerai Tazawa and Shintani; E, external latex cast (E1, E2) and internal mould (E3) of ventral valve, NU-B1882; F, external latex cast (F1) and internalmould (F2) of ventral valve, NU-B1884; G, external latex cast of dorsal valve, NU-B1893; H, I, Terrakea sp.; H, external latex cast (H1, H2) and internallatex cast (H3) of ventral valve, NU-B1894; I, external latex cast (I1) and internal mould (I2) of ventral valve, NU-B1895. Scale bars are 1 cm.

24

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIof Illinois in the USA. The Kitakami species more like theRussian species than the American species in size, shape andexternal ornament of the dorsal valve. However, identification isdifficult for the present material.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Superfamily ECHINOCONCHOIDEA Stehli, 1954
Family ECHINOCONCHIDAE Stehli, 1954
Subfamily ECHINOCONCHINAE Stehli, 1954
Genus ECHINARIA Muir-Wood and Cooper, 1960
Type species.—Productus semipunctatus Shepard, 1838.
Echinaria sp.
(Fig. 20A)
Echinaria sp. Tazawa and Shintani, 2015, p. 44, fig. 3.1.
Material.—One specimen from locality SSK27, external andinternal moulds of a ventral valve, NU-B1899.
Remarks.—This specimen was described by Tazawa andShintani (2015, p. 44, fig. 3.1) as Echinaria sp. on account of its
large, elongate ventral valve (length more than 37 mm, widthabout 28 mm), with narrow and moderately deep sulcus, andornamented by numerous broad concentric lamellae with 3–4
rows of spine bases on each lamella. The species identificationis, however, difficult owing to the poorly preserved specimen.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Subfamily JURESANIINAE Muir-Wood and Cooper, 1960
Genus JURESANIA Fredericks, 1928
Type species.—Productus juresanensis Tschernyschew,
1902.
Juresania sp.
(Fig. 20B)
Juresania sp. Tazawa and Shintani, 2015, p. 45, fig. 3.2.
Material.—One specimen from locality SSK27, internalmould of a dorsal valve, NU-B1900.
Remarks.—The single dorsal valve specimen was previously
described by Tazawa and Shintani (2015, p. 45, fig. 3.2) as
Juresania sp. on account of its small size (length more than 24mm, width 27 mm) and flat visceral disc, covered internally by
numerous concentrically arranged endospines, and in having along, thin breviseptum, short converging buttress plates
enclosing end of breviseptum and the more posterior antron. The
Nagaiwa–Sakamotozawa species somewhat resembles
Juresania juresanensis (Tschernyschew, 1902, p. 276, 620, pl.

29, figs. 1, 2; pl. 47, figs. 1, 2; pl. 53, fig. 4), from the Cora
Limestone of Timan, northern Russia and from the Schwagerina
Limestone (Sakmarian) of the Urals, central Russia, in size andshape of the dorsal valve, but accurate comparison is difficult
due to the poorly preserved specimen.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Family WAAGENOCONCHIDAE Muir-Wood and Cooper,
1960
Subfamily WAAGENOCONCHINAE Muir-Wood and Cooper,
1960
Genus WAAGENOCONCHA Chao, 1927
Type species.—Productus humboldti d’Orbigny, 1842.
Waagenoconcha humboldti (d’Orbigny, 1842)(Fig. 20C, D)
Productus humboldti d’Orbigny, 1842, p. 54, pl. 5, figs. 4–7;
Tschernyschew, 1902, p. 275, 620, pl. 53, figs. 1–3;
Kozlowski, 1914, p. 40, pl. 7, figs. 7–9; Fredericks, 1925, p.
19, pl. 2, fig. 84.
Waagenoconcha humboldti (d’Orbigny). Chao, 1927, p. 86, pl.
15, figs. 2, 3; Sarytcheva in Sarytcheva and Sokolskaya,
1952, p. 98, pl. 15, fig. 109; Chronic, 1953, p. 86, pl. 15, figs.
4–7; Muir-Wood and Cooper, 1960, p. 252, pl. 89, figs. 6–10;
Samtleben, 1971, p. 60, pl. 2, figs. 17–19; Ifanova, 1972, p.
102, pl. 3, figs. 11–13; Tazawa, 1974, p. 125, pl. 1, figs. 2, 3;
pl. 2, fig. 1; pl. 4, fig. 6; Duan and Li, 1985, p. 108, pl. 35,
figs. 2, 3; Minato et al., 1979a, pl. 65, figs. 3, 4; Wang andZhang, 2003, p. 94, pl. 9, figs. 5–7; pl. 15, figs. 8–10; Tazawa
and Shintani, 2010, p. 56, fig. 4.1–4.5; Tazawa and Shintani,
2015, p. 45, fig. 4.1.
Material.—Six specimens from locality SSK27: (1) internalmould of a conjoined shell, with external mould of the dorsal
valve, NU-B1281; (2) external mould of a ventral valve,
NU-B1282; (3) external and internal moulds of a dorsal valve,
NU-B1283; (4) external mould of a dorsal valve, with umbonal
region of the opposite valve, NU-B1284; and (5) externalmoulds of two dorsal valves, NU-B1285, 1872.
Remarks.—These specimens were described by Tazawa andShintani (2010, p. 56, fig. 4.1–4.5; 2015, p. 45, fig. 4.1) as
Waagenoconcha humboldti (d’Orbigny, 1842) on account ofsize, shape and external ornament of both valves, especially thestrong rugae on the ventral valve. Waagenoconcha irginae(Stuckenberg, 1898, p. 220, pl. 2, fig. 16), from the lower
Permian (Asselian) of the Urals, is similar in general shape, butdiffers from W, humboldti in its larger size and finer spine bases
on both ventral and dorsal valves.
Occurrence.—SK1 Unit.
Distribution.—Gzhelian–Capitanian: northeastern Japan(Nagaiwa ‒ Sakamotozawa and Kamiyasse–Imo in the SouthCISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

25

FIGURE 22. Brachiopods of the SK1 assemblage (5). A, Terrakea sp., external latex cast (A1, A2) and internal mould (A3) of dorsal valve, NU-B1897; B,
Cyclacantharia sp., external latex cast (B1) and internal mould (B2, B3) of ventral valve, NU-B1870; C–F, Derbyia crassa (Meek and Hayden); C, externallatex cast (C1, C2) of ventral valve, NUB1223; D, external latex cast (D1) and internal mould (D2) of ventral valve, NU-B1225; E, external latex cast (E1)and internal mould (E2) of dorsal valve, NU-B1232; F, external latex cast (F1) and internal mould (F2) of dorsal valve, NU-B1239. Scale bars are 1 cm.

26

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

Kitakami Belt), northern Russia (Timan, Pechora Basin andnorthern Urals), western Russia (Moscow Basin), northern
China (Inner Mongolia), eastern Russia (South Primorye) and
Bolivia.
Superfamily AULOSTEGOIDEA Muir-Wood and Cooper,
1960
Family ECHINOSTEGIDAE Muir-Wood and Cooper, 1960
Subfamily ECHINOSTEGINAE Muir-Wood and Cooper, 1960
Genus EDRIOSTEGES Muir-Wood and Cooper, 1960
Type species.—Edriosteges multispinosus Muir-Wood andCooper, 1960.
Edriosteges cf. multispinosus Muir-Wood and Cooper, 1960
(Fig. 20E)
Edriosteges cf. multispinosus Muir-Wood and Cooper. Tazawa
and Shintani, 2015, p. 47, fig. 4.2.
Material.—Two specimens from locality SSK28: (1) external
and internal moulds of a ventral valve, NU-B1868; and (2)internal mould of a ventral valve, NU-B1869.
Remarks.—These specimens were described by Tazawa andShintani (2015, p. 47, fig. 42) as Edriosteges cf. multispinosus
Muir-Wood and Cooper, 1960. The Nagaiwa–Sakamotozawa
species resembles well the type species, Edriosteges
multispinosus Muir-Wood and Cooper, 1960, from the upper
Leonard Formation of Texas, in the medium size (length 33 mm,
width about 34 mm in the larger specimen, NU-B1868) andequidimensional subquadrate outline of the ventral valve, but
accurate comparison is difficult owing to lack of the dorsal
valve. Edriosteges sp. A, described by Tazawa and Araki (2014,
p. 47, fig. 3.5) from the upper Permian (Changhsingian)Nabekoshiyama Formation of the Kesennuma area, South
Kitakami Belt, differs from the present species in subtriangularoutline of the ventral valve.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Subfamily AGELESIINAE Cooper and Grant, 1975
Genus XENOSTEGES Muir-Wood and Cooper, 1960
Type species.—Xenosteges adherens Muir-Wood andCooper, 1960.
Xenosteges adherens Muir-Wood and Cooper, 1960
(Fig. 30G)
Xenosteges adherens Muir-Wood and Cooper, 1960, p. 112, pl.
10, figs. 1–13; Cooper and Grant, 1975, p. 855, pl. 232, figs.
1–48; pl. 233, fig. 40; pl. 234, figs. 28–32.
Material.—One specimen from locality SSK24, externalmould of a dorsal valve, NU-B2321.
Remarks.—This specimen can be referred to Xenosteges
adherens Muir-Wood and Cooper, 1960, redescribed by Cooper
and Grant (1975, p. 855, pl. 232, figs. 1–48; pl. 233, fig. 40; pl.
234, figs. 28–32), from the Cathedral Mountain, Cibolo andBone Spring Formations of Texas, by the medium-sized,
transverse dorsal valve (length 10 mm, width 17 mm), having
large ears and ornamented with irregular concentric lamellae.
Xenosteges magnus Cooper and Grant (1975, p. 858, pl. 234,
figs. 1–27), from the Road Canyon Formation of Texas, is also alarge, transverse Xenosteges species, but differs from X.
adherens in having low dorsal fold. Xenosteges anomalus
Cooper and Grant (1975, p. 859, pl. 235, figs. 1–60; pl. 236,
figs. 1–18), from the Cherry Canyon Formation of Texas, differsfrom the present species in having much smaller ears.
Occurrence.—SK4 Unit.
Distribution.—Sakmarian–Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt) and theUSA (Texas).
Family SCACCHINELLIDAE Licharew, 1928
Subfamily SCACCHINELLINAE Licharew, 1928
Genus SCACCHINELLA Gemmellaro, 1891
Type species.—Scacchinellsa variabilis Gemmellaro, 1897.
Scacchinella sp.
(Fig. 21A)
Scacchinella sp. Tazawa and Shintani, 2010, p. 58, fig. 5.2.
Materilal.—One specimen from locality SSK27, external andinternal moulds of a ventral valve, NU-B1286.
Remarks.—This specimen was previously described by
Tazawa and Shintani (2010, p. 58, fig. 5.2) as Scacchinella sp.
The ventral valve is large (length about 28 mm, width about 38mm), conical shape, having a high and flat interarea which isboth horizontally and longitudinally striated, and ornamented
with strong irregular rugae and numerous fine spine bases on theanterolateral slopes. The Nagaiwa–Sakamotozawa species
somewhat resembles Scacchinella gigantea Schellwien (1900b,
p. 35, pl. 4, figs. 1–3; pl. 5, figs. 1–8, text-figs. 5, 6, 8) from the
Trogkofel Formation (Sakmarian) of the Carnian Alps. The
present material is, however, too imperfect for comparison.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Superfamily LINOPRODUCTOIDEA Stehli, 1954
Family LINOPRODUCTIDAE Stehli, 1954
Subfamily LINOPRODUCINAE Stehli, 1954
Genus LINOPRODUCTUS Chao, 1927
Type species.—Productus cora d’Orbigny, 1842.

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

27

FIGURE 23. Brachiopods of the SK1 assemblage (6). A, Derbyia crassa (Meek and Hayden), external latex cast (A1) and internal mould (A2) of dorsal
valve, NU-B1240; B–F, Derbyia dorsosulcata Liu and Waterhouse; B, ventral (B1) and posterior (B2) views of external latex cast and internal mould (B3)of ventral valve, NU-B1261; C, external latex cast (C1) and internal latex cast (C2) of ventral valve, NU-B1262; D, internal mould of dorsal valve,
NU-B1272; E, ventral (E1) and posterior (E2, E3) views of external latex cast and internal mould (E4) of dorsal valve, NU-B1268; F, external latex cast (F 1)
and internal mould (F2) of dorsal valve, NU-B1269. Scale bars are 1 cm.

28

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

Linoproductus simensis (Tschernyschew, 1902)(Fig. 21B–D)
Productus simensis Tschernyschew, 1902, p. 286, 626, pl. 35,
fig. 7; pl. 55, figs. 2–5.
Linoproductus simensis (Tschernyschew). Volgin, 1960, p. 72,
pl. 8, fig. 1; Zhao, 1965, p. 425, pl. 1, figs. 6, 7; Bamber andWaterhouse, 1971, pl. 16, figs. 8, 11; Sergunkova andZhizhilo, 1975, p. 62, pl. 9, figs. 9, 10; pl. 10, figs. 8, 9; Lee
and Gu, 1976, p. 258, pl. 139, figs. 9–12; Tong, 1978, p. 231,
pl. 81, fig. 6; Kalashnikov, 1980, p. 47, pl. 10, figs. 8, 9 only;
Lee et al., 1980, p. 376, pl. 152, fig. 11; Tazawa et al., 2001,
p. 38, fig. 2D–J; Tazawa and Shintani, 2015, p. 48, fig.
4.3–4.5.
Linoproductus neimongolensis Lee and Gu, 1976, p. 258, pl.
178, figs. 1–10.
Material.—Seven specimens from locality SSK27: (1)external and internal moulds of two ventral valves, NU-B1875,
1876; (2) external and internal moulds of three dorsal valves,
NU-B1877–1879; (3) external mould of a dorsal valve,
NU-B1880; and (4) internal mould of a dorsal valve,
NU-B1881.
Remarks.—These specimens were previously described by
Tazawa and Shintani (2015, p. 48, figs. 4.3–4.5) as
Linoproductus simensis (Tschernyschew, 1902) on account of
its small size (length 37 mm, width 35 mm in the largest dorsal
valve specimen, NU-B1881; length 25 mm, width 23 mm in thebest preserved ventral valve specimen, NU-B1875), slightly
elongate oval outline and external ornament consisting of
numerous costellae and some rugae on the ventral valve.
Linoproductus neimongolensis Lee and Gu (1976, p. 258, pl.
178, figs. 1–10), from the lower Permian of the Dongujimqinqi,
Inner Mongolia, northern China, is deemed to be a juniorsynonym of L. simensis. The shells described by Chao (1927, p.
137, pl. 14, figs. 6–8) as Linoproductus simensis (Tschernyschew,1902), from the Visean of Guizhou, southwestern China, differsfrom L. simensis in having stronger and fewer costellae.
Linoproductus hayasakai Tazawa (1979, p. 26, pl. 4, figs. 5–11),
from the lower Kamiyasse Formation (Wordian) of Wayama in
the Kesennuma area, South Kitakami Belt, differs from L.
simensis in its larger, transverse shell and in having stronger
costellae on the ventral valve.
Occurrence.—SK1 Unit.
Distribution.—Bashkirian–Roadian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt), northern
Canada (northern Yukon Territory), northern Russia (northern
Urals), central Russia (southern Urals), Uzbekistan (Fergana),
northern China (Inner Mongolia) and southwestern China(Sichuan).
Genus COSTATUMULUS Waterhouse in Waterhouse andBriggs, 1986
Type species.—Auriculispina tumida Waterhouse in

Waterhouse et al., 1983.
Costatumulus pseudotruncata (Ustritsky, 1960)(Fig. 30D, E)
Cancrinella pseudotruncata Ustritsky, 1960, p. 36, pl. 6, figs.
10–12; pl. 7, figs. 1–3; Lee and Gu, 1976, p. 262, pl. 158, fig.
7; pl. 166, figs. 4, 7; Zhang et al., 1983, p. 296, pl. 130, figs.
13–16.
Costatumulus pseudotruncata (Ustritsky). Chen, 2004, p. 24, pl.
4, fig. 9; Chen and Shi, 2006, p. 159, pl. 12, figs. 13, 17–19;
pl. 15, fig. 4.
Material.—Two specimens from locality SSK24, external
and internal moulds of two ventral valves, NU-B2311, 2312.
Remarks.—These specimens are referred to Costatumulus
pseudotruncata (Ustritsky, 1960, p. 36, pl. 6, figs. 10–12; pl. 7,
figs. 1‒3), from the upper Sakmarian–lower Artinskian of thewestern Kunlun Mountains, Xinjiang, northwestern China, inthe medium size (length 19 mm, width 18 mm in the larger
specimen, NU-B2311) and in having irregular rugae and finenumerous costellae (numbering 4–5 in 2 mm at about
midlength) with quincuncially arranged elongate spine bases.
The type species, Costatumulus tumida (Waterhuse in
Waterhouse et al., 1983, p. 133, pl. 3, figs. 2–4, 6, 7), from theTiverton Formation (Artinskian) of the north Bowen Basin,
Queensland, eastern Australia, differs from C. pseudotruncata in
the larger size and in having more elongate and sparsely
arranged spine bases. Costatumulus cancriniformis(Tschernyschew, 1889, p. 283, 373, pl. 7, figs. 32, 33), from theArtinskian of the northern Urals, is distinguished from thepresent species by the strongly inflated ventral valve, which isornamented with numerous undulated concentric rugae.
Occurrence.—SK4 Unit.
Distribution.—Sakmarian–Wordian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt), northern
China (Inner Mongolia) and northwestern China (Xinjiang).
Subfamily ANIDANTHINAE Waterhouse, 1968b
Genus ANIDANTHUS Hill, 1950
Type species.—Linoproductus springsurensis Booker, 1932.
Anidanthus sp.
(Fig. 32G, H)
Material.—Two specimens from locality SSK17, externalmoulds of two dorsal valves, NU-B2301, 2302.
Remarks.—These specimens are safely assigned to the genus
Anidanthus Booker, 1932 on the basis of lamellose rugae on thevisceral disc of the dorsal valve and large, prominent ears withrugae on the valve. Anidanthus boikowi (Stepanov, 1946),
redescribed by Grigorjeva and Kotlyar (in Sarytcheva, 1977, p.
57, pl. 5, figs . 4–13), from the low er Permian of the
Verkhoyansk Range, northern Russia, somewhat resembles theCISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

29

FIGURE 24. Brachiopods of the SK1 assemblage (7). A, B, Meekella striatocostata (Cox); A, ventral (A 1), posterior (A2) and lateral (A3) views ofexternal latex cast of ventral valve, NU-B1212; B, external latex cast (B 1, B2) and internal mould (B 3) of dorsal valve, NU-B1213; C–E, Meekelladepressa Schellwien; C, external latex cast (C1) and internal mould (C2) of ventral valve, NU-B2379: D, external latex cast (D1) and internal mould (D2) ofventral valve, NU-B2380; E, external latex cast (E1, E2) and internal mould (E3) of dorsal valve, NU-B2881; F, G, Derbyia sakamotozawensis Shintani; F,
external latex cast (F1), internal latex cast (F2) and internal mould (F3) of ventral valve, NU-B1273; G, external latex cast (G1, G2) and internal mould (G3)of ventral valve, NU-B1274. Scale bars are 1 cm.

30

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

Kitakami species in having large ears but differs in the larger
and less transverse dorsal valve. The type species, Anidanthus
springsurensis (Booker, 1932, p. 67, pl. 3, figs. 1–6; pl. 4, figs.
1–7), from the lower Bowen Series of Queensland, eastern
Australia, differs from the present species in being lesstransverse outline. The Kitakami species is probably a new
species, although the material is poorly preserved and notadequate for the establishment.
Occurrence.—KN1 Unit.
Distribution.—Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Family KANSUELLIDAE Muir-Wood and Cooper, 1960
Subfamily AURICULISPININAE Waterhouse in Waterhouse
and Briggs, 1986
Genus AURICURLISPINA Waterhouse, 1975
Type species.—Cancrinella levis Maxwell. 1964.
Auriculispina kanmerai Tazawa and Shintani, 2015
(Figs. 21E–G, 30F)
Auriculispina kanmerai Tazawa and Shintani, 2015, p. 49, fig.
5.1–5.3.
Material.—Thirteen specimens from locality SSK24 andSSK27: (1) external and internal moulds of five ventral valves,
NU-B1882 (holotype), NU-B1883–1886; (2) external mould ofa ventral valve, NU-B2322; (3) internal moulds of three ventral
valves, NU-B1887–1889; (4) external and internal moulds oftwo dorsal valves, NU-B1890, 1891; and (4) external mould of
two dorsal valves, NU-B1892, 1893.
Remarks.—These specimens were described by Tazawa andShintani (2015, p. 49, figs. 5.1–5.3) as Auriculispina kanmerai
Tazawa and Shintani, 2015. This species is small in size (length
7 mm, width about 9 mm in the holotype, NU-B1882) for the
genus and transversely subquadrate in outline, and mostresembles Auriculispina capillata (Waterhouse, 1988, p. 156,
fig. 8), from the Grant Formation (Asselian–Sakmarian) of theCanning Basin, western Australia, in size and shape of the shell,
but differs from the Australian species in having finer capillaeon the ventral valve. The type species, Auriculispina levis(Maxwell, 1964, p. 34, pl. 6, figs. 15–18), from the upper
Carboniferous–lower Permian (Sakmarian) of the Yarrol Basin,eastern Australia, is readily distinguished from A. kanmerai by
the larger and more transverse shell and in having coarser
capillae on the ventral valve.
Occurrence.—SK1 and SK4 units.
Distribution.—Sakmarian–Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Subfamily PAUCISPINAURIINAE Waterhouse in Waterhouse
and Briggs, 1986
Genus TERRAKEA Booker, 1930

Type species.—Productus brachythaerus Morris in deStrezelecki, 1845.
Terrakea sp.
(Figs. 21H, I, 22A)
Terrakea sp. Tazawa and Shintani, 2015, p. 51, figs. 5.4–5.6.
Material.—Five specimens from localities SSK14 andSSK28: (1) external and internal moulds of two ventral valves,
NU-B1894, 1895; (2) internal mould of a ventral valve,
NU-B1896; and (3) external and internal moulds of two dorsal
valves, NU-B1897, 1898.
Remarks.—These specimens were described by Tazawa andShintani (2015, p. 51, fig. 5.4–5.6) as Terrakea sp. The
Nagaiwa–Sakamotozawa species is a large, transverse species(length 28 mm, width 38 mm in the largest specimen,
NU-B1895), and somewhat resembles the type species,
Terrakea brachythaera (Morris in de Strezelecki, 1845),
redescribed and refigured by Briggs (1998, p. 176, fig. 87A–I),
from the Broughton Formation of the Sydney Basin, eastern
Australia, in outer configuration, but accurate comparison isdifficult due to the ill-preserved specimens. Terrakea
nabekoshiyamensis Tazawa (2012, p. 26, fig. 4.13, 4.14), fromthe upper Permian (Changhsingian) Nabekoshiyama Formationof the Kesennuma area, South Kitakami Belt, is also transverse
in outline, but it differs from the present species by the muchsmaller dimensions.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Superfamily RICHTHOFENIOIDEA Waagen, 1885
Family TEGULIFERINIDAE Muir-Wood and Cooper, 1960
Subfamily CYCLACANTHARIINAE Cooper and Grant, 1975
Genus CYCLACANTHARIA Cooper and Grant, 1969
Type species.—Cyclacantharia kingorum Cooper and Grant,
1969.
Cyclacantharia sp.
(Fig. 22B)
Cyclacantharia sp. Tazawa and Shintani, 2015, p. 52, figs. 5.7, 5.8.
Material.—Two specimens from locality SSK14: (1) external
and internal moulds of a ventral valve, NU-B1870; and (2)internal mould of a ventral valve, NU-B1871.
Remarks.—These specimens were described by Tazawa andShintani (2015, p. 52, fig. 5.7, 5.8) as Cyclacantharia sp. on
account of the highly cone-shaped ventral valve (height about 40
mm in the larger specimen, NU-B1870), without a median
septum and with numerous strong spine bases, all around theinside of the cup aperture. The Nagaiwa–Sakamotozawa species
resembles the type species, Cyclacantharia kingorum Cooper

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

31

FIGURE 25. Brachiopods of the SK1 assemblage (8). A, B, Meekella nagaiwensis Shintani; A, ventral (A1), lateral (A2) and posterior (A3) views ofexternal latex cast of ventral valve, NU-B1216; B, external latex cast (B1, B2) and internal mould (B3) of dorsal valve, NU-B1217. Scale bars are 1 cm.

and Grant (1969, p. 7, pl. 5, figs. 13–16), from the Word
Formation of Texas, in size and shape of the ventral valve.
However, accurate comparison is difficult because of ill
preservation of the present material.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Suborder LYTTONIIDINA Williams, Harper and Grant, 2000
Superfamily LYTTONIOIDEA Waagen, 1883

Family LYTTONIIDAE Waagen, 1883
Subfamily POIKILOSAKINAE Williams, 1953
Genus PSEUDOLEPTODUS Stehli, 1956
Type species.—Pseudoleptodus getawayensis Stehli, 1956.
Pseudoleptodus sp.
(Fig. 32J, K)

32JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

Material.—Two specimens from locality SSK17: (1) external
and internal moulds of a ventral valve, NU-B2299; and (2)internal mould of a ventral valve, NU-B2300.
Remarks.—These specimens can be assigned to the genus
Pseudoleptodus Stehli, 1956 in having regularly arranged, low
and wide lateral septa, with flattened top. The
Nagaiwa–Sakamotozawa species is probably an advanced-formof the genus, and somewhat resembles Pseudoleptodus
getawayensis Stehli (1956), redescribed by Cooper and Grant(1974, p. 395, pl. 130, figs. 18–34) from the Cherry Canyon
Formation of the Guadalupe Mountains, Texas, in size andshape of the shell. But the Texan species differs from the present
species in having lateral septa extending anteriorly at steeper
angle to the median septum.
Occurrence.—KN1 Unit.
Distribution.—Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Superfamily PERMIANELLOIDEA He and Zhu, 1979
Family PERMIANELLIDAE He and Zhu, 1979
Genus DICYSTOCONCHA Termier and Termier in Termier etal., 1974
Type species.—Dicystoconcha lapparenti Termier andTermier in Termier et al., 1974.
Dicystoconcha lapparenti Termier and Termier in Termier et al., 1974
(Fig. 32I)
Dicystoconcha lapparenti Termier and Termier in Termier et al.,
1974, p. 123, pl. 22, figs. 1, 2, text-fig. 22; Wang and Jin,
1991, p. 495, pl. 1, figs. 1–9; pl. 3, figs. 1–7; Shen andTazawa, 2014, p. 248, fig. 3.1–3.5; Tazawa et al., 2014, p.
383, fig. 2.6; Tazawa, 2015, p. 73, fig. 6.6; Tazawa and Araki,
2018, p. 16, fig. 4.2 .
Dipunctella contricta Liang in Wang et al., 1982, p. 229, pl. 102,
fig. 3.
Guangjiayanella guangjiayanensis Yang, 1984, p. 212, pl. 31,
figs. 11–16, text-fig. 5.9.
Guangdongina xiamaoensis Mou and Liu, 1989, p. 458, pl. 1,
figs. 1–9; pl. 2, figs. 1– 7; text-fig. 5.
Guandongina leguminiformis Mou and Liu, 1989, p. 458, pl. 3,
figs. 4–6.
Guangdongina perforates Mou and Liu, 1989, p. 459, pl. 2, fig.
8; pl. 3, figs. 1–3.
Guangdongina sp. Mou and Liu, 1989, p. 459, pl. 2, fig. 9.
Peritisteges latesulcata Liang, 1990, p. 380, pl. 42, figs. 1, 2.
Fabulasteges planata Liang, 1990, p. 381, pl. 42, figs. 3, 4.
Material.—One specimen from locality SSK17, internalmould of a ventral valve, NU-B2280.
Remarks. —The single specimen from the lower part of the
Kanokura Formation (KN1 Unit) in the Nagaiwa–Sakamotozawa
area is referred to Dicystoconcha lapparenti Termier andTermier (in Termier et al., 1974, p. 123, pl. 22, figs. 1, 2,

text-fig. 22), from the lower Murgabian of Wardak, central
Afghanistan, in the small, ovate and bilobate ventral valve(length more than 11 mm, width about 13 mm), with a shallow
incision and a distinct central platform. As discussed by Shen
and Tazawa (2014, p. 248), the following six species from the
Kungurian–Capitanian of South China are junior synonyms ofDicystoconcha lapparenti: Guangjiayanella guangjiayanensis
Yang, 1984, Guangdongina xiamaoensis Mou and Liu, 1989, G.
leguminiformis Mou and Liu, 1989, G. perforatus Mou and Liu,
1989, Guangdongina sp. Mou and Liu, 1989 and Fabulasteges
planata Liang, 1990. Moreover, Paritisteges latesulcata Liang(1990, p. 380, pl. 42, figs. 1, 2), from the Wordian of Zhejiang,
eastern China, is also considered to be a junior synonym of thepresent species.
Occurrence.—KN1 Unit.
Distribution.—Kungurian‒Wuchiapingian: northeasternJapan (Nagaiwa–Sakamotozawa, Kamiyasse–Imo and Hitachi in
the South Kitakami Belt), Afghanistan, northern China (Inner
Mongolia), eastern China (Zhejiang) and central-southern China(Hubei and Guangdong).
Order ORTHOTETIDA Waagen, 1884
Suborder ORTHOTETIDINA Waagen, 1884
Superfamily ORTHOTETOIDEA Waagen, 1884
Family DERBYIIDAE Stehli, 1954
Genus DERBYIA Waagen, 1884
Type species.—Derbyia regularis Waagen, 1884.
Derbyia crassa (Meek and Hayden, 1858)(Figs. 22C‒F, 23A)
Orthisina crasa Meek and Hayden, 1858, p. 261.
Derbyia crassa (Meek and Hayden). Girty, 1915, p. 54, pl. 7,
fig. 1; Kelly, 1930, p. 138, pl. 11, fig. 4; Sayre, 1930, p. 93,
pl. 4, figs. 3–5; Dunbar and Condra, 1932, p. 79, pl. 3, figs.
1–12, text-fig. 3; Hoare, 1961, p. 27, pl. 1, figs. 17–23;
Sturgeon and Hoare, 1968, p. 26, pl. 3, figs. 1–4; Sutherlandand Harlow, 1973, p. 21, pl. 2, figs. 8–12; Brew and Beus,
1976, p. 894, pl. 1, figs. 14–18; Shintani, 2011, p. 81, figs.
4.4–4.8, 5.1–5.3.
Derbyia sp. C. Nakamura, 1972, p. 401, pl. 8, figs. 2, 4–6.
Derbyia sp. Minato et al., 1979a, pl. 46, figs. 7, 8; Tazawa andNakamura, 2015, p. 167, fig. 7.11–7.13.
Derbyia buchi (d’Orbigny). Tazawa and Shintani, 2014, p. 21,
fig. 3.5, 3.6.
Material.—Forty specimens from localities SSK14 andSSK28; (1) external and internal moulds of two conjoined
shells, NU-B1221, 1222; (2) external and internal moulds ofseven ventral valves, NU-B1223–1229; (3) internal moulds of
two ventral valves, NU-B1230, 1231; (4) external and internalmoulds of twenty dorsal valves, NU-B1232–1251; (5) externalmoulds of two dorsal valves, NU-B1252, 1253; and (6) internalmoulds of seven dorsal valves, NU-B1254–1260.

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

33

FIGURE 26. Brachiopods of the SK1 assemblage (9). A, Schuchertella cooperi Grant, external latex cast (A1, A2) and internal latex cast (A3) of ventral
valve, NU-B2433; B, C, Schuchertella sp., B, external latex cast (B1, B2) and internal latex cast (B3) of ventral valve, NU-B2436; C, external latex cast(C1) and internal latex cast (C2) of dorsal valve, NU-B2437; D, Streptorhynchus sibiricus Zavodowsky, external latex cast (D1, D2) and internal latex cast(D3) of dorsal valve, NU-B2434. Scale bars are 1 cm.

34

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

Remarks.—The specimens from Nagaiwa–Sakamotozawa
were previously described by Shintani (2011, p. 81, figs.
4.4–4.8, 5.1–5.3) as Derbyia crassa (Meek and Hayden, 1858)
on account of the small size (length 24 mm, width 26 mm in the
average- sized ventral valve specimen, NU-B1221), subquadrate
outline, and in having a low interarea, and ornamented withnumerous fine costellae (25–28 in 10 mm at midlength). Two
species, Derbyia sp. C Nakamura, 1972 and Derbyia sp. Minato
et al., 1979a, both from the Hosoo Formation (Kungurian) of
Nakadaira in the South Kitakami Belt, are assigned to thepresent species. Derbyia buchi (d’Orbigny, 1842), redescribedby Kozlowski (1914, p. 57, pl. 8, figs. 1–6) from the upper
Carboniferous of Bolivia, resembles D. crassa, but differs in its
larger size and in having coarser costellae on the ventral anddorsal valves.
Occurrence.—SK1 Unit.
Distribution.—Moscovian–Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa, Nakadaira and Kamiyasse in theSouth Kitakami Belt) and the USA (Michigan, Ohio, Missouri,
Nebraska, Kansas, Oklahoma, New Mexico and Arizona).
Derbyia dorsosulcata Liu and Waterhouse, 1985
(Fig. 23B–F)
Magniderbyia sp. Nakamura, 1972, p. 403, pl. 9, fig. 2.
Derbyia dorsosulcata Liu and Waterhouse, 1985, p. 11, pl. 1,
figs. 1, 7, 8, 10; Wang and Zhang, 2003, p. 121, pl. 25, figs. 1,
2; pl. 26, figs. 2–6; Shintani, 2011, p. 84, figs. 5.4–5.6,
6.1–6.4; Tazawa and Shintani, 2014, p. 21, fig. 3.8.
Material.—Twelve specimens from localities SSK14 andSSK28: (1) external and internal moulds of four ventral valves,
NU-B1261–1264; (2) external mould of a ventral valve,
NU-B1265; (3) internal moulds of two ventral valves,
NU-B1266, 1267; (4) external and internal moulds of four
dorsal valves, NU-B1268–1271; and (5) internal mould of adorsal valve, NU-B1272.
Remarks.—These specimens were described by Shintani(2011, p. 84, figs. 5.1–5.6, 6.1–6.4) as Derbyia dorsosulcata Liu
and Waterhouse, 1985 on account of the large (length 44 mm,
width 64 mm in the best preserved ventral valve specimen,
NU-B1261), subrectangular shell, with a low, transversely
subtriangular interarea and moderately deep dorsal sulcus, theexternal ornament consisting of a few rugae and numerous
costellae with broad interspaces, and the large muscle scars in
the ventral valve. Comparison with the other Derbyia species isfully discussed by Shintani (2011, p. 86).
Occurrence.—SK1 Unit.
Distribution.—Sakmarian–Wordian: northeastern Japan(Nagaiwa–Sakamotozawa and Kamiyasse in the South KitakamiBelt) and northern China (Inner Mongolia).
Derbyia sakamotozawensis Shintani, 2011
(Fig. 24F, G)

Derbyia sakamotozawensis Shintani, 2011, p. 86, fig. 7.1–7.4.
Material.—Seven specimens from locality SSK14: (1)external and internal moulds of four ventral valves, NU-B1273(holotype), 1274–1276; (2) internal moulds of two ventral
valves, NU-B1277, 1278; and (3) internal mould of a dorsal
valve, NU-B1279.
Remarks.—Derbyia sakamotozawensis Shintani, 2011, fromthe lower part (SK1 Unit) of the Sakamotozawa Formation in
Nagaiwa–Sakamotozawa, is large, semicircular species (length
49 mm, width 61 mm in the holotype, NU-B1273), with external
ornament consisting of numerous fine costellae (17–20 in 10mm at about midlength of ventral valve), and a large muscle
scar in the ventral valve. Derbyia wabaunsensis Dunbar andCondra (1932, p. 95, pl. 7, figs. 1–6), from the Wabaunsee
Group of Nebraska in the USA, is similar to D.
sakamotozawensis in size and outline of the shell, but theAmerican species differs from the Nagaiwa‒Sakamotozawa
species in having strongly concave ventral valve and smallerve ntra l mus c le s c a r. The pre c e ding s pe c ie s , D erbyi adorsosulcata Liu and Waterhouse, differs from D.
sakamotozawensis in the subrectangular outline and in having
smaller muscle scar in the ventral valve.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Family MEEKELLIDAE Stehli, 1954
Subfamily MEEKELLINAE Stehli, 1954
Genus MEEKELLA White and St. John, 1867
Type species.—Plicatula striatocostata Cox, 1857.
Meekella striatocostata (Cox, 1857)(Fig. 24A, B)
Plicatula striatocostata Cox, 1857, p. 568, pl. 8, fig. 7.
Meekella striatocostata (Cox). Tschernyschew, 1902, p. 211,
582, pl. 24, figs. 7, 8; pl. 26, fig. 4; pl. 51, fig. 3; Girty, 1909,
p. 54, pl. 6, fig. 6; Hamlet, 1928, p. 9, pl. 1, fig. 5; Dunbar andCondra, 1932, p. 125, pl. 16, figs. 1–10; pl. 17, fig. 3;
Sturgeon and Hoare, 1968, p. 24, pl. 2, figs. 1–3; West, 1977,
p. 741, text-fig. 3; Li (Lee) et al., 1986, p. 218, pl. 1, fig. 6;
Shintani, 2011, p. 77, fig. 3.1–3.3.
Meekella cf. striatocostata (Cox). Nakamura, 1972, p. 385, pl.
5, fig. 1; Minato et al., 1979a, pl. 59, figs. 3, 4.
Meekella cf. striatocostata (Cox). Sutherland and Harlow, 1973,
p. 20, pl. 1, fig. 18.
Material.—Six specimens from localities SSK14, SSK27 andSSK28: (1) external and internal moulds of a ventral valve,
NU-B2377; (2) external mould of a ventral valve, NU-B1212;(3) external and internal moulds of two dorsal valves,
NU-B1213, 1214; (4) internal mould of a ventral valve,
NU-B2378; and (5) external mould of a dorsal valve,

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

35

FIGURE 27. Brachiopods of the SK1 assemblage (10). A, Streptorhynchus sp., external latex cast (A1, A2) and internal latex cast (A3) of dorsal valve,
NU-B2435; B, Rhynchopora sp., internal mould (B1, B2) of ventral valve, NU-B1280; C–E, Hustedia ratburiensis Waterhouse and Piyasin; C, ventral(C1, C2) and dorsal (C3) views of external latex cast of conjoined shell, NU-B2371; D, ventral (D 1) and dorsal (D2) views of internal mould of conjoined
shell, NU-B2374; E, internal mould of dorsal valve, NU-B2376; F, G, Squamularia asiatica Chao; F, external latex cast (F1, F2) of ventral valve,
NU-B2429; G, ventral (G1, G2) and dorsal (G3) views of external latex cast, and ventral (G4) and dorsal (G5) views of internal mould of conjoined shell,
NU-B2424; H, Pinegathyris royssiana (von Keyserling), external latex cast (H1, H2) and internal mould (H3) of ventral valve, NU-B2285. Scale bars are 1
cm.

36

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANINU-B1215.
Remarks.—Most of the specimens from
Nagaiwa–Sakamotozawa were previously described by Shintani(2011, p. 77, fig. 3.1–3.3) as Meekella striatocostata (Cox,1857). This species is medium in size for the genus (length 29mm, width 42 mm in the best preserved and average-sized
dorsal valve specimen, NU-B1213), subtriangular in outline,with the greatest width near midlength of ventral valve; externalsurface of both valves are ornamented with regular, rounded
costae and fine costellae, numbering 13–14 costae and 15–17
costellae in 5 mm at midlength. Meekella skenoides Girty (1908,
p. 206, pl. 30, figs. 8, 9), from the Delaware Mountain
Formation of Texas, is similar in size and trigonal outline, butdiffers in having more strongly convex ventral valve and rather
angular costae. Meekella eximia (von Eichwald, 1840, p. 157,
pl.17, figs. 4–8), from the Kasimovian of Kasimov, western
Russia, differs from M. striatocostata in having less convex
ventral valve with flat interarea.
Occurrence.—SK1 Unit.
Distribution.—Bashkirian–Wuchiapingian: northeasternJapan (Nagaiwa–Sakamotozawa and Imo in the South KitakamiBelt), the USA (Ohio, Kentucky, Iowa, Missouri, Nebraska,
Kansas and New Mexico), central Russia (southern Urals),
central-southern China (Guangxi) and Indonesia (Timor).
Meekella depressa Schellwien, 1900b(Fig. 24C–E)
Meekella depressa Schellwien, 1900b, p. 23, pl. 3, figs. 3, 4;
Gauri, 1965, p. 74, pl. 10, fig. 8; Ramovs, 1965, pl. 15, fig. 4.
Material.—Three specimens from locality SSK27: (1)external and internal moulds of two ventral valves, NU-B2379,
2380; and (2) external and internal moulds of a dorsal valve,
NU-B2381.
Description.—Shell small in size for genus, transverselly
subrectangular in outline, hinge long but shorter than thegreatest width at about midlength; length 15 mm, width 25 mm
in the largest specimen (NU-B2379). Ventral valve moderately
convex in lateral and anterior profiles; sulcus absent. Dorsal
valve gently convex in both lateral and anterior profiles; no fold.
External surface of both valves ornamented with costae andnumerous fine costellae; costae occur anterior one third of valve,
numbering 7–8 on both valves; costellae rounded, numbering
8–9 in 2 mm at about midlength of dorsal valve. Ventral interior
with long, subparallel dental plates. Dorsal interior with a pair oflong, widely divergent socket plates. Other internal structures ofboth valves not well preserved.
Remarks.—These specimens can be referred to Meekelladepressa Schellwien (1900b, p. 23, pl. 3, figs. 3, 4), from the
Trogkofel Formation of the Carnian Alps, Austria, in the small
size, transverse outline and in having costae on anterior one
third of both ventral and dorsal valves. Meekella bisculpta Grant(1976, p. 58, pl. 10, figs. 1–35), from the Ratburi Formation ofKo Muk, southern Thailand, is also characterized by the costae

beginning relatively far forward, but the Thailand species differsfrom M. depressa in being less transverse outline.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt), Austriaand Slovenia.
Meekella nagaiwensis Shintani, 2011
(Fig. 25A, B)
Meekella nagaiwensis Shintani, 2011, p. 79, figs. 3.4–3.5,
4.1–4.3.
Material.—Seven specimens from localities SSK14, SSK27
and SSK28; (1) external and internal moulds of a ventral valve,
NU-B1216 (holotype); (2) external and internal moulds of two
dorsal valves, NU-B1217, 1218; (3) internal mould of a ventral
valve, NU-B2382; (4) external mould of two dorsal valves,
NU-B1219, 2383; and (5) internal mould of a dorsal valve,
NU-B1220.
Remarks.—Meekella nagaiwensis Shintani, 2011 wasdescribed by Shintani (2011, p. 79, figs. 3.4–3.5, 4.1–4.3) fromthe lower part (SK1 Unit) of the Sakamotozawa Formation in
Nagaiwa–Sakamotozawa, South Kitakami Belt. This species islarge (length 58 mm, width 82 mm in the largest specimen,
NU-B1216), transverse Meekella, having strongly convex
ventral valve and slightly convex to nearly flat dorsal valve, andornamented with somewhat irregular costae and fine numerous
costellae (14–16 in 5 mm at midlength) on both valves.
Meekella kueichowensis Huang (1933, p. 27, pl. 3, figs. 19–21;
pl. 4, figs. 1–4), from the Lungtan Formation of Guizhou,
southwestern China, resembles M. nagaiwensis in the large size,
but differs from the latter in the less transverse outline and inhaving more regular costae on the both valves. Meekella grandis
King (1931, p. 54, pl. 6, figs. 5–7), from the Leonard Formationof Texas, is also a large-sized species, but differs from M.
nagaiwensis in the less transverse outline and in having coarser
costellae. Meekella gigantea Hayasaka (1933, p. 26, pl. 6, fig. 2;
pl. 9, figs. 3, 4; pl. 10, figs. a, b; pl. 11, figs. a, b; pl. 12, fig. 1,
text-fig. 4) from the Nabeyama Formation (Kungurian) of Kuzu,
central Japan, is readily distinguished from the present species
by the larger size and much coarser costae on both valves.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa‒Sakamotozawa in the South Kitakami Belt).
Family SCHUCHERTELLIDAE Williams, 1953
Subfamily SCHUCHERTELLINAE Williams, 1953
Genus SCHUCHERTELLA Girty, 1904
Type species.—Streptorhynchus lens White, 1862.
Schuchertella cooperi Grant, 1976
(Fig. 26A)CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

37

FIGURE 28. Brachiopods of the SK1 assemblage (11). A–E, Choristites sp.; A, external latex cast (A 1) and internal mould (A 2) of ventral valve,
NU-B2386; B, external latex cast of ventral valve, NU-2387; C, internal mould of dorsal valve, NU-B2390; D, internal mould of dorsal valve, NU-B2391;
E, external latex cast of ventral valve, NU-B2388. Scale bars are 1 cm.

Schuchertella cooperi Grant, 1976, p. 42, pl. 4, figs. 1–42; pl. 6,
figs. 1–12.
Material.—One specimen from locality SSK27, external andinternal moulds of a ventral valve, NU-B2433.
Remarks.—This specimen can be referred to Schuchertella
cooperi Grant (1976, p. 42, pl. 4, figs. 1–42; pl. 6, figs. 1–12),from the Ratburi Formation of Ko Muk, southern Thailand, inthe medium size (length 20 mm, width 21 mm), semicircular,
flattened ventral valve, externally ornamented with numerous
costellae (numbering 12–14 in 5 mm at about midlength) andseveral irregular concentric lamellae, and the interior with nomedian septum. Schuchertella semiplana (Waagen, 1884, p.
608, pl. 55, figs. 1, 2), from the Chhidru Formation of the Salt
Range, Pakistan, differs from Schuchertella cooperi in theslightly larger size and in having coarser costellae on the ventral
valve. Schuchertella debaisiensis Wang and Zhang (2003, p.
122, pl. 27, figs. 1–9; pl. 29, figs. 1, 2), from the Dashizhai andZhesi Formations of Inner Mongolia, northern China, differsfrom the present species in the larger size and more transverse

outline.
Occurrence.—SK1 Unit
Distribution.—Sakmarian ‒ Kungurian: northeastern Japan(Nagaiwa ‒ Sakamotozawa in the South Kitakami Belt) andsouthern Thailand (Ko Muk).
Schuchertella sp.
(Fig. 26B, C)
Material.—Two specimens from locality SSK27: (1) external
and internal moulds of a ventral valve, NU-B2436; and (2)external and internal moulds of a dorsal valve, NU-B2437.
Description.—Shell large in size for genus, transversely
subrectangular in outline, hinge slightly shorter than greatest
width at about midlength; length 23 mm, width about 38 mm in
the ventral valve specimen (NU-B2436); length 28 mm, widthabout 43 mm in the dorsal valve specimen (NU-B2437). Dorsal
valve nearly flat, ornamented with numerous costellae,
numbering 9–10 in 5 mm at about midlength. Dorsal interior
with a pair of strong socket ridges and a short, thin median

38

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

ridge. Muscle scar not well preserved.
Remarks.—These specimens are safely assigned to the genus
Schuchertella Girty, 1904 by the presence of median ridge in thedorsal valve. The Nagaiwa–Sakamotozawa species resembles
Schuchertella bassa Grant (1995, p. 658, fig. 4.1–4.16), fromthe upper Permian (Dorashamian) of Hydra Island, Greece, insize and shape of the dorsal valve, but differs from the Greek
species in having more regular and slightly coarser costellae.
The preceding species, Schuchertella cooperi Grant, is readily
distinguished from the present species in much smaller size andless transverse outline. The present species may be a new
species, although the material is inadequate for establishment.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotoawa in the South Kitakami Belt).
Subfamily STREPTORHYNCHINAE Stehli, 1954
Genus STREPTORHYNCHUS King, 1850
Type species.—Terebratulites pelargonatus von Schlotheim,
1816.
Streptorhynchus sibiricus Zavodowsky, 1968
(Fig. 26D)
Streptorhynchus sibiricus Zavodowsky, 1968, p. 89, pl. 32, fig.
13; Zavodowsky and Stepanov, 1970, p. 73, pl. 1, figs. 6, 7.
Material.—One specimen from locality SSK14, external andinternal moulds of a dorsal valve, NU-B2434.
Remarks.—The single specimen from
Nagaiwa–Sakamotozawa is lacking the ventral valve, but can bereferred to Streptorhynchus sibiricus Zavodowsky (1968, p. 89,
pl. 32, fig. 13), from the lower Permian (Asselian) of theKolyma Basin, northern Russia, in the medium size (length 31mm, width about 32 mm), moderately convex dorsal valve,ornamented with numerous costellae (numbering 11–12 in 5 mmnear anterior margin of the valve) and irregularly developed
strong concentric rugae, and the internal structure, consisting ofstrong socket plates and large cardinal process. Streptorhynchus
kayseri Schellwien, 1900a, redescribed by Grabau (1931, p. 241,
pl. 24, fig. 1) from the Zhesi Formation of Zhesi, Inner
Mongolia, northern China, differs from S. sibiricus in the muchtransverse dorsal valve. The type species, Streptorhynchus
pelargonatus von Schlotheim, 1816, redescribed by Waagen(1884, p. 579, pl. 50, figs. 3–5, 7), from the Wargal Formationof the Salt Range, Pakistan, is readily distinguished from thepresent species in being much smaller size.
Occurrence.—SK1 Unit.
Distribution.—Asselian–Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt) andnorthern Russia (Kolyma).
Streptorhynchus sp.
(Fig. 27A)

Material.—One specimen from locality SSK27, external andinternal moulds of a dorsal valve, NU-B2435.
Remarks.—This specimen is safely assigned to the genus
Streptorhynchus King, 1850 by the medium-sized (length about
22 mm, width about 22 mm), subcircular and moderately convex
dorsal valve, which is externally ornamented with numerous finecostellae (18–19 in 5 mm near the anterior margin of the valve)and irregular strong concentric rugae, and internally provided
with prominent muscle scars and large cardinal process. The
Nagaiwa–Sakamotozawa species most resembles Streptorhynchus
zhesiensis Duan and Li (1985, p. 105, 202, pl. 32, figs. 7, 8),from the Yihewusu Formation of Inner Mongolia, northern
China, in size, shape and external ornament of the dorsal valve.
But accurate comparison is difficult owing to lack of the ventral
valve. Streptorhynchus khwaense Grant (1976, p. 49, pl. 7, figs.
1–41), from the Ratburi Formation of Ban Kao and Khao Phrik,southern Thailand, differs from the present species in having
slightly irregular and stronger costellae on the dorsal valve. The
preceding species, Streptorhynchus sibiricus Zavodowsky,
1968, is distinguished from the present species in the larger size
and in having coarser costellae on the dorsal valve.
Occurrence.—SK1 Unit.
Distribution.—Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt) .
Order RHYNCHONELLIDA Kuhn, 1949
Superfamily STENOSCISMATOIDEA Oehlert, 1887
Family STENOSCISMATIDAE Oehlert, 1887
Subfamily STENOSCISMATINAE Oehlert, 1887
Genus STENOSCISMA Conrad, 1839
Type species.—Terebratula schlottheimii von Buch, 1834.
Stenoscisma sp.
(Fig. 32L)
Material.—One specimen from locality SSK17, external andinternal moulds of a ventral valve, NU-B2306.
Remarks.—This specimen is safely assigned to the genus
Stenoscisma Conrad, 1839 by the rhynchonellid-formed
shell, with a spondylium in the ventral valve. The
Nagaiwa–Sakamotozawa species is medium in size (length 15mm, width 27 mm), transversely subtrigonal in shape, andexternal surface of the ventral valve is ornamented with strong
costae, numbering 4 on sulcus and 5 on each flank. Stenoscisma
hueconianum (Girty, 1929), redescribed by Cooper and Grant(1976a, p. 2096, pl. 563, figs. 1–54), from the upper
Wolfcampian of Texas, resembles the present species in shapeand external ornament of the ventral valve, but differs from thelatter in the much smaller size. Stenoscisma mutabilis(Tschernyschew, 1902, p. 81, 491, pl. 22, fig. 18; pl. 23, fig. 10;
pl. 45, figs. 1–15; pl. 46, fig. 14), from the Schwagerina Horizonof the Urals and Timan, resembles the present species in size,
shape and external ornament of the ventral valve, particularly inCISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

39

FIGURE 29. Brachiopods of the SK1 assemblage (12). A, B, Callispirina ornata (Waagen); A, external latex cast (A1, A2) and internal mould (A3) ofventral valve, NU-B2392; B, external latex cast (B 1 , B 2 ) and internal mould (B 3 ) of dorsal valve, NU-B2393; C–F, Spiriferellina cristata (vonSchlotheim); C, ventral (C1, C2) and dorsal (C4) views of external latex cast, and ventral (C3) and dorsal (C5) views of internal mould of conjoined shell,
NU-B2394; D, ventral view (D1, D2) of external latex cast, and ventral (D3) and dorsal (D4) views of internal mould of conjoined shell, NU-B2395; E,
ventral (E1, E2) and dorsal (E4) views of internal mould, and dorsal view (E3) of external latex cast of conjoined shell, NU-B2396; F, external latex cast(F1, F2) and internal mould (F3) of ventral valve, NU-B2400. Scale bars are 1 cm.

40

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

the medium-sized, transversely subtrigonal specimen (illustrated
by Tschernyschew, 1902, pl. 45, fig. 14). However, an accuratecomparison is difficult for this poorly preserved specimen.
Occurrence.—KN1 Unit.
Distribution.—Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Superfamily RHYNCHOPOROIDEA Muir-Wood, 1955
Family RHYNCHOPORIDAE Muir-Wood, 1955
Subfamily RHYNCHOPORINAE Muir-Wood, 1955
Genus RHYNCHOPORA King, 1865
Type species.—Terebratula geinitziana de Verneuil, 1845.
Rhynchopora sp.
(Fig. 27B)
Rhynchopora sp. Tazawa and Shintani, 2010, p. 58, fig. 5.1.
Material.—Two spcimens from localities SSK5 and SSK28:(1) external and internal moulds of a ventral valve, NU-B2432;
and (2) internal mould of a ventral valve, NU-B1280.
Remarks.—One of the specimens (NU-B1280) was
previously described by Tazawa and Shintani (2010, p. 58, fig.
5.1) as Rhynchopora sp. The ventral valve is small in size forthe genus (length about 8 mm, width about 12 mm), transversely
subpentagonal in outline, having a broad and deep sulcus,
ornamented with simple costae numbering 5 on sulcus and 4 oneach lateral slope, and possessing thin, short dental plates in thevalve. The Nagaiwa–Sakamotozawa species resembles
Rhynchopora tschernyshae Koczyrkevicz (1979, p. 47, pl. 11,
figs. 1–4, text-fig. 4) from the lower Barabaschevka Formation(Wordian) of South Primorye, eastern Russia in size, shape andexternal ornament of the ventral valve. However, the single
imperfect specimen does not allow specific assignment.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Superfamily RHYNCHOTETRADOIDEA Licharew, 1956
Family RHYNCHOTETRADIDAE Licharew, 1956
Subfamily RHYNCHOTETRADINAE Licharew, 1956
Genus TRASGU Martinez Chacon, 1979
Type species.—Trasgu minor Martinez Chacon, 1979.
Trasgu confinensis (Schellwien, 1892)(Fig. 31E, F)
Rhynchonella confinensis Schellwien, 1892, p. 54, pl. 8, figs.
11, 12; Schellwien, 1900b, p. 93, pl. 14, figs. 8–10.
Material.—Four specimens from locality SSK24: (1)external and internal moulds of two ventral valves, NU-B2307,
2308; and (2) internal mould of two ventral valves, NU-B2309,

2310
Description.—Shell large in size for genus, subtriangular inoutline, widest at two-thirds length from umbo; length 23 mm,
width 22 mm in the largest specimen (NU-B2307). Ventral
valve gently and unevenly convex in lateral profile, strongly
convex at umbo, gently convex to almost flattened at broad
venter, and strongly convex near anterior margin; sulcus wideand shallow on anterior half of valve. Dorsal valve moderately
convex, with wide and low fold. External surface of ventral
valve ornamented with strong costae, occurring nearantero-lateral margins, numbering five on each lateral flank.
Interior of ventral valve, distinct spondylium with median
septum in posterior portion. Interior of dorsal vale not wellpreserved.
Remarks.—These specimens are referred to Trasgu
confinensis (Schellwien, 1892), originally described bySchellwien (1892, p. 54, pl. 8, figs. 11, 12) as Rhynchonellaconfinensis Schellwien, 1892, from the Trogkofel Formation of
the Karavanke Mountains, Slovenia, in the large size. The typespecies, Trasgu minor Martinez Chacon (1979, p. 252, pl. 30,
figs. 8–23, text-fig. 13), from the lower Bashkirian of theCantabrica Mountains, northern Spain, differs from T.
confidensis in the much smaller size.
Occurrence.—SK4 Unit.
Distribution.—Sakmarian–Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt) andSlovenia (Karavanke Mountains).
Order ATHYRIDIDA Boucot, Johnson and Staton, 1964
Suborder ATHYRYDIDINA Boucot, Johnson and Staton, 1964
Superfamily ATHYRIDOIDEA Davidson, 1881
Family ATHYRIDIDAE Davidson, 1881
Subfamily CLEIOTHYRIDININAE Alvarez, Rong and Boucot,
1998
Genus PINEGATHYRIS Grunt, 1980
Type species.—Terebratula royssiana von Keyserling, 1846.
Pinegathyris royssiana (von Keyserling, 1846)(Figs. 27H)
Terebratula royssiana von Keyserling, 1846, p. 237.
Athyris (Cliothyris) royssiana (von Keyserling). Tschernyschew,
1902, p. 103, 511, pl. 43, figs. 11, 12.
Athyris royssiana (von Keyserling). Netschajew, 1911, p. 93,
153, pl. 13, figs. 1–7.
Athyris royssiana Tschernyschew. Wiman, 1914, p. 30, pl. 1,
figs. 21–55; pl. 2, figs. 1–13; Frebold. 1950, p. 69, pl. 6, figs. 6, 7.
Cleiothyris royssiana aberr. typica Netschaew. Fredericks, 1934,
p. 23, pl. 5, figs. 1–10.
Athyris (Cleiothyridina) royssiana (von Keyserling). Stepanov,1937, p. 156, 181, pl. 9, figs. 8–10.
Cleiothyridina royssiana (von Keyserling). Gobbett, 1963, p.
161, pl. 21, figs. 13–16; pl. 22, figs. 1, 2; Grigorjeva, 1967,
pl. 8, figs. 1–3; Stepanov et al., 1975, pl. 3, figs. 20, 21.

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

41

FIGURE 30. Brachiopods of the SK4 assemblage (1). A–C, Anemonaria kitakamiense sp. nov.; A, external latex cast (A1), external mould (A2), internalmould (A3) and lateral view of external latex cast A4) of ventral valve, NU-B2325 (holotype); B, internal mould of ventral valve, NU-B2326; C, internalmould of ventral valve, NU-B2329; D, E, Costatumulus pseudotruncata (Ustritsky); D, external latex cast (D1) and internal mould (D2) of ventral valve,
NU-B2311; E, external latex cast (E1, E2), and internal mould (E3) of ventral valve, NU-B2312; F, Auriculispina kanmerai Tazawa and Shintani, externallatex cast (F1) and external mould (F2, F3) of ventral valve, NU-B2322; G, Xenosteges adherens Muir-Wood and Cooper, external latex cast (G1, G2) andinternal mould (G3 ) of dorsal valve, NU-B2321; H, I, Hustedia ratburiensis Waterhouse and Piyasin; H, internal mould (H 1 , H 2 ) of dorsal valve,
NU-B2323; I, internal mould (I1, I2) of dorsal valve, NU-B2324. Scale bars are 1 cm.

Pinegathyris royssiana roysiana (von Keyserling). Grunt, 1980,
p. 91, pl. 10, figs. 1–7; pl. 11, figs. 1, 2, text-figs. 44–46;
Kalashnikov, 1986, pl. 130, figs. 1, 2.
Pinegathyris royssiana (von Keyserling). Grunt, 2006, p. 166, pl.
17, figs. 4, 8.
Material.—One specimen from locality SSK14, external andinternal moulds of a ventral valve, NU-B2385.
Remarks.—This specimen can be referred to Pinegathyris
royssiana (von Keyserling, 1846), redescribed by Grunt (2006,
p. 166, pl. 17, figs. 4, 8) from the upper Kazanian of the Kanin
Peninsula, northern Russia, in the large, transverse ventral valve(length about 25 mm, width about 50 mm), with a narrow anddeep sulcus, and external ornament consisting of growth42

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

lamellae bearing numerous flat spines (numbering 5–6 in 2 mm
at about midlength of the ventral valve). Pinegathyris alata(Grunt, 1980, p. 92, pl. 10, fig. 8), from the Kazanian of Pinega,
northern Russia, differs from P. royssiana in its more transverse
outline.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian–Wordian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt), northern
Russia (Kanin Peninsula, Pinega and Timan), Greenland,
Svalbard (Spitsbergen), and central Russia (southern Urals).
Suborder RETZIIDINA Boucot, Johnson and Staton, 1964
Superfamily RETZIOIDEA Waagen, 1883
Family NEORETZIIDAE Dagys, 1972
Subfamily HUSTEDIINAE Grunt, 1986
Genus HUSTEDIA Hall and Clarke, 1893
Type species.—Terebratula mormoni Marcou, 1858.
Hustedia ratburiensis Waterhouse and Piyasin, 1970
(Figs. 27C–E, 30H, I, 33A, B)
Hustedia ratburiensis Waterhouse and Piyasin, 1970, p. 138, pl.
23, figs. 15–30; Grant, 1976, p. 241, pl. 66, figs. 1–69: pl. 67,
figs. 51–58; Yanagida and Nakornsri, 1999, p. 118, pl. 32,
figs. 11–16; Archbold, 1999, figs. 5E–H; Tazawa, 2001, p.
299, fig. 8.6; Tazawa, 2008b, p. 53, fig. 8.2–8.6; Tazawa in
Tazawa et al., 2015, p. 44, fig. 6.7; Tazawa and Nakamura,2015, p. 169, fig. 7.1–7.7.
Hustedia nakornsrii Yanagida, 1970, p. 79, pl. 14, fig. 9.
Material.—Eleven specimens from localities SSK17, SSK24,
SSK27 and SSK28: (1) external and internal moulds of aconjoined shell, NU-B2371; (2) internal moulds of two
conjoined shells, with external moulds of the ventral valves,
NU-B2372, 2373; (3) internal moulds of two conjoined shells,with external mould of the dorsal valves, NU-B 2303, 2374; (4)external and internal moulds of a dorsal valve, NU-B 2375; (5)external moulds of a dorsal valve, NU-B 2304; and (6) internalmoulds of four dorsal valves, NU-B2305, 2323, 2324, 2376.
Remarks.—These specimens are referred to Hustedia
ratburiensis Waterhouse and Piyasin (1970, p. 138, pl. 23, figs.
15–30), from the Wordian of Khao Phrik, southern Thailand, by
the medium size (length 11 mm, width 7 mm in the best
preserved specimen, NU-B2303; length 8 mm, width 7 mm inan average-sized specimen, CU40) and in having rounded costae
which occur three close-set medianly and four pairs laterally on
the dorsal valve. Hustedia nakornsrii Yanagida (1970, p. 79, pl.
14, fig. 9), from the Ratburi Limestone of Khao Phrik, isdeemed to be conspecific with the present species. Hustedia
indica (Waagen, 1883, p. 493, pl. 35, figs. 1, 2), from theWargal Formation of the Salt Range, differs from H .
ratburiensis in having fewer and broader costae on both valves.
Occurrence.—SK1, SK4 and KN1 units.
Distribution.—Sakmarian–Wuchiapingian: northeasternJapan (Nagaiwa–Sakamotozawa, Nakadaira and Takakurayama
in the South Kitakami Belt), central Japan (Hida Gaien Belt),southwestern Japan (Mizukoshi in central Kyushu Kyushu),north-central Thailand (Khao Hin King) and southern Thailand(Khao Phrik, Khao Tok Nam and Ko Muk).
Order SPIRIFERIDA Waagen, 1883
Suborder SPIRIFERIDINA Waagen, 1883
Superfamily MARTINIOIDEA Waagen, 1883
Family MARTINIIDAE Waagen, 1883
Subfamily MARTINIINAE Waagen, 1883
Genus MARTINIA M’Coy, 1844
Type species.—Spirifer glaber Sowerby, 1820.
Martinia lata Grabau, 1936
(Fig. 33C–E)
Martinia semiplana var, lata Grabau, 1936, p. 239, pl. 21, figs.
1–3; Hayasaka and Minato, 1956, p. 146, pl. 23, fig. 3.
Martinia lata Grabau. Tazawa, 2008a, p. 38, fig. 5.7–5.14.
Material.—Three specimens from locality SSK17, external
and internal moulds of three ventral valves, NU-B2294–2296.
Remarks.—These specimens are referred to Martinia lata
Grabau, 1936, originally described by Grabau (1936, p. 239, pl.
21, figs. 1–3) as Martinia semiplana Waagen var. lata Grabau,
1936, from the Maping Formation of Guangxi, central-southern
China, by the medium-sized, transversely subelliptical shell(length 17 mm, width 24 mm in the best preserved specimen,
NU-B2294) and in having a shallow ventral sulcus. Martinia
semiplana Waagen (1883, p. 536, pl. 43, fig. 4), from theWargal Formation of the Salt Range, differs from M. lata in its
smaller size and less transverse outline.
Occurrence.—KN1 Unit.
Distribution.—Asselian–Wuchiapingian: northeastern Japan(Nagaiwa–Sakamotozawa, Kamiyasse–Imo and Takakurayama
in the South Kitakami Belt) and central-southern China(Guangxi).
Genus JILINMARTINIA Lee and Gu, 1980
Type species.—Brachythyris shansiensis Chao, 1929.
Jilinmartinia sp.
(Fig. 33H)
Material.—One specimen from locality SSK17, external andinternal moulds of a conjoined shell, NU-B2293.
Remarks.—The single specimen from Nagaiwa–Sakamotozawa
in the South Kitakai Belt is safely assigned to the genus
Jilinmartinia Lee and Gu, 1980 by the large size (length about
33 mm, width about 85 mm), wider subcircular outline,
moderately developed sulcus, external ornament consisting offine concentric growth lines, and some radial vascular markings

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

43

FIGURE 31. Brachiopods of the SK4 assemblage (2). A–D, Crenispirifer sagus Cooper and Grant; A, external latex cast (A1), external mould (A2, A3) andinternal mould (A4) of ventral valve, NU-B2313; B, external latex cast (B1), external mould (B2, B3) and internal mould (B4) of dorsal valve, NU-B2318;
C, external latex cast of ventral valve, NUB2315; D, external latex cast (D1) and internal mould (D2) of dorsal valve, NU-B2319; E, F, Trasgu confinensis(Schellwien); E, external latex cast (E1), internal mould (E2) and internal latex cast (E3) of ventral valve, NU-B2308; F, external latex cast (F1), internalmould (F2) and internal latex cast (F3, F4) of ventral valve, NU-B2307. Scale bars are 1 cm.

in the ventral valve. The Kitakami species resembles the typespecies, Jilinmartinia shansiensis (Chao, 1929), originallydescribed by Chao (1929, p. 55, pl. 9, figs. 1–3) as Brachythyris
shansiensis Chao, 1929, from the Lichiachuan Formation(Asselian) of Gansu, northwestern China, in size and shape ofthe shell and external ornament of the ventral valve. But the
Chinese species differs from the present species in the lesstransverse outline and in having narrower and deeper sulcus on
the ventral valve. Jilinmartinia sokolovi (Tschernyschew, 1902,
p. 166, pl. 8, fig. 3; pl. 39, fig. 4), from the Asselian of theUrals, is also a transverse Jilinmartinia species, but the Russian
species differs from the Kitakami species in having two costae
in the ventral sulcus.
Occurrence.—KN1 Unit.
Distribution.—Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Family INGELARELLIDAE Campbell, 1959a
Subfamily INGELARELLINAE Campbell, 1959a
Genus MARTINIOPSIS Waagen, 1883

44

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANIType species.—Martiniopsis inflata Waagen, 1883.
Martiniopsis sp.
(Fig. 33F)
Material.—One specimen from locality SSK17, external andinternal moulds of a ventral valve, NU-B2297.
Remarks.—The sole specimen from Nagaiwa–Sakamotozawa
can be assigned to the genus Martiniopsis Waagen, 1883 bylong, slender subparallel dental adminicula in the ventral valve
and in having no impressions of costae on the valve. The
Kitakami species closely resembles Martiniopsis inflata Waagen(1883, p. 525, pl. 41, figs. 7, 8, text-fig. 9), from the Chhidru
Formation of the Salt Range, in the medium size (length morethan 17 mm, width about 35 mm) and transverse outline.
Martiniopsis cathaysiensis Grabau (1936, p. 242, pl. 21, figs. 7,
8; pl. 24, fig. 9), from the Maping Limestone of Guangxi,
central-southern China and Guizhou, southwestern China, isalso transversely wider in outline but much smaller in size. An
accurate comparison is difficult for this poorly preserved
specimen.
Occurrence.—KN1 Unit.
Distribution.— Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Superfamily SPIRIFEROIDEA King, 1846
Family CHORISTITIDAE Waterhouse, 1968a
Subfamily CHORISTINAE Waterhouse, 1968a
Genus CHORISTITES Fischer de Waldheim, 1825
Type species.—Choristites mosquensis Buckman, 1908.
Choristites sp.
(Fig. 28A‒E)
Material.—Six specimens from localities SSK14 and SSK27:(1) external and internal moulds of two ventral valves,
NU-B2386, 2387; (2) external moulds of two ventral valves,
NU-B2388, 2389; (3) external and internal moulds of a dorsal
valve, NU-B2390; and (4) internal mould of a dorsal valve,
NU-B2391.
Remarks.—These specimens can be assigned to the genus
Choristites Fischer de Waldheim, 1825 by the large, transverseshell (length about 53 mm, width about 70 mm in the largest
ventral valve specimen, NU-B2388; length about 48 mm, widthabout 85 mm in the largest dorsal valve specimen, NU-B2391),
with a narrow, deep sulcus and a narrow, high fold, andornamented with numerous flattened simple costae (7–9 in 10mm at midlength) over both valves. The Nagaiwa–Sakamotozawa
species resembles well Choristites pavlovi (Stuckenberg, 1905),
redescribed by Chao (1929, p. 36, pl. 4, figs. 9, 10; pl. 5, figs.
1–4; pl. 6, figs. 5, 6) from the Taiyuan Formation of Shanxi,
northern China, in size, shape and external ornament of shell.
But accurate comparison is difficult for the ill-preserved

specimens.
Occurrence.—SK1 Unit.
Distribution.—Sakmarian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt).
Suborder DELTHYRIDINA Ivanova, 1972
Superfamily RETICULARIOIDEA Waagen, 1883
Family ELYTHIDAE Fredericks, 1924
Subfamily PHRICODOTHYRIDINAE Caster, 1939
Genus PHRICODOTHYRIS George, 1932
Type species.—Phricodothyris lucerna George, 1932.
Phricodothyris asiatica (Chao, 1929)(Fig. 27F, G)
Reticularia lineata Martin. Tschernyschew, 1902, p. 193, 574, pl
20, figs. 9–13.
Squamularia asiatica Chao, 1929, p. 91, pl. 11, figs. 12–14;
Ozaki, 1931, p. 76, pl. 8, figs. 15–19, pl. 9, figs. 2–4; Grabau,
1934, p. 71, pl. 5, figs. 3, 4.
Neophricodothyris asiatica (Chao). Licharew, 1939, p. 109, pl.
27, fig. 6.
Phricodothyris asiatica (Chao). Mironova, 1967, p. 51, pl. 5,
figs. 9, 10; Yanagida, 1967, p. 75, pl. 14, figs. 1, 2, 5, 7,
text-fig. 11; Pavlova, 1969, p. 95, pl. 8, fig. 4; pl. 9, figs. 1, 2,
text-figs. 60, 61; Ifanova, 1972, p. 141, pl. 13, figs. 5–10; Jinet al., 1974, p. 311, pl. 163, figs. 4–6; Lee and Gu, 1976, p.
298, pl. 136, figs. 1, 2; pl. 147, figs. 8–13; pl. 149, fig. 8; pl.
177, fig. 11; Yang et al., 1977, p. 450, pl. 179, fig. 5; Tong,
1978, p. 261, pl. 90, fig. 9; Zhan and Wu, 1987, p. 229, pl. 62,
figs. 20–25; Kotlyar and Zakharov, 1989, pl. 18, fig. 6; pl. 19,
fig. 12; Liang, 1990, p. 285, pl. 62, figs. 1–10; pl. 63, figs.
6–11; pl. 65, fig. 18, text-fig. 38; He et al., 1995, pl. 66, figs.
45–49; Carter and Poletaev, 1998, p. 172, fig. 27.19–27.32;
Wang and Yang, 1998, p. 127, pl. 23, figs. 5,6, 9.
Material.— Ten specimens from locality SSK27: (1) external
and internal moulds of three conjoined shells, NU-B2422–2424;(2) external and internal moulds of four ventral valves,
NU-B2425–2428; and (3) external moulds of three ventral
valves, NU-B2429–2431.
Description.—Shell medium in size for genus, slightly
transverse suboval in outline; cardinal extremities rounded;hinge approximately half or less than maximum width atmidlength; length 10 mm, width 13 mm in the best preserved
specimen (NU-B2424). Ventral valve gently to moderately
convex in lateral profile; sulcus absent or very weakly
expressed. Dorsal valve slightly less convex than opposite
valve; fold absent or very low. External surface of both valvesornamented with narrow distinct concentric lamellae, fringed byclosely spaced, fine uniramous spines or spine bases; numbering
7–8 lamellae in 5 mm. Internal structures of both valves not wellpreserved.
Remarks.—These specimens are referred to Phricodothyris

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

45

FIGURE 32. Brachiopods of the KN1 assemblage (1). A–C, Transennatia insculpta (Grant); A, external latex cast (A1, A2) of ventral valve, NU-B2282;
B, external latex cast (B1, B2), external mould (B3), internal latex cast (B4) and internal mould (B5) of dorsal valve, NU-B2283; C, external mould (C1, C2)of dorsal valve, NU-B2284; D–F, Echinauris sp.; D, external mould (D1, D2) and internal mould (D3) of ventral valve, NU-B2287; E, external mould (E1,
E2) of dorsal valve, NU-B2292; F, external mould (F1, F2) and internal mould (F3) of dorsal valve, NU-B2289; G, H, Anidanthus sp.; G, external mould(G1, G2) of dorsal valve, NU-B2302; H, external mould (H1, H2) of dorsal valve, NU-B2301; I, Dicystoconcha lapparenti Ternier and Termier, internalmould (I1, I2) of ventral valve, NU-B2280; J, K, Pseudoleptodus sp.; J, internal mould (J1, J2) of ventral valve, NU-B2300; K, internal mould (K1, K2) andinternal latex cast (K3) of ventral valve, N(U-B2299; L, Stenoscisma sp., external latex cast (L1, L2), internal mould (L3) and internal latex cast (L4) ofventral valve, NU-B2306. Scale bars are 1 cm.

46

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

asiatica (Chao), Originally described by Chao (1929, p. 91, pl.
11, figs. 12–14) as Squamularia asiatica Chao, 1929 from thelower Permian of Guizhou, southwestern China, in the small,transversely suboval shell, ornamented with distinct lamellae,
which are fringed with numerous fine uniramous spines.
Phricodothyris echinata Chao (1929, p. 86, pl. 8, figs. 17–19),
from the Penchi and Taiyuan series of Gansu and Shanxi,
northern China, differs from P. asiatica in the less transverse
outline and in having broader and less strong concentriclamellae on the both valves. The type species, Phricodothyris
lucerna George (1932, p. 546, pl. 35, fig. 2), from the upper
Visean of Northumberland, England, differs from the present
species in the larger size and less transverse outline.
Occurrence.—SK1 Unit.
Distribution.—Moscovian–Wuchiapingian: northeasternJapan (Nagaiwa–Sakamotozawa in the South Kitakami Belt),
northern Canada (Ellesmere Island), northern Russia (Pechora
Basin), western Russia (Transcaucasus), central Russia(southern Urals), northwestern China (Xinjiang), northern China(Inner Mongolia and Shanxi), eastern China (Shandong andZhejiang), central-southern China (Hunan and Guangxi),
southwestern China (Guizhou and Sichuan) and north-central
Thailand (Thum Nam Maholan).

width 13 mm in the largest specimen, NU-B1517), coarse
rounded costae (numbering 3–4 on each lateral flank) andclosely spaced, imbricate lamellae and numerous pustules overthe both valves. The present specimens were previously
described by Tazawa and Shintani (2014, p. 35, fig. 6.10) as
Callispirina sp. from the basal part of the Nakadaira Formation(Sakmarian) of Kamiyasse in the South Kitakami Belt.
Callispirina rotunda Cooper and Grant (1976, p. 2743, pl. 705,
figs. 66–82), from the Bell Canyon Formation of Texas, differsfrom C. ornata in larger size and in having more numerous
costae on lateral flanks.
Occurrence.—SK1 Unit.
Distribution.—Kasimovian–Changhsingian: northeasternJapan (Nagaiwa–Sakamotozawa and Kamiyasse in the South
Kitakami Belt), northern Russia (Timan, Vaygach Island andnorthern Urals), western Russia (Moscow Basin), central Russia(southern Urals), northeastern China (Heilongjiang and Jilin)
and Pakistan (Salt Range).

Order SPIRIFERINIDA Ivanova, 1972
Suborder SPIRIFERINIDINA Ivanova, 1972
Superfamily PENNOSPIRIFERINOIDEA Dagys, 1972
Family PARASPIRIFERINIDAE Cooper and Grant, 1976
Genus CALLISPIRINA Cooper and Muir-Wood, 1951

Spiriferellina cristata (von Schlotheim, 1816)(Fig. 29C–F)

Type species.—Spiriferina ornata Waagen, 1883.
Callispirina ornata (Waagen, 1883)(Fig. 29A, B)
Spiriferina ornata Waagen, 1883, p. 505, pl. 50, figs. 1, 2;
Tschernyschew, 1902, p. 113, 515, pl. 12, figs. 8–10: pl. 37,
figs. 8–11; Sokolskaya in Sarytcheva and Sokolskaya, 1952,
p. 226, pl. 66, fig. 383.
Callispirina ornata (Waagen). Mironova, 1967, p. 49, pl. 5, figs.
4, 5; Kulikov, 1974, p. 102, pl. 7, fig. 6; Kalashnikov, 1980,
p. 94, pl. 36, fig. 9; Kalashnikov, 1998, p. 75, pl. 26, fig. 5; pl.
31, figs. 1–5.
Punctospirifer ornata (Waagen). Lee et al., 1980, p. 420, pl. 155,
fig. 22; pl. 179, figs. 1, 19, 20.
Callispirina sp. Tazawa and Shintani, 2014, p. 35, fig. 6.10.
Material.—Two specimens from localities SSK27 andSSK28: (1) external and internal moulds of a ventral valve,
NU-B2392; and (2) external and internal moulds of a dorsal
valve, NU-B2393.
Remarks.—These specimens are referred to Callispirina
ornata (Waagen, 1883), originally described by Waagen (1883,
p. 505, pl. 50, figs. 1, 2) from the Chhidru Formation of the Salt
Range, Pakistan, by the small, transverse shell (length 11 mm,

Family SPIRIFERELLINIDAE Ivanova, 1972
Genus SPIRIFERELLINA Fredericks, 1924
Type species.—Terebratulites cristatus von Schlotheim, 1816.

Terebratulites cristatus von Schlotheim, 1816, p. 28, pl. 1, fig. 3.
Spirifer cristatus (Schlotheim). de Koninck, 1843, p. 240, pl. 15,
fig. 5.
Spiriferina cristata (Schlotheim). Tschernyschew, 1902, p. 115,
517, pl. 37, figs. 1, 2; Ozaki, 1931, p. 172, pl. 15, fig. 14;
Malzahn, 1937, p. 40, pl. 3, figs. 26, 27.
Spiriferellina cristata (Schlotheim). Heritsche, 1935, p. 364, pl.
2, fig. 22; Heritsche, 1938, p. 133, pl. 7, figs. 15–19; Campbell,
1959b, p. 358, pl. 59, figs. 1–9; pl. 60, fig. 3, text-fig. 5;
Schréter, 1963, p. 144, pl. 8, figs. 11–14; Alexandrov andEinor, 1979, p. 91, pl. 38, figs. 2, 3; Lee et al., 1980, p. 422,
pl. 179, figs. 3, 6–8; Kalashnikov, 1998, p. 75, pl. 30, fig. 3;
pl. 32, figs. 1, 2; Wang and Yang, 1998, p. 125, pl. 22, figs. 9,13, 14; Fan and He, 1999, p. 146, pl. 33, figs. 11–20; Wang
and Zhang, 2003, p. 168, pl. 34, figs. 12–16; pl. 50, figs. 14,
16; Tazawa, 2012, p. 42, figs. 3.17–3.19.
Punctospirifer cristata (Schlotheim). Dunbar, 1955, p. 149, pl.
29, figs. 13–20.
Material.—Twenty-eight specimens from localities SSK14,
SSK27 and SSK28: (1) external and internal moulds of aconjoined shell, NU-B2394; (2) internal mould of a conjoined
shell, with external mould of the ventral valve, NU-B2395; (3)internal mould of a conjoined shell, with external mould of thedorsal valve, NU-B2396; (4) internal moulds of three conjoined
shells, NU-B2397–2399; (5) external and internal moulds ofeleven ventral valves, NU-B2400–2410; (6) internal moulds of
two ventral valves, NU-B2411, 2412; (7) external and internal

CISURALIAN BRACHIOPODS FROM NAGAIWA–SAKAMOTOZAWA

47

FIGURE 33. Brachiopods of the KN1 assemblage (2). A, B, Hustedia ratburiensis Waterhouse and Piyasin; A, ventral internal mould (A1, A2), dorsalinternal mould (A3) and dorsal external latex cast (A4) of conjoined shell, NU-B2303; B, external latex cast (B1, B2) of dorsal valve, NU-B2304; C–E,
Martinia lata Grabau; C, internal mould of ventral valve, NU-B2295; D, internal mould (D 1, D2) and external latex cast (D3) of ventral valve, NU-B2296;
E, internal mould (E1, E2) and external latex cast (E3) of ventral valve, NUB2294; F, Martiniopsis sp., internal mould (F1, F2) of ventral valve, NU-B2297;
G, Crenispirifer sagus Cooper and Grant, internal mould (G1, G2) and external latex cast (G3) of dorsal valve, NU-B2298; H, Jilinmartinia sp., dorsal
external latex cast (H1), ventral internal mould (H2) and dorsal internal mould (H3) of conjoined shell, NUB2293. Scale bars are 1 cm.

48

JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI

moulds of seven dorsal valves, NU-B2413–2419; (8) externalmould of a dorsal valve, NU-B2420; and (9) internal mould of adorsal valve, NU-B2421.
Remarks.—These specimens are referred to Spiriferellina
cristata (von Schlotheim, 1816), redescribed and refigured byCampbell (1959b, p. 358, pl. 59, figs. 1–9; pl. 60, fig. 3, text-fig.
5) on the syntype and lectotype specimens, from the Zechsteinof Thuringia, Germany, in the small, transverse shell (length 16mm, width 24 mm in the largest specimen, NU-B2400), and 4–5
pairs of rounded costae on both ventral and dorsal valves.
Spiriferellina fredericksi Tazawa (2014, p. 19, fig. 3.5–3.7),
from the lower part of the Kamiyasse Formation (Wordian) of
Kamiyasse–Imo, South Kitakami Belt, is readily distinguishedfrom S. cristata in its much larger size.
Occurence.—SK1 Unit.
Distribution.—Kasimovian–Changhsingian: northeasternJapan (Nagaiwa–Sakamotozawa and Kesennuma in the South
Kitakami Belt), northern Russia (Timan and northern Urals),
Greenland, Germany, Austria (Carnian Alps), Hungary, central
Russia (southern Urals), northwestern China (Xinjiang),
northern China (Inner Mongolia and Shanxi) and northeastern
China (Heilongjiang).
Genus CRENISPIRIFER Stehli, 1954
Type species.—Spiriferina angulata King, 1931.
Crenispirifer sagus Cooper and Grant, 1976
(Figs. 31A–D, 33G)
Crenispirifer sagus Cooper and Grant, 1976, p. 2715, pl. 718,
figs. 1–15.
Material.—Nine specimens from localities SSK17 andSSK24: (1) external and internal moulds of a ventral valve,
NU-B2313; (2) external moulds of three ventral valves,
NU-B2314–2316; (3) internal mould of a ventral valve,
NU-B2317, (4) external and internal moulds of three dorsal
valves, NU-B2298, 2318, 2319; and (5) internal mould of adorsal valve, NU-B2320.
Description.—Shell small in size for genus, transversely
subelliptical in outline, with greatest width at slightly anterior tohinge; cardinal extremities rounded; length 8 mm, width about
11 mm in the largest ventral valve specimen (NU-B2313);
length 6 mm, width 12 mm in the largest dorsal valve specimen(NU-B2318). Ventral valve moderately convex in lateral profile,most convex at umbonal region; umbo small, incurved; sulcus
narrow, deep; lateral slopes gently convex. Dorsal valve flatly
convex; umbo slightly swollen; fold narrow, moderately high,with rounded crest. External surface of both valves ornamented
with strong costae, numbering 4–5 on each side; numerous veryfine pustules over the valves; concentric rugae or growthlaminae not observed. Internal structures of both valves not wellpreserved, except for a short, thin median septum in the ventral valve.
Remarks.—These specimens are referred to Crenispirifer

sagus Cooper and Grant (1976, p. 2715, pl. 718, figs. 1–15),
from the Bone Spring Formation (lower Leonardian) of Texas,by the small-sized, flatly biconvex shell, ornamented with rather
numerous costae on the both valves. Crenispirifer nakamurai
Tazawa and Shintani (2014, p. 36, figs. 6.11, 6.12), from the
Nakadaira Formation (Sakmarian) of Nakadaira, South Kitakami
Belt, is readily distinguished from the present species by its
fewer and broader costae on both ventral and dorsal valves. Twosmall-sized Crenispirifer species, C. effrenus Cooper and Grant(1976, p. 2712, pl. 718, figs. 16–29) from the Cherry Canyon
Formation of Texas, and C. myllus Cooper and Grant (1976, p.
2714, pl. 718, figs. 30–85; pl. 719, figs. 36–40) from the Bell
Canyon Formation of Texas, differ from C. sagus in the morestrongly biconvex shell and in having growth laminae on theboth valves.
Occurrence.—SK4 and KN1 units.
Distribution.—Artinskian–Kungurian: northeastern Japan(Nagaiwa–Sakamotozawa in the South Kitakami Belt) and USA(Texas).
ACKNOWLEDGEMENTS
We sincerely thank Kenshiro Ogasawara (Professor Emeritusof the University of Tsukuba, Tsukuba) and Soichiro Kawabe(Fukui Prefectural Dinosaur Museum, Katsuyama) for editing
the manuscript; Shuzhong Shen (Nanjing University, Nanjing)
and an anonymous reviewer for their constructive comments;
and Yousuke Ibaraki (Fossa Magna Museum, Itoigawa) for his
help in drawing the figures.
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JUN-ICHI TAZAWA AND TOMOHIKO SHINTANI＜
Endosawa
エンドウ沢Hikami Granite
氷上花崗岩
Hikoroichi-cho
日頃市町
Hosoo Formation
細尾層
Ichinoseki
一関Imahorazawa Valley
イマホラ沢
Imo
飯森
Ishibashi
石橋
Kabayama Stage
樺山階Kamaishi釜石
Kamisakamotozawa
上坂本沢
Kamiishibashi
上石橋
Kamijo上条
Kamiyasse–Imo
上八瀬‒飯森Kanokura Formation
叶倉層
Kawaguti (Kawaguchi) Stage
川口階
Kesengawa River
気仙川

地名・地層名＞Kesennuma
気仙沼
Maiya
米谷
Mizukoshi
水越Motoiwazawa Sandstone Member 元岩沢砂岩部層
Mt. Goyosan
五葉山
Mt. Takamoriyama
高森山
Nabekoshiyama Formation
鍋越山層
Nagaiwa Formation
長岩層
Nagaiwa–Sakamotozawa 長岩‒坂本沢
Nakadaira
中平Nakazato Formation
中里層
Nedamo Belt
根田茂帯
西郡層
Nishigori (Nishikori) Formation
大船渡市
Ofunato City
雄勝
Ogatsu鬼丸層
Onimaru Formation
大野層
Ono Formation

六郎峠
Rokuro-toge
坂本沢層
Sakamotozawa Formation
盛川
Sakarigawa River世田米
Setamai
白石峠
Shiraishi-toge
Shiratorizawa Limestone Member
白鳥沢石灰岩部層
Sumita-cho
住田町
Takakurayama
高倉山
Takougawa River鷹生川
Tashiroyama Limestone Member
田代山石灰岩部層
Tashiroyashiki
田代屋敷
Tassobe達曽部
Tono
遠野
Toyoma Series (Formation)
登米統（層）
Usuginu-type conglomerate 薄衣型（式）礫岩
Yubanosawa Slate Member 湯場沢粘板岩部層
Yukizawa (Yukisawa) Series
雪沢統