ARTICLE

Memoir of the Fukui Prefectural Dinosaur Museum 19: 11–88 (2020)© by the Fukui Prefectural Dinosaur Museum

EARLY CARBONIFEROUS (MISSISSIPPIAN) BRACHIOPODSFROM THE SHITTAKAZAWA, ARISU AND ODAIRAFORMATIONS, SOUTH KITAKAMI BELT, JAPANJun-ichi TAZAWA
Hamaura-cho 1-260-1, Chuo-ku, Niigata 951-8151, Japan

ABSTRACTThis study describes 56 species of early Carboniferous (Mississippian) brachiopods in 39 genera (including a new
species, Levitusia elongata sp. nov.) from the Shittakazawa, Arisu and Odaira Formations in the central part
(Shimoarisu, Yokota, Yahagi, Nisawa, Okuhinotsuchi and Shizu areas) of the South Kitakami Belt (southern
Kitakami Mountains), northeastern Japan. On the basis of brachiopod biostratigraphy, the Shittakazawa Formation iscorrelated with the upper Tournaisian, the Arisu Formation with the lower Visean, and the Odaira Formation with theupper Visean. In terms of palaeobiogeography, the late Tournaisian (Shittakazawa) fauna is allied with those of the
USA (Oklahoma), the UK (England), Poland, Belgium, central Russia (Kuznetsk Basin), Kazakhstan andnorthwestern China (Xinjiang); the early Visean (Arisu) fauna has a close affinity with those of the UK (England),Belgium, central Russia (southern Urals and Kuznetsk Basin), Kazakhstan and northwestern China (Xinjiang),
particularly with Kazakhstan and northwestern China (Xinjiang); and the late Visean (Odaira) fauna exhibits
affinities with assemblages of the UK (England), Belgium, central Russia (southern Urals), Kazakhstan andnorthwestern China (Xinjiang), particularly with Kazakhstan. Thus, the early Carboniferous (late Tournaisian–lateVisean) brachiopod faunas of the central part of the South Kitakami Belt are allied with those of Kazakhstan andnorthwestern China (Xinjiang). The South Kitakami region was probably the eastern extension of the North China
Provine, and was located at the eastern end of the Central Asian Orogenic Belt (CAOB), near and to the east of theNorth China Block as a shallow sea bordering a microcontinent during the early Carboniferous.
Key words: Arisu Formation, Brachiopoda, early Carboniferous, Odaira Formation, Shittakazawa Formation, South
Kitakami Belt
田沢純一 (2020) 南部北上帯の尻高沢層，有住層，大平層から産出する前期石炭紀 (ミシシッピー亜紀 ) 腕足類．福井県立恐竜博物館紀要 19：11‒88.
南部北上帯中央部（下有住・横田・矢作・荷沢・奥火の土・清水地域）の下部石炭系尻高沢層，有住層，大平層から産出する 39 属 56 種（1 新種 Levitusia elongata sp. nov. を含む）の前期石炭紀（ミシシッピー亜紀）腕足類を記載した . これらの腕足類により，尻高沢層は Tournaisian 上部に，有住層は Visean 下部に，そして大平層は Visean 上部に対比される . 古生物地理学的には，尻高沢フォー ナはアメリカ（オハイオ），
イギリス，ポーランド，ベルギー，ロシア中央部（クズネツク盆地），カザフスタン，中国西北部（新疆）の，有住フォーナはイギリス，ベルギー，ロシア中央部（ウラル山脈南部，クズネツク盆地），カザフスタンおよび中国西北部（新疆），とくにカザフスタンと中国西北部（新疆）の，また，大平フォーナはイギリス，
ベルギー，ロシア中央部（ウラル山脈南部），カザフスタン，中国西北部（新疆），とくにカザフスタンの前期石炭紀腕足類フォーナと類縁がある．このことから，前期石炭紀（Tournaisian 後期 ~Visean 後期）に，
南部北上地域は前期石炭紀腕足動物区の一つである北中国区に属し，おそらく中央アジア造山帯の東縁，
北中国地塊の東方に存在したマイクロコンチネントの縁辺浅海域であったと推定される．
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や はぎ

にさわ

おく ひ

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しっ たか ざわ

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INTRODUCTION

understanding the geotectonic history of the South Kitakami
Belt and the Japanese Islands in general. Brachiopoda is one of
the most useful taxa for the Palaeozoic biogeography owing to
its very abundant fossil records and being benthos. The
Brachiopoda is useful also for biostratigraphy, because there areso numerous and diverse that it has been recorded more than two
centuries. The method “assemblage zone” is useful for the
Palaeozoic brachiopod biostratigraphy, although most of thebrachiopod species have rather long ranges.

Palaeozoic geography and biogeography of the South
Kitakami Belt, northeastern Japan are important for
Received February 20, 2020. Accepted October 21, 2020
E-mail: j1025-tazawa＊memoad.jp

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FIGURE 1. Map showing the location and geology of the Shimoarisu, Yokota, Yahagi, Nisawa, Okuhinotsuchi and Shizu areas in the South Kitakami
Belt. (A) Geotectonic map of the Japanese Islands, showing the distribution of the South Kitakami Belt (after Tazawa, 2018c); (B) geotectonic map of thenortheastern part of Honshu, Japan, showing the distribution of the Carboiferous rocks in the South Kitakami Belt (modified from Tazawa, 2018c).

Brachiopods are extremely common in the early
Carboniferous (Mississippian) marine fauna of the South
Kitakami Belt (southern Kitakami Mountains). Since thepioneering studies of Minato (1951, 1952), about 120
brachiopod species (in 65 genera) were described by Tachibana(1956, 1963, 1964, 1969, 1981), Minato and Kato (1977),
Tazawa and Katayama (1979), Tazawa (1980, 1981b, 1984a,
1984b, 1985, 1989, 2006, 2017, 2018a), Tazawa and Kurita(1986, 2019a, 2019b), Tazawa and Miyake (2002), Chen andTazawa (2007), Tazawa and Ibaraki (2009, 2019), Tazawa and
Iryu (2019) and Tazawa et al. (2019). However, most of thespecies described in 1950–1980s need redescription on therevised stratigraphy of the lower Carboniferous formations of
the South Kitakami Belt and on the new taxonomy ofBrachiopoda.
The South Kitakami Belt contains well-exposed sections oflower Carboniferous (Mississippian) rocks. Brachiopods arecommon in the early Carboniferous marine fauna of the belt.
The central part of the belt, including the Shimoarisu(Sumita-cho, Kesen-gun, Iwate Prefecture), Yokota(Yokota-cho, Rikuzentakata City, Iwate Prefecture), Yahagi(Yahagi-cho, Rikuzentakata City, Iwate Prefecture), Nisawa(Otomo-cho, Tono City, Iwate Prefecture), Okuhinotsuchi(Sumita-cho, Kesen-gun, Iwate Prefecture) and Shizu(Sumita-cho, Kesen-gun, Iwate Prefecture) areas (Figs. 1 and 2),
is an important and classical area for the lower Carboniferous
stratigraphy and palaeontology of Japan as well as the South
Kitakami Belt. Since the work of Endo (1924), the stratigraphy
and palaeontology of the lower Carboniferous rocks and fossils
in the central part of the belt have been extensively investigated(e.g., Minato, 1941, 1951, 1952; Minato et al., 1953, 1979b;
Onuki, 1969; Minato and Kato, 1979; Tazawa and Katayama,
1979; Tazawa et al., 1981; Kawamura, 1985a, 1985b, 1985c;
Kawamura et al., 1985, Tazawa, 2017; Tazawa and Kurita,
2019a; Tazawa and Iryu, 2019).
This paper is the third contribution of the series on the early
Carboniferous brachiopods of the South Kitakami Belt. In thefirst of the series (Tazawa, 2018c), I described early
Carboniferous brachiopods from the Hikoroichi Formation of
the Hikoroichi area in the eastern part of the belt; the second
contribution (Tazawa et al., 2019) addressed the brachiopods
from the Karaumedate Formation in the Nagasaka area, westernpart of the belt. In the present study, I describe the early
Carboniferous brachiopods from the Shittakazawa, Arisu andOdaira Formations in the central part of the South Kitakami
Belt. I also discuss the stratigraphy and correlation of the lower

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FIGURE 2. A distant view (A) and a close view (B) of locality KF159, outcrop of calcareous shale of the OD3 Unit, upper Odaira Formation in Tairagai,
Yokota-cho, Rikuzentakata City, Iwate Prefecture.

Carboniferous formations, and the specific composition, age andpalaeobiogeography of the brachiopod faunas.

PREVIOUS WORK
Stratigraphy
Previous studies on the stratigraphy of the lower

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FIGURE 3. Comparison of stratigraphic schemes proposed for the lower Carboniferous formations in the Shimoarisu, Yokota and Yahagi areas, South
Kitakami Belt.

Carboniferous formations in the central part of the South
Kitakami Belt are summarized in Figs. 3 and 4. The modern
stratigraphic studies started in 1941 when M. Minato publisheda paper, entitled “On the Lower Carboniferous deposits at
Setamai, Kesen-gori, Iwate Prefecture”. Since then, manystudies have been investigated by Minato (1951, 1952), Minato
et al. (1953, 1979b), Hirokawa and Yoshida (1954), Onuki(1956, 1969, 1981), Takeda (1960), Saito (1966, 1968), Moriai(1972), Minato and Kato (1979), Tazawa and Katayama (1979),
Tazawa (1979, 1981b, 1985, 2006, 2017), Tazawa et al. (1981),
Kawamura (1985a, 1985b, 1985c), Kawamura et al. (1985),
Tazawa and Kurita (1986, 2019a, 2019b), Kawamura andKawamura (1989), Tazawa and Ibaraki (2019) and Tazawa and
Iryu (2019).
Minato (1950, 1955, 1966) emphasized the pre-Onimaru
hiatus, a long hiatus between the upper Visean Onimaru
Formation and the underlying formations, as evidence of the“Shizu Folding”, one of the Palaeozoic orogenic events recordedin the South Kitakami Belt. But the Minato’s proposal was
denied litho- and bio-stratigraphically by Tazawa and Katayama(1979), Tazawa (1979, 1981b), Tazawa et al. (1981), Kawamura(1985a, 1985b, 1985c) and Kawamura et al. (1985) in the central
part (Shimoarisu, Yokota, Yahagi, Nisawa, Okuhinotsuchi and
Shizu areas) of the belt as well as the eastern part (Hikoroichi
area; Mori and Tazawa, 1980; Tazawa, 1981a, 1984a, 1984b,
2018c; Kawamura, 1983), the western part (Nagasaka area;
Tazawa and Osawa, 1979; Tazawa, 1980; Tazawa et al., 2019)
and the southern part (Soma area; Tazawa et al., 1984; Tazawa,
1996) of the belt.
The lithostratigraphy of the lower Carboniferous formationsin the central part of the South Kitakami Belt was almost
concluded by Kawamura (1985a, 1985b, 1985c) and Kawamura
et al. (1985). However, the biostratigraphy of the formations
was poor and remained in long time at the concept of Minato etal. (1953, 1979b) and Minato and Kato (1979), because of apaucity of fossil evidence. During the last few years, Tazawa
and his collaborators clarified the age of the Shittakazawa
Formation (Tazawa and Kurita, 2019a), the Arisu Formation(Tazawa and Kurita, 2019b; Tazawa and Ibaraki, 2019; Tazawa
and Iryu, 2019) and the Odaira Formation (Tazawa, 2017) basedon the brachiopods from the formations. Kawamura et al. (1985)
also found that the uppermost part of the Odaira Formation wascorrelated with the upper Visean based on the corals, although
the coral specimens have been described yet.
Palaeontology
Studies on the invertebrate fossils from the Shittakazawa,
Arisu and Odaira Formations are poor, except for thebrachiopods. Until now only a few species of the corals,
trilobites, crinoids and blastoids have been described: Amplexus
nipponensis Minato and Amplexus sp. from the Arisu and Odaira
Formations (Minato, 1952); Sugiyamaella carbonarium Yabe
and Minato from the Odaira Formation (Yabe and Minato, 1944;
Minato, 1951, 1952); Palaeosmilia membiensis Minato andOgata from the Odaira Formation (Minato and Ogata, 1977);
Syringopora sp. and Phyllipsia? sp. from the Odaira Formation(Minato, 1951); and Platycrinus asiatica Minato andNucleocrinidae gen. et sp. indet. from the Arisu Formation(Minato, 1951). Kawamura et al. (1985) reported, but not

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FIGURE 4. Comparison of stratigraphic schemes proposed for the lower Carboniferous formations in the Nisawa, Okuhinotsuchi and Shizu areas, South
Kitakami Belt.

described, eight rugose coral species (Pseudouralinia
tangpakouensis Yü, Siphonophyllia sp., Sugiyamaella sp.,Caninophyllum? sp., Kueichouphyllum heishihkuanense Yü,Yuanophyllum kansuense Yü, “Lophophyllum” sp. and“Caninia” sp.) from the Odaira Formation.
In contrast, taxonomic studies on the brachiopods areplentiful. Minato (1951, 1952) described the following
brachiopods (of 27 species in 17 genera) from the Arisu andOdaira Formations: Planoproductus gigantoides Minato,
Pustula cf. tenuipustulata Thomas, Productus sp., Orthotetes
keokuk (Hall), Orthotetes sp., Schellwienella izirii Minato,
Schellwienella? sp., Derbyia depressa var. transversa Minato,
Schizophoria resupinata (Martin), Actinoconchus lamellosa(Léveillé), A. cf. lamellosa (Léveillé), Cleiothyridina royssii(Léveillé), Spirifer kozuboensis Minato, Brachythyrina nagaoi
Minato, Fusella nipponotrigonalis Minato, F. nipponotrigonalis
var. minor Minato, Brachythyris aff. pinguis (Sowerby),
Brachythyris? sp., Delthyris aff. clarksvillensis (Winchell),
Kitakamithyris tyoanjiensis Minato, K. semicircularis Minato,
Syringothyris jumonjiensis Minato, S. transversa Minato, S.
kitakamiensis Minato, Syringothyris sp., Plicatosyrinx singulare

Minato and P.? kumanoi Minato. After that, brachiopods of 41
species in 27 genera were described from the Shittakazawa,
Arisu and Odaira Formations by Minato and Kato (1977),
Tazawa and Katayama (1979), Tazawa (1981b, 1985, 2006,
2017, 2018a), Tazawa and Kurita (1986, 2019a, 2019b), Tazawa
and Ibaraki (2019) and Tazawa and Iryu (2019).
The brachiopods are: Leptagonia analoga (Phillips),
Rugosochonetes extensus (Chao), Levitusia humerosa(Sowerby), Tomiproductus elegantulus (Tolmatchoff),
Marginicinctus marginicinctus (Prout), Marginicinctus sp.,
Marginatia burlingtonensis (Hall), Marginatia sp.,
Echinoconchus punctatus (Sowerby), Echinaria sp., Pustula
pustulosa (Phillips), Ovatia elongata Muir-Wood and Cooper,
Fluctuaria cf. undata (Defrance), Striatifera striata (Fischer),Orthotetes cf. australis (Campbell), Schellwienella radialis(Phillips), Rhipidomella michelini (Léveillé), R. kusbassica
Beznossova, Rhipidomella sp., Schizophoria resupinata(Martin), S. pinguis Demanet, S. woodi Bond, S. mayesensis
Carter, Cleiothyridina fimbriata (Phillips), C. harkeri Carter,
Martinia aff. glabra (Sowerby), Spirifer liangchowensis Chao,“S.” mundulus Rowley, Grandispirifer mylkensis Yang,

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Unispirifer striatoconvolutus (Dun and Benson), U. kozuboensis(Minato), Unispirifer sp., Imbrexia cf. incertus (Hall),Kitakamithyris hikoroitiensis Minato, Kitakamithyris sp.,
Syringothyris texta (Hall), Asyrinxia nipponotrigonalis(Minato), Asyrinxia sp., Pseudosyrinx jumonjiensis (Minato),
Pseudosyrinx sp. and Dimegelasma sp.
In terms of biostratigraphy, the lower Carboniferous
formations, except for the upper Visean Onimaru Formation, inthe central part of the South Kitakami Belt were considered tobe correlated with the lower Tournaisian to the lower Visean(Minato, 1941, 1951, 1952; Minato et al., 1953, 1979b; Minato
and Kato, 1979) based mainly on the brachiopods, Leptagonia
analoga, Spirifer kozuboensis (= Unispirifer kozuboensis),
Imbrexia cf. incertus (= Imbrexia incertus), Kitakamithyris
semicircularis (= K. hikoroitiensis), Syringothyris transversa (=S. texta), S. jumonjiensis (= Pseudosyrinx jumonjiensis) andFusella nipponotrigonalis (= Asyrinxia nipponotrigonalis) andcorals, Palaeosmilia membiensis, Amplexus nipponensis andSugiyamaella carbonarium. After that, the middle part of the
Arisu Formation was correlated with the lower Visean by thebrachiopods, Marginatia burlingtonensis, Syringothyristransversa (= S. texta) and Unispirifer sp. (= U. minnewankensis)(Tazawa, 1985, 2006); and the Odaira Formation was correlatedwith the upper Visean by the corals, Pseudouralinia
tangpakouensis, Siphonophyllia sp., Sugiyamaella sp.,Caninophyllum? sp., Kueichouphyllum heishikuanensis,
Yuanophyllum kansuense, “Lophophyllum” sp. and “Caninia”
sp. (Kawamura et al., 1985) and brachiopods, Leptagonia
analoga, Rugosochonetes extensus, Marginatia burlingtonensis,
Echinoconchus punctatus, Echinaria sp., Pustula pustulosa,
Schellwienella radialis, Cleiothyridina fimbriata, Spirifer
liangchowensis, Kitakamithyris sp. and Pseudosyrinx sp.
(Tazawa e t al. , 1 9 8 1 ; T aza w a, 1 9 8 1 b ). Recen tly , theShittakazawa Formation was correlated with the upper
Tournaisian by four brachiopod species, Rhipidomellakusbassica, Schizophoria pinguis, S. mayesensis and Unispirifer
kozuboensis (Tazawa and Kurita, 2019a); the Arisu Formation
with the lower Visean by brachiopods (17 species in 13 genera),
Levitusia humerosa, Tomiproductus elegantulus, Marginatia
burlingtonensis, Ovatia elongata, Rhipidomella michelini,
Schizophoria resupinata, S. pinguis, S. woodi, Cleiothyridina
harkeri, Grandispirifer mylkensis, Unispirifer striatoconvolutus,
Unispirifer sp., Kitakamithyris hikoroitiensis, Syringothyris
texta, S. platypleura, Asyrinxia nipponotrigonaris andPseudosyrinx jumonjiensis (Tazawa and Ibaraki, 2019; Tazawa
and Iryu, 2019; Tazawa and Kurita, 2019b); and the uppermost
part of the Odaira Formation with the upper Visean bybrachiopods (11 species in 11 genera), Leptagonia analoga,
Rugosochonetes extensus, Marginatia burlingtonensis,
Echinoconchs punctatus, Echinaria sp., Pustula pustulosa,
Schellwienella radialis, Cleiothyridina fimbriata, Spirifer
liangchowensis, Kitakamithyris sp. and Pseudosyrinx sp.
(Tazawa, 2017).
In palaeobiogeography, Minato (1956) explained in short thatthe early Carboiferous marine fauna of the South Kitakami Belt

has a close affinity with those of eastern Australia and theMongolian geosynclinal region (central Asia), and quite
different from those of southern China. Recently, Tazawa andhis collaboraters (Tazawa, 2017; Tazawa and Iryu, 2019;
Tazawa and Kurita, 2019b) proposed that early and late Viseanbrachiopod faunas of the central part of the South Kitakami Belt
resemble those of western Europe and central Asia (Kazakhstan,
Uzbekistan, Kyrgyzstan and northwestern China), and the South
Kitakami region probably belonged to the North China Provinceof Yang (1983) and was located near and to the east of the North
China Block during the Visean.
STRATIGRAPHY
The lower Carboniferous succession of the central part
(Shimoarisu, Yokota, Yahagi, Nisawa, Okuhinotsuchi amd
Shizu areas; Figs. 5‒9) of the South Kitakami Belt is divided
into four formations: the Shittakazawa, Arisu, Odaira andOnimaru Formations in stratigraphic ascending order.
Furthermore, each formation, except for the Onimaru
Formation, is subdivided into three units in stratigraphic
ascending order: the Shittakazawa Formation is subdivided into
the ST1, ST2 and ST3 units; the Arisu Formation into the AR1,AR2 and AR3 units; and the Odaira Formation into the OD1,OD2 and OD3 units (Figs. 10‒13). In this paper, the upper
Visean volcaniclastic sequence in the Nisawa, Okuhinotsuchi
and Shizu areas is assigned to the Odaira Formation (named by
Onuki, 1956; defined by Minato et al., 1979b), rather than the
Karoyama Formation (Tazawa et al., 1981) or the middle andupper parts of the Karosawa Formation (Kawamura, 1985c).
Shittakazawa Formation
The stratigraphy of the Shittakazawa Formation was studied
by Kawamura (1985a) and Tazawa and Iryu (2019) in the
Shimoarisu area and by Tazawa and Kurita (1986, 2019a) in the
Okuhinotsuchi area (Fig. 5). According to Tazawa and Kurita(2019a), the Shittakazawa Formation in the Okuhinotuchi area
has a total thickness of 986 m and is subdivided into a lower
part (ST1 Unit: alternating light grey to light greenish-greyrhyolitic tuff and conglomerate, with thin layers of limestone
and green to dark green andesitic tuff, 257 m thick), a middle
part (ST2 Unit: rhyolitic tuff, 500 m thick) and an upper part
(ST3 Unit: alternating rhyolitic tuff, sandstone and shale, with athin limestone layer, 229 m thick; Fig. 13). Brachiopod fossils
were collected from one locality (KAR1), an outcrop of lightgrey rhyolitic tuffaceous shale of the ST3 Unit.
Arisu Formation
Stratigraphy of the Arisu Formation was studied by
Kawamura (1985a) and Tazawa and Iryu (2019) in the
Shimoarisu area and by Tazawa and Katayama (1979), Tazawa(1979), Kawamura (1985b) and Tazawa and Ibaraki (2019) in
the Yokota area (Fig. 6). According to Tazawa and Ibaraki

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FIGURE 5. Map showing the fossil localities KAR1, KAR6 and KAR11 in the Okuhinotuchi area, and IST1 in the Shimoarisu area (using the electronic
topographical map of GSI).

(2019), the Arisu Formation in the Yokota area has a total
thickness of 330 m and is subdivided into a lower part (AR1
Unit: green to dark green andesitic lapilli tuff, 40 m thick), a
middle part (AR2 Unit: sandstone with subordinate andesitic
tuff and limestone, 260 m thick) and an upper part (AR3 Unit:
black shale with a thin limestone bed as the topmost bed, 130 m
thick; Fig. 11). Brachiopod fossils occur from the AR2 and AR3
units, although the AR1 Unit is barren of fossils.
Odaira Formation
Stratigraphy of the Odaira Formation was studied by
Kawamura (1985a) and Tazawa and Iryu (2019) in the
Shimoarisu area, by Tazawa and Katayama (1979), Tazawa(1979, 2017), Kawamura (1985b) and Kawamura et al. (1985) in
the Yokota area, by Tazawa et al. (1981) and Kawamura(1985c) in the Nisawa area (Fig. 8), and by Kawamura (1985c)

and Tazawa and Kurita (2019a) in the Shizu area (Fig. 9).
According to Tazawa and Katayama (1979) and Tazawa (1979,2017), the Odaira Formation in the Yokota area has a total
thickness of about 550 m and is subdivided into a lower part
(OD1 Unit: andesitic tuff and tuff breccia, 125 m thick), a
middle part (OD2 Unit: alternating sandstone‒tuff, intercalatedwith thin limestone layers, 318 m thick) and an upper part (OD3
Unit: black shale and sandy limestone, 107 m thick; Fig. 11).
Brachiopod fossils occur from the OD2 and OD3 units, although
the OD1 Unit is barren of fossils.
MATERIALS
Most of the brachiopod specimens described herein, except
for the specimens numbered UHR prefix, were collected from
42 localities (stations) in the Shimoarisu, Yokota, Yahagi,
Nisawa, Okuhinotsuchi and Shizu areas of the South Kitakami

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FIGURE 6. Map showing the fossil localities AOD1, IYK1, IYK2, IYK3, KF158, KF159, KF175, KF177, KF179, KF180, KF181, KF207, KF208,
KYT1, KYT2, KYT3 and KYT4 in the Yokota area (using the electronic topographical map of GSI).

Belt. The topographic and stratigraphic locations, lithologies andbrachiopod species of the 42 fossil localities are indicated in
Figs. 5‒13, and summarized in Fig. 14 and Appendix. The
materials were collected by T. Katayama, F. Itabashi, Y. Iryu,
M. Yamazaki, Y. Kurita, Y. Ibaraki, H. Araki and the present
author. In addition, some specimens were prepared from thecollections of the Hokkaido University Museum, Sapporo. The
specimens described herein are registered and stored in the
Department of Geology, Niigata University, Niigata (NU-B

prefix), the Tohoku University Museum, Sendai (IGPS prefix),
the Hokkaido Univerity Museum, Sapporo (UHR prefix) and theKesennuma Board of Education, Kesennuma (KCG prefix).
BRACHIOPOD FAUNAS
The brachiopods described herein are the following 56
species in 39 genera, including a new species (Levitusia
elongata sp. nov.): Orbiculoidea sp. Leptagonia analoga

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FIGURE 7. Map showing the fossil localities KF210 and YUK1 in the Yahagi area (using the electronic topographical map of GSI).

(Phillips, 1836), Rugosochonetes extensus (Chao, 1928),
Rugosochonetes sp., Plicatifera plicatilis (Sowerby, 1824), P.
pseudoplicatilis (Muir-Wood, 1928), Absenticosta uldzejtuensis
Suursuren and Lazarev in Lazarev, 1991, Argentiproductus sp.,
Alitaria konincki (Muir-Wood and Cooper, 1960), Levitusia
humerosa (Sowerby, 1822), L. elongata sp. nov., Tolmatchoffia
robusta (Tolmatchoff, 1924), Tomiproductus elegantulus(Tolmatchoff, 1924), Marginicinctus marginicinctus (Prout,1857), Marginatia burlingtonensis (Hall, 1858), Echinoconchus
punctatus (Sowerby, 1822), Echinaria sp., Pustula pustulosa(Phillips, 1836), Ovatia elongata Muir-Wood and Cooper, 1960,
Fluctuaria undata (Defrance, 1826), Striatifera angusta(Yanishevsky, 1910), Orthotetes keokuk (Hall, 1858),
Schellwienella radialis (Phillips, 1836), Rhipidomella michelini(Léveillé, 1835), R. kusbassica Beznossova in Sarytcheva et al.,1963, Schizophoria resupinata (Martin, 1809), S. pinguis
Demanet, 1934, S. woodi Bond, 1941, S. mayesensis Carter,
1999, Rotaia hikoroichiensis Tazawa, 2006, Cleiothyridina
fimbriata (Phillips, 1836), C. harkeri Carter, 1987, Martinia
georgei Tazawa in Tazawa et al., 2019, Spirifer liangchowensis
Chao, 1929, Grandispirifer mylkensis Yang, 1959, Unispirifer
striatoconvolutus (Dun and Benson in Benson et al., 1920), U.
minnewankensis (Shimer, 1926), U. kozuboensis (Minato,
1952), Unispirifer sp., Brachythyrina sp., Imbrexia forbesi(Norwood and Pratten, 1855), I. incertus (Hall, 1858),
Acuminothyris triangularis Roberts, 1963, Brachythyris
chouteauensis (Weller, 1909), Kitakamithyris hikoroitiensis

Minato, 1951, Kitakamithyris sp., Torynifer asiatica Beznossova
in Beznossova et al., 1962, Syringothyris texta (Hall, 1857), S.
platypleura Weller, 1914, Syringothyris sp., Asyrinxia
nipponotrigonalis (Minato, 1951), Asyrinxia sp., Pseudosyrinx
jumonjiensis (Minato, 1951), Pseudosyrinx sp., Dimegelasma
sp. and Punctospirifer sp.
The brachiopod species from the Shittakazawa, Arisu andOdaira Formations are classified into three faunas: theShittakazawa fauna from the Shittakazawa Formation, the Arisu
fauna from the Arisu Formatin, and the Odaira fauna from the
Odaira Formation. Moreover, the three faunas are subdivided
into five asemblages: the ST3 assemblage from the ST3 Unit
(upper part of the Shittakazawa Formation), the AR2
assemblage from the AR2 Unit (middle part of the Arisu
Formation), the AR3 assemblage from the AR3 Unit (upper partof the Arisu Formation), the OD2 assemblage from the OD2Unit (middle part of the Odaira Formation), and the OD3
assemblage from the OD3 Unit (upper part of the Odaira
Formation).
Shittakazawa Fauna
ST3 assemblage
The ST3 assemblage, from the ST3 Unit of the Shittakazawa
Formation, includes four species in three genera: Rhipidomellakusbassica, Schizophoria pinguis, S. mayesensis and Unispirifer
kozuboensis. Of these species, Schizophoria mayesensis is20

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FIGURE 8. Map showing the fossil localities INS1, INS2, INS3, INS4, INS5, INS6, INS7, INS8, INS9, INS10, INS11, INS12 and INS13 in the Nisawa
area (using the electronic topographical map of GSI).

common; and the other species are rare (Fig. 14).
Arisu Fauna
AR2 assemblage
The AR2 assemblage, from the AR2 Unit of the Arisu
Formation, includes 22 species in 17 genera: Levitusia humerosa,
Tolmatchoffia robusta, Tomiproductus elegantulus, Marginatia
burlingtonensis, Ovatia elongata, Orthotetes keokuk,
Rhipidomella michelini, Schizophoria resupinata, S. pinguis, S.

woodi, Cleiothyridina harkeri, Grandispirifer mylkensis,
Unispirifer striatoconvolutus, U. kozuboensis, Unispirifer sp.,
Acuminothyris triangularis, Brachythyris chouteauensis,
Kitakamithyris hikoroitiensis, Syringothyris texta, S. platypleura,
Asyrinxia nipponotrigonalis and Pseudosyrinx jumonjiensis. Of
these species, Schizophoria resupinata is abundant; Tolmatchoffia
robusta, Marginatia burlingtonensis, Rhipidomella michelini,
Schizophoria pinguis, Unispirifer striatoconvolutus, U.
kozuboensis and Kitakamithyris hikoroitiensis are common; andthe other species are rare (Fig. 14).

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

21

FIGURE 9. Map showing the fossil localities KAR4, YKA12, YKA16, YKA18, YKA20 and YKA21 in the Shizu area (using the electronic topographicalmap of GSI).

AR3 assemblage
The AR3 assemblage, from the AR3 Unit of the Arisu
Formation, includes five species in five genera: Orbiculoideasp., Marginatia burlingtonensis, Unispirifer minnewankensis,
Brachythyrina sp. and Punctospirifer sp. All the species are rare
(Fig. 14).
Odaira Fauna
OD2 assemblage
The OD2 assemblage, from the OD2 Unit of the Odaira
Formation, includes 17 species in 16 genera, with a new species(Levitusia elongata sp. nov.): Absenticosta uldzejtuensis,
Levitusia elongata sp. nov., Marginicinctus marginicinctus,
Fluctuaria undata, Striatifera angusta, Orthotetes keokuk,
Rhipidomella michelini, Schizophoria resupinata, Rotaia
hikoroichiensis, Martinia georgei, Spirifer liangchowensis,
Imbrexia incertus, Brachythyris chouteauensis, Syringothyrissp., Asyrinxia nipponotrigonalis, Asyrinxia sp. andDimegelasma sp. Of these species, Rhipidomella michelini and
Schizophoria resupinata are abundant; Absenticosta
uldzejtuensis, Martinia georgei, Spirifer liangchowensis,

Syringothyris sp. and Dimegelasma sp. are common; and theother species are rare (Fig. 14).
OD3 assemblage
The OD3 assemblage, from the OD3 Unit of the Odaira
Formation, includes 25 species in 22 genera: Leptagonia
analoga, Rugosochonetes extensus, Rugosochonetes sp.,
Plicatifera plicatilis, P. pseudoplicatilis, Argentiproductus sp.,
Alitaria konincki, Marginicinctus marginicinctus, Marginatia
burlingtonensis, Echinoconchus punctatus, Echinaria sp.,
Pustula pustulosa, Fluctuaria undata, Orthotetes keokuk,
Schellwienella radialis, Rhipidomella michelini, Cleiothyridina
fimbriata, Martinia georgei, Spirifer liangchowensis, Imbrexia
forbesi, I. incertus, Brachythyris chouteauensis, Kitakamithyrissp., Torynifer asiatica and Pseudosyrinx sp. Of these species,
Spirifer liangchowensis is abundant; Alitaria konincki,
Marginatia burlingtonensis, Imbrexia forbesi and I. incertus arecommon; and the other species are rare (Fig. 14).
AGE AND CORRELATION
Shittakazawa Fauna

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ST3 assemblage
The stratigraphic distributions of the brachiopod species of
the ST3 assemblage are described in the section “Systematic
descriptions”, and summarized in Fig. 15. Of the brachiopods
listed above, Rhipidomella kusbassica is known from the lower
Tournaisian–lower Visean, Schizophoria pinguis is known fromthe upper Tournaisian–upper Visean, Schizophoria mayesensis
occurs in the upper Tournaisian, and Unispirifer kozuboensis
occurs in the upper Tournaisian–lower Visean. In summary, theST3 assemblage is identified as late Tournaisian in age. Thus,
the upper part (ST3 Unit) of the Shittakazawa Formation iscorrelated with the upper Tournaisian. This conclusion isconsistent with the finding of Tazawa and Kurita (2019a).
Arisu Fauna

FIGURE 10. Generalized columnar section of the Carboniferous
formations in the Shimoarisu area, showing the fossil horizon IST1 in
the AR2 Unit of the Arisu Formation (modified from Tazawa and Iryu,2019). 1, shale; 2, alternating shale and sandstone; 3, conglomerate; 4,limestone of the Nagaiwa Formation; 5, 6, limestone of the Arisu andOdaira Formations; 7, andesitic tuff; 8, andesitic lapilli tuff; 9,alternating shale and rhyolitic tuff; 10, rhyolitic tuff.

AR2 assemblage
The stratigraphic distributions of the brachiopod species of
the AR2 assemblage are described in the section “Systematic
descriptions”, and summarized in Fig. 16. Of the brachiopods
listed above, Levitusia humerosa, Syringothyris texta andPseudosyrinx jumonjiensis are known only from the lower
Visean, Tolmatchoffia robusta, Kitakamithyris hikoroitiensis
and Syringothyris platypleura have a stratigraphic range of thelower Tournaisian— lower Visean, and four species(Cleiothyridina harkeri, Grandispirifer mylkensis, Unispirifer
kozuboensis and Acuminothyris triangularis) are known fromthe upper Tournaisian–lower Visean. In contrast, Asyrinxia
nipponotrigonalis occurs in the lower and upper Visean, three
species (Tomiproductus elegantulus, Ovatia elongata andBrachythyris chouteauensis) are known from the lower
Tournaisian–upper Visean, and five species (Marginatia
burlingtonensis, Orthotetes keokuk, Schizophoria resupinata, S.
pinguis and Unispirifer striatoconvolutus) have a stratigraphic
range of the upper Tournaisian–upper Visean. Two species(Rhipidomella michelini and Schizophoria woodi) are
long-ranging species, from the upper Tournaisian to the
Serpukhovian. To summarize, the age of the AR2 assemblage isidentified as early Visean; thus, the middle part (AR2 Unit) of
the Arisu Formation is correlated with the lower Visean. Thisconclusion is consistent with the age determination andcorrelation by Tazawa and Ibaraki (2019), Tazawa and Iryu
(2019) and Tazawa and Kurita (2019b).
AR3 assemblage
The stratigraphic distributins of the brachiopod species of theAR3 assemblage are described in the section “Systematic
descriptions”, and summarized in Fig. 17. Of the brachiopods
listed above, Marginatia burlingtonensis is known from theupper Tournaisian–upper Visean, and Unispirifer minnewankensis
is known from the upper Tournaisian–lower Visean. The otherspecies (Orbiculoidea sp., Brachythyrina sp. and Punctospirifersp.) are obscure in age, because of their indeterminate state at
species level. In summary, the age of the AR3 assemblage isidentified as early Visean, not to be late Tournaisian–early
Visean, because this assemblage is stratigraphically younger

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

23

FIGURE 11. Generalized columnar section of the Carboniferous formations in the Yokota‒Yahagi area, showing the fossil horizons in the AR2 and AR3
units in the Arisu Formation and the OD2 and OD3 units in the Odaira Formation (modified and adapted from Tazawa, 2017). 1, shale; 2, sandstone; 3,
limestone of the Nagaiwa Formation; 4, limestone of the Onimaru Formation; 5, limestone of the Arisu and Odaira Formations; 6, andesitic tuff; 7,
andesitic tuff breccia; 8, andesitic lapilli tuff; 9, rhyolitic tuff.

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FIGURE 12. Generalized columnar section of the Carboniferous
formations in the Nisawa area, showing the fossil horizons in the OD2
and OD3 units in the Odaira Formation (modified and adapted from
Tazawa et al., 1981). 1, shale; 2, sandstone; 3, conglomerate; 4,limestone of the Nagaiwa Formation; 5, limestone of the Onimaru
Formation; 6, limestone of the Odaira Formation; 7, andesitic tuff.

FIGURE 13. Generalized columnar section of the Carboniferous
formations in the Okuhinotsuchi‒Shizu area, showing the fossil horizonsin the ST3 Unit in the Shittakazawa Formation, the AR2 Unit in the
Arisu Formation, and the OD2 Unit in the Odaira Formation (modified
and adapted from Tazawa and Kurita, 2019a). 1, shale; 2, sandstone; 3,alternating shale and sandstone; 4, conglomerate; 5, limestone of theNagaiwa Formation; 6, limestone of the Onimaru Formation; 7,
limestone of the Shittakazawa, Arisu and Odaira Formations; 8,
andesitic tuff; 9, andesitic lapilli tuff; 10, rhyolitic tuff. F = fault.

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

25

FIGURE 14. Occurrence of brachiopod species from the Shittakazawa, Arisu and Odaira Formations in the Shimoarisu, Yokota, Yahagi, Nisawa,Okuhinotsuchi and Shizu areas, South Kitakami Belt. A, abundant; C, common; R, rare.

26

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FIGURE 15. Stratigraphic distributions of brachiopod species of the
ST3 assemblage (Shittakazawa fauna).

than the previously discussed AR2 assemblage (early Visean).
Thus, the upper part (AR3 Unit) of the Arisu Formation iscorrelated with the lower Visean. This conclusion is consistent
with the age determination and correlation by Tazawa (1985).
Odaira Fauna
OD2 assemblage
The stratigraphic distributins of the brachiopod species of the
OD2 assemblage are described in the section “Systematic
descriptions”, and summarized in Fig. 18. Of the brachiopods
listed above, Absenticosta uldzejtuensis, Levitusia elongata sp.
nov. and Marginicinctus marginicinctus are known only fromthe upper Visean, two species (Rotaia hikoroichiensis andSpirifer liangchowensis) are known from the lower–upper
Visean, Brachythyris chowteauensis is known from the lower
Tournaisian–upper Visean, and three species (Orthotetes
keokuk, Schizophoria resupinata and Imbrexia incertus) are
know n f r o m t h e u p p e r T o u rn a is ia n – u p p e r V is e a n. In
contrast, Striatifera angusta is known from the upper
Visean–Serpukhovian. Three species are long-ranging species:
Rhipidomella michelini ranges from the upper Tournaisian to the
Bashkirian, and both Fluctuaria undata and Martinia georgei
range from the lower Visean to the Bashkirian. At thegeneric level, Syringothyris has a long range from theFamennian–Serpukhovian (Carter, 2006), Asyrinxia is known
from the upper Tournaisian–upper Visean (Carter, 2006;
Tazawa, 2018a), and Dimegelasma is known from thelower–upper Visean (Carter, 2006). In summary, the age of the
OD2 assemblage is identified as late Visean. Thus, the middle
part (OD2 Unit) of the Odaira Formation is correlated with theupper Visean.
OD3 assemblage
The stratigraphic distributins of the brachiopod species of the
OD3 assemblage are described in the section “Systematic

FIGURE 16. Stratigraphic distributions of brachiopod species of theAR2 assemblage in the Arisu fauna. Broken lines show those of genera.

descriptions” , and summarized in Fig. 19. Of the brachiopods
listed above, Plicatifera plicatilis and Marginicinctus
marginicinctus are known only from the upper Visean,
Leptagonia analoga and Brachythyris chouteauensis are known
from the lower Tournaisian–upper Visean, five species(Marginatia burlingtonensis, Orthotetes keokuk, Schellwienellaradialis, Imbrexia forbesi and I. incertus) are known from theupper Tournaisian–upper Visean, and four species(Rugosochonetes extensus, Plicatifera pseudoplicatilis,
Cleiothydina fimbriata and Spirifer liangchowensis) are
known from the lower–upper Visean. In contrast, Alitaria
konincki and Pustula pustulosa are known from the lower

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

27

FIGURE 17. Stratigraphic distributions of brachiopod species of theAR3 assemblage in the Arisu fauna. Broken lines show those of genera.

Visean–Serpukhovian, and Fluctuaria undata is known from thelower Visean–Bashkirian. Four species are long-ranging species:
Torinifer asiatica ranges from the lower Tournaisian to the
Serpukhovian, Rhipidomela michelini ranges from the upper
Tournaisian–Bashkirian, Echinoconchus punctatus ranges fromthe upper Tournaisian to the Asselian, and Martinia georgei
ranges from the lower Visean to the Bashkirian. At thegeneric level, Rugosochonetes has a long range from theTournaisian–Gzhelian (Racheboeuf, 2000), Argentiproductus
occurs in the lower–upper Visean (Brunton et al., 2000),
Echinaria is known from the upper Visean–Cisuralian (Tazawa,
1981a; Brunton et al., 2000; this study), Kitakamithyris is
known from the Famennian–upper Visean (Tazawa, 2017,2018b), and Pseudosyrinx is known from the upper
Tournaisian–upper Visean (Carter, 2006; this study). In
summary, the age of the OD3 assemblage is identified as late
Visean. Thus, the upper part (OD3 Unit) of the Odaira
Formation is correlated with the upper Visean. This conclusion
is consistent with the age determination and correlation by
Tazawa (1985, 2017) and Kawamura et al. (1985).
PALAEOBIOGEOGRAPHY
Late TournaisianThe geographic distributions of the late Tournaisianbrachiopod species of the ST3 assemblage are described in thesection “Systematic descriptions”, and summarized in Figs. 20
and 21. Among the four species of the ST3 assemblage, one
species also occurs in the USA (Oklahoma), the UK (England),
Poland, Belgium, central Russia (Kuznetsk Basin), Kazakhstan
and northwestern China (Xinjiang). Thus, the ST3 assemblage
has a close affinity with those of the USA (Oklahoma), the UK(England), Poland, Belgium, central Russia (Kuznetsk Basin),

FIGURE 18. Stratigraphic distributions of brachiopod species of the
OD2 assemblage in the Odaira fauna. Broken lines show those ofgenera.

Kazakhstan and northwestern China (Xinjiang). This conclusion
is consistent with the late Tournaisian biogeography based on
the brachiopod faunas of the South Kitakami Belt: the HK1
asseblage from the lower part (HK1 Unit) of the Hikoroichi
Formation in the Hikoroichi area (Tazawa, 2018c); and the KU1
assemblage from the lower part (KU1 Unit) of the Karaumedate
Formation in the Nagasaka area (Tazawa et al., 2019).
Early Visean
The geographic distributions of the early Visean brachiopod
species of the AR2 and AR3 assemblages are described in thesection “Systematic descriptions”, and summarized in Figs. 22
and 23. Of the 22 species of the two assemblages, six species
also occur in Kazakhstan and northwestern China (Xinjiang);
five species are found in the UK (England), Belgium and central
Russia (southern Urals and Kuznetsk Basin); four species havebeen reported from the USA (Missouri), Canada (Alberta) and
Poland; and three species also occur in northern Russia(Verkhoyansk Range and Pechora Basin), western Russia(Moscow Basin and Donetz Basin), Iran, Uzbekistan,

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FIGURE 20. Geographic distributions of brachiopod species of the ST3
assemblage (Shittakazawa fauna).

(Tazawa, 2018c): and the Shimoarisu fauna from the middle part(AR2 Unit) of the Arisu Formation in the Shimoarisu area
(Tazawa and Iryu, 2019).
Late Visean

FIGURE 19. Stratigraphic distributions of brachiopod species of the
OD3 assemblage in the Odaira fauna. Broken lines show those ofgenera.

Kyrgyzstan, northwestern China (Qinghai) and southwestern
China (Yunnan). To summarize, the Arisu fauna (AR2 and AR3
as semblag es) ex h ib its a clo s e affinity w ith the early
Carboniferous (Mississippian) brachiopod faunas of the UK(England), Belgium, central Russia (southern Urals andKuznetsk Basin), Kazakhstan and northwestern China(Xinjiang), particularly with Kazakhstan and northwestern
China (Xinjiang). This conclusion is consistent with the early
Visean biogeography inferred from the brachiopod faunas of theSouth Kitakami Belt: the HK2 asseblage from the middle part
(HK2 Unit) of the Hikoroichi Formation in the Hikoroichi area

The geographic distributions of the late Visean brachiopod
species of the OD2 and OD3 assemblages are described in thesection “Systematic descriptions”, and are summarized in Figs.
24 and 25. Among the 26 species of the two assemblages, 11
species also occur in Kazakhstan; 10 species also occur in the
UK (England), Belgium, central Russia (southern Urals) andnorthwestern China (Xinjiang); seven species also occur inwestern Russia (Moscow Basin) and Kyrgyzstan; six species also
occur in the UK (northern Ireland), central Russia (Kuznetsk
Basin) and northwestern China (Gansu); five species also occur
in the USA (Iowa), northern Russia (Verkhoyansk Range),
Germany and Uzbekistan; and four species also occur in the USA(Missouri), northern Russia (Pechora Basin), the UK (Wales),Spain and western Russia (Donetz Basin). In summary, theOdaira fauna (OD2 and OD3 assemblages) exbits affinities withthe early Carboniferous (Mississippian) brachiopod faunas of the
UK (England), Belgium, central Russia (southern Urals),
Kazakhstan and northwestern China (Xinjiang), particularly with
Kazakhstan. This conclusion is consistent with the late Viseanbiogeography based on the brachiopod faunas of the South
Kitakami Belt: the Tairagai fauna from the upper part of the
Odaira Formation in the Yokota area (Tazawa, 2017); the HK4
asseblage from the upper part (HK4 Unit) of the Hikoroichi
Formation in the Hikoroichi area (Tazawa, 2018c); and the KU4
assemblage from the upper part (KU4 Unit) of the Karaumedate
Formation in the Nagasaka area (Tazawa et al., 2019).

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

29

FIGURE 21. Late Tournaisian reconstruction map of the world (adapted from Qiao and Shen, 2014), showing the geographic distributions of brachiopod
species of the ST3 assemblage (Shittakazawa fauna). Solid circles indicate numbers of brachiopod species listed in the ST3 assemblage. Station numbers
are same as in Fig. 22.

DISCUSSION: EARLY CARBONIFEROUS GEOGRAPHY
AND BIOGEOGRAPHY Of THE SOUTH KITAKAMI BELT
Yang (1980, 1983, 1985) discussed the early Carboniferous
brachiopod biogeography of China and the surrounding regions.
He proposed the existence of three provinces: the North China
Province, the South China Province and the Himalayan
Province. The North China Province covers a broad area
between the North China Block and the Siberian Block,including Kazakhstan (Altay Mountains), Kyrgyzstan (Tien
Shan), northwestern China (Xinjiang), northern China (Inner
Mongolia) and northeasten China (Heilongjiang). The fauna of
the province is closely related to those of North America,
Siberia and western Europe, and is characterized by Marginatia,
Tolmatchoffia, Rotaia, Syringothyris, Pseudosyrinx,
Grandispirifer and Balakhonia. The South China Province
covers most areas of the South China Block, including theYangtze River and Pearl River regions. The brachiopod fauna ispeculiar to southern China and eastern Asia, and is characterized
by Yanguania, Hunanoproductus, Eochoristites, Martiniella,
Vitiliproductus, Datangia, Lochengia and Gondolina. The
Himalayan Province covers the area south of the Yarlung
Zangbo River, including Xizang (Tibet). The fauna ischaracterized by Fusella, Ovatia, Marginirugus, Syringothyris
and Balakhonia. After that, Yang (1990) proposed three realms
in the Early Carboniferous brachiopod biogeography in China

and the adjacent regions, i.e., the Boreal Realm, the Tethyan
Realm and the Himalayan Realm. He treated that northern Japan(South Kitakami) belonged to the Verkhoyansk–Northeast Japan
Subprovince in the Boreal Realm.
The early Carboniferous brachiopod faunas of the central partof the South Kitakami Belt are characterized by Tolmatchoffia,
Marginatia, Rhipidomella, Schizophoria, Rotaia,
Grandispirifer, Unispirifer, Kitakamithyris, Syringothyris andPseudosyrinx. The faunas display close affinities with those of
North America, western Europe, northern, western and central
Russia, Kazakhstan, Uzbekistan, Kyrgyzstan and norhwestern
China, particularly with those of Kazakhstan and northwestern
China. Thus, the early Carboniferous brachiopod faunas of theSouth Kitakami Belt are allied with those of the North China
Province of Yang (1983). The South Kitakami region wasprobably the eastern extension of the province, and was locatednear and to the east of the North China Block during the early
Carboniferous. This conclusion is consistent with the works of
Tazawa (2002, 2006, 2017, 2018c), Tazawa and Iryu (2019),
Tazawa and Kurita (2019b) and Tazawa et al. (2019). In thesecond contribution of the present series, Tazawa et al. (2019)
proposed that the South Kitakami Belt was probably a shallow
sea bordering a microcontinent, which consisted of the
Hida–Oki Terrane (Suwa, 1990) and the East China Sea region,
and was situated near and to the east of the North China Block at
the eastern end of the Central Asian Orogenic Belt (CAOB)

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FIGURE 22. Geographic distributions of brachiopod species of the AR2 and AR3 assemblages in the Arisu fauna.

during the late Tournaisian and Visean. In the present paper, I
consider that the microcontinent includes the Hida–Oki Terraneand the South Kitakami Terrane (Tazawa, 2004; revised by
Tazawa, 2018d), but uncertain about the East China Sea region.
Minato (1956) briefly discussed the early Carboniferous(Hikoroichi–Arisu–Odaira ages, late Tournaisian–late Visean)
biogeography and geography of the South Kitakami Belt based
mainly on brachiopods. In that paper Minato proposed that theearly Carboniferous brachiopod fauna of the South Kitakami
Belt exhibited a close affinity with those of eastern Australia
and the Mongolian geosynclinal region (central Asia), and wasquite different from that of South China. Subsequently, Minato
and Kato (1984) proposed that the early Carboniferous(Hikoroichi–Arisu–Odaira ages) faunas, consisting mainly ofbrachiopods and corals, of the South Kitakami Belt showedstrong affinity with those of the United States, Russia andnorthern China.
Kobayashi and Hamada (1980) proposed two realms and fiveprovinces in the Carboniferous trilobite biogeography, the Old
World Realm (including the West–Central European, theEuropean Russia–Central Asiatic, the Oriental and theAustralian provinces) and the New World Province (including
the North American Mid-Continent Province). They treated theSouth Kitakami fauna as a member of the Mongolian
Geosynclinal Region in the European Russia–Central Asiatic

Province.
Qiao and Shen (2014) discussed the Misissippian global
biogeography on the basis of quantitative analysis of theMississipian brachiopod faunas of the world. It is noteworthy
that South Kitakami was most closely related to northern
Tianshan in Xinjiang, northwestern China (Qiao and Shen,
2014, fig. 6B) and located near and to the east of the North
China Block (Qiao and Shen, 2014, figs. 3B) during the Visean.
In contrast to the above studies, Kawamura et al. (1999),
Ehiro (2001), Okawa et al. (2013) and Isozaki et al. (2014,2017) considered that South Kitakami was located near theSouth China Block during the Carboniferous. However, thepalaeontological (taxonomical) data are insufficient or lacking inthese studies. That hypothesis is not supported by the early
Carboniferous brachiopod biogeography discussed above.
CONCLUSIONSIn this study, early Carboniferous brachiopods are describedfrom the Shittakazawa, Arisu and Odaira Formations in thecentral part (Shimoarisu, Yokota, Yahagi, Nisawa,Okuhinotsuchi and Shizu areas) of the South Kitakami Belt,
northeastern Japan. A total of 56 species in 29 genera aredescribed, of which one (Levitusia elongata sp. nov.) is new. In
terms of brachiopod biostratigraphy, the Shittakazawa

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

31

FIGURE 23. Early Visean reconstruction map of the world (adapted from Qiao and Shen, 2014), showing the geographic distributions of brachiopod
species of the AR2 and AR3 assemblages in the Arisu fauna. Solid circles indicate numbers of brachiopod species listed in the AR2 and AR3 assemblages.
Station numbers are same in Fig. 24.

FIGURE 4. Comparison of stratigraphic schemes proposed for the lower Carboniferous formations in the Nisawa, Okuhinotsuchi and Shizu areas, South
Kitakami Belt.

FIGURE 24. Geographic distributions of brachiopod species of the OD2 and OD3 asemblages in the Odaira fauna.

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JUN-ICHI TAZAWA

FIGURE 25. Late Visean reconstruction map of the world (adapted from Qiao and Shen, 2014), showing the geographic distributions of brachiopod
species of the OD2 and OD3 assemblages in the Odaira fauna. Solid circles indicate numbers of brachiopod species listed in the OD2 and OD3
assemblages. Station numbers are same in Fig. 26.

Formation is correlated with the upper Tournaisian; the Arisu
Formation is correlated with the lower Visean; and the Odaira
Formation is correlated with the upper Visean.
Palaeobiogeographically, the late Tournaisian (Shittakazawa)
fauna resembles brachiopod assemblages of the USA(Oklahoma), the UK (England), Poland, Belgium, central Russia(Kuznetsk Basin), Kazakhstan and northwestern China(Xinjiang). The early Visean (Arisu) fauna has a close affinity
with those of the UK (England), Belgium, central Russia(southern Urals and Kuznetsk Basin), Kazakhstan andnorthwestern China (Xinjiang), particularly with Kazakhstan
and northwestern China (Xinjiang). The late Visean (Odaira)fauna displays affinities with those of the UK (England),Belgium, central Russia (southern Urals), Kazakhstan andnorthwestern China (Xinjiang), particularly with Kazakhstan. I
conclude that the early Carboniferous (late Tournaisian–lateVisean) brachiopod faunas of the central part of the South
Kitakai Belt are allied with those of Kazakhstan andnorthwestern China (Xinjiang), and belonged to the North China
Province of Yang (1983). The South Kitakami region wasprobably located in the eastern part of the province. During theearly Carboniferous, the area was sited in the eastern part of theCentral Asian Orogenic Belt and near and to the east of theNorth China Block as a shallow sea bordering a nicrocontinent.

SYSTEMATIC DESCRIPTIONSThe 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 LINGULIDA Waagen, 1885
Superfamily DISCINOIDEA Gray, 1840
Family DISCINIDAE Gray, 1840
Genus ORBICULOIDEA d’Orbigny, 1847
Type species.―Orbicula forbesii Davidson, 1848.
Orbiculoidea sp.
(Fig. 36A)
Material.—One specimen from locality KF207, externalmould of a ventral valve, NU-B2266.
Remarks.—This specimen is safely assigned to the genus
Orbiculoidea by the slightly concave, subcircular ventral valve,with a narrow pedicle track near apex, and the external ornamentconsisting of numerous fine concentric growth lines. The ventral
valve is slightly elongate subcircular in outline, gently concaveand large in size for genus (length 26 mm, width 23 mm), andMISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

33

FIGURE 26. Brachiopds of the ST3 assemblage. A, Rhipidomella kusbassica Beznossova, internal mould of dorsal valve, IGPS111760; B, C,
Schizophoria pinguis Demanet; dorsal (B1), anterior (B2), posterior (B3) and lateral (B4) views of internal mould of dorsal valve, IGPS99007; dorsal (C1)and lateral (C 2 ) views of internal mould of dorsal valve, IGPS99006; D–F, Schizophoria mayesensis Carter; D, internal mould of ventral valve,
IGPS111762; E, internal mould of dorsal valve, IGPS111763; internal mould (F 1) and external latex cast (F 2) of dorsal valve, IGPS111764; G, H,
Unispirifer kozuboensis (Minato); G, internal mould of ventral valve, IGPS99008; H, internal mould of ventral valve, IGPS111761. Scale bars are 1 cm.

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JUN-ICHI TAZAWA

having a narrow, relatively short pedicle track. The single
specimen from Yokota most resembles Orbiculoidea
portlockiana Demanet (1934, p. 24, pl. 1, figs. 14–17, text-fig.
6), from the upper Tournaisian of Tournai, Belgium, in size andoutline of the ventral valve. But accurate comparison is difficult
for the poorly preserved specimen.
Occurrence.—AR3 Unit (Yokota).
Order STROPHOMENIDA Öpik, 1934
Superfamily STROPHOMENOIDEA King, 1846
Family RAFINESQUINIDAE Schuchert, 1893
Subfamily LEPTAENINAE Hall and Clarke, 1894
Genus LEPTAGONIA M’Coy, 1844
Type species.—Producta analoga Phillips, 1836.
Leptagonia analoga (Phillips, 1836)(Fig. 43A)
Producta analoga Phillips, 1836, p. 116, pl. 7, fig. 10.
Strophomena rhomboidalis var. analoga Phillips. Davidson,
1861, p. 119, pl. 28, figs. 1–6, 9–13; Etheridge, 1872, p. 333,
pl. 18, fig. 1.
Leptaena analoga (Phillips). Weller, 1914, p. 49, pl. 2, figs.
1–10; Frech, 1916, p. 237, pl. 2, figs. 2, 3d; Girty, 1920, pl.
54, fig. 3; Tolmatchoff, 1924, p. 209, 569. pl. 13, fig. 8; Girty,
1927, pl. 22, figs. 6–8; Demanet, 1934, p. 61, pl. 5, figs. 1–14,
text-figs. 11–14; Branson, 1938, p. 24, pl. 5, fig. 31; Nelson,
1961, pl. 4, fig. 26; Sokolskaya in Sarytcheva et al., 1963, p.
80, pl. 4, figs. 9–14.
Leptaena rhomboidalis Wilckens. Sommer, 1909, p. 626, pl. 29,
fig. 14.
Leptagonia cf. analoga (Phillips). Cvancara, 1958, p. 860, pl.
110, figs. 6–13, text-figs. 3, 4.
Leptaenella analoga (Phillips). Yang, 1964, p. 61, pl. 1, fig. 5;
Grechishnikova, 1966, p. 94, pl. 1, figs. 19, 20; pl. 2, figs.
1–6; Abramov, 1970, p. 108, pl. 1, figs. 11, 12; Aisenverg and
Poletaev, 1971, pl. 60, figs. 2, 3; Nalivkin and Fotieva, 1973,
p. 20, pl. 1, figs. 9–13.
Leptagonia analoga (Phillips). Brunton, 1968, p. 29, pl. 3, figs.
26–31; pl. 4, figs. 1–9, text-figs. 6–17; Gaetani, 1968, p. 688,
pl. 47, fig. 3; Thomas, 1971, p. 30, pl. 18, figs. 1–8, text-fig.
11; Bublichenko, 1971, p. 37, pl. 3, figs. 1–5; Brand, 1972, p.
59, pl. 8, figs. 1–6, text-figs. 1a, 3; Kalashnikov, 1974, p. 23,
pl. 3, fig. 5; Litvinovich et al., 1975, p. 53, pl. 16, fig. 11;
Bublichenko, 1976, p. 22, pl. 1, fig. 10; Yang et al., 1977, p.
316, pl. 131, fig. 2; Nalivkin, 1979, p. 18, pl. 3, figs. 1–3, 5, 6;
Ding and Qi, 1983, p. 251, pl. 89, figs. 9, 10, 12; Zhang et al.,1983, p. 271, pl. 107, fig. 13; pl. 106, fig. 3; Tazawa et al.,1984, p. 350, pl. 67, figs. 2–4; Yang, 1984, p. 205, pl. 29, fig.
11; Xu and Yao, 1988, p. 274, pl. 67, figs. 4, 6–10; Carter,
1999, p. 96, figs. 1A–E; Shi et al., 2005, p. 39, figs. 3A, E;
Tazawa et al., 2016, p. 51, fig. 5.5; Tazawa, 2017, p. 333, fig.
6.1; Tazawa, 2018c, p. 40, figs. 16B–F, 22A, B, 30A.

Material.—One specimen from locality KF159, externalmould of a ventral valve, NU-B2154.
Remarks.—This specimen was described by Tazawa (2017,p. 333, fig. 6.1) as Leptagonia analoga (Phillips, 1836). The
single specimen from Tairagai is referred to L. analoga(Phillips, 1836), redescribed by Brunton (1968, p. 29, pl. 3, figs.
26–31; pl. 4, figs. 1–9, text-figs. 6–17) from the Visean ofEngland and northern Ireland, in the transversely trapezoidal andflattened ventral valve, which is ornamented with numerous finecostellae (numbering 13–15 in 5 mm at about midlength) andregularly but slightly flexuous concentric rugae. The Tairagai
specimen, being smaller in size (length 23 mm, width about 34mm) than the type specimens of L. analoga, may be a juvenile
shell.
Occurrence.—OD3 Unit (Yokota).
D i s t r i b u t i o n . — L o w e r T o u r n a i s i a n –u p p e r V i s e a n :northeastern Japan (Hikoroichi, Yokota and Soma in the South
Kitakami Belt), USA (Illinois, Iowa, Missouri, Oklahoma, Utah
and New Mexico), western Canada (Alberta), northern Russia(Verkhoyansk Range and Pechora Basin), UK (England, lsle of
Man and northern Ireland), Germany, Belgium, western Russia(Donetz Basin), Turkey (Taurus Mountains), Iran (Elburz
Range), central Russia (southern Urals and Kuznetsk Basin),
Kazakhstan, northwestern China (Xinjiang and Gansu),
central-southern China (Hubei, Guangdong and Guangxi),
southwestern China (Yunnan), western Australia (Bonaparte
Gulf Basin) and eastern Australia (Queensland and New South
Wales).
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 RUGOSOCHONETES Sokolskaya, 1950
Type species.—Orthis hardrensis Phillips, 1841.
Rugosochonetes extensus (Chao, 1928)(Fig. 43B, C)
Chonetes extensa Chao, 1928, p. 9, pl. 1, figs. 7–10; Fang inYang et al., 1962, p. 39, pl. 12, figs. 1–6; Ding and Qi, 1983,
p. 262, pl. 93, fig. 2.
Rugosochnetes extensus (Chao). Tazawa, 2017, p. 334, fig. 6.2;
Tazawa in Tazawa et al., 2019, p. 51, fig. 16A.
Rugosochonetes sp. Tazawa, 1980, p. 361, pl. 41, fig. 1.
Material.—Three specimens from localities KF159 andYUK1: (1) external and internal moulds of a ventral valve,
KCG76; (2) external and internal moulds of a dorsal valve,
NU-B2172; and (3) external mould of a dorsal valve, KCG77.
Remarks.—One of the specimens (NU-B2172) was describedby Tazawa (2017, p. 334, fig. 6.2) as Rugosochonetes extensus(Chao, 1928). The specimens from the upper Odaira Formation

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

35

FIGURE 27. Brachiopods of the AR2 assemblage (1). A, Levitusia humerosa (Sowerby), ventral (A1), anterior (A2) and lateral (A3, A4) views of externallatex cast of ventral valve, IGPS111744; B, C, Grandispirifer mylkensis Yang; external latex cast (B1, B2) and external mould (B4) of ventral valve, andenlarged external latex cast of ventral valve (B3) showing fine growth lines over the valve. IGPS111742; C, internal mould of ventral valve, IGPS111743.
Scale bars are 1 cm, except for B3.

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of Yahagi are referred to R. extensus (Chao, 1928), from the
Chouniugou Formation of Gausu, northwestern China, by thetransverse outline (length 7 mm, width 14 mm in the best
preserved specimen, NU-B2172), acute cardinal extremities andexternal ornament consisting of numerous fine capillae(numbering 7–8 in 2 mm at about midlength). Rugosochonetessp., described by Tazawa (1980, p. 361, pl. 41, fig. 1) from theupper part of the Karaumedate Formatin in the Nagasaka area,
South Kitakami Belt, is conspecific with the present species.
Rugosochonetes transversalis Brunton (1968, p. 65, pl. 9, figs.
16–25), from the Visean of County Fermanagh, northern Irelandin the UK, differs from R. extensus in having coarser capillae onboth ventral and dorsal valves.
Occurrence.—OD3 Unit (Yahagi).
Distribution.—Lower–upper Visean: northeastern Japan(Yahagi and Nagasaka in the South Kitakami Belt) andnorthwestern China (Gansu).
Rugosochonetes sp.
(Fig. 43D)
Material.—One specimen from locality YUK1, externalmould of a ventral valve, KCG78.
Remarks.—This specimen resembles Rugosochonetes hindi
Muir-Wood (1962, p. 70, pl. 6, figs. 1–7, text-fig. 2A), from the
Serpukhovian of the UK (Scotland, England and Wales), in themedium-sized, slightly transverse shell (length 11 mm, width 13mm) and fine capillae (numbering 6–7 in 1 mm at midlength) on
the ventral valve. Accurate comparison is, however, difficult forthe poorly preserved specimen. Rugosochonetes hardlensis(Phillips, 1841), refigured by Muir-Wood (1962, pl. 2, figs.
7–10, 13) from the upper Visean of Yorkshire, England, differsfrom the present species in being more transverse outline. The
preceding species, Rugosochonetes extensus (Chao, 1928), isreadily distinguished from the present species in the muchtransverse outline and in having coarser capillae on the ventral
valve.
Occurrence.—OD3 Unit (Yahagi).
Suborder PRODUCTIDINA Waagen, 1883
Superfamily PRODUCTELLOIDEA Schuchert in Schuchertand LeVene, 1929
Family OVERTONIIDAE Muir-Wood and Cooper, 1960
Subfamily PLICATIFERINAE Muir-Wood and Cooper, 1960
Genus PLICATIFERA Chao, 1927
Type species.—Productus plicatilis Sowerby, 1824.
Plicatifera plicatilis (Sowerby, 1824)(Fig. 43F)
Productus plicatilis Sowerby, 1824, p. 85, pl. 459, fig. 2;
Thomas, 1914, p. 331, pl. 20, fig. 22, text-fig. 13.
Plicatifera plicatilis (Sowerby). Muir-Wood and Cooper, 1960,
pl. 56, figs. 13–23; Galitskaya, 1977, p. 22, pl. 8, figs. 6, 7;

Zakowa, 1985, p. 306, pl. 1, figs. 5–7; pl. 2, figs. 1–3; pl. 5,
fig. 5; Brunton and Mundy in Brunton et al., 1993, p. 108,
figs. 29–35.
Productus-Plicatifera-plicatilis Sowerby. Pareyn, 1961, p. 209,
pl. 26, fig. 5.
Material.—Two specimens from locality YUK1, externalmoulds of two ventral valves, KCG65, 66.
Remarks.—The fragmentarily preserved specimens from
Yahagi can be referred to Plicatifera plicatilis (Sowerby, 1824),
redescribed by Brunton and Mundy in Brunton et al. (1993, p.
108, figs. 19–35) from the upper Visean of Derbyshire andYorkshire, England, by the medium-sized, transverse and gently
convex ventral valve (length about 25 mm, width about 50 mm
in the better preserved specimen, KCG65), and external
ornament of the ventral valve consisting of numerous regularconcentric rugae on visceral disc and numerous small spines
scattered on rugae. Plicatifera pseudoplicatilis (Muir-Wood,
1928, p. 189, pl. 11, fig. 17, text-fig. 34), from the upper Viseanof Staffordshire, England, differs from P. plicatilis in having
numerous costellae on the visceral disc and trail of the ventral
valve.
Occurrence.—OD3 Unit (Yahagi).
Distribution.—Upper Visean: northeastern Japan (Yahagi in
the South Kitakami Belt), UK (England), Poland, Belgium,
Algeria and Kyrgyzstan.
Plicatifera pseudoplicatilis (Muir-Wood, 1928)(Fig. 43E)
Productus pseudoplicatilis Muir-Wood, 1928, p. 189, pl. 11, fig.
17, text-fig. 34; Kalashnikov, 1974, p. 69, pl. 20, figs. 9–11.
Plicatifera plicatilis (non Sowerby). Brunton, 1966, p. 204, pl.
6, figs. 6–15; pl. 7, figs. 1–18, text-fig. 8.
Plicatifera pseudoplicatilis (Muir-Wood). Brunton and Mundy
in Brunton et al., 1993, p. 109, figs. 36–46.
Material.—One specimen from locality YUK1, externalmould of a dorsal valve, KCG70.
Remarks.—This specimen can be referred to Plicatifera
pseudoplicatilis (Muir-Wood, 1928), from the upper Visean ofStaffordshire and Yorkshire, England, by the presence ofnumerous fine costellae over the visceral disc and trail of thedorsal valve. The preceding species, Plicatifera plicatilis(Sowerby, 1824), differs from P. pseudoplicatilis in the lacking
costellae on the dorsal valve.
Occurrence.—OD3 Unit (Yahagi).
Distribution.—Lower–upper Visean: northeastern Japan(Yahagi in the South Kitakami Belt), northern Russia (northern
Urals) and UK (England).
Genus ABSENTICOSTA Lazarev, 1991
Type species.—Absenticosta uldzejtuensis Suursuren andLazarev in Lazarev, 1991

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FIGURE 28. Brachiopods of the AR2 assemblage (2). A–D, Tomiproductus elegantulus (Tolmatchoff); ventral (A1, A2), anterior (A3), posterior (A4) andlateral (A5) views of internal mould of ventral valve, IGPS112115; ventral (B1, B2), anterior (B3), posterior (B4) and lateral (B5) views of internal mould ofventral valve, IGPS112118; dorsal (C1, C2) and anterior (C3) views of external mould of dorsal valve, IGPS112119; D1, D2, internal mould of ventral
valve, NU-B2262; E, F, Tolmatchoffia robusta (Tolmatchoff), external latex cast (E1), external mould (E2) and internal mould (E3) of dorsal valve,
IGPS112098; external latex cast (F1) and internal mould (F2) of ventral valve, IGPS112097; G, Ovatia elongata Muir-Wood and Cooper, external mouldof dorsal valve, IGPS111725. Scale bars are 1 cm.

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Absenticosta uldzejtuensis Suursuren and Lazarev in Lazarev,
1991(Fig. 38A, B)
Plicatifera plicatilis (Sowerby). Kotlyar in Kotlyar and Popeko,
1967, p. 95, pl. 15, figs. 16–18; pl. 16, figs. 1–3.
Absenticosta uldzejtuensis Suursuren and Lazarev in Lazarev,
1991, p. 58, pl. 4, figs. 1–6; Lazarev and Suursuren, 1992, p.
64, pl. 14, figs. 14–17.
Material.—Five specimens from localities AOD1 andYKA20: (1) internal mould of a ventral valve, IGPS112133; and(2) external moulds of four dorsal valves, IGPS112134‒112136,
NU-B2242.
Remarks.—These specimens are referred to Absenticosta
uldzejtuensis Suursuren and Lazarev in Lazarev, 1991, from theupper Visean of the Khangay Highland, Mongolia, in the small,slightly transverse and gently convex ventral valve (length about
11 mm, width about 14 mm in the sole ventral valve specimen,
IGPS112133), ornamented with numerous thin, somewhat
irregular concentric rugae (numbering 5–6 in 2 mm at about
midlength) on entire surface of the valve. Absenticosta
bruntoneileenae Taboada and Shi (2011, p. 105, fig. 12), fromthe upper Visean of the Uspallata-Iglesia Basin, western
Argentina, is distinguished from A. uldzejtuensis in the larger
and less transverse shell.
Occurrence.—OD2 Unit (Yokota and Shizu).
Distribution.—Upper Visean: northeastern Japan (Yokota
and Shizu in the South Kitakami Belt), eastern Russia(Transbaykal) and Mongolia (Khangay Highland).
Family PRODUCTINIDAE Muir-Wood and Cooper, 1960
Subfamily PRODUCTININAE Muir-Wood and Cooper, 1960
Genus ARGENTIPRODUCTUS Cooper and Muir-Wood, 1951
Type species.—Productus margaritacea Phillips, 1836.
Argentiproductus sp.
(Fig. 43G)
Material.—Two specimens from locality YUK1, externalmoulds of two dorsal valves, KCG63, 64.
Remarks.—These specimens are safely assigned to the genus
Argentiproductus by the external ornament of dorsal valve,consisting of numerous irregular rounded costae and closely
placed concentric lamellae. The type species, Argentiproductus
margaritaceus (Phillips, 1836), refigured by Muir-Wood andCooper (1960, pl. 123, figs. 11–17) from the Visean of northern
Ireland and Wales, the UK, is more transverse in outline.
Argentiproductus auriculatus Carter (1988, p. 38, figs.
6.33–6.50), from the Glen Park Formation of Illinois, is asmaller and more transverse species. Specific identification isdifficult for the poorly preserved specimens.
Occurrence.—OD3 Unit (Yahagi).

Superfamily MARGINIFEROIDEA Stehli, 1954
Family PAUCISPINIFERIDAE Muir-Wood and Cooper, 1960
Subfamily PAUCISPINIFERINAE Muir-Wood and Cooper,
1960
Genus ALITARIA Cooper and Muir-Wood, 1967
Type species.—Alifera konincki Muir-Wood and Cooper, 1960.
Alitaria konincki (Muir-Wood and Cooper, 1960)(Fig. 43H–J)
Productus expansus de Koninck, 1842, p. 159, pl. 7, fig. 4.
Alifera konincki Muir-Wood and Cooper, 1960, p. 208, pl. 66,
figs. 6–18; Galitskaya, 1977, p. 50, pl. 13, figs. 12–17.
Alitaria konincki (Muir-Wood and Cooper). Cooper andMuir-Wood, 1967, p. 808.
Material.—Three specimens from locality YUK1: (1)internal mould of a conjoined valve, with external mould of thedorsal valve, KCG71; and (2) internal moulds of two ventral
valves, KCG72, 73.
Remarks.—These specimens are fragmentarily preserved,
but can be referred to Alitaria konincki (Muir-Wood andCooper, 1960, p. 208, pl. 66, figs. 6–18), from the Visean ofVisé, Belgium, by the small and transverse shell (length 13 mm,
width more than 18 mm in the best preserved specimen,
KCG71), with a prominent marginal ridge in the ventral valve.
Alitaria minuta (Muir-Wood, 1928, p. 195, pl. 12, fig. 3), fromthe upper Tournaisian (?)–upper Visean of England and Ireland,differs from A. konincki in the smaller size and finer costation.
Occurrence.—OD3 Unit (Yahagi).
Distribution.—Lower Visean–Serpukhovian: northeasternJapan (Yahagi in the South Kitakami Belt), Belgium andKyrgyzstan.
Superfamily HORRIDONIOIDEA Muir-Wood and Cooper,
1960
Family LEIOPRODUCTIDAE Muir-Wood and Cooper, 1960
Subfamily LEVITUSIINAE Muir-Wood and Cooper, 1960
Genus LEVITUSIA Muir-Wood and Cooper, 1960
Type species.—Productus humerosus Sowerby, 1822.
Levitusia humerosa (Sowerby, 1822)(Fig. 27A)
Productus humerosus Sowerby, 1822, p. 21, fig. 322; Davidson,
1861, p. 147, pl. 36, figs. 1, 2; Yanishevsky, 1918, p. 38, pl.
3, figs. 1–5.
Productus sublaevis de Koninck. Davidson, 1861, p. 177, pl. 31,
figs. 1, 2; pl. 32, fig. 1; pl. 51, figs. 1, 2; Rotai, 1931, p. 50, pl.
1, figs. 7, 8; pl. 2, figs. 1, 5.
Productus (Plicatifera) humerosus Sowerby. Paeckelmann,
1931, p. 120, pl. 6, fig. 5; pl. 7, figs. 1–4; pl. 8, figs. 1–3; pl.

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

39

FIGURE 29. Brachiopods of the AR2 assemblage (3). A–F, Rhipidomella michelini (Léveillé); ventral (A1, A2) and dorsal (A3) views of internal moud ofconjoined shell, IGPS111721; B, internal mould of dorsal valve, IGPS111724; C1, C2, internal mould of dorsal valve, NU-B2256; D1, D2, internal mouldof ventral valve, IGPS112100; E1, E2, internal mould of dorsal valve, IGPS112103; F1, F2, internal mould of dorsal valve, IGPS112104; G, H, Marginatia
burlingtonensis (Hall); G, internal mould of vental valve, NU-B2258; ventral (H1, H2) and dorsal (H3, H4) views of internal mould of conjoined shell,
NU-B2260; I, J, Orthotetes keokuk (Hall); I, external latex cast of ventral valve, IGPS112120; J, internal mould of dorsal valve, IGPS112121. Scale bars
are 1 cm.

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JUN-ICHI TAZAWA

11, fig. 1; Rotai, 1941, p. 96, pl. 17, figs. 3–6.
Productus (Plicatifera) humerosus var. longa Paeckelmann,
1931, p. 128, pl. 9, fig. 1.
Productus (Plicatifera) humerosus var. christiani de Koninck.
Paeckelmann, 1931, p. 131, pl. 10, fig. 1; pl. 12, fig. 4; pl. 13,
fig. 1.
Levitusia humerosa (Sowerby). Muir-Wood and Cooper, 1960,
pl. 109, figs. 2–5; pl. 110, figs. 1–5; Nalivkin and Fotieva,
1973, p. 43, pl. 9, figs. 4, 5: Garanj et al., 1975, p. 181, pl. 78,
fig. 2; pl. 80, fig. 5; Galitskaya, 1977, p. 123, pl. 12, figs. 4–7;
pl. 38, fig. 1; Brunton, 1979, p. 7, pl. 1, figs. 1–6; pl. 2, figs.
1–5; pl. 4, figs. 1–4; Nalivkin, 1979, p. 59, pl. 18, fig. 6;
Tazawa and Kurita, 2019b, p. 194, fig. 3.
Levitusia christiani uralica Fotieva in Nalivkin and Fotieva,
1973, p. 44, pl. 10, figs. 1, 2.
Material.—One specimen from locality KAR11, externalmould of a ventral valve, IGPS111744.
Remarks.—This specimen was described by Tazawa andKurita (2019b, p. 194, fig. 3) as Levitusia humerosa (Sowerby,1822). The specimen, from the middle part of the Arisu
Formation (AR2 Unit) of Okuhinotsuchi in the South Kitakami
Belt, is poorly preserved, but can be referred to L. humerosa(Sowerby, 1822), redescribed by Brunton (1979, p. 7, pl. 1, figs.
1–6; pl. 2, figs. 1–5; pl. 4, figs. 1–4) on the type specimens fromthe lower Visean of Belgium and Staffordshire, England, in thelarge, strongly convex ventral valve (length more than 45 mm;
width more than 35 mm), with long sulcated trail, and theexternal ornament consisting of irregular rugae on visceralportion and several strong longitudinal flutings on trail.
Levitusia sublaevis (de Koninck, 1843), redescribed by Brunton(1979, p. 12, pl. 3, figs. 1–9; pl. 4, figs. 5, 6) from the lower
Visean of Belgium and Lancashire, England, differs from L.
humesosa in having weaker posterior rugation and shallower
ventral sulcus.
Occurrence.—AR2 Unit (Okuhinotsuchi).
Distribution.—Lower Visean: northeastern Japan(Okuhinotsuchi in the South Kitakami Belt), the UK (England),Germany, Belgium, western Russia (Donetz Basin), central
Russia (western and southern Urals), Uzbekistan (Fergana) andKyrgyzstan (northern Tian-Shan).
Levitusia elongata sp. nov.
(Fig. 38F, G)
Etymology.—In reference to elongate outline of the species.
Material.— Four specimens from localities INS7, INS11,
YKA12 and YKA21: (1) external and internal moulds of aconjoined shell, IGPS112155; (2) external and internal moulds
of a ventral valve, IGPS112154 (holotype); and (3) internalmoulds of two ventral valves, IGPS112156, 112157.
Diagnosis.—Medium-sized, elongate Levitusia, with awell-developed ventral sulcus and strong reticulate ornament onventral valve.
Description.—Shell medium in size for genus, elongate

subrectangular in outline, widest at hinge; length about 67 mm,
width about 32 mm in the holotype (IGPS112154). Ventral
valve strongly convex in lateral profile, with a long trail; umbo
small, strongly incurved; ears large, trigonal and nearlyflattened; sulcus narrow and deep. External surface of ventral
valve ornamented with strong rugae and fine costae on visceraldisc, but costae only on trail; numbering 4 rugae in 10 mm, 7–8
costae in 5 mm on visceral disc, and 7 costae in 5 mm on trail.
Internal structures of both valves not well-preserved, although
ventral valve having widely diverging, anterior ridged diductor
scars, and dorsal valve having large trilobate cardinal process
and elevated adductor scars.
Remarks.—Levitusia elongata sp. nov. is distinguished fromthe type species, Levitusia humerosa (Sowerby, 1822), in thesmaller size, less transverse outline and in having
well-developed sulcus on the ventral valve. Levitusia spinauris
Brunton (1979, p. 14, pl. 2, figs. 6–8), from the lower Visean ofStaffordshire, England, differs from the present new species inhaving shallower ventral sulcus, and finer costellae on ventral
valve. Levitusia sublaevis (de Koninck, 1843), redescribed by
Brunton (1979, p. 12, pl. 3, figs. 1–9; pl. 4, figs. 5, 6), from thelower Visean of Belgium (Visé) and, England (Lancashire),differs from L. elongata sp. nov. in its much transverse outline.
Occurrence.—OD2 Unit (Nisawa and Shizu).
Distribution.—Upper Visean: northeastern Japan (Nisawa
and Shizu in the South Kitakami Belt).
Superfamily PRODUCTOIDEA Gray, 1840
Family BUXTONIIDAE Muir-Wood and Cooper, 1960
Subfamily BUXTONIINAE Muir-Wood and Cooper, 1960
Genus TOLMATCHOFFIA Fredericks, 1933
Type species.― Productus robustus Tolmatchoff, 1924.
Tolmatchoffia robusta (Tolmatchoff, 1924)（ Fig. 28E, F）
Productus robustus Tolmatchoff, 1924, p. 230, 572, pl. 13, fig.
17; pl. 14, figs. 1–3.
Dictyoclostus robustus (Tolmatchoff). Benediktova, 1955, p.
309, pl. 85, fig. 6; Yang, 1964, p. 96, pl. 9, figs. 6, 7.
Tolmatchoffia robusta (Tolmatchoff). Muir-Wood and Cooper,
1960, pl. 95, figs. 1–3; Sarytcheva in Sarytcheva et al., 1963,
p. 171, pl. 21, figs. 1–3; pl. 22, figs. 1–3, text-figs. 71, 72;
Galitskaya, 1977, p. 74, pl. 34, fig. 3; Zhang et al., 1983, p. 302,
pl. 118, figs. 4, 5; Tazawa, 2018c, p. 44, figs. 17A‒F, 22G‒J.
Buxtonia sp. Tazawa, 1989, p. 60, pl. 1, figs. 6–9.
Material.—Five specimens from locality KF179 and KF181:(1) internal mould of a conjoined shell with external mould of
the dorsal valve, IGPS112122; (2) external and internal moulds
of a ventral valve, IGPS112097; (3) external and internalmoulds of two dorsal valves, IGPS112098, 112123; and (4)external mould of a dorsal valve, IGPS112099.
Remarks.—These specimens are referred to Tolmatchoffia

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

41

FIGURE 30. Brachiopods of the AR2 assemblage (4). A, Schizophoria woodi Bond, ventral (A1), dorsal (A2), anterior (A3), posterior (A4) and lateral (A5,
A6) views of internal mould of conjoined shell, IGPS111728; B–E, Schizophoria pinguis Demanet; ventral (B1), anterior (B2) and posterior (B3) views ofinternal mould of dorsal valve, IGPS111730; C, internal mould of ventral valve, IGPS112085; dorsal (D1), anterior (D2), posterior (D3) and lateral (D4)views of internal mould of dorsal valve, IGPS112087; external latex cast (E1) and internal mould (E2) of dorsal valve, IGPS111729; F– I, Schizophoria
resupinata (Martin); F, internal mould of ventral valve, IGPS112106; ventral (G 1 ) and dorsal (G 2 ) views of internal mould of conjoined shell,
IGPS111716; ventral (H1), dorsal (H2), anterior (H3), posterior (H4) and lateral (H5) views of internal mould of conjoined shell, IGPS111715; I, internalmould of ventral valve, IGPS112105. Scale bars are 1 cm.

42

JUN-ICHI TAZAWA

robusta (Tolmatchoff, 1924), redescribed by Sarytcheva in
Sarytcheva et al. (1963, p. 171, pl. 21, figs. 1–3; pl. 22, figs.
1–3, text-figs. 71, 72) from the lower and upper Tournaisian of
the Kuznetsk Basin, central Russia, in the large size (length 45mm, width 68 mm in the ventral valve specimen, IGPS112097),
distinct sulcus and numerous costae (numbering 7 in 5 mm in
the midlength) on the ventral valve. Tolmatchoffia barzassica
Sarytcheva in Sarytcheva et al. (1963, p. 174, pl. 23, figs. 1, 2,
text-fig. 73), from the lower and upper Tournasian of theKuznetsk Basin, differs from T. robusta in its smaller size and inhaving no ventral sulcus.
Occurrence.—AR2 Unit (Yokota).
D i s t r i b u t i o n . — L o w e r T o u r n a i s i a n –l o w e r V i s e a n :northeastern Japan (Yokota in the South Kitakami Belt), central
Russia (Kuznetsk Basin), Kyrgyzstan and northwestern China(Xinjiang).
Genus TOMIPRODUCTUS Sarytcheva in Sarytcheva,
Sololskaya, Besnossova and Maksimova, 1963
Type species.—Productus elegantulus Tolmatchoff, 1924.
Tomiproductus elegantulus (Tolmatchoff, 1924)(Fig. 28A–D)
Productus elegantulus Tolmatchoff, 1924, p. 244, 579, pl. 14,
figs. 5–7.
Tomiproductus elegantulus (Tolmatchoff). Sarytcheva in
Sarytcheva et al., 1963, p. 202, pl. 31, figs. 1–11; pl. 32,
figs. 1–7, text-figs. 88, 89; Nalivkin and Fotieva, 1973, p. 42,
pl. 8, figs. 12, 13; Kalashnikov, 1974, p. 55, pl. 13, fig. 6;
Bublichenko, 1976, p. 54, pl. 3, fig. 9; pl. 5, figs. 1, 2;
Bahrammanesh et al., 2011, p. 145, fig. 5k-q; Tazawa and
Ibaraki, 2019, p. 13, fig. 4A.
Material.—Seven specimens from localities IYK1, IYK2,
KF179, KF180 and KF181: (1) external and internal moulds of aventral valve, IGPS112115; (2) internal moulds of three ventral
valves, IGPS112116–112118; (3) external mould of a dorsal
valve, IGPS112119; and (4) internal moulds of two ventral
valves, NU-B2262, 2263.
Remarks.—Two specimens (NU-B2262, 2263) from the
middle Arisu Formation of Yokota were described by Tazawa and
Ibaraki (2019, p. 13, fig. 4A) as Tomiproductus elegantulus(Tolmatchoff, 1924). All the other specimens are also referred toTomiproductus elegantulus (Tolmatchoff, 1924), redescribed by
Sarytcheva in Sarytcheva et al. (1963, p. 202, pl. 31, figs. 1–11;
pl. 32, figs. 1–7, text-figs, 88, 89) from the lower–upper
Tournaisian of the Kuznetsk Basin, central Russia, in their small
size (length 10 mm, width 17 mm in the largest specimen,
IGPS112115; length 15 mm, width 11 mm in the best preserved
ventral valve specimen, IGPS112117), strongly inflated ventral
valve with a long trail, dorsal valve with flat disc, strongly
geniculated and followed by a long trail, numerous fine costellae
on entire surface of both valves (10–12 in 5 mm on ventral trail),

and weak concentric rugae on discs of both valves.
Tomiproductus minimus (Demanet, 1921), redescribed by
Nalivkin (1979, p. 91, pl. 13, fig. 26; pl. 31, figs. 1–11) from theupper Tournaisian–middle Visean of the northern Urals, is also asmall-sized Tomiproductus, but it differs from T. elegantulus inhaving stronger rugae on dorsal disc. Tomiproductus kollari
Carter (1990, p. 224, figs. 3.1–3.14), from the middle–upper
Keokuk Limestone of Missouri, USA, differs from T. elegantulus
in the slightly larger size and coarser rugae on ventral valve.
Occurrence.—AR2 Unit (Yokota).
D i s t r i b u t i o n . — L o w e r T o u r n a i s i a n –u p p e r V i s e a n :northeastern Japan (Yokota in the South Kitakami Belt),
northern Russia (Pechora Basin), Iran, central Russia (western
Urals and Kuznetsk Basin) and Kazakhstan (Altay Mountains).
Genus MARGINICINCTUS Sutton, 1938
Type species.—Productus marginicinctus Prout, 1857.
Marginicinctus marginicinctus (Prout, 1857)(Figs. 38C, D, 44A)
Productus marginicinctus Prout, 1857, p. 43, pl. 2, figs. 1–6;
Weller, 1914, p. 127, pl. 13, figs. 18–23.
Marginicinctus marginicinctus (Prout). Sutton, 1938, p. 63, figs.
5–8; Muir-Wood and Cooper, 1960, pl. 75, figs. 10–12; pl.
76, figs. 1–11, 17–20.
Marginicinctus sp. Tazawa, Itabashi and Mori, 1981, pl. 1, figs.
7–10; Tazawa, 1981b, p. 71, pl. 5, fig. 2.
Marginicinctus cf. marginicinctus (Prout). Tazawa, 1981b, p.
70, pl. 5, fig. 1.
Material.—Four specimens from localities INS3, INS13 andKF210: (1) internal mould of a conjoined shell, with externalmould of the dorsal valve, IGPS97656; and (2) internal mouldof three ventral valves, IGPS97657, 97658, NU-B2330.
Remarks.—Most of the specimens from Nisawa werepreviously figured by Tazawa et al. (1981, pl. 1, figs. 7–10) andTazawa (1981b, p. 71, pl. 5, fig. 2) as Marginicinctus sp., anddescribed by Tazawa (1981b, p. 70, pl. 5, fig. 1) as
Marginicinctus cf. marginicinctus (Prout, 1857). But the Nisawa
specimens are referred to Marginicinctus marginicinctus (Prout,1857), redescribed and refigured by Weller (1914, p. 127, pl. 13,
figs. 18–23) from the St. Louis Limestone of Missouri, in size,
shape and external ornament of the both ventral and dorsal
valves. In the present article a ventral valve specimen(NU-B2330), from the uppermost part of the Odaira Formation
at Yukisawa, is added to the material. Marginicinctus reticulatus
Roberts (1976, p. 66, pl. 11, figs. 1–12), from the upper Viseanof eastern Australia, differs from M. marginicinctus in the larger
size and the stronger rugae on the both ventral and dorsal valves.
Occurrence.—OD2 Unit (Nisawa) and OD3 Unit (Yahagi).
Distribution.—Upper Visean: northeastern Japan (Yahagi andNisawa in the South Kitakami Belt) and USA (Iowa and Missouri).

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS43

FIGURE 31. Brachiopods of the AR2 assemblage (5). A, Cleiothyridina harkeri Carter, external latex cast (A1, A2) and internal mould (A4) of dorsal
valve, and internal mould (A3) of ventral valve of conjoined shell, NU-B2265; B–D, Unispirifer striatoconvolutus (Dun and Benson); ventral (B1) andposterior (B2) views of internal mould of ventral valve, IGPS111733; external latex cast of dorsal valve (C1) and posterior view of internal mould (C2) ofconjoined shell, IGPS112094; D, external latex cast of dorsal valve, IGPS112095; E, Unispirifer sp., ventral view (E1, E2) of internal mould of ventral
valve, IGPS111734; F–I, Unispirifer kozuboensis (Minato); ventral (F 1), dorsal (F 2) and posterior (F 3) views of internal mould of conjoined shell,
IGPS112091; G, internal mould of dorsal valve, IGPS112113; H, internal mould of dorsal valve, IGPS112093; I, internal mould of dorsal valve,
IGPS112127. Scale bars are 1 cm.

44

JUN-ICHI TAZAWA

Subfamily MARGINATIINAE Waterhouse, 2002
Genus MARGINATIA Muir-Wood and Cooper, 1960
Type species.—Productus fernglenensis Weller, 1909.
Marginatia burlingtonensis (Hall, 1858)(Figs. 29G, H, 36B, C, 44B–D)
Productus flemingi var. burlingtonensis Hall, 1858, p. 598, pl.
12, fig. 3.
Productus burlingtonensis Hall. Weller, 1914, p. 104, pl. 9, figs.
1–10; Frech, 1916, p. 239, pl. 6, fig. 1; Tolmatchoff, 1924, p.
237, 575, pl. 14, figs. 8–11; Girty, 1929, p. 85, pl. 9, figs.
20–24.
Productus (Productus) burlingtonensis Hall. Nalivkin, 1937, p.
66, pl. 7, figs. 7–11.
Productus sp. Minato, 1951, p. 366, pl. 1, fig. 4.
Productus (Dictyoclostus) burlingtonensis Hall. Simorin, 1956,
p. 136, pl. 9, figs. 1–3.
Marginatia burlingtonensis (Hall). Sarytcheva in Sarytcheva etal., 1963, p. 191, pl. 28, figs. 5–8, text-figs. 81, 82; Grechishnikova,
1966, p. 116, pl. 8, figs. 11–13; Aksenova in Litvinovich etal., 1969, p. 213, pl. 35, figs. 2– 4; Nalivkin and Fotieva,
1973, p. 39, pl. 8, fig. 1; Bublichenko, 1976, p. 50, pl. 2, fig.
12; pl. 4, fig. 6; pl. 5, figs. 4–6; pl. 6, fig. 9; Galitskaya, 1977,
p. 83, pl. 22, figs. 6–10; Nalivkin, 1979, p. 94, pl. 32, figs.
1–10; pl. 34, figs. 3, 4; Lee et al., 1980, p. 368, pl. 148, fig.
10; Jin, 1985, p. 77, pl. 1, figs. 20–22; Carter, 1987, p. 39, pl.
9, figs. 1–8; Shi et al., 2005, p. 44, figs. 5D, I–K, M; Tazawa,
2006, p. 132, figs. 6.1–6.8; Tazawa, 2017, p. 335, figs.
6.3–6.5; Tazawa, 2018c, p. 44, figs. 23A–D; Tazawa and
Ibaraki, 2019, p. 14, figs. 4C, D.
Dictyoclostus sp. Hase and Yokoyama, 1975, pl. 18, fig. 1.
Marginatia sp. Tazawa, 1985, p. 459, figs. 2.3–2.7; Tazawa,
1989, p. 60, pl. 1, fig. 1; Tazawa, 2002, figs. 7.1, 7.2.
Material.—Ten specimens from localities IYK2, IYK3,KF159, KF207, KF208 and KYT3: (1) external and internalmoulds of a ventral valve, NU-B630; (2) external mould of two
ventral valves, IGPS112132, NU-B2160; (3) internal moulds offour ventral valves, NU-B650, 2161, 2258, 2259; (4) external
and internal moulds of a dorsal valve, NU-B2260; and (5)external moulds of two dorsal valves, NU-B2162, 2261.
Remarks.—Most of the specimens were described by
Tazawa (2006, 2017) and Tazawa and Ibaraki (2019)as Marginatia burlingtonensis (Hall). These specimens are
fragmentarily preserved or severely squashed, but can bereferred to Marginatia burlingtonensis (Hall, 1858), from the
Burlington Limestone of Illinois and Iowa, by the medium size(length about 20 mm, width about 46 mm in the best preserved
ventral valve specimen, NU-B2160), strongly geniculated
ventral valve, reticulate ornament on visceral discs of both
valves, and two symmetrically arranged strong halteroid spines
on ventral trail. Marginatia patersonensis Roberts (1965, p. 63,
pl. 10, figs. 1–5), from the lower and upper Visean of New

South Wales, eastern Australia, differs from the present species
in having finer costae and larger number of spines on the ventral
valve. The type species, Marginatia fernglenensis Weller (1909,
p. 299, pl. 12, figs. 14–17) from the Fern Glen Formation of
Missouri, differs from M. burlingtonensis in having shallower
ventral sulcus.
Occurrence.—AR2, AR3 and OD3 units (Yokota).
Distribution.—Upper Tournaisian–upper Visean: northeasternJapan (Hikoroichi and Yokota in the South Kitakami Belt),southwestern Japan (Hina in the Akiyoshi Belt), USA (Illinois,
Iowa and Arkansas), western Canada (Alberta), Turkey (Taurus
Mountains), central Russia (southern Urals and Kuznetsk
Basin), Kazakhstan, Kyrgyzstan and northeastern China(Liaoning).
Superfamily ECHINOCONCHOIDEA Stehli, 1954
Family ECHINOCONCHIDAE Stehli, 1954
Subfamily ECHINOCONCHINAE Stehli, 1954
Genus ECHINOCONCHUS Weller, 1914
Type species.—Productus punctatus Sowerby, 1822.
Echinoconchus punctatus (Sowerby, 1822)(Fig. 44G, H)
Productus punctatus Martin. Sowerby, 1822, p. 22, pl. 323,
lower right figure; Davidson, 1861, p. 172, pl. 44, figs. 9–11,
16, 17.
Pustula punctata (Martin). Thomas, 1914, p. 303, pl. 17, figs.
16–19, text-fig. 11; Tolmatchoff, 1924, p. 256, 584, pl. 16,
fig. 9; Rotai, 1931, p. 58, pl. 4, figs. 1, 11.
Productus (Pustula) punctatus (Martin). Yanishevsky, 1918, p.
47, pl. 3, figs. 7, 9.
Echinoconchus punctatus (Martin). Chao, 1927, p. 67, pl. 6,
figs. 7, 8, 15, 16; Sarytcheva in Sarytcheva and Sokolskaya,
1952, p. 103, pl. 18, fig. 120; Dedok and Tschernjak, 1960, p.
53, pl. 1, fig. 6; Pareyn, 1961, p. 197, pl. 23, figs. 1–4; Ding
in Yang et al., 1962, p. 51, pl. 19, figs. 1–4; Yang, 1964, p.
81, pl. 4, figs. 5, 6, 9, 10, text-fig. 7; Abramov, 1965, p. 38,
pl. 3, fig. 2; Aksenova in Litvinovich et al., 1969, p. 164, pl.
9, figs. 5, 6; pl. 10, fig. 1; Abramov, 1970, p. 117, pl. 9, fig. 4;
Alexandrow and Solomina, 1973, p. 93, pl. 22, figs. 1– 3;Volgin and Kushnar, 1975, p. 46, pl. 4, fig. 1; Donakova,
1978, p. 208, pl. 1, figs. 5, 6; Nalivkin, 1979, p. 78, pl. 24,
figs. 8, 9; Zhang et al., 1983, p. 288, pl. 127, fig. 111; pl. 128,
fig. 2; Jin et al., 1985, p. 192, pl. 9, figs. 11, 12; Zhan andWu, 1987, p. 207, pl. 48, fig. 38; Archbold and Stojanović-Kuzenko,
1995, pl. 62, fig. 10; Wang and Yang, 1998, p. 77, pl. 9, figs.
17, 18.
Productus (Echinoconchus) punctatus (Martin) em. Thomas.
Paeckelmann, 1931, p. 152, pl. 15, figs. 7–10.
Productus (Echinoconchus) punctatus (Martin). Nalivkin, 1937,
p. 64, pl. 9, fig. 5.
Echinoconchus punctatus (Sowerby). Muir-Wood, 1951, p. 102,
pl. 4, fig. 2; Muir-Wood and Cooper, 1960, pl. 66, figs. 1, 2;

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS45

FIGURE 32. Brachiopods of the AR2 assemblage (6). A–C, Acuminothyris triangularis Roberts; A1, A2, internal mould of dorsal valve, IGPS112114; B,internal moud of dorsal valve, IGPS112128; C, internal mould of dorsal valve, IGPS112129; D, E, Brachythyris chouteauensis (Weller); D1, D2, internalmould of ventral valve, UHR16252; E, internal mould of dorsal valve, UHR16012; F–I, Kitakamithyris hikoroitiensis Minato; external latex cast of dorsal
valve (F1), internal mould of ventral valve (F2) and internal mould of dorsal valve (F3) of conjoined shell, IGPS112107; G, external latex cast of dorsal
valve, IGPS112109; external latex cast (H1) and external mould (H2) of ventral valve, IGPS111727; external mould (I1, I2) and enlarged concentric rows ofbiramous spine bases (I3) of external mould of dorsal valve, NU-B2257. Scale bars are 1 cm, except for I3.

46

JUN-ICHI TAZAWA

pl. 82, figs. 8–10; pl. 83, figs. 1–4; pl. 88, fig. 11; pl. 125, fig.
5; Winkler Prins, 1968, p. 89, pl. 3, figs. 12–14; Nalivkin andFotieva, 1973, p. 35, pl. 6, fig. 8; Kalashnikov, 1974, p. 48,
pl. 9, figs. 1–3; Martinez Chacon and Legrand-Blain, 1992, p.
110, pl. 3, figs. 15–18; Tazawa, 2017, p. 335, figs. 6.6, 6.7;
Tazawa, 2018c, p. 46, figs. 23G, H.
Echinoconchus (Echinoconchus) punctatus (Sowerby). Galitskaya,
1977, p. 62, pl. 16, figs. 1–5; pl. 18, fig. 1, text-fig. 7; Kalashnikov,
1980, p. 34, pl. 5, fig. 1.
Echinoconchus aohanensis Lee et al., 1980, p. 363, pl. 147, figs.
1, 2.
Material.—Two specimens from KF159: (1) external andinternal moulds of a ventral valve, NU-B2155; and (2) external
and internal moulds of a dorsal valve, NU-B2156.
Remarks.—These specimens were described by Tazawa(2017, p. 335, figs. 6.6, 6.7) as Echinoconchus punctatus(Sowerby, 1822). The specimens from Tairagai are referred toE. punctatus (Sowerby, 1822), refigured by Muir-Wood andCooper (1960, pl. 66, figs. 1, 2; pl. 82, figs. 8–10; pl. 83, figs.
1–4; pl. 88, fig. 11; pl. 125, fig. 5) from the Visean of the UK(Scotland and England) and Belgium, by the large size (length
about 62 mm, width about 82 mm in the larger specimen,
NU-B2155), regular broad bands on the ventral valve,prominent bilobate cardinal process and elongate smooth
adductor scars in the dorsal valve. The smaller dorsal valve
specimen may be a young shell. Anomites punctatus, describedby Martin (1809), were declared nomenclatorially invalid in
1948 by the International Commission on Zoological
Nomenclature, and the species have been referred to subsequentauthors, Sowerby, 1822 (Muir-Wood and Cooper, 1960, p. 243).
Echinoconchus aohanensis Lee, Gu and Su (1980, p. 363, pl.
147, figs. 1, 2), from the lower Pennsylvanin of Liaoning,northeastern China, is regarded as a junior synonym of thepresent species. Echinoconchus alternatus (Norwood andPratten, 1855), redescribed by Weller (1914, p. 138, pl. 17, figs.
1–7), from the Osagean and Meramecian of the Mississippi
Valley, USA, differs from E. punctatus in having longer hinge.
Occurrence.—OD3 Unit (Yokota).
Distribution.—Upper Tournaisian–lower Permian (Asselian):
northeastern Japan (Hikoroichi and Yokota in the South
Kitakami Belt), northern Russia (Verkhoyansk Range, Taimyr
Peninsula and northern Urals), UK (Scotland, England andWales), Germany, Belgium, Spain, western Russia (Moscow
Basin), central Russia (southern Urals), Kyrgyzstan,northwestern China (Xinjiang and Qinghai) and northern China(Shanxi).
Genus ECHINARIA Muir-Wood and Cooper, 1960
Type species.—Productus semipunctatus Shepard, 1838.
Echinaria sp.
(Fig. 45C)

Echinaria sp. Tazawa, 2017, p. 336, fig. 7.2.
Material.—One specimen from locality KF159, external andinternal moulds of a dorsal valve, NU-B2157.
Remarks.—The single dorsal valve specimen from Tairagai
was described by Tazawa (2017, p. 336, fig. 7.2) as Echinaria
sp. This pecimen can be safely assigned to the genus Echinaria
by its external ornament consisting of closely-spaced regularbands with numerous spine bases of two series, larger posterior
one and smaller anterior one and elongate adductor scars ofdendritic posterior one and smooth anterior one. The Kitakami
species is large in size (length more than 60 mm, width morethan 55 mm), and closely resembles two species, Echinaria
semipunctata (Shepard, 1838), redescribed by Dunbar andCondra (1932, p. 205, pl. 24, fig. 6; pl. 25, figs. 1–3), from theupper Pennsylvanian of Nebraska, and Echinaria rara
(Nasikanova in Sarytcheva, 1968, p. 91, pl. 6, figs. 11– 16,
text-figs. 33–35), from the Bashkirian of eastern Kazakhstan, in
size and shape of the dorsal valve, although accurate comparison
is difficult owing to ill-preservation of the present material.
Echinaria minatoi Tazawa (1981a, p. 54, pl. 4, figs. 3–11), fromthe lower part of the Nagaiwa Formation (upper Visean–
Serpukhovian) of the Hikoroichi area, South Kitakami Belt, isreadily distinguished from the Tairagai species by its muchsmaller dimensions.
Occurrence.—OD3 Unit (Yokota).
Distribution.—Upper Visean: northeastern Japan (Yokota in
the South Kitakami Belt).
Family WAAGENOCONCHIDAE Muir-Wood and Cooper,
1960
Subfamily PUSTULINAE Waterhouse, 1981
Genus PUSTULA Thomas, 1914
Type species.—Producta pustulosa Phillips, 1836.
Pustula pustulosa (Phillips, 1836)(Fig. 44E, F)
Producta pustulosa Phillips, 1836, p. 216, pl. 7, fig. 15.
Productus pustulosus Phillips. Krenkel, 1913, p. 18, 43, pl. 1,
fig. 7.
Pustula pustulosa (Phillips). Thomas, 1914, p. 261, pl. 17, figs.
24–28; pl. 18, fig. 1; Muir-Wood and Cooper, 1960, pl. 59,
fig. 4; pl. 84, figs. 1– 7; pl. 85, figs. 6– 10; Nalivkin andFotieva, 1973, p. 36, pl. 6, figs. 11–13; Kalashnikov, 1974, p.
51, pl. 10, figs. 6–9; pl. 11, figs. 1, 2; pl. 13, figs. 4, 5; pl. 30,
fig. 6; Garanj et al., 1975, p. 163, pl. 65, figs. 3, 4; Volgin andKushnar, 1975, p. 45, pl. 3, figs. 10–12; Galitskaya, 1977, p.
67, pl. 19, figs. 1–5; pl. 20, fig. 1; Zhang et al., 1983, p. 289,
pl. 109, fig. 4; Tazawa, 2017, p. 336, figs. 7.3, 7.4; Tazawa,
2018c, p. 47, fig. 30F.
Productus (Pustula) pustulosus Phillips emend. Thomas.
Paeckelmann, 1931, p. 138, pl. 13, figs. 2, 3.
Pustula cf. pustulosa (Phillips). Mori and Tazawa, 1980,

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS47

FIGURE 33. Brachiopods of the AR2 assemblage (7). A, B, Syringothyris texta (Hall); ventral (A1), dorsal (A2), anterior (A3) and posterior (A4) views ofinternal mould, and external latex cast of ventral interarea (A5) of conjoined shell, UHR16925; posterior (B1) and ventral (B2) views of internal mould ofventral valve, IGPS111710. Scale bars are 1 cm.

text-fig. 3.2; Tazawa, 1984b, p. 306, pl. 61, fig. 8.
Material.—Two specimens from KF159: (1) external andinternal moulds of a ventral valve, NU-B2158; and (2) external
and internal moulds of a dorsal valve, NU-B2159.
Remarks.—The fragmentary specimens from Tairagai were
described by Tazawa (2017, p. 336, figs. 7.3, 7.4) as Pustula
pustulosa (Phillips, 1836). The Tairagai specimens are referred
to P. pustulosa (Phillips, 1836), redescribed by Thomas (1914,
p. 261, pl. 17, figs. 24–28; pl. 18, fig. 1) from the upper Viseanof England, by the large size (length more than 47 mm, width
more than 42 mm in the ventral valve specimen, NU-B2158),

subrectangular outline, well-developed ventral sulcus and dorsal
fold, and external ornament of both valves consisting of regularconcentric bands and numerous, quincunxially arranged
elongate spine bases. Pustula cf. pustulosa (Phillips, 1836),
figured by Mori and Tazawa (1980, text-fig. 3.2) andsubsequently described by Tazawa (1984b, p. 306, pl. 61, fig. 8)
from the upper part of the Hikoroichi Formation of the
Hikoroichi area, South Kitakami Belt, is regarded as a synonymof the present species. Pustula tenuipustulosa Thomas (1914, p.
288, pl. 20, figs. 10, 11, text-fig. 7) differs from P. pustulosa inhaving smaller spine bases and less-marked bands.
Occurrence.—OD3 Unit (Yokota).

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JUN-ICHI TAZAWA

Distribution.—Lower Visean–Serpukhovian: northeasternJapan (Hikoroichi and Yokota in the South Kitakami Belt),
northern Russia (Pechora Basin), UK (England and northern
Ireland), Germany, central Russia (western and southern Urals),
Uzbekistan (Fergana), Kyrgyzstan and northwestern China(Xinjiang).
Superfamily LINOPRODUCTOIDEA Stehli, 1954
Family LINOPRODUCTIDAE Stehli, 1954
Subfamily LINOPRODUCTINAE Stehli, 1954
Genus OVATIA Muir-Wood and Cooper, 1960
Type species.—Ovatia elongata Muir-Wood and Cooper,
1960.
Ovatia elongata Muir-Wood and Cooper, 1960
(Fig. 28G)
Ovatia elongata Muir-Wood and Cooper, 1960, p. 312, pl. 114,
figs. 1–4, 7, 11, 12: Jin et al., 1979, p. 93, pl. 28, figs. 8, 9,
11; Zhang et al., 1983, p. 297, pl. 108, fig. 9; Zong et al.,2012, p. 423, pl. 1, figs. 41–43; Tazawa and Iryu, 2019, p. 99,
fig. 6H.
Material.—Two specimens from locality IST1, externalmoulds of two dorsal valves, IGPS111725, 111726.
Remarks.—These specimens were described by Tazawa and
Iryu (2019, p. 99, fig. 6H) as Ovatia elongata Muir-Wood andCooper, 1960. The specimens from Shimoarisu are referred toO. elongata Muir-Wood and Cooper (1960, p. 312, pl. 114, figs.
1– 4, 7, 11, 12), from the lower Fayetteville Formation of
Oklahoma, on account of the external characters, the large,
elongate shell (length 43 mm, width 24 mm in the larger dorsal
valve specimen, IGPS111725) and external ornament of thedorsal valve consisting of numerous fine costellae and irregularrugae, numbering 12–13 costellae in 5 mm at midlength. Ovatia
ovata (Hall, 1858), redescribed by Sarytcheva (1937, p. 72, 110,
pl. 6, fig. 5) from the upper Visean of the Moscow Basin,
western Russia, differs from O. elongata in the smaller and lesselongate shell and the finer costellae on the dorsal valve. Ovatia
laevicosta (White, 1860), redescribed by Carter (1999, p. 109,
figs. 4I–N) from the St. Joe Formation of Oklahoma, differsfrom O. elongata in having coarser costellae on both ventral
and dorsal valves. Ovatia nascens Carter (1988, p. 42, figs.
7.32–7.37), from the Glen Park Formation of Illinois, differsfrom the present species in having finer costellae.
Occurrence.—AR2 Unit (Shimoarisu).
Distribution.—Famennian–upper Visean: northeastern Japan(Shimoarisu in the South Kitakami Belt), USA (Oklahoma) andnorthwestern China (Xinjiang and Qinghai).
Genus FLUCTUARIA Muir-Wood and Cooper, 1960
Type species.—Productus undatus Defrance, 1826.

Fluctuaria undata (Defrance, 1826)(Figs. 38E, 45D)
Productus undatus Defrance, 1826, p. 354; Koninck, 1847, p.
59, pl. 5, fig. 3; Davidson, 1861, p. 161, pl. 34, figs. 10, 11
only; Gröber, 1909, p. 374, pl. 1, fig. 6; Yanishevsky, 1918,
p. 45, pl. 5, fig. 10; pl. 8, fig. 12.
Striatifera undata (Defrance). Chao, 1927, p. 114, pl. 9, figs.
7–9.
Productus (Linoproductus) undatus Defrance. Paeckelmann,
1931, p. 217, pl. 19, fig. 9; Rotai, 1941, p. 99, pl. 20, fig. 3;
Gladchenko, 1955, p. 14, pl. 5, figs. 2, 3.
Productus (Thomasella) undatus Defrance. Nalivkin, 1937, p.
64, pl. 10, fig. 16.
Cancrinella undata (Defrance). Sarytcheva, 1937, p. 82, 111, pl.
7, figs. 1‒6, text-figs. 18‒20; Sarytcheva in Sarytcheva and
Sokolskaya, 1952, p. 110, pl. 20, fig. 136; Lapina, 1957, p.
40, pl. 3, figs. 6–8; Yang, 1964, p. 85, pl. 5, fig. 3.
Fluctuaria undata (Defrance). Muir-Wood and Cooper, 1960,
pl. 115, figs. 11‒20: Sarytcheva in Sarytcheva et al., 1963, p.
229, pl. 37, figs. 6‒11, text-figs. 101, 102; Abramov, 1965, p.
43, pl. 4, fig. 12; Nalivkin and Fotieva, 1973, p. 47, pl. 11,
figs. 1, 2; Donakova, 1978, p. 212, pl. 1, figs. 11, 12;
Martinez Chacon, 1979, p. 223, pl. 25, figs. 8–11; Chen andShi, 2003, p. 154, pl. 5, figs. 3, 16.
Productus-Cancrinella-undatus Defrance. Pareyn, 1961, p. 205,
pl. 25, fig. 10.
Fluctuaria cf. undata (Defrance). Tazawa et al., 1981, pl. 1,
figs. 11–13; Tazawa, 1981b, p. 72, pl. 5, fig. 7.
Material.—Two specimens from localities INS2 and KF210:(1) internal mould of a conjoined shell, with external mould of
the dorsal valve, IGPS97659; and (2) external mould of a ventral
valve, NU-B2331.
Remarks.—The better preserved specimen (IGPS97659) was
previously described by Tazawa (1981b, p. 72, pl. 5, fig. 7) as
Fluctuaria cf. undata (Defrance, 1826). But the Nisawa species
is referred to Fluctuaria undata (Defrance, 1826), refigured by
Muir-Wood and Cooper (1960, pl. 115, figs. 11–20) from theupper Visean of Belgium, in size, shape and external ornamentof the ventral and dorsal valves, particularly the small-sizedshell, and strongly and somewhat irregularly developed
concentric rugae on both valves. Fluctuaria campbelli Roberts(1964, p. 184, pl. 4, figs. 5–13), from the middle Visean ofGreenhills, New South Wales, eastern Australia, differs from F.
undata in having more numerous finer costellae on the ventral
valve. The shells described by Yang (1978, p. 113, pl. 30, fig. 3)
as Fluctuaria undata (Defrance, 1826) from the Serphkhovian
of Guizhou, southwestern China, are not referred to F. undata inbeing much transverse outline.
Occurrence.—OD2 Unit (Nisawa) and OD3 Unit (Yahagi).
Distribution.—Lower Visean– lower Bashkirian:northeastern Japan (Yahagi and Nisawa in the South KitakamiBelt), central Japan (Hida Gaien Belt), northern Russia(Verkhoyansk Range and northern Urals), UK (Scotland,

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS49

FIGURE 34. Brachiopods of the AR2 assemblage (8). A, B, Syringothyris platypleura Weller; ventral (A1), posterior (A2) and lateral (A3) views ofinternal mould of ventral valve, IGPS111711; ventral (B1) and posterior (B2) views of internal mould of ventral valve, IGPS111712; C, Syringothyris texta(Hall), ventral (C1) and dorsal (C2) views of external latex cast, and ventral (C3), dorsal (C4), anterior (C5) and posterior (C6) views of internal mould ofconjoined shell, IGPS112110. Scale bars are 1 cm.

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JUN-ICHI TAZAWA

England, Isle of Man and northern Ireland), Germany, Belgium,
Spain, Algeria, western Russia (Moscow Basin), central Russia(southern Urals and Kuznetsk Basin), Kazakhstan, Uzbekistan,
Kyrgyzstan and northwestern China (Xinjiang and Gansu).
Subfamily STRIATIFERINAE Muir-Wood and Cooper, 1960
Genus STRIATIFERA Chao, 1927
Type species.—Mytilus striatus Fischer de Waldheim, 1837.
Striatifera angusta (Yanishevsky, 1910)(Fig. 40A)
Productus striatus (Fischer de Waldheim). Gröber, 1909, p. 369,
pl. 1, fig. 4; Yanishevsky, 1918, p. 29, pl. 1, fig. 8; pl. 6, fig. 8.
Productus striatus var. angusta Yanishevsky, 1910, p. 43, pl. 4,
figs. 3, 6.
Striatifera striata var. angusta Yanishevsky. Sarytcheva, 1937,
p. 37, pl. 1, fig. 6.
Striatifera angusta (Yanishevsky). Kalashnikov, 1974, p. 80, pl.
23, fig. 4; pl. 26, fig. 4; Galitskaya, 1977, p. 138, pl. 50, figs.
1–6; Jin et al., 1979, p. 93, pl. 28, fig. 3; Zhang et al., 1983, p.
300, pl. 111, fig. 6.
Striatifera cf. striata (Fischer de Waldheim). Tazawa et al.,
1981, pl. 1, figs. 14–16.
Striatifera striata (Fischer de Waldheim). Tazawa, 1981b, p. 73,
pl. 5, fig. 6.
Material.—One specimen from locality INS12, externalmould of a conjoined shell, IGPS97660.
Remarks.—This specimen was figured by Tazawa et al.
(1981, pl. 1, figs. 14–16) as Striatifera cf. striata (Fischer deWaldheim, 1837), and described by Tazawa (1981b, p. 73, pl. 5,
fig. 6) as Striatifera striata (Fischer de Waldheim, 1837). But
the Nisawa species can be referred to Striatifera angusta(Yanishevsky, 1910), redescribed by Kalashnikov (1974, p. 80,
pl. 23, fig. 4; pl. 26, fig. 4) from the upper Visean–
Serpukhovian of the northern Urals, northern Russia, by theelongate subcyrindrical ventral valve with very small ears. The
type species, Striatifera striata (Fischer de Waldheim, 1837),
redescribed by Sarytcheva (1937, p. 24, 107, pl. 1, figs.
1– 5; pl. 2, figs. 1– 3, text-figs. 4– 7) from the upper
Visean–Serpukhovian of the Moscow Basin, western Russia,
differs from S. angusta in the larger, wider and flattened ventral
valve with larger ears.
Occurrence.—OD2 Unit (Nisawa).
Distribution.—Upper Visean–Serpukhovian: northeasternJapan (Nisawa in the South Kitakami Belt), northern Russia(northern Urals), western Russia (Moscow Basin), Uzbekistan,
Kyrgyzstan and northwestern China (Xinjiang and Qinghai).
Order ORTHOTETIDA Waagen, 1884
Suborder ORTHOTETIDINA Waagen, 1884
Superfamily ORTHOTETOIDEA Waagen, 1884
Family ORTHOTETIDAE Waagen, 1884

Genus ORTHOTETES Fischer de Waldheim, 1829
Type species.—Orthotetes radiata Fischer de Waldheim, 1850.
Orthotetes keokuk (Hall, 1858)(Figs. 29I, J, 39H, I)
Orthis keokuk Hall, 1858, p. 640, pl. 19, fig. 5.
Orthotetes keokuk (Hall). Weller, 1914, p. 75, pl. 7, figs. 1–4;
Nalivkin, 1937, p. 40, pl. 3, figs. 1, 2; Minato, 1952, p. 151,
pl. 11, figs. 1, 2, 4; Simorin, 1956, p. 94, pl. 1, figs. 12, 13;
Armstrong, 1962, p. 45, pl. 6, figs. 18–24; Litvinovich, 1962,
p. 181, pl. 1, fig. 6; pl. 2, fig. 1; Abramov, 1965, p. 36, pl. 2,
fig. 6; pl. 3, fig. 1; Litvinovich in Litvinovich et al., 1969, p.
135, pl. 3, fig. 5; pl. 4, figs. 1–4, text-fig. 40; Abramov, 1970,
p. 109, pl. 2, figs. 4, 5; pl. 3, fig. 1; Tazawa, 2018c, p. 52,
figs. 25D, 30H.
Derbyia depressa var. transversa Minato, 1952, p. 153, pl. 8,
fig. 1.
Orthotetes cf. australis (Campbell). Tazawa, 1981b, p. 69, pl. 5,
fig. 8.
Orthotetes sp. Tazawa, 1984b, p. 305, pl. 61, fig. 11.
Material.—Six specimens from localities INS7, KF179,
KF181, YKA20 and YUK1, and from Otsubosawa, Yokota-cho(exact location unknown): (1) external and internal moulds of aventral valve, IGPS97655; (2) external mould of two ventral
valves, IGPS112120, KCG69; (3) external and internal moulds
of a dorsal valve, IGPS112162; (4) nxternal mould of a ventral
valve, UHR15970: and (5) internal mould of a dorsal valve,
IGPS112121.
Remarks.—One of the specimens (IGPS97655) was
described by Tazawa (1981b, p. 69, pl. 5, fig. 8) as Orthotetes
cf. australis (Campbell, 1957). But all the specimens available
are referred to Orthotetes keokuk (Hall, 1858), redescribed by
Weller (1914, p. 75, pl. 7, figs. 1–4) from the Keokuk
Limestone of the Mississippi Valley, in the large size (length
about 75 mm, width about 104 mm in the largest specimen,
IGPS112162), transverse outline and external ornament (7–8
costellae in 5 mm at anterior margin) of the dorsal valve,
particularly, being widest at about midlength. Orthotetes
australis (Campbell, 1957, p. 45, pl. 11, figs. 1–7), from theupper part of the lower Burindi Group of Babbinboon, NewSouth Wales, eastern Australia, differs from O. keokuk in thewidest part at or slightly anterior to hinge. The type species,
Orthotetes radiata Fischer de Waldheim, 1850, redescribed by
Sokolskaya (in Sarytcheva and Sokolskaya, 1952, p. 52, pl. 7,
fig. 39) from the Moscovian of the Moscow Basin, western
Russia, is distinguished from O. keokuk by the smaller size andless transverse outline. A ventral valve specimen, described by
Minato (1952, p. 153, pl. 8, fig. 1) as Derbyia depressa
Demanet var. transversa Minato, 1952 from the Jumonji Stage(= AR2 Unit) of the Arisu Formatin, Otsubosawa, Yokota, isidentical with the present species in all external and internal
characters of the ventral valve.

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

51

FIGURE 35. Brachiopods of the AR2 assemblage (9). A, B, Asyrinxia nipponotrigonalis (Minato); internal mould (A1) and external latex cast (A2) of
dorsal valve, UHR16016; dorsal (B1), posterior (B2) and anterior (B3) views of internal mould of conjoined shell, UHR16018; C, D, Pseudosyrinx
jumonjiensis (Minato); external mould (C1) and external latex cast (C2) of ventral interarea, IGPS111714; external mould (D1) and external latex cast (D2)of conjoined shell, UHR15995. Scale bars are 1 cm.

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JUN-ICHI TAZAWA

Occurrence.—AR2 Unit (Yokota), OD2 Unit (Nisawa and
Shizu) and OD3 Unit (Yahagi).
Distribution.—Upper Tournaisian–upper Visean: northeasternJapan (Hikoroichi, Yahagi, Nisawa and Shizu in the South
Kitakami Belt), USA (Iowa, Missouri, New Mexico andArizona), northern Russia (Verkhoyansk Range) andKazakhstan.
Family PULSIIDAE Cooper and Grant, 1974
Genus SCHELLWIENELLA Thomas, 1910
Type species.—Spirifera crenistria Phillips, 1836.
Schellwienella radialis (Phillips, 1836)(Fig. 45A)
Spirifera radialis Phillips, 1836, p. 220, pl. 11, fig. 5.
Streptorhynchus crenistria var. radialis (Phillips). Davidson,
1861, p. 129, pl. 25, figs. 16–18.
Schuchertella radialis (Phillips). Paeckelmann, 1930, p. 199, pl.
11, figs. 5, 6; pl. 12, figs. 1, 2; pl. 14, fig. 10; Sokolskaya in
Sarytcheva and Sokolskaya, 1952, p. 43, pl. 4, fig. 29; Yang,
1964, p. 63, pl. 2, fig. 1; Zhang et al., 1983, p. 275, pl. 106,
fig. 8; Tazawa, 2017, p. 338, fig. 7.1.
Schellwienella aspis mut. radialiformis Demanet, 1934, p. 85,
pl. 7, figs. 6–12.
Schuchertella aff. radialis (Phillips). Zhang in Yang et al., 1962,
p. 23, pl. 2, fig. 1.
Schellwienella radialis (Phillips). Brunton, 1968, p. 42, pl. 6,
figs. 13–24, text-figs. 27–35; Nalivkin, 1979, p. 21, pl. 4,
fig. 4; Tazawa. 2017, p. 338, fig. 7.1.
Material.—One specimen from locality KF159, external andinternal moulds of a ventral valve, NU-B2152.
Remarks.—This specimen was described by Tazawa (2017,p. 338, fig. 7.1) as Schellwienella radialis (Phillips, 1836). The
Tairagai specimen is poorly preserved but can be referred to S.
radialis (Phillips, 1836), redescribed by Brunton (1968, p. 42,
pl. 6, figs. 13–24, text-figs. 27–35) on the type specimens fromthe upper Visean of Fermanagh, northern Ireland, by themedium-sized (length about 32 mm, width about 33 mm),flattened ventral valve, with strong divergent dental plates, andthe parvicostellate ornament with concentric growth lines
forming serrations along crests of the costellae. S. radialis isdistinguished from other species of the genus Schellwienella by
the strong parvicostellate ornament. Schellwienella izirii Minato(1951, p. 363, pl. 5, fig. 3), from the Hikoroichi Series (= Odaira
Formation) of Okuhinotsuchi, South Kitakami Belt, is not
assigned to the genus Schellwienella by the presence of median
septum in the ventral valve. The Okuhinotsuchi species is
identical with Schizophoria resupinata (Martin, 1809) on
account of its size, shape, external ornament and internal
structure of the ventral valve.
Occurrence.—OD3 Unit (Yokota).
Distribution.—Upper Tournaisian–upper Visean: northeasternJapan (Yokota in the South Kitakami Belt), UK (Scotland,
England and northern Ireland), Germany, Belgium, western
Russia (Moscow Basin), central Russia (southern Urals) andnorthwestern China (Xinjiang and Gansu).
Order ORTHIDA Schuchert and Cooper, 1932
Suborder DALMANELLIDINA Moore, 1952
Superfamily DALMANELLOIDEA Schuchert, 1913
Family RHIPIDOMELLIDAE Schuchert, 1913
Subfamily RHIPIDOMELLINAE Schuchert, 1913
Genus RHIPIDOMELLA Oehlert, 1890
Type species.—Terebratula michelini Léveillé, 1835.
Rhipidomella michelini (Léveillé, 1835)(Figs. 29A–F, 39A–F, 45B)
Terebratula michelini Léveillé, 1835, p. 39, pl. 2, figs. 14–17.
Orthis michelini (Léveillé). Davidson, 1861, p. 132, pl. 30, figs.
6–12.
Dalmanella michelini (Léveillé). Frech, 1900, p. 201, pl. 16, fig. 15.
Rhipidomella michelini (Léveillé). Rotai, 1931, p. 44, pl. 1, fig.
3; Demanet, 1934, p. 37, pl. 2, figs. 1–9; Sarytcheva in
Sarytcheva and Sokolskaya, 1952, p. 26, pl. 1, fig. 7:
Litvinovich, 1962, p. 177, pl. 1, fig. 1; Zang in Yang et al.,1962, p. 19, pl. 1, figs. 1–7; Ustritsky and Tschernjak, 1963,
p. 68, pl. 1, figs. 11, 12; Yang, 1964, p. 58, pl. 1, fig. 1;
Brunton, 1968, p. 17, pl. 3, figs. 1–25, text-fig. 5; Litvinovich
in Litvinovich et al., 1969, p. 127, pl. 1, figs. 9, 10; Bublichenko,
1971, p. 29, pl. 2, figs. 9–12; Alexandrow and Solomina,
1973, p. 87, pl. 21, fig. 1; Kalashnikov, 1974, p. 21, pl. 3,
figs. 7–9; Volgin and Kushnar, 1975, p. 21, pl. 1, figs. 1, 2;
Lee and Gu, 1976, p. 231, pl. 131, figs. 1–6; Martinez Chacon,
1979, p. 63, pl. 3, figs. 12–15; pl. 4, figs. 1–15, text-figs. 6, 7;
Lee et al., 1980, p. 330, pl. 145, fig. 4; Ding and Qi, 1983, p.
250, pl. 88, fig. 13; Zakowa, 1989, p. 115, pl. 3, fig. 5; pl. 7,
fig. 7; Harper and Jeffrey, 1996, fig. 3a. Legrand-Blain in
Legrand-Blain et al., 1996, p. 180, pl. 28, figs. 21, 22. Jiang,
1997, pl. 1, fig. 3; Bassett and Bryant, 2006, p. 502, pl. 1,
figs. 1–4; pl. 6, figs. 11–17; Sun and Baliński, 2008, p. 519,
fig. 26; Bahrammanesh et al., 2011, p. 149, fig. 5aa–dd,
7a–e; Tazawa, 2018c, p. 52, figs. 26A–C, 30G; Tazawa andIryu, 2019, p. 101, figs. 6A, B; Tazawa and Ibaraki, 2019, p.
16, fig. 4B.
Rhipidomella sp. Tazawa and Katayama, 1979, p. 170, pl. 11,
figs. 1–7; Mori and Tazawa, 1980, text-figs. 3.4–3.6; Tazawa,
1984b, p. 305, pl. 61, figs. 5–7.
Material.—Twenty-one specimens from localities IST1,
IYK1, KF158, KF175, KF179, KF180, KF181, YKA16 andYUK1: (1) internal mould of a conjoined shell, with externalmould of the dorsal valve, IGPS112158; (2) internal mould of
three conjoined shells, IGPS111721, 111722, KCG67; (3)internal moulds of six ventral valves, IGPS96102, 96116,
96126, 111723, 112100, NU-B2249; (4) external and internal

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

53

FIGURE 36. Brachiopods of the AR3 assemblage (1). A, Orbiculoidea sp., external latex cast (A1 , A 2 ) and external mould (A3 ) of ventral valve,
NU-B2266; B, C, Marginatia burlingtonensis (Hall); ventral (B1, B2), anterior (B3), posterior (B4) and lateral (B5) views of internal mould of ventral valve,
NU-B630; ventral (C1, C2) and posterior (C3) views of internal mould of ventral valve, NU-B650. Scale bars are 1 cm.

54

JUN-ICHI TAZAWA

moulds of two dorsal valves, IGPS112101, KCG68; (5) externalmoulds of two dorsal valves, IGPS112102, NU-B2256; and (6)internal moulds of seven dorsal valves, IGPS96103, 96111,
96121, 111724, 112103, 112104, NU-B2250.
Remarks.—Some of the specimens were previously
described by Tazawa and Katayama (1979, p. 170, pl. 11, figs.
1‒7) as Rhipidomella sp. But the Odaira species is referred to
Rhipidomella michelini (Léveillé, 1835), redescribed by Brunton(1968, p. 17, pl. 3, figs. 1–25, text-fig. 5) from the Visean of
Fermanagh, northern Ireland, in the small size (length about 24
mm, width about 16 mm in the largest specimen, IGPS96102),
ovate outline with short hinge and being widest at slightly
anterior to midlength. The other specimens from the Arisu andOdaira formations of Shimoarisu, Yokota and Shizu areas arealso referred to the present species. Rhipidomella altaica
Tolmatchoff (1924, p. 213, 569, pl. 13, figs. 5–7, 9, 10), fromthe Tournaisian of the Kuznetsk Basin, central Russia, differsfrom R. michelini in having wider hinge and the widest part at
midlength of the shell.
Occurrence.—AR2 Unit (Shimoarisu and Yokota), OD2 Unit
(Yokota and Shizu) and OD3 Unit (Yahagi).
Distribution.—Upper Tournaisian–lower Bashkirian:northeastern Japan (Hikoroichi, Shimoarisu, Yokota, Yahagi and
Shizu in the South Kitakami Belt), northern Russia (Taimyr
Peninsula and Pechora Basin), UK (England, Wales andnorthern Ireland), Ireland, Poland, Belgium, France (French
Pyrenees), Spain (Cantabrian Mountains), western Russia(Moscow Basin and Donetz Basin), Iran, central Russia(southern Urals), Kazakhstan, Uzbekistan, northwestern China(Xinjiang, Qinghai, Gansu and Ningxia), northern China (Inner
Mongolia), northeastern China (Liaoning) and southwestern
China (Guizhou and Yunnan).
Rhipidomella kusbassica Beznossova in Sarytcheva et al., 1963
(Fig. 26A)
Rhipidomella michelini (non Léveillé). Tolmatchoff, 1924, p.
212, 569, pl. 13, fig. 4; Nalivkin, 1937, p. 36, pl. 3, figs. 6, 7.
Rhipidomela kusbassica Beznossoba in Sarytcheva et al., 1963,
p. 74, pl. 2, figs. 9–11; Grechishnikova, 1966, p. 91, pl. 1,
figs. 5–10; Zhang et al., 1983, p. 264, pl. 106, fig. 9; Tazawa
and Kurita, 2019a, p. 221, fig. 4A; Tazawa in Tazawa et al.,2019, p. 56, fig. 12G, H.
Material.—One specimen from locality KAR1, internalmould of a dorsal valve, IGPS111760.
Remarks.—This specimen was described by Tazawa andKurita (2019a, p. 221, fig. 4A) as Rhipidomella kusbassica
Beznossova in Sarytcheva et al., 1963. The Okuhinotsuchi
specimen is poorly preserved but can be referred to Rhipidomelakusbassica Beznossoba in Sarytcheva et al. (1963, p. 74, pl. 2,
figs. 9–11), from the lower Tournaisian–lower Visean of theKuznetsk Basin, central Russia, in the small size (length morethan 14 mm, width about 15 mm), relatively wide hinge andstrong brachiophores diverging anteriorly. Rhipidomella altaica

Tolmatchoff (1924, p. 213, 569, pl. 13, figs. 5–7, 9, 10), fromthe Tournaisian of the Kuznetsk Basin, is the most similar
species having a wide hinge. But R. altaica differs from R.
kusbassica in more transverse outline and in having slightly
shorter hinge. The preceding species, Rhipidomella michelini(Léveillé), differs from R. kusbassica in having much shorter
hinge.
Occurrence.—ST3 Unit (Okuhinotsuchi).
Distribution.—Lower Tournaisian–lower Visean: northeasternJapan (Nisawa, Okuhinotsuchi and Nagasaka in the South
Kitakami Belt), central Russia (Kuznetsk Basin), Kazakhstan
and northwestern China (Xinjiang).
Superfamily ENTELETOIDEA Waagen, 1884
Family SCHIZOPHORIIDAE Schuchert and LeVene, 1929
Genus SCHIZOPHORIA King, 1850
Type species.—Conchyliolithus (Anomites) resupinatus
Martin, 1809.
Schizophoria resupinata (Martin, 1809)(Figs. 30F–I, 40B–L, 41I–L)
Conchiliolithus (Anomites) resupinatus Martin, 1809, pl. 49,
figs. 13, 14.
Orthis resupinata (Martin). Davidson, 1861, p. 130, pl. 29, figs.
1–4; pl. 30, figs. 1–5.
Schizophoria resupinata (Martin). Yanishevsky, 1918, p. 19, pl.
1, figs. 4, 12; pl. 4, fig. 2; pl. 6, fig. 16; Demanet, 1934, p. 45,
pl. 3, figs. 1–5, text-fig. 9; Miloradovich, 1935, p. 6, pl. 1,
figs. 11, 12; Bond, 1941, p. 289, pl. 21, figs. A–C, text-figs.
33, 34; Minato, 1952, p. 150, pl. 5, fig. 3; pl. 6, fig. 4;
Sarytcheva in Sarytcheva and Sokolskaya, 1952, p. 29, pl. 2,
fig. 12; Parkinson, 1954, p. 368, text-figs. 1, 2; Litvinovich,
1962, p. 178, pl. 1, fig. 2; Besnossova in Sarytcheva et al.,1963, p. 77, pl. 3, figs. 5–8, text-fig. 24; Ustritsky andTschernjak, 1963, p. 69, pl. 1, figs. 13–16; Yang, 1964, p. 59,
pl. 1, figs. 2, 3; Abramov, 1965, p. 35, pl. 2, fig. 3; Brunton,
1968, pl. 2, figs. 1–6; Pocock, 1968, p. 80, pl. 18, fig. 7,
text-figs. 13–15; Besnossova et al. in Sarytcheva, 1968, p. 53,
pl. 1, figs. 11–13; Lazarev, 1969, pl. 10, figs. 1–5, text-figs. 1,
2; Litvinovichi in Litvinovich et al., 1969, p. 129, pl. 2, fig. 1;
Abramov, 1970, p. 107, pl. 1, figs. 5–7; Aisenverg and
Poletaev, 1971, pl. 60, fig. 1; Nalivkin and Fotieva, 1973, p.
20, pl. 1, figs. 6–8; Yanagida, 1973, p. 101, pl. 16, figs. 3—9;
Kalashnikov, 1974, p. 22, pl. 3, figs. 1–3; Garanj et al., 1975,
p. 155, pl. 62, fig. 2; Volgin and Kushnar, 1975, p. 23, pl. 1,
figs. 3–5; Litvinovich et al., 1975, p. 52, pl. 16, fig. 7;
Lazarev, 1976, pl. 2, figs. 3, 4; pl. 3, figs. 1–5, text-fig. 58,
table 11; Lee and Gu, 1976, p. 229, pl. 131, figs. 7, 9–11;
Martinez Chacon, 1979, p. 54, pl. 2, figs. 1–15; pl. 3, figs.
1–10, text-figs. 3–5; Minato et al., 1979a, pl. 22, figs. 1, 2;
Tazawa and Katayama, 1979, p. 169, pl. 11, figs. 8–14;
Kalashnikov, 1980, p. 24, pl. 2, figs. 2, 3; Mori and Tazawa,
1980, text-fig. 3.3; Tazawa, 1981b, p. 67, pl. 5, figs. 3–5; Tazawa

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

55

FIGURE 37. Brachiopods of the AR3 assemblage (2). A, B, Brachythyrina sp.; external latex cast (A 1, A2) and internal mould (A3) of dorsal valve,
UHR16178; internal mould (B 1, B2) of dorsal valve, UHR15993; C, Punctospirifer sp., internal mould (C 1, C2) of dorsal valve, NU-B2255; D, E,
Unispirifer minnewankensis (Shimer); external latex cast (D1, D2) of ventral valve, NU-B2254; ventral (E1, E2) and dorsal (E3) views of internal mould,
and dorsal view of external latex cast (E4) of conjoined shell, NU-B2253. Scale bars are 1 cm.

et al., 1981, pl. 1, figs. 4–6; Ding and Qi, 1983, p. 245, pl. 88,
fig. 7; Zhang et al., 1983, p. 265, pl. 107, figs. 1–3; Tazawa,
1984b, p. 304, pl. 61, fig. 9; Abramov and Grigorjeva, 1986,
p. 74, pl. 1, figs. 15–18; Yanai et al., 1988, pl. 1, figs. 9, 10;
Zakowa, 1989, p. 103, pl. 1, figs. 1–5; pl. 2, figs. 1–5; pl. 3,
figs. 1–4, text-figs. 2–10, table 2; Jiang, 1997, pl. 1, figs. 1, 2;
Bassett and Bryant, 2006, p. 504, pl. 6, figs. 1–10; pl. 7, figs.
1–16, text-figs. 5–7; Butts, 2007, p. 55, figs. 5.3–5.10; Ibaraki
et al., 2014, p. 73, figs. 4.1, 4.2; Tazawa, 2018c, p. 54, figs.
18A, B, 26G, H, 31A–E; Tazawa and Iryu, 2019, p. 101, fig. 6C, D.

Schellwienella izirii Minato, 1951, p. 363, pl.5, fig. 3.
Schellwienella? sp. Minato, 1951, p. 364.
Schizophoria aff. resupinata (Martin). Yanagida, 1962, p. 122,
pl. 21, figs. 4–13, text-fig. 22; Hase and Yokoyama, 1975, pl.
16, figs. 6, 7.
Schizophoria (Schizophoria) resupinata (Martin). Sun andBaliński, 2008, p. 521, figs. 27F‒L; Bahrammanesh et al.,2011, p. 153, fig. 7f, g.
Material.—Thirty-nine specimens from localities INS2,

56

JUN-ICHI TAZAWA

INS7, INS9, INS10, INS11, INS13, IST1, KF158, KF177,
YKA16, YKA18, YKA20 and YKA21: (1) internal moulds ofnine conjoined shells, IGPS111715, 111716, 112138–112144;(2) external and internal moulds of three ventral valves,
IGPS112145, 112146, NU-B2251; (3) internal moulds ofnineteen ventral valves, IGPS96078, 96080, 96081,
97646–97650, 111717–111720, 112105, 112106,
112147–112150, UHR16142; and (4) internal moulds of eightdorsal valves, IGPS96099, 97651‒97654, 112151–112153.
Remarks.—Some of the specimens were described by
Tazawa and Katayama (1979, p. 169, pl. 11, figs. 8–14) as
Schizophoria resupinta (Martin, 1809). All the specimens
avairable are referred to Schizophoria resupinata (Martin,
1809), redescribed by Pocock (1968, p. 80, pl. 18, fig. 7,
text-figs. 13–15) from the upper Tournaisian‒upper Visean ofBelgium and the Britain, in the large size (length more than 40
mm, width about 67 mm in the largest specimen, IGPS112138),transversely subrectangular outline, respinate shell, and themoderately incised and flabellate ventral muscle field.
Schizophoria connivens (Phillips, 1836), redescribed by Pocock(1968, p. 64, pl. 18, figs. 1, 2, text-figs. 1–4) from theTournaisian‒Visean of Belgium and the Britain, differs from S.
resupinata in the smaller size and in having deeper sulcus andmore strongly incised, elliptical muscle field in the ventral
valve. Two species, described by Minato (1951) as
Schellwienella izirii Minato, 1951 and Schellwienella? sp. fromthe Hikoroiti Series (= Odaira Formation) of “the 808 m hill” ,Nisawa, are identical with S. resupinata (Martin) in size, shape,
external ornament and internal structure of the ventral valves.
Occurrence.—AR2 Unit (Shimoarisu and Yokota) and OD2Unit (Yokota, Nisawa and Shizu).
Distribution.—Upper Tournaisian—upper Visean: northeasternJapan (Hikoroichi, Shimoarisu, Yokota, Nisawa and Shizu in
the South Kitakami Belt), central Japan (Kanto Mountains),
southwestern Japan (Akiyoshi Belt), USA (Idaho), northern
Russia (Verkhoyansk Range, Taimyr Peninsula, Novaya
Zemlya and Pechora Basin), UK (England and Wales),Poland, Belgium, Spain, western Russia (Moscow Basin andDonetz Basin), Iran, central Russia (southern Urals andKuznetsk Basin), Kazakhstan, Uzbekistan (Fergana),
northwestern China (Xinjiang and Shaanxi), northern China(Inner Mongolia) and southwestern China (Guizhou andYunnan).

Schizophoria (Schizophoria) resupinata forma pinguis Demanet.
Zakowa, 1989, p. 109, pl. 6, figs. 1–10; pl. 7, figs. 1–4,
text-figs. 2–9, table 6.
Material.—Twelve specimens from localities IST1, KAR1,
KF179, KF180 and KF181: (1) external and internal moulds of adorsal valve, IGPS111729; and (2) internal moulds of threeventral valves, IGPS112084–112086; and (3) internal moulds ofeight dorsal valves, IGPS 99006, 99007, 111730–111732,
112087–112089.
Remarks.—Most of the specimens were previously describedby Tazawa and Kurita (1986, p. 167, figs. 2.2, 2.3) as
Schizophoria resupinata pinguis Demanet, 1934, and by Tazawa
and Iryu (2019, p. 102, fig. 6E, F) and Tazawa and Kurita(2019a, p. 222, fig. 4B, C) as Schizophoria pinguis Demanet(1934). The Shimoarisu and Okuhinotsuchi specimens arereferred to Schizophoria pinguis Demanet (1934, p. 59, pl. 4,
figs. 9–11), from the upper Visean of Visé, Belgium, in themedium to large size (length 48 mm, width 28 mm in the largest
specimen, IGPS111729; length 23 mm, width 46 mm in the best
preserved specimen, IGPS111730), highly inflated dorsal valve
and weakly uniplicate anterior commissure. Schizophoria
gibbera (Portlock, 1843), redescribed by Pocock (1968, p. 69,
pl. 18, fig. 3, text-figs. 5–7) from the upper Tournaisian–lower
Visean of England and Ireland, has also strongly convex dorsal
valve, but differs from S. pinguis in having a concentric fold on
the dorsal valve. The type species, Schizophoria resupinata(Martin, 1809), is readily distinguished from the present species
in having gently convex dorsal valve.
Occurrence.—ST3 Unit (Okuhinotsuchi) and AR2 Unit
(Shimoarisu and Yokota).
Distribution.—Upper Tournaisian–upper Visean: northeasternJapan (Shimoarisu, Yokota and Okuhinotsuchi in the South
Kitakami Belt), UK (England), Poland and Belgium.
Schizophoria woodi Bond, 1941
(Fig. 30A)

Schizophoria pinguis Demanet, 1934
(Figs. 26B, C, 30B–E)

Schizophoria resupinata var. gibbera (Portlock). Demanet,
1934, p. 55, pl. 4, figs. 1–3 only.
Schizophoria woodi Bond, 1941, p. 299, pl. 22, figs. f, g, text-fig.
37; Pocock, 1968, p. 86, pl. 18, fig. 8, text-figs. 16–19;
Lazarev, 1976, pl. 6, fig. 2, text-fig. 64; Tazawa and Iryu,
2019, p. 102, fig. 6G.
Schizophoria (Paraschizophoria) woodi Bond. Zakowa, 1989, p.
113, pl. 8, figs. 1, 2, table 10.

Schizophoria resupinata var. pinguis Demanet, 1934, p. 59, pl.
4, figs. 9–11; Bond, 1941, figs. 33, 34; Pocock, 1968, text-fig.
21.
Schizophoria pinguis Demanet. George and Ponsford, 1938,
figs. 8, 9; Tazawa and Iryu, 2019, p. 102, fig. 6E, F; Tazawa
and Kurita, 2019a, p. 222, fig. 4B, C.
Shizophoria resupinata pinguis Demanet. Tazawa and Kurita,
1986, p. 167, figs. 2.2, 2.3.

Material.—One specimen from IST1, internal mould of aconjoined shell, IGPS111728.
Remarks.—The single specimen from Shimoarisu wasdescribed by Tazawa and Iryu (2019, p. 102, fig. 6G) as
Schizophoria woodi Bond, 1941. The Shimoarisu specimen isreferred to Schizophoria woodi Bond (1941, p. 299, pl. 22, figs.
F, G, text-fig. 37), from the upper Visean of Craven, Yorkshire,
England, in the small size (length about 33 mm, width about 20

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

57

FIGURE 38. Brachiopods of the OD2 assemblage (1). A, B, Absenticosta uldzejtuensis Suursuren and Lazarev; A, internal mould (A1, A2) of ventral
valve, IGPS112133; B, internal mould (B1, B2) of dorsal valve, IGPS112134; C, D, Marginicinctus marginicinctus (Prout); C, ventral (C1, C2) and anterior(C3) views of internal mould of ventral valve, IGPS97657; D, ventral internal mould (D1, D2), dorsal internal mould (D3), dorsal external mould (D4) anddorsal external latex cast (D5) of conjoined shell, IGPS97656; E, Fluctuaria undata (Defrance); ventral internal mould (E1, E2), dorsal internal mould (E3)and dorsal external mould (E4) of conjoined shell, IGPS97659; F, G, Levitusia elongata sp. nov.; F, external latex cast (F1) and internal mould (F2) ofventral valve, IGPS112154; G, ventral (G1) and dorsal (G2) views of internal mould of conjoined shell, IGPS112155. Scale bars are 1 cm.

58

JUN-ICHI TAZAWAmm) and the strongly dorsibiconvex shell, with angular
uniplicate anterior commissure. Schizophoria palliata Demanet(1934, p. 58, pl. 4, figs. 7, 8), from the upper Tournaisian ofBelgium, differs from S. woodi in having a rounded anterior
commissure.
Occurrence.—AR2 Unit (Shimoarisu).
D i s t r i b u t i o n . —U p p e r T o u r n a i s i a n –S e r p u k h o v i a n :northeastern Japan (Shimoarisu in the South Kitakami Belt), UK(England, Isle of Man and northern Ireland), Poland, Belgium
and central Russia (southern Urals).
Schizophoria mayesensis Carter, 1999
(Fig. 26D‒F)
Schizophoria mayesensis Carter, 1999, p. 109, figs. 6A‒nn, tt, 7;
Tazawa and Kurita, 2019a, p. 223, fig. 4D‒F.
Material.—Three specimens from locality KAR1: (1)internal mould of a ventral valve, IGPS111762; and (2) external
and internal moulds of two dorsal valves, IGPS111763, 111764.
Remarks.—These specimens were described by Tazawa andKurita (2019a, p. 223, fig. 4D‒F) as Schizophoria mayesensis
Carter, 1999. The Okuhinotsuchi specimens can be referred to
Schizophoria mayesensis Carter (1999, p. 109, figs. 6A‒nn, tt,7), from the St. Joe Formation of Mayes County, northeastern
Oklahoma, in being medium size (length 21 mm, width morethan 25 mm in the largest specimen, IGPS111763), transversely
subelliptical outline with the greatest width at slightly posterior
to midlength, and thin-bodied shell with flattened ventral valve
and gently convex dorsal valve. S. mayesensis resembles
Schizophoria chouteauensis Weller (1914, p. 163, pl. 23, figs.
6–19), from the Chouteau Limestone of Missouri, but the latter
differs from S. mayesensis in having less transverse outline andthe maximum width at about midlength.
Occurrence.—ST3 Unit (Okuhinotsuchi).
Distribution.—Upper Tournaisian: northeastern Japan(Okuhinotsuchi in the South Kitakami Belt) and USA(Oklahoma).
Order RHYNCHONELLIDA Kuhn, 1949
Superfamily RHYNCHOTETRADOIDEA Licharew, 1956
Family TETRACAMERIDAE Licharew, 1956
Genus ROTAIA Rzhonsnitskaya, 1956
Type species.—Rhynchonella subtrigona Meek and Worthen,
1860.
Rotaia hikoroichiensis Tazawa, 2006
(Fig. 41M)
Rotaia sp. Tazawa, 1996, p. 8, fig. 4; Tazawa, 2002, fig.
7.3–7.5.
Rotaia hikoroichiensis Tazawa, 2006, p. 135, fig. 8.1–8.3;
Tazawa, 2018c, p. 56, figs. 26E, F, 30I.

Material.—One specimen from locality YKA20, external andinternal moulds of a ventral valve, IGPS112137.
Remarks.—This specimen can be referred to Rotaia
hikoroichiensis Tazawa, 2006, originally described by Tazawa(2006, p. 135, fig. 8.1–8.3) from the middle part (HK2 Unit) of
the Hikoroichi Formation in the Hikoroichi area, South Kitakami
Belt, in the large, transverse ventral valve (length about 46 mm,
width about 57 mm), with shallow sulcus and ornamented withcoarse costae on the valve. Comparison with the other species ofRotaia has been discussed by Tazawa (2006, p. 136).
Occurrence.—OD2 Unit (Shizu).
Distribution.—Lower–upper Visean: northeastern Japan(Hikoroichi and Shizu in the South Kitakami Belt).
Order ATHYRIDIDA Boucot, Johnson and Staton, 1964
Suborder ATHYRIDIDINA Boucot, Johnson and Staton, 1964
Superfamily ATHYRIDOIDEA Davidson, 1881
Family ATHYRIDIDAE Davidson, 1881
Subfamily CLEIOTHYRIDININAE Alvarez, Rong and Boucot,
1998
Genus CLEIOTHYRIDINA Buckman, 1906
Type species.—Atrypa pectinifera Sowerby, 1840.
Cleiothyridina fimbriata (Phillips, 1836)(Fig. 46H)
Spirifer fimbriata Phillips, 1836, p. 220.
Terebratula plano-sulcata (Phillips). de Koninck, 1843, p. 301,
pl. 21, figs. 1e, f only.
Athyris roysii Léveillé. Davidson, 1861, p. 84, pl. 18, figs. 8–11
only.
Cleiothyridina fimbriata (Phillips). Brunton and Champion,
1974, p. 829, pl. 109, figs. 22, 23; Brunton, 1980, p. 227, figs.
19–22; Brunton, 1984, p. 53, figs. 53–66; Tazawa, 2017, p.
338, fig. 8.8.
Material.—Two specimens from locality KF159, externalmoulds of two dorsal valves, NU-B2164, 2165.
Remarks.—These specimens were described by Tazawa(2017, p. 338, fig. 8.8) as Cleiothyridina fimbriata (Phillips,
1836). The specimens from Tairagai can be referred to
Cleiothyridina fimbriata (Phillips, 1836), redescribed by
Brunton (1980, p. 227, figs. 19–22), from the upper Visean ofCounty Fermanagh, northern Ireland, by the absence of dorsal
fold and the external ornament of concentric lamellae withspine-like frilly outgrowths. Cleiothyridina deroissyi (Léveillé,1835), from the Lower Carboniferous (Tournaisian?) of Tournai,
Belgium, is readily distinguished from C. fimbriata by thepresence of distinct ventral sulcus and dorsal fold.
Occurrence.—OD3 Unit (Yokota).
Distribution.—Lower–upper Visean: northeastern Japan(Yokota in the South Kitakami Belt), UK (England and northern
Ireland) and Belgium.

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS59

FIGURE 39. Brachiopods of the OD2 assemblage (2). A‒F, Rhipidomella michelini (Léveillé); A‒D, internal moulds of ventral valves, IGPS96102 (A1,
A2), IGPS96116 (B), IGPS96126 (C) and NU-B2249 (D); E‒G, internal moulds of dorsal valves, IGPS96103 (E), IGPS96111 (F) and IGPS96121 (G); H,
I, Orthotetes keokuk (Hall); H, external latex cast (H1) and internal mould (H2) of ventral valve, IGPS97655; I, external latex cast (I1) and internal mould(I2) of dorsal valve, IGPS11216. Scale bars are 1 cm.

Cleiothyridina harkeri Carter, 1987
(Fig. 31A)
Cleiothyridina obmaxima (McChesney)? Nelson, 1961, pl. 4,
figs. 5, 6; pl. 7, fig. 14; Abramov and Grigorjeva, 1986, pl.
15, figs. 17, 18.
Cleiothyridina obmaxima (McChesney). Abramov, 1970, p.
157, pl. 38, figs. 1–3.
Cleiothyridina harkeri Carter, 1987, p. 60, pl. 19, figs. 24–27;
Tazawa, 2018c, p. 63, fig. 19C; Tazawa and Ibaraki, 2019, p.

19, fig. 5A; Tazawa in Tazawa et al. 2019, p. 58, fig. 12E.
Material.—One specimen from locality IYK2, internal mouldof a conjoined shell, with external mould of the dorsal valve,
NU-B2265.
Remarks.—This specimen was described by Tazawa and
Ibaraki (2019, p. 19, fig. 5A) as Cleiothyridina harkeri Carter,
1987, on the basis of its large, transverse shell (length about 25
mm, width about 36 mm), weakly uniplicate anterior
commissure with shallow ventral sulcus and low dorsal fold, and60

JUN-ICHI TAZAWA

external ornament consisting of numerous dense growthlamellae. Cleiothyridina obmaxima (McChesney), redescribedby Weller (1914, p. 475, pl. 79, figs. 1–11) from the Keokuk andBurlington Formations of the Mississippi Valley, is also a large,transverse Cleiothyridina, but the American species differs fromC. harkeri in having well developed sulcus and fold.
Occurrence.—AR2 Unit (Yokota).
Distribution.—Upper Tournaisian–lower Visean: northeasternJapan (Hikoroichi, Yokota and Nagasaka in the South KitakamiBelt), western Canada (Alberta) and northern Russia(Verkhoyansk Range).
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 georgei Tazawa in Tazawa et al., 2019
(Fig. 41A‒E)
Martinia aff. glabra (Martin). George, 1927, p. 112, text-figs.
3‒11; Demanet, 1938, p. 103, pl. 9, figs. 24‒29.
Spirifer (Martinia) glaber Martin. Einor, 1936, p. 40, pl. 7, figs.
4, 5.
Martinia glabra (Martin). Yang, 1964, p. 141, pl. 21, figs. 3, 4,
text-fig. 32A; Litvinovich in Litvinovich et al., 1969, p. 275,
pl. 68, fig. 1; Donakova, 1983, pl. 3, fig. 8.
Martinia glabra (Sowerby). Poletaev, 1975, p. 43, pl. 9, figs. 1,
2; Chen and Shi, 2003, p. 168, pl. 11, figs. 1‒8, 12‒17, 37, 39.
Martinia sp. Tazawa and Osawa, 1979, p. 775, text-fig. 2.
Martinia aff. glabra (Sowerby). Tazawa, 1980, p. 366, pl. 42,
figs. 6‒11; Tazawa et al., 1981, pl. 1, figs. 1‒3; Tazawa,
1981b, p. 74, pl. 5, figs. 9‒13.
Martinia georgei Tazawa in Tazawa et al., 2019, p. 59, figs.
19E‒J.
Material.—Twenty-six specimens from localities INS1,
INS2, INS3, INS4, INS5, INS6, INS7, INS8, INS10, INS13 andYKA12: (1) external and internal moulds of a ventral valve,
IGPS112159; and (2) internal moulds of twenty-five ventral
valves, IGPS97662‒97684, 112160, 112161.
Remarks.—These specimens were figured by Tazawa et al.
(1981, pl. 1, figs. 1‒3) and described by Tazawa (1981b, p. 74,
pl. 5, figs. 9‒13) as Martinia aff. glabra (Sowerby, 1820). But
the specimens from Nisawa are referred to Martinia georgei
Tazawa (in Tazawa et al., 2019, p. 59, figs. 19E‒J), from theupper part of the Karaumedate Formation in the Nagasaka area,
South Kitakami Belt, in the medium-sized ventral valve (length
23 mm, width about 44 mm in the largest specimen,
IGPS97662), with a shallow sulcus. Martinia glabra (Sowerby,1820), redescribed by Muir-Wood (1951, p. 109, pl. 3, fig. 2)

from the upper Visean of Derbyshire, England, differs from M.
georgei in much larger dimensions.
Occurrence.—OD2 Unit (Nisawa and Shizu) and OD3 Unit
(Nisawa).
Distribution.—Lower Visean–Bashkirian: northeastern Japan(Nisawa, Shizu and Nagasaka in the South Kitakami Belt), UK(Wales), Belgium, western Russia (Donetz Basin), central
Russia (western and southern Urals), Kazakhstan andnorthwestern China (Xinjiang).
Superfamily SPIRIFEROIDEA King, 1846
Family SPIRIFERIDAE King, 1846
Subfamily SPIRIFERINAE King, 1846
Genus SPIRIFER Sowerby, 1816
Type species.—Conchyliolithus (Anomia) striatus Martin,
1793.
Spirifer liangchowensis Chao, 1929
(Figs. 41F‒H, 46A‒E)
Spirifer liangchowensis Chao, 1929, p. 6, pl. 1, figs. 1–7,
text-fig. 1; Yang in Yang et al., 1962, p. 99, pl. 39, fig. 1;
Tazawa, 2017, p. 340, figs. 8.1–8.5; Tazawa, 2018c, p. 63,
fig. 31G.
Neospirifer liangchowensis (Chao, 1929): Wang et al., 1964, p.
508, pl. 89, figs. 14–16; Ding and Qi, 1983, p. 401, pl. 135,
fig. 5.
Spirifer sp. Tazawa, 1984b, p. 307, pl. 61, fig. 1.
Material.—Thirteen specimens from localities KF158,
KF159 and YKA21: (1) internal moulds of a conjoined shell,
with external mould of the ventral valve, IGPS112163; (1)external and internal moulds of a ventral valve, NU-B2166; (2)external mould of a ventral valve, NU-B2167; (3) internalmoulds of seven ventral valves, IGPS112164, 112165,
NU-B2168, 2169, 2243–2245; (4) external and internal moulds
of a dorsal valve, NU-B2170; and (5) internal moulds of two
dorsal valves, IGPS112166, NU-B217.
Remarks.—The specimens from Tairagai were described by
Tazawa (2017, p. 340, figs. 8.1–8.5) as Spirifer liangchowensis
Chao, 1929. The other specimens from Shizu are also referred to
S. liangchowensis Chao, 1929, from the Chouniugou Formationof Gansu, northwestern China, on account of the middle-sized,
less transverse shell, with deep ventral sulcus and high dorsal
fold. Spirifer karagai (Litvinovich, 1962, p. 273, pl. 33, fig. 2),
from the upper Visean of central Kazakhstan, resembles S.
liangchouensis in size and outline of the shell, but differs fromthe latter in having shallower ventral sulcus and lower dorsal
fold. Spirifer gregeri Weller (1914, p. 359, pl. 55, figs. 1–8),
from the lower Burlington Limestone of the Mississippi Valley,differs from S. liangchowensis in its longer shell with shorter
hinge. The type species, Spirifer striata (Martin, 1793), from the
Visean of Derbyshire, England, is readily distinguished from thepresent species by the much larger size, transverse outline, andMISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

61

FIGURE 40. Brachiopods of the OD2 assemblage (3). A, Striatifera angusta (Yanishevsky), ventral (A1), lateral (A2) and posterior (A3) views of externallatex cast of ventral valve, and external latex cast of dorsal valve (A4), IGPS97660; B‒L, Schizophoria resupinata (Martin); B, ventral (B1) and dorsal (B2)views of internal mould of conjoined shell, IGPS112139; C, internal mould of ventral valve, IGPS96081; D, internal mould of dorsal valve, IGPS96099;
E, internal mould of ventral valve, NU-B2251; F, internal mould of ventral valve, IGPS96080; G, internal mould of ventral valve, UHR16142; H, internalmould of ventral valve, IGPS97646; I, internal mould of ventral valve, IGPS97647; J, ventral (J1) and dorsal (J2) views of internal mould of conjoined
shell, IGPS112140; K, internal mould of dorsal valve, IGPS97652; L, ventral (L 1 ) and dorsal (L 2 ) views of internal mould of conjoined shell,
IGPS112138. Scale bars are 1 cm.

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JUN-ICHI TAZAWA

more acute cardinal extremities.
Occurrence.—OD2 Unit (Yokota and Shizu) and OD3 Unit
(Yokota).
Distribution.—Lower–upper Visean: northeastern Japan(Hikoroichi, Yokota and Shizu in the South Kitakami Belt) andnorthwestern China (Gansu).

Type species.—Spirifer striatoconvolutus Dun and Benson in
Benson, Dun and Browne, 1920.

Genus GRANDISPIRIFER Yang, 1959

Spirifera striato-convoluta Dun and Benson in Benson, Dun and
Browne, 1920. p. 350, pl. 20, figs. 7, 8.
Unispirifer striatoconvolutus (Dun and Benson). Campbell,
1957, p. 68, pl. 14, figs. 1–9, text-figs. 10–12; Jiang, 1993,
p. 216, pl. 1, figs. 1–14, text-figs. 1, 2; Jiang, 1997, pl. 3, figs.
12–14; pl. 4, fig. 13; Tazawa, 2018c, p. 64, fig. 27D–G.
Tazawa and Iryu, 2019, p. 102, fig. 7A.
Unispirifer (Unispirifer) striatoconvoluts (Dun and Benson). Shi
et al., 2005, p. 55, figs. 12G–K.

Type species.—Grandispirifer mylkensis Yang, 1959.
Grandispirifer mylkensis Yang, 1959
(Fig. 27B, C)
Grandispirifer mylkensis Yang, 1959, p. 113, 118, pl. 1, figs. 1,
2; pl. 2, figs. 1–3; Yang, 1964, p. 121, pl. 16, figs. 4, 5; pl.
17, figs. 1–3, text-figs. 18, 19; Zhang et al., 1983, p. 350, pl.
121, fig. 1; Shi et al., 2016, p. 586, figs. 4, 5A, B; Tazawa andKurita, 2019b. p. 196. fig. 4.
Spirifer subaequalis Hall. Yang in Yang et al., 1962, p. 99, pl.
39, figs. 2, 3.
Material.—Three specimens from localities KAR4, KAR6
and KF181: (1) external and internal moulds of a ventral valve,
IGPS111742; and (2) internal moulds of two ventral valves,
IGPS111743, 112125.
Remarks.—Two specimens from the middle part of the Arisu
Formation (AR2 Unit) of Okuhinotsuchi were described by
Tazawa and Kurita (2019b, p. 196. fig. 4) as Grandispirifer
mylkensis Yang, 1959. All the specimens from the AR2 Unit of
Yokota and Okuhinotsuchin are referred to G. mylkensis Yang,
1959, from the lower Visean of Mt. Borohoro, Xinjiang,
northwestern China, in the large, transverse ventral valve (length
36 mm, width 77 mm in the best preserved specimen,
IGPS111742) with weakly developed sulcus and roundedcardinal extremities. Shells described by Yang (in Yang et al.,1962, p. 99, pl. 39, figs. 2, 3) as Spirifer subaequalis (Hall) fromthe Chengqianggou Formation of the southern Qilianshan,
Qinghai, northwestern China, are considered to be conspecificwith the present species. Grandispirifer qaidamensis Lee, Shi
and Chen in Shi et al. (2016, p. 586, figs. 5C, 6, 7), from theupper part of the Huaitoulata Formation of Qinghai,
northwestern China, is readily distinguished from G. mylkensis
by the strongly transverse outline.
Occurrence.—AR2 Unit (Yokota and Okuhinotsuchi).
Distribution.—Upper Tournaisian–lower Visean: northeasternJapan (Yokota and Okuhinotsuchi in the South Kitakami Belt)and northwestern China (Xinjiang and Qinghai).
Suborder SPIRIFERIDINA Waagen, 1883
Superfamily SPIRIFEROIDEA King, 1846
Family SPIRIFERIDAE King, 1846
Subfamily PROSPIRIFERINAE Carter, 1974
Genus UNISPIRIFER Campbell, 1957

Unispirifer striatoconvolutus (Dun and Benson in Benson, Dun
and Browne, 1920)(Fig. 31B‒D)

Material.—Four specimens from localities IST1 and KF179:(1) internal mould of a conjoined shell, with external mould of
the ventral valve, IGPS112094; (2) external moulds of aconjoined shell, IGPS112095; and (3) internal moulds of two
ventral valves, IGPS111733, 112096.
Remarks.—One of the specimens (IGPS111733) was
described by Tazawa and Iryu (2019, p. 102, fig. 7A) as
Unispirifer striatoconvolutus (Dun and Benson in Benson, Dun
and Browne, 1920), on account of the large, transverse andmucronate ventral valve (length about 26 mm, width about 59mm), with very shallow sulcus. The other three specimens(IGPS112094–112096) from Yokota are also referred to thepresent species. Unispirifer fluctuosus (Glenister, 1955, p. 68,
pl. 7, figs. 1–14; pl. 8, figs. 1–8), from the Moogooree
Limestone of the North-West Basin, northwestern Australia,differs from U. striatoconvolutus in the smaller size and inhaving more acute cardinal extremities.
Occurrence.—AR2 Unit (Shimoarisu and Yokota).
D i s t r i b u t i o n . —U p p e r T o u r n a i s i a n –u p p e r V i s e a n :northeastern Japan (Hikoroichi, Shimoarisu and Yokota in theSouth Kitakami Belt), southwestern China (Yunnan), andeastern Australia (New South Wales).
Unispirifer minnewankensis (Shimer, 1926)(Fig. 37D, E)
Spirifer centronatus var. minnewankensis Shimer, 1926, p. 52,
pl. 1, fig. 8.
Spirifer minnewankensis Shimer. Brown, 1952, p. 100, pl. 4, fig.
6; Nelson, 1961, pl. 7, figs. 5, 6.
Spirifer forbesi Norwood and Pratten. Nelson, 1961, pl. 7, figs.
1–4.
Unispirifer sp. Tazawa, 1985, p. 460, figs. 2.1, 2.2.
Unispirifer minnewankensis (Shimer). Carter, 1987, p. 75, pl.
23, figs. 1–13, 22–25; Chen and Archbold, 2000, p. 195, figs.
5.7, 5.8.

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FIGURE 41. Brachiopods of the OD2 assemblage (4). A‒E, Martinia georgei Tazawa; A, internal mould of ventral valve, IGPS97662; B, internal mouldof ventral valve, IGPS97663; C, internal mould of ventral valve, IGPS97664; D, internal mould of ventral valve, IGPS97668, E, internal mould of ventral
valve, IGPS112159; F‒H, Spirifer liangchowensis Chao; external latex cast of ventral valve (F1), and ventral (F2) and dorsal (F3) views of internal mouldof conjoined shell, IGPS112163; G, internal mould of ventral valve, NU-B2243; H, ventral (H1), anterior (H2), posterior (H3) and lateral (H4) views ofinternal mould of conjoined shell, NU-B2244; I‒L, Schizophoria resupinata (Martin); I, internal mould of ventral valve, IGPS112147; J, internal mould of
dorsal valve, IGPS112153; K, ventral (K1) and dorsal (K2) views of internal mould of conjoined shell, IGPS112143; L, external latex cast (L1) and internalmould (L 2 ) of ventral valve, IGPS112145; M, Rotaia hikoroichiensis Tazawa; external latex cast (M 1 ) and internal mould (M 2 ) of ventral valve,
IGPS112137. Scale bars are 1 cm.

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JUN-ICHI TAZAWA

Material.—Two specimens from locality KYT1: (1) external
and internal moulds of a conjoined shell, NU-B2253; (2)external mould of a ventral valve, NU-B2254.
Remarks.—These specimens were described by Tazawa(1985, p. 460，figs. 2.1) as Unispirifer sp. from the upper part of
the Arisu Formation (SR3 Unit) of the Yokota area. But the
Yokota specimens are referred to Unispirifer minnewankensis(Shimer, 1926, p. 52, pl. 1, fig. 8), from the lower part of the
Runde Formation of the Lake Minnewanka region, western
Alberta, Canada, in the medium-sized and transverse shell(length about 20 mm, width about 40 mm in the better preserved
specimen, NU-B2253), with numerous simple rounded costae
on both valves (numbering 5–6 in 5 mm near anterior margin of
the ventral valve). Unispirifer platynotus (Weller, 1914, p. 317,
pl. 39, figs. 1–10), from the Kinderhookian of the Mississippi
Valley, differs from U. minnewankensis in the smaller size,
more alate outline, and in having fewer and stronger costae onboth valves.
Occurrence.—AR3 Unit (Yokota).
Distribution.—Upper Tournaisian‒ lower Visean:
northeastern Japan (Hikoroichi and Yokota in the South
Kitakami Belt), western Canada (Alberta) and northwestern
China (Xinjiang).

dental plates. Dorsal interior not well preserved and obscure.
Remarks.—This species was originaly described by Minato(1952, p. 155, pl. 5, fig. 7) as Spirifer kozuboensis Minato, 1952.
The Kitakami species is assigned to the genus Unispirifer,because of the mucronate cardinal extremities and numerous
simple costae on the ventral valve. Unispirifer tornacensis (deKoninck, 1883), redescribed by Sartenaer and Plodowski (1996,
p. 60, pl. 1, figs. 1–10; pl. 2, figs. 11–19; pl. 3, figs. 27–41; pl. 4,
figs. 42–58; pl. 5, figs. 60–73) as Atylephorus tornacensis (deKoninck, 1883), from the upper Tournaisian of Belgium,
somewhat resembles U. kozuboensis; but the Belgian species isdistinguished from the latter by the larger dimensions and inhaving less numerous, stronger costae on the ventral valve.
Occurrence.—ST3 Unit (Okuhinotsuchi) and AR2 Unit
(Yokota).
Distribution.—Upper Tournaisian–lower Visean: northeasternJapan (Hikoroichi, Yokota, Okuhinotsuchi and Nagasaka in theSouth Kitakami Belt).

Unispirifer kozuboensis (Minato, 1952)(Figs. 26G, H, 31F‒I)

Material.—Two specimens from locality IST1, internalmoulds of two ventral valves, IGPS111734, 111735.
Remarks.—These specimens were described by Tazawa and
Iryu (2019, p. 104, fig. 7B) as Unispirifer sp. The specimens from
Shimoarisu are safely assigned to the genus Unispirifer on thebasis of alate cardinal extremities of the ventral valves. The
Shimoarisu species resembles Unispirifer mediocris(Tolmatchoff, 1924, p. 181, 561, pl. 11, figs. 13, 14), from theTournaisian of the Kuznetsk Basin, central Russia, in the small
size (length 19 mm, width about 25 mm in the larger specimen,
IGPS111734) and transverse outline. However, accuratecomparison is difficult for the poorly preserved specimens.
Unispirifer kozuboensis (Minato, 1952, p. 155, pl. 5, fig. 7), fromthe D0 Zone of the Jumonji Stage (= middle part of the Arisu
Formation) of the Yokota area, South Kitakami Belt, differs fromthe present species in the larger size and more transverse outline.
Occurrence.—AR2 Unit (Shimoarisu).

Spirifer kozuboensis Minato, 1952, p. 155, pl. 5, fig. 7; Minato
et al., 1979a, pl. 18, fig. 5.
Unispirifer kozuboensis (Minato). Tazawa and Kurita, 1986, p.
167, fig. 2.1; Tazawa, 2018c, p. 66, fig. 28A‒E; Tazawa andKurita, 2019a, p. 223, fig. 4G, H; Tazawa in Tazawa et al.,2019, p. 64, fig. 15A, B.
Material.—Eight specimens from localities KAR1, KF179,
KF181, KYT2 and Otsubosawa (exact location unknown) in
Yokota: (1) internal mould of a conjoined shell, with externalmould of the ventral valve, UHR15991 (holotype); (2) internalmould of a conjoined shell, IGPS112091; (3) external andinternal moulds of a ventral valve, IGPS112126; (4) internalmoulds of two ventral valves, IGPS99008, 111761; and (5)internal moulds of three dorsal valves, IGPS112093, 112113,
112127.
Description.—Shell medium in size for genus, wider thanlong, with greatest width at hinge; cardinal extremities
mucronate; length 28 mm, width about 50 mm in the holotype(UHR15991). Ventral valve moderately and unevenly convex inlateral profile, most convex at umbonal region; umbo small,incurved; sulcus narrow and shallow, not clearly demarcatedfrom lateral flanks; ears moderately large, triangular andproduced; interarea low, apsacline. External surface of ventral
valve ornamented with numerous simple costae, irregularconcentric rugae and very fine closely spaced growth lines;
costae becoming broad and flat anteriorly and numbering 7–8 in
10 mm on anterior margin. Ventral interior with stout divergent

Unispirifer sp.
(Fig. 31E)
Unispirifer sp. Tazawa and Iryu, 2019, p. 104, fig. 7B.

Family CHORISTITIDAE Waterhouse, 1968
Subfamily ANGIOSPIRIFERINAE Legrand-Blain, 1985
Genus BRACHYTHYRINA Fredericks, 1929
Type species.—Spirifer strangwaysi de Verneuil, 1845.
Brachythyrina sp.
(Fig. 37A, B)
Brachythyrina nagaoi Minato, 1951, p. 371, pl. 1, figs. 2, 10; pl.
3, fig. 2; pl. 4, fig. 4; Minato et al., 1979a, pl. 20, fig. 3.

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

65

FIGURE 42. Brachiopods of the OD2 assemblage (5). A‒C, Syringothyris sp.; A, external latex cast (A1) and external mould (A2) of posterior portion ofventral valve, NU-B2246; B, internal mould of ventral valve, NU-B2247; C, internal mould of ventral valve, NU-B2248; D, Asyrinxia nipponotrigonalis(Minato), external latex cast of dorsal valve, NU-B2264; E, Asyrinxia sp., posterior (E 1) and ventral (E2) views of internal mould of ventral valve,
IGPS97661; F‒H, Dimegelasma sp.; internal moulds of ventral valves, NU-B2241 (F), NU-B2240 (G) and IGPS96136 (H); I, Brachythyris chouteauensis(Weller), external latx cast (I1) and internal mould (I2) of ventral valve, UHR16140; J, Imbrexia incertus (Hall), ventral (J1) and dorsal (J2) views ofexternal latx cast, and ventral (J3) and dorsal (J4) views of internal mould of conjoined shell, UHR30191. Scale bars are 1 cm.

Material.—Two specimens from locality KF207: (1)external and internal moulds of a dorsal valve, UHR16178; and(2) internal mould of a dorsal valve, UHR15993.
Remarks.—These specimens were described by Minato (1951,
p. 371, pl. 1, figs. 2, 10; pl. 3, fig. 2; pl. 4, fig. 4) as Brachythyrina
nagaoi Minato, 1951, from the E 0 Zone (in the AR3 Unit) of
Yokota. However, the specimens from Yokota are inadequate forestablishing the new species because of ill preservation. Minato(1951, p. 371) described as follows, “the Japanese form differsfrom the genotype in having a narrow fold, and simpler mediancomplex of the fold”. The dorsal fold is wide and having three orfour faint sulci. The Kitakami species somewhat resembles
Brachythyrina lata Chao (1929), described by Chao (1929, p. 63,
pl. 8, figs. 9–14) as Brachythyrina strangwaysi var. lata Chao,

1929, from the Moscovian–Asselian of Gansu, northwestern
China and from Shanxi, northern China, in having medium,transverse dorsal valve (length 14 mm, width 31 mm in the better
preserved specimen, UHR16178). The type species, Spirifer
strangwaysi de Verneuil, 1845, redescribed by Sokolskaya (inSarytcheva and Sokolskaya, 1952, p. 189, pl. 53, fig. 305) from
the Moscovian—Gzhelian of the Moscow Basin, western Russia,
differs from the present species in having distinct mucronate
cardinal extremities.
Occurrence.—AR3 Unit (Yokota).
Family IMBREXIIDAE Carter, 1992
Genus IMBREXIA Nalivkin, 1937

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JUN-ICHI TAZAWA

Type species.—Spirifer imbrex Hall, 1858.
Imbrexia forbesi (Norwood and Pratten, 1855)(Fig. 47A, B)
Spirifer forbesi Norwood and Pratten, 1855, p. 73, pl. 9, fig. 3;
Weller, 1914, p. 331, pl. 42, figs. 1–3; pl. 43, fig. 16; pl. 83,
figs. 1, 2.
Spirifer (Imbrexia) forbesi Norwood and Pratten. Nalivkin,
1937, p. 106, pl. 30, fig. 5.
Imbrexia forbesi (Norwod and Pratten). Armstrong, 1962, p. 52,
pl. 7, figs. 18–21, text-fig. 30; Grechishnikova, 1966, p. 149,
pl. 15, figs. 9, 10; Nalivkin, 1979, p. 138, pl. 49, fig. 6; Lee etal., 1980, p. 410, pl. 154, fig. 14.
Material.—Three specimens from locality YUK1: (1)internal mould of a ventral valve, KCG80; (2) external andinternal moulds of a dorsal valve, KCG81; and (3) internalmould of a dorsal valve, KCG82.
Remarks.—These specimens are referred to Imbrexia forbesi(Norwood and Pratten, 1855), redescribed by Armstrong (1962, p.
52, pl. 7, figs. 18–21, text-fig. 30), from the Keating Formatin(lower Osagean) of New Mexico and Arizona, in the medium,much transverse shell (length about 15 mm, width about 40 mm
in the largest specimen, KCG80), with wide ventral sulcus anddorsal fold, and external ornament consisting of numerous simple
or bifurcate costae (numbering 5–6 in 5 mm at midlength of
dorsal valve) and imbricate growth lamellae. Imbrexia ussuilensis
Nalivkin (in Garanj et al., 1975, p. 188, pl. 84, figs. 8, 9), from theKosvinsky Horizon (lower Visean) of the southern Urals, central
Russia, differs from I. forbesi in the less transverse outline.
Occurrence.—OD 3 Unit (Yahagi).
Distributin.—Upper Tournaisian–upper Visean: northeasternJapan (Yahagi in the South Kitakami Belt), USA (New Mexicoand Arizona), central Russia (southern Urals), Kazakhstan andnortheastern China (Heilongjiang).
Imbrexia incertus (Hall, 1858)(Figs. 42J, 47C)
Spirifer incertus Hall, 1858, p. 602, pl. 13, fig. 3; Weller, 1914,
p. 355, pl. 41, figs. 6–14.
Spirifer (Imbrexia) incertus Hall. Nalivkin, 1937, p. 107, pl. 30,
fig. 4.
Imbrexia cf. incertus (Hall). Minato and Kato, 1977, p. 614, pl.
1, figs. 2–11.
Material.—Three specimens from locality YUK1 and north of
Mt. Karoyama (exact location unknown), Nisawa: (1) external
and internal moulds of a conjoined shell, UHR30191; (2) internalmould of a conjoined shell, with external mould of the dorsal
valve KCG83; and (3) external mould of a ventral valve, KCG84.
Remarks.—The best preserved specimen (UHR30191), fromthe Odaira Formation of Nisawa, was described by Minato andKato (1977, p. 614, pl. 1, figs. 2–11) as Imbrexia cf. incertus(Hall, 1858). But this specimen can be referred to Imbrexia
incertus (Hall, 1858), redescribed by Weller (1914, p. 355, pl.
41, figs. 6– 14.), from the Burlington Limestone of theMississippi Valley, the USA, in size, shape and external
ornament of both valves, particularly in having numerous
relatively fine costae (numbering 5 in 5 mm at midlength) on theventral valve. The type species, Imbrexia imbrex (Hall, 1858),
redescribed by Carter (1974, p. 687, pl. 1, figs. 1–7; pl. 4, figs.
1–4, text-fig. 3) from the Burlington Limestone of Iowa, differsfrom I. incertus in having coarser costae on the ventral valve.
Occurrence.—OD2 Unit (Nisawa) and OD3 Unit (Yahagi).
Distribution.—Upper Tournaisian–upper Visean: northeasternJapan (Yahagi and Nisawa in the South Kitakami Belt), USA(Iowa) and Kazakhstan.
Superfamily PAECKELMANELLOIDEA Ivanova, 1972
Family STROPHOPLEURIDAE Carter, 1974
Subfamily STROPHOPLEURINAE Carter, 1974
Genus ACUMINOTHYRIS Roberts, 1963
Type species.—Acuminothyris triangularis Roberts, 1963.
Acuminothyris triangularis Roberts, 1963
(Fig. 32A–C)
Acuminothyris triangularis Roberts, 1963, p. 15, pl. 5, figs. 1–8,
text-figs. 7, 8; Tazawa, 2018c, p. 68, fig. 28L.
“Spirifer” mundulus Rowley. Minato and Kato, 1977, p. 615, pl.
1, fig. 1; Minato et al., 1979a, pl. 14, fig. 2.
Material.—Three specimens from localities KF180, KYT3
and KYT4, internal moulds of three dorsal valves, IGPS112114,
112128, 112129.
Remarks.—These specimens can be referred to Acuminothyris
triangularis Roberts (1963, p. 15, pl. 5, figs. 1–8, text-figs. 7,8), from the Bingleburra Formation (upper Tournaisian) of
Lewinsbrook, New South Wales, eastern Australia, by the large
size (length 13 mm, width about 57 mm in the largest specimen,
IGPS112114), very transverse outline, moderately broad simple
fold, numerous costae on lateral slopes, and fine numerous
concentric lamellae on anterior portion of fold. A single ventral
valve specimen, described by Minato and Kato (1977, p. 615, pl.
1, fig. 1) as “Spirifer” mundulus Rowley (1893) from the lower
part of the Hikoroichi Formation in the Hikoroichi area, isidentical with the present species, although the Hikoroichi
specimen is smaller in size.
Occurrence.—AR2 Unit (Yokota).
D i s t r i b u t i o n . —U p p e r T o u r n a i s i a n – l o w e r V i s e a n :northeastern Japan (Hikoroichi and Yokota in the South
Kitakami Belt) and eastern Australia (New South Wales)
Superfamily BRACHYTHYRIDOIDEA Fredericks, 1924
Family BRACHYTHYRIDIDAE Fredericks, 1924
Genus BRACHYTHYRIS M’Coy, 1844

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

67

FIGURE 43. Brachiopods of the OD3 assemblage (1). A, Leptagonia analoga (Phillips), external latex cast (A1, A2) of ventral valve, NU-B2154; B, C,
Rugosochonetes extensus (Chao); B, external mould (B1, B2) and internal mould (B3) of dorsal valve, NU-B2172; C, external latex cast (C1, C2) andinternal mould (C3) of ventral valve, KCG76; D, Rugosochonetes sp.; external latex cast (D1, D2) of ventral valve, KCG78; E, Plicatifera pseudoplicatilis(Muir-Wood); external mould (E1, E2) of dorsal valve, KCG70; F, Plicatifera plicatilis (Sowerby); external latex cast (F1, F2) of ventral valve, KCG65; G,
Argentiproductus sp.; external latex cast (G1, G2) and external mould (G3) of dorsal valve, KCG63; H‒J, Alitaria konincki (Muir-Wood and Cooper); H,
ventral (H1, H2) and dorsal (H3) views of internal mould, and dorsal external mould (H4) of conjoined shell, KCG71; I, internal mould of ventral valve,
KCG73; J, internal mould of ventral valve, KCG72. Scale bars are 1 cm.

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JUN-ICHI TAZAWA

Type species.—Spirifera ovalis Phillips, 1836.
Brachythyris chouteauensis (Weller, 1909)(Figs. 32D, E, 42I, 47D)
Spirifer chouteauensis Weller, 1909, p. 305, pl. 13, fig. 11.
Brachythyris chouteauensis (Weller). Weller, 1914, p. 373, pl.
57, figs. 4–11; Branson, 1938, p. 65, pl. 6, figs. 1–4; Sokolskayain Sarytcheva and Sokolskaya, 1952, p. 193, pl. 54, fig. 312;
Carter, 1967, p. 394, pl. 39, figs. 1–9, text-figs. 37, 38;
Bublichenko, 1976, p. 123, pl. 14, figs. 1–5; Carter, 1987, p.
82, pl. 26, figs. 7, 8; Liu, 1988, p. 367, pl. 2, fig. 11; Carter,
1999, p. 129, figs. 18M–X.
Sirifer ovalis Phillips. Tolmatchoff, 1924, p. 186, 562, pl. 11,
figs. 17, 18.
Brachythyris? sp. Minato, 1951, p. 368, pl. 5, fig. 9.
Brachythyris aff. pinguis (Sowerby). Minato, 1951, p. 369, pl. 1,
figs. 14, 15; pl. 3, fig. 8; Minato et al., 1979a, pl. 21, figs. 9, 10.
Brachythyris krapivnensis Beznossova, 1959, p. 123, pl. 10, fig.
6; pl. 11, fig. 14, text-figs. 54, 55; Besnssova in Sarytcheva etal., 1963, p. 297, pl. 57, fig. 4.
Brachythyris sp. Minato et al, 1979a, pl. 17, fig. 9.
Material.—Seven specimens from localities YUK1, 808 mhill in Nisawa and Otsubosawa in Yokota (exact location
unknown): (1) internal mould of a conjoined shell, KCG74; (2)external and internal moulds of a ventral valve, UHR16140; (3)internal moulds of three ventral valves, UHR16011, 16250,
16252; (4) external and internal moulds of a dorsal valve,
KCG75; and (5) internal mould of a dorsal valve, UHR16012.
Remarks.—Most of the specimens, except for two specimens(KCG74, 75) from the uppermost part of the Odaira Formatin in
Yahagi, were described by Minato (1951, p. 368–369) as
Brachythyris? sp. and Brachythyris aff. pinguis (Sowerby).
However, all the specimens are referred to Brachythyris
chouteauensis (Weller, 1909), redescribed by Carter (1967, p.
394, pl. 39, figs. 1–9, text-figs. 37, 38) from the Chappel
Limestone of Texas, in the medium size (length 26 mm, width
34 mm in the largest specimen, UHR16140), rounded outline
and in having one or two pairs of weak costae in the ventral
sulcus and a distinct median groove in the dorsal fold.
Brachythyris suborbicularis (Hall, 1858), redescribed by Weller(1914, p. 374, pl. 61, figs. 1–8; pl. 62, figs. 1–12) from the
Burlington and Keokuk limestones of Iowa, Missouri andIndiana in the USA, differs from B. chouteauensis in the larger
dimensions. Brachythyris pinguis (Sowerby, 1820), redescribedby George (1927, p. 108, text-fig. 1) from the upper Visean ofGower, South Wales, is readily distinguished from the present
species by the much larger and more transverse shell withbroader costae on lateral flanks of both valves.
Occurrence.—AR2 Unit (Yokota), OD2 Unit (Nisawa) andOD3 Unit (Yahagi).
Distribution.—Lower Tournaisian–upper Visean: northeasternJapan (Yokota, Yahagi and Nisawa in the South Kitakami Belt),
USA (Missouri, Oklahoma and Texas), Canada (Alberta),

western Russia (Moscow Basin), central Russia (Kuznetsk
Basin), Kazakhstan and northeastern China (Jilin).
Suborder DELTHYRIDINA Ivanova, 1972
Superfamily RETICULARIOIDEA Waagen, 1883
Family ELYTHIDAE Fredericks, 1924
Subfamily ELYTHINAE Fredericks, 1924
Genus KITAKAMITHYRIS Minato, 1951
Type species.—Torynifer (Kitakamithyris) tyoanjiensis
Minato, 1951.
Kitakamithyris hikoroitiensis Minato, 1951
(Fig. 32F‒I)
Torynifer (Kitakamithyris) hikoroitiensis Minato, 1951, p. 375,
pl. 1, fig. 1.
Kitakamithyris hikoroitiensis Minato. Minato, 1952, p. 171, pl.
7, fig. 3; pl. 8, fig. 6; Minato et al., 1979a, pl. 16, fig. 1; Tazawa,
2018c, p. 72, figs. 20B, C, 21B, 29A, B; Tazawa and Iryu,
2019, p. 104, figs. 7C, 8; Tazawa and Ibaraki, 2019, p. 20, fig. 5C.
Kitakamithyris semicircularis Minato. Minato, 1952, p. 171, pl.
7, fig. 6; pl. 8, fig. 5; pl. 10, fig. 3; Minato et al., 1979a, pl.
15, fig. 1.
Material.— Five specimens from localities IST1, IYK1 and
KF179: (1) internal mould of a conjoined shell, with externalmould of the dorsal valve, IGPS112107; (2) extermal andinternal moulds of a ventral valve, IGPS112108; (3) externalmould of a ventral valve, IGPS111727; and (4) external mouldof two dorsal valves, IGPS112109, NU-B2257.
Remarks.—Two specimens, a ventral valve specimen(IGPS111727) from Shimoarisu and a dorsal valve specimen(NU-B2257) from Yokota, were described by Tazawa and Iryu
(2019, p. 104, figs. 7C, 8) and by Tazawa and Ibaraki (2019, p.
20, fig. 5C), as Kitakamithyris hikoroitiensis Minato, 1951. The
other three specimens (IGPS112107–112109) from Yokota arealso referred to Kitakamithyris hikoroitiensis Minato (1951, p.
375, pl. 1, fig. 1), originally described from the lower part of the
Hikoroiti Series (= Choanji Formation of Tazawa and Niikawa,2018) of Tyoanji (Choanji), South Kitakami Belt, on account of
the relatively large and sporadically arranged spine bases(numbering 5–6 in 5 mm at about midlength) on both ventral
and dorsal vaves. Kitakamithyris semicircularis Minato (1952,
p. 171, pl. 7, fig. 6; pl. 8, fig. 5; pl. 10, fig. 3), from the Choanji,
Hikoroichi and Arisu Formations of the South Kitakami Belt, isdeemed to be a junior synonym of K. hikoroitiensis. The typespecies, Kitakamithyris tyoanjiensis Minato (1951, p. 374, pl. 1,
fig. 3; pl. 4, fig. 7), from the Choanji Formation (Upper
Devonian) of Choanji is readily distinguished from K.
hikoroitiensis by the much smaller and tightly arranged spine
bases on the ventral valve.
Occurrence.—AR2 Unit (Shimoarisu and Yokota).
Distribution.—Upper Devonian (upper Famennian)–lower
Visean: northeastern Japan (Choanji, Hikoroichi, Shimoarisu

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS69

FIGURE 44. Brachiopods of the OD3 assemblage (2). A, Marginicinctus marginicinctus (Prout), internal mould (A1, A2) of ventral valve, NU-B2330; B‒
D, Marginatia burlingtonensis (Hall); B, ventral (B1) and anterior (B2) views of external latex cast of ventral valve, NU-B2160; C, external latex cast of
dorsal valve, NU-B2162; D, anterior view (D1, D2) of internal mould of ventral valve, NU-B2161; E, F, Pustula pustulosa (Phillips); E, external latex castof ventral valve, NU-B2158; F, external latex cast of dorsal valve, NU-B2159; G, H, Echinoconchus punctatus (Sowerby); G, internal mould (G1) andexternal latex cast (G2) of ventral valve, NU-B2155; H, external mould (H1) and internal latex cast (H2, H3) of dorsal valve, NU-B2156. Scale bars are 1
cm.

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JUN-ICHI TAZAWA

and Yokota in the South Kitakami Belt).
Kitakamithyris sp.
(Figs. 46G, 47E).
Kitakamithyris sp. Tazawa, 1979, p. 11, fig. 11.4; Tazawa, 2017,p. 340, fig. 8.7.
Material.—Two specimens from locality KF159 and YUK1:(1) external and internal moulds of a ventral valve, KCG79; and(2) internal mould of a ventral valve, NU-B2153.
Remarks.—One of the specimens (NU-B2153) was
described by Tazawa (2017, p. 340, fig. 8.7) as Kitakamithyris
sp. The fragmentarily preserved specimen from Tairagai can be
assigned to the genus Kitakamithyris by its medium (length
about 18 mm, width about 32 mm), subelliptical and gently
convex ventral valve, ornamented with numerous radiating
grooves on the internal surface, and the presence of a long
median septum and a pair of divergent adminicula. The anotherspecimen (KCG79) from Yukisawa in the Yahagi area, also canbe assigned to the genus Kitakamithyris by the presence ofnumerous concentric rows of biramous spine bases (numbering
9–10 spine bases in 5 mm at midlength) on the ventral valve.
These specimens may be a new species of Kitakamithyris, but
the poorly preserved specimens preclude that determination.
Occurrence.—OD3 Unit (Yokota and Yahagi).
Subfamily TORYNIFERINAE Carter in Carter et al., 1994
Genus TORYNIFER Hall and Clarke, 1894
Type species.—Spirifer pseudolineatus Hall, 1858.
Torynifer asiatica Beznossova in Beznossova et al., 1962
(Fig. 47F)
Reticularia lineata Martin. Tolmatchoff, 1924, p. 191, 563, pl.
12, figs. 1–3, 5.
Reticularia sinuata Tolmatchoff, 1924, p. 194, 564, pl. 12, figs. 4, 7.
Reticularia pseudolineata Hall. Nalivkin, 1937, p. 115, pl. 2,
figs. 5, 6; pl. 33, fig. 6; Rotai, 1941, p. 114, pl. 28, figs. 2–4;
Simorin, 1956, p. 198, pl. 17, figs. 10, 11.
Torynifer pseudolineatus asiaticus Beznossova in Beznossova etal., 1962, p. 176, pl. C-23, figs. 4–5; Beznossova in Sarytcheva
et al., 1963, p. 303, pl. 58, figs. 1–6, text-figs. 133‒135;
Kotlyar and Popeko, 1967, p. 173, pl. 48, figs. 3–6; Plodowski,
1970, p. 106, pl. 8, fig. 2; Yang and Fan, 1983, p. 279, pl. 4,
fig. 17; Zhang et al., 1983, p. 368, pl. 124, fig. 7.
Torynifer pseudolineata (Hall). Litvinovich in Litvinovich et al.,1969, p. 270, pl. 66, figs. 7, 8.
Torynifer asiatica Beznossova. Pavlova, 1969, p. 63, pl. 4, figs.
1–6; pl. 11, figs. 6, 7; pl. 13, fig. 3, text-figs. 37–39; Tazawa,
2018c, p. 72, figs. 32A, B.
Material.—One specimen from locality KF210, external andinternal moulds of a ventral valve, NU-B2332.

Remarks.—This specimen is referred to Torynifer asiatica
Beznossova in Beznossova et al. (1962, p. 176, pl. C-23, figs.
4–5), from the upper Tournaisian of the Kuznetsk Basin, central
Russia, on account of the external ornament of the ventral valve
consisting of slightly irregular concentric lamellae bearing a row
of fine biramous spine bases (numbering 7 in 5 mm at about
midlength) and the long thin median septum in the ventral valve.
The type species, Torynifer pseudolineata (Hall, 1858),
redescribed by Weller (1914, p. 429, pl. 74, figs. 1–11; pl. 75,
fig. 20) from the Keokuk Limestone of the Mississippi Valley,differs from T. asiatica in having more prominent ventral
sulcus.
Occurrence.—OD3 Unit (Yahagi).
Distribution.—Lower Tournaisian–Serpukhovian: northeasternJapan (Hikoroichi and Yahagi in the South Kitakami Belt),
Afghanistan, central Russia (Kuznetsk Basin and Transbaykal),
Kazakhstan and northwestern China (Xinjiang).
Order SPIRIFERINIDA Ivanova, 1972
Suborder SPIRIFERINIDINA Ivanova, 1972
Superfamily SYRINGOTHYRIDOIDEA Fredericks, 1926
Family SYRINGOTHYRIDIDAE Fredericks, 1926
Subfamily SYRINGOTHYRIDINAE Fredericks, 1926
Genus SYRINGOTHYRIS Winchell, 1863
Type species.—Syringothyris typa Winchell, 1863.
Syringothyris texta (Hall, 1857)(Figs. 33A, B, 34C)
Spirifer textus Hall, 1857, p. 169.
Syringothyris textus (Hall). Weller, 1914, p. 399, pl. 69, figs.
6–9; pl. 70, figs. 1–4; pl. 71, figs. 1, 2.
Syringothyris transversa Minato, 1951, p. 377, pl. 5, fig. 1;
Minato, 1952, p. 167, pl. 11, fig. 5 only; Minato et al., 1979a,
pl. 21, fig. 11; Tazawa, 2002, fig. 7.6; Tazawa, 2006, p. 134,
figs. 7.1, 7.2.
Syringothyris texta (Hall). Tazawa, 2018a, p. 2, fig. 1.1; Tazawa
and Iryu, 2019, p. 105, fig. 7H.
Material.—Three specimens from localities IST1 and KF181,
and from the Nashirozawa Valley (exact location unknown) in
Shimoarisu: (1) external and internal moulds of a conjoined
shell, IGPS112110; (2) internal mould of a conjoined shell, withexternal mould of the ventral interarea, UHR16925; and (3)internal mould of a ventral valve, IGPS111710.
Remarks.—One of the specimens (UHR16925) was
described by Minato (1951, p. 377, pl. 5, fig. 1) as the holotype
of Syringothyris transversa Minato, 1951. But the Kitakami
species is, as discussed by Tazawa (2018a, p. 3), deemed to be ajunior synonym of Syringothyris texta (Hall, 1857), from the
Keokuk Limestone of Missouri and from the Knobstone
Formation of Indiana, in the large, transverse shell (length 50
mm, width about 84 mm) and in having a relatively low andslightly concave ventral interarea with a syrinx in the

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71

FIGURE 45. Brachiopods of the OD3 assemblage (3). A, Schellwienella radialis (Phillips), external latex cast (A1, A2) and internal mould (A3) of ventral
valve, NU-B2152; B, Rhipidomella michelini (Léveillé), ventral (B1, B2) and dorsal (B3) views of internal mould of conjoined shell, KCG67; C, Echinariasp., external mould (C1), external latex cast (C2) and internal mould (C3, C4) of dorsal valve, NU-B2157; D, Fluctuaria undata (Defrance), external latex
cast (D1, D2) of ventral valve, NU-B2331; E, Orthotetes keokuk (Hall), external latex cast of ventral valve, KCG69. Scale bars are 1 cm.

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JUN-ICHI TAZAWA

delthyrium. Syringothyris altaica Tolmatchoff (1924, p. 162,
555, pl. 8, figs. 9‒11; pl. 9, fig. 1), from the upper Tournaisianof the Kuznetsk Basin, central Russia, differs from S. texta inhaving blunt cardinal extremities.
Occurrence.—AR2 Unit (Shimoarisu and Yokota).
Distribution.—Lower Visean: northeastern Japan (Shimoarisu
and Yokota in the South Kitakami Belt) and USA (Indiana andMissouri).
Syringothyris platypleura Weller, 1914
(Fig. 34A, B)
Syringothyris platypleurus Weller, 1914, p. 397, pl. 72, figs.
1–4.
Syringothyris sp. Minato, 1952, p. 165, pl. 5, fig. 2; pl. 9, fig. 1.
Syringothyris platypleura Weller. Tazawa and Iryu, 2019, p.
105, fig. 7D, E.
Material.—Two specimens from locality IST1, internalmoulds of two ventral valves, IGPS111711, 111712.
Remarks.—These specimens were described by Tazawa and
Iryu (2019, p. 105, fig. 7D, E) as Syringothyris platypleura
Weller, 1914. The specimens from Shimoarisu are referred to S.
platypleura Weller (1914, p. 397, pl. 72, figs. 1–4), from the
Burlington Limestone of the Mississippi Valley, in the
subpyramidal ventral valve with a very high, flat interarea andnearly flat flanks. Syringothyris sp. Minato (1952, p. 165, pl. 5,
fig. 2; pl. 9, fig. 1), from the Jumonji Stage (= middle part of the
Arisu Formation) of the Yokota area, South Kitakami Belt, isdeemed to be a junior synonym of the present species.
Syringothyris kitakamiensis Minato (1952, p. 165, fig. 5), fromthe Arisu Formation of the Yokota area, South Kitakami Belt,
differs from S. platypleura in having a high but slightly concave
ventral interarea.
Occurrence.—AR2 Unit (Shimoarisu and Yokota).
Distribution.—Upper Tournaisian‒lower Visean: northeasternJapan (Shimoarisu and Yokota in the South Kitakami Belt) andUSA (Missouri).
Syringothyris sp.
(Fig. 42A–C)
Material.—Three specimens from locality KF158: (1)external mould of a ventral interarea, NU-B2246; and (2)internal moulds of two ventral valves, NU-B2247, 2248.
Remarks.—These specimens are fragmentarily preserved,
but can be assigned to the genus Syringothyris by the large size,
high and flat ventral interarea, and well-developed divergent
dental plates in the ventral valve. However, specificidentification is difficult for the poorly preserved specimens.
Occurrence.—OD2 Unit (Yokota).
Subfamily PERMASYRINXINAE Waterhouse, 1986
Genus ASYRINXIA Campbell, 1957

Type species.—Spirifera lata M’Coy, 1847.
Asyrinxia nipponotrigonalis (Minato, 1951)(Figs. 35A, B, 42D)
Fusella nipponotrigonalis Minato, 1951, p. 372, pl. 2, fig. 5;
Minato, 1952, p. 160, pl. 5, fig. 1; pl. 6, fig. 6; pl. 11, fig. 3;
Minato et al., 1979a, pl. 22, fig. 4.
Fusella nipponotrigonalis var. minor Minato, 1952, p. 160, pl.
6, fig. 3.
Asyrinxia sp. Tazawa, 1981b, p. 74, pl. 5, fig. 14.
Asyrinxia nipponotrigonalis (Minato). Tazawa, 2018a, p. 4, figs.
3.1, 3.2; Tazawa and Ibaraki, 2019, p. 20, fig. 5B.
Material.—Five pecimens from localities IYK3 and KYT3
and from the Otsubosawa Valley (exact location unknown) in
Yokota: (1) internal moulds of a conjoined shell, UHR16018(holotype); (2) external and internal moulds of a dorsal valve,
UHR16016; and (3) internal moulds of three dorsal valves,
IGPS112130, 112131, NU-B2264.
Remarks.—Most of the specimens were described by Minato(1951, p. 372, pl. 2, fig. 5) as the type specimens of Fusella
nipponotrigonalis Minato, 1951. However, as suggested byCampbell (1957, p. 83–84), the Kitakami species is assigned to
the genus Asyrinxia, in the large size and in having moderately
high, concave ventral interarea, and lacking syrinx. Twosyringothyridid species from the Lower Carbniferous of theSouth Kitakami Belt, Fusella nipponotrigonalis var. minor
Minato (1952, p. 160, pl. 6, fig. 3), from the Jumonji and Maide
series (= middle and upper parts of the Arisu Formation) of
Yokota and Asyrinxia sp. Tazawa (1981b, p. 74, pl. 5, fig. 14),from the Karoyama Formation (= Odaira Formation) of Nisawa,
are junior synonyms of the present species. The type species,
Asyrinxia lata (M’Coy, 1847), redescribed by Campbell (1957,
p. 81, pl. 16, figs. 1–9) from the middle Visean of Babbinboon,
N ew South Wales , eas tern A us tralia, differs fr om A.
nipponotrigonalis by the more transverse outline.
Occurrence.—AR2 Unit (Yokota) and OD2 Unit (Nisawa).
Distribution.—Lower–upper Visean: northeastern Japan(Shimoarisu, Yokota and Nisawa in the South Kitakami Belt).
Asyrinxia sp.
(Fig. 42E)
Asyrinxia sp. Tazawa, 1981b, p. 74, pl. 5, fig. 14.
Material.—One specimen from locality INS3, internal mouldof a ventral valve, IGPS97661.
Remarks.—This specimen was described by Tazawa (1981b,
p. 74, pl. 5, fig. 14) as Asyrinxia sp. The Nisawa specimen
somewhat resembles Asyrinxia lata (M’Coy, 1847), redescribedby Campbell (1957, p. 81, pl. 16, figs. 1–9) from the middle
Visean of New South Wales, eastern Australia, in size, shapeand internal structure of the ventral valve. But accuratecomparison is difficult for the poorly preserved specimen.

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS73

FIGURE 46. Brachiopods of the OD3 assemblage (4). A‒E, Spirifer liangchowensis Chao; A, external latex cast (A1) and internal mould (A2) of ventral
valve, NU-B2166; B, external latex cast (B1) and internal mould (B2) of dorsal valve, NU-B2170; C, external latex cast of ventral valve, NU-B2167; D,internal mould of ventral valve, NU-B2168; E, internal mould of dorsal valve, NU-B2171; F, Pseudosyrinx sp., external latex cast of ventral interarea,
NU-B2163; G, Kitakamithyris sp., internal mould (G1, G2) of ventral valve, NU-B2153; H, Cleiothyridina fimbriata (Phillips), external latex cast (H1, H2)and enlarged microornamentation (H3) of dorsal valve, NU-B2164. Scale bars are 1 cm, except for H3.

Occurrence.—OD2 Unit (Nisawa).
Genus PSEUDOSYRINX Weller, 1914
Type species.—Pseudosyrinx missouriensis Weller, 1914.
Pseudosyrinx jumonjiensis (Minato, 1951)(Fig. 35C, D)

Syringothyris jumonjiensis Minato, 1951, p. 376, pl. 2, fig. 1;
Minato et al., 1979a, pl. 21, fig. 12.
Pseudosyrinx jumonjiensis (Minato). Tazawa, 2018a, p. 4, fig.
2.1; Tazawa and Iryu, 2019, p. 105, fig. 7F, G.
Material.—Four specimens from localities IST1, KF181 andfrom Shimoarisu (exact location unknown): (1) external mouldof a ventral valve, IGPS111713; and (2) external mould of threedorsal valves, IGPS111714, 112090, UHR15995 (holotype).

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JUN-ICHI TAZAWA

Remarks.—The holotype (UHR15995) of this species was
described by Minato (1951) as Syringothyris jumonjiensis
Minato, 1951. But the genus of this species, as suggested by
Tazawa (2018a, p. 4), should be replaced to Pseudosylinx, forthe presence of high, flat ventral interarea and lacking syrinx inthe delthyrium. The present species most resembles
Pseudosyrinx missouriensis Weller (1914, p. 406, pl. 65, figs.
5–9; pl. 66, figs. 11–13), from the Burlington Limestone of theMississippi Valley, in general shape, but the American species
is much smaller in size. Pseudosyrinx sampsoni (Weller, 1909,
p. 311, pl. 14, fig. 4), from the Fern Glen Formation of
Missouri, is also a large-sized Pseudosyrinx species, but differsfrom P. jumonjiensis in having fewer and stronger costae on thedorsal valve.
Occurrence.—AR2 Unit (Shimoarisu and Yokota).
D i s t r i b u t i o n . —L o w e r V i s e a n : n o r t h e a s t e r n J a p a n(Shimoarisu and Yokota in the South Kitakami Belt).
Pseudosyrinx sp.
(Fig. 46F)
Pseudosyrinx sp. Tazawa, 2017, p. 340, fig. 8.6.
Material.—One specimen from locality KF159, externalmould of ventral interarea, NU-B2163.
Remarks.—The single specimen from Tairagai wasdescribed by Tazawa (2017, p. 340, fig. 8.6) as Pseudosyrinx sp.
This specimen can be assigned to the genus Pseudosyrinx
Weller, 1914 in having high interarea with narrowly triangulardelthyrium and the absence of syrinx. The Tairagai species
somewhat resembles the type species, Pseudosyrinx missouriensis
Weller (1914, p. 65, figs. 5–9; pl. 66, figs. 11–13), from the
Burlington Limestone of the Mississippi Valley, by the highinterarea with narrowly triangular delthyrium in the ventral
valve. But accurate comparison is difficult in the fragmentarily
preserved specimen.
Occurrence.—OD3 Unit (Yokota).
Family DIMEGELASMIDAE Carter in Carter et al., 1994
Genus DIMEGELASMA Cooper, 1942
Type species.—Spirifer neglectus Hall, 1858.
Dimegelasma sp.
(Fig. 42F‒H)
Dimegelasma sp. Tazawa and Katayama, 1979, p. 171, pl. 11,
figs. 15, 16; Tazawa, 2018b, p. 74, fig 33A, B.
Tomiopsis sp. Tazawa, 1984b, p. 308, pl. 61, fig. 10.
Material.—Four specimens from locality KF158, internalmoulds of three ventral valves, IGPS96135, 96136, NU-B2240,
2241.
Remarks.—Most of the specimens were previously describedby Tazawa and Katayama (1979, p. 171, pl. 11, figs. 15, 16) as
Dimegelasma sp. In this paper, two specimens (NU-B2240,
2241) from the same locality (KF158) are added to the material.
These specimens can be assigned to the genus Dimegelasma by
the medium size (length more than 18 mm, width more than 30
mm in the largest but fragmentarily preserved specimen,
IGPS96136), gently convex ventral valve, with rounded cardinal
extremities, and by the presence of very long thin dental plates
in the ventral valve. Tomiopsis sp. Tazawa (1984b, p. 308, pl.
61, fig. 10), from the upper part of the Hikoroichi Formation in
the Hikoroichi area, South Kitakami Belt, is conspecific with thepresent species. The Odaira species superficially resembles
Dimegelasma grandicosta (Abramov and Grigorjeva, 1986, p.
162, pl. 29, figs. 4– 6; pl. 30, figs. 1, 2, 10), from the
middle–upper Visean of the Verkhoyansk Range, northern
Russia. Specific identification, however, remains difficult due tothe poor preservation of the material.
Occurrence.—OD2 Unit (Yokota).
Suborder SPIRIFERINIDINA Ivanova, 1972
Superfamily PENNOSPIRIFERINOIDEA Dagys, 1972
Family PUNCTOSPIRIFERIDAE Waterhouse, 1975
Genus PUNCTOSPIRIFER North, 1920
Type species.—Punctospirifer scabricosta North, 1920.
Punctospirifer sp.
(Fig. 37C)
Material.—One specimen from locality KYT1, internalmould of a ventral valve, NU-B2255.
Remarks.—The fragmentarily preserved dorsal valve
specimen from Yokota is safey assigned to the genus
Punctospirifer by the small size, transverse outline (length 8mm, width about 12 mm), smooth, rounded fold andmicroornament of closey spaced imbricated growth laminae.
This specimen resembles shells of Punctospirifer pauciplicatus
Roberts (1971, p. 265, pl. 58, figs. 17–30), from the Utting
Calcarenite of the Bonaparte Gulf Basin, northwestern Australia,
in having 3–4 rounded costae on each lateral slope of the dorsal
valve. But accurate comparison is difficult for the poorly
preserved specimen.
Occurrence.—AR3 Unit (Yokota).
ACKNOWLEDGEMENTS
I sincerely thank Soichiro Kawabe (Fukui Prefectural
Dinosaur Museum, Katsuyama) for editing the manuscript;
Shuzhong Shen (Nanjing University, Nanjing) and an
anonymous reviewer for their constructive comments; Hiroki
Echizenya (Hokkaido University Museum, Sapporo) and Jun
Nemoto (Tohoku University Museum, Sendai) for loan of thetype specimens; Yousuke Ibaraki (Fossa Magna Museum,
Itoigawa) for his help in drawing the figures; late Toshio
Katayama (Nakatsugawa, Gifu Prefecture), Fumio Itabashi

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONS

75

FIGURE 47. Brachiopods of the OD3 assemblage (5). A, B, Imbrexia forbesi (Norwood and Pratten); A, internal mould (A1, A2) of ventral valve, KCG80;
B, external latex cast (B1, B2) and internal mould (B3, B4) of dorsal valve, KCG81; C, Imbrexia incertus (Hall), external latex cast (C1, C2) of ventral valve,
KCG84; D, Brachythyris chouteauensis (Weller), ventral (D1) and dorsal (D2) views of internal mould of conjoined shell, KCG74; E, Kitakamithyris sp.,external mould (E1), internal mould (E2) and enlarged concentric rows of biramous spine bases (E 3) of ventral valve, KCG79; F, Torynifer asiatica
Beznossova, external latex cast (F1), internal mould (F2) and enlarged concentric rows of biramous spine bases (F3) of ventral valve, NU-B2332. Scalebars are 1 cm, except for E3 and F3.

76

JUN-ICHI TAZAWA(Ashikaga, Tochigi Prefecture), Yasufumi Iryu (Tohoku
University, Sendai), Madoka Yamazaki (Aomori) and Yuji
Kurita (Niigata University, Niigata) for help in the field; andHideo Araki (Kesennuma, Miyagi Prefecture) for providing partof the brachiopod specimens.
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有住層
Choanji Formation
長安寺層
Dobasawa
堂場沢“808 m hill”“ 808 m 高地 ”
Hikoroichi Formation
日頃市層
Hina
日南
Hinotsuchigawa River
火の土川
Jyumonji
十文字
Kaisenosawa
カイセノ沢Karaumedate Formation
唐梅館層
Karosawa Formation
加労沢層
Karoyama Formation
加労山層
Kesengawa River気仙川
Kesen-gun
気仙郡
Komatagawa River
小股川
Kominekurasawa
小峰倉沢
Konikurazawa
小仁倉沢
Maide
舞出
Mt. Karoyama
加労山

Surveying Team of Xinjiang, Institute of Geoscience ofXinjiang, and Geological Surveying Group of Petroleum
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Palaeontologica Sinica 51: 416–435.*** : in Japanese** : in Chinese*** : in Russian

地名・地層名Mt. Odairayama
Myojinsawa
Nagaiwa Formation
Nagasaka
Nashirozawa
Nekonosawa
NisawaNisawa Pass
North China Province
Odaira Formation
Okuhinotsuchi
Omatagawa River
Onimaru Formation
Otomo-choOtsubosawa
Rikuzentakata City
Sasanohora Formation
Senbakaya Formation
Shiba Formation

＞
大平山
明神沢
長岩層
長坂
ナシロ沢
猫ノ沢
荷沢
荷沢峠
北中国区
大平層
奥火の土
大股川
鬼丸層
小友町
小坪沢
陸前高田市
笹ノ洞層
仙婆巌層
芝層

Shimoarisu
Shittakazawa Formation
Shizu
Shokarosawa
Soma
Suesawa
Suisawa
Sumita-cho
TairagaiTakaseTono City
YahagiYahagigawa River
Yasumotozawa
Yokokawa River
YokotaYonesato Formation
Yukisawa

下有住
尻高沢層
清水
小加労沢
相馬
末沢
スイ沢
住田町
平貝
高瀬
遠野市
矢作
矢作川
ヤスモト沢
横川
横田
米里層
雪沢

MISSISSIPPIAN BRACHIOPODS FROM SHITTAKAZAWA, ARISU AND ODAIRA FORMATIONSAPPENDIX
AOD1: Yokota area (39°05′
17″N, 141°34′
20″E), south-facing
slope, 400 m N of junction of the Otsubosawa andKaisenosawa valleys, light greenish grey coarse-grained
tuffaceous sandstone, lower OD2 Unit of the Odaira
Formation, with Absenticosta uldzejtuensis.
I N S 1 : Nisawa area (39°13′28″N, 141°28′06″E), middle
Dobasawa Valley; dark grey shale, upper OD2 Unit of
the Odaira Formation, with Martinia georgei.
I NS2 : Nisawa area (39° 13′
24″N, 141° 28′
15″E), middle Dobasawa
Valley, dark grey shale, upper OD2 Unit of the Odaira
Formation, with Fluctuaria undata, Schizophoria
resupinata and Martinia georgei.
I NS3 : Nisawa area (39°13′
24″N, 141°28′
19″E), middle Dobasawa
Valley; greenish grey tuff, middle OD2 Unit of the
Odaira Formation, with Marginicinctus marginicinctus,
Martinia georgei and Asyrinxia sp.
I NS4 : Nisawa area (39°12′22″N, 141°28′02″E), western slopeof “808 m hill” , 750 m NE of Nisawa Pass, greenish
grey tuff, middle OD2 Unit of the Odaira Formation,with Martinia georgei.
I NS5 : Nisawa area (39°12′18″N, 141°28′
24″E), upper Karosawa
Valley, greenish grey tuff, middle OD2 Unit of the
Odaira Formation, with Martinia georgei.
I NS6 : Nisawa area (39°11′54″N, 141°28′
15″E), ridge between
the Karosawa Valley and the Komatagawa River, grey
fine-grained calcareous sandstone, OD3 Unit of the
Odaira Formation, with Martinia georgei.
I NS7 : Nisawa area (39°11′39″N, 141°28′
23″E), middle Karosawa
Valley, grey limestone, middle OD2 Unit of the Odaira
Formation, with Levitusia elongata sp. nov., Orthotetes
keokuk, Schizophoria resupinata and Martinia georgei.
27″N, 141°28′
30″E), middle Shokarosawa
I NS8 : Nisawa area (39°11′
Valley, greenish grey tuff, middle OD2 Unit of the
Odaira Formation, with Martinia georgei.
INS9: Nisawa area (39°11′35″N, 141°28′53″E), upper Shokarosawa
Valley, black shale, lower OD2 Unit of the Odaira
Formation, with Schizophoria resupinata.
INS10: Nisawa area (39°11′37″N, 141°28′
59″E), upper Shokarosawa
Valley, greenish grey tuff, lower OD2 Unit of the Odaira
Formation, with Schizophoria resupinata and Martinia
georgei.
INS11: Nisawa area (39°11′22″N, 141°29′04″E), upper Nekonosawa
Valley, greenish grey tuff, lower OD2 Unit of the Odaira
Formation, with Levitusia elongata sp. nov. and
Schizophoria resupinata.
INS12: Nisawa area (39°11′01″N, 141°28′42″E), middle Nekonosawa
Valley, grey medium-grained sandstone, upper OD2
Unit of the Odaira Formation, with Striatifera angusta.
INS13: Nisawa area (39°10′52″N, 141°28′42″E), middle Nekonosawa
Valley, grey medium-grained sandstone, upper OD2
Unit of the Odaira Formation, with Marginicinctus
marginicinctus, Schizophoria resupinata and Martinia
georgei.

87

IST1: Shimoarisu area (39°11′06″N, 141°32′
06″E), road cutting
beside the lower Hinotsuchigawa River, 125 m SE ofjunction of the Hinotsuchigawa River and Shittakazawa
Valley, dark grey calcareous shale, lower AR2 Unit of
the Arisu Formation, with Ovatia elongata, Rhipidomellamichelini, Schizophoria resupinata, S. pinguis, S. woodi,
Unispirifer striatoconvolutus, Unispirifer sp.,
Kitakamithyris hikoroitiensis, Syringothyris texta, S.
platypleura and Pseudosyrinx jumonjiensis.
IYK1: Yokota area (39° 06′
41″N, 141° 33′53″E), northeastern
slope of Mt. Odairayama, dark grey calcareous
sandstone, upper AR2 Unit of the Arisu Formation, with
Tomiproductus elegantulus, Rhipidomella michelini andKitakamithyris hikoroitiensis.
IYK2: Yokota area (39° 06′
41″N, 141° 33′47″E), northeastern
slope of Mt. Odairayama, dark grey calcareous
sandstone, upper AR2 Unit of the Arisu Formation, with
Tomiproductus elegantulus, Marginatia burlingtonensis
and Cleiothyridina harkeri.
IYK3: Yokota area (39° 06′
42″N, 141° 33′42″E), northeastern
slope of Mt. Odairayama, dark grey calcareous
sandstone, upper AR2 Unit of the Arisu Formation, with
M a rg i n a t i a b u r l i n g t o n e n s i s a n d A s y r i n x i anipponotrigonalis.
KAR1: Okuhinotsuchi area (39° 11′
36″N, 141° 30′
47″E), small
tributary on the northern bank of the Hinotsuchigawa
River, 1.4 km NW of junction of the Hinotsuchigawa
and Yokokawa rivers, light grey tuffaceous shale, lower
ST3 Unit of the Shittsakazawa Formation, with
Rhipidomella kusbassica, Schizophoria pinguis, S.
mayesensis and Unispirifer kozuboensis.
KAR4: Shizu area (39°09′
54″N, 141°31′
18″E), middle Yasumotozawa
Valley, greenish-grey medium-grained tuffaceous
sandstone, middle AR2 Unit of the Arisu Formation,with Grandispirifer mylkensis.
KAR6: Okuhinotsuchi area (39°10′26″N, 141°31′05″E), upper
Yasumotozawa Valley, greenish grey coarse-grained
tuffaceous sandstone, lower AR2 Unit of the Arisu
Formation, with Grandispirifer mylkensis.
KAR11: Okuhinotsuchi area (39°11′52″N, 141°30′
53″E), ridge
between the Yokokawa and Hinotsuchigawa rivers, 0.8
km N of Okuhinotsuchi, greenish grey coarse-grained
tuffaceous sandstone, middle AR2 Unit of the Arisu
Formation, with Levitusia humerosa.
KF158: Yokota area (39° 05′
05″N, 141° 34′09″E), road cutting
at 30 m W of junction of the Otsubosawa and Kaisenosawa
valleys, light greenish grey coarse-grained tuffaceous
sandstone, lower OD2 Unit of the Odaira Formation,with Rhipidomella michelini, Schizophoria resupinata,
Spirifer liangchowensis, Syringothyris sp. andDimegelasma sp.
KF159: Yokota area (39°04′38″0N, 141°34′30″E), road cutting
along the lower Tairagaisawa Valley; dark grey
calcareous shale, OD3 Unit of the Odaira Formation,with Leptagonia analoga, Rugosochonetes extensus,

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JUN-ICHI TAZAWA

Marginatia burlingtonensis, Echinoconchus punctatus,
Echinaria sp., Pustula pustulosa, Schellwienellaradialis, Cleiothyridina fimbriata, Spirifer
liangchowensis, Kitakamithyris sp. and Pseudosyrinx sp.
KF175: Yokota area (39° 05′
07″N, 141° 33′
15″E), south-facing
slope 350 m W of junction of the Otsubosawa and
Myojinsawa valleys, light greenish-grey coarse-grained
tuffaceous sandstone, lower OD2 Unit of the Odaira
Formation, with Rhipidomella michelini.
KF177: Yokota area (39°05′05″N, 141°33′
16″E), middle Otsubosawa
Valley, 350 m W of junction of the Otsubosawa and
Myojinsawa valleys, light greenish grey coarse-grained
tuffaceous sandstone, lower OD2 Unit of the Odaira
Formation, with Schizophoria resupinata.
KF179: Yokota area (39° 05′
21″N, 141° 33′
23″E), middle Myojinsawa
Valley, light greenish grey coarse-grained tuffaceous
sandstone, lower AR2 Unit of the Arisu Formation, with
Tolmatchoffia robusta, Tomiproductus elegantulus,
Orthotetes keokuk, Rhipidomella michelini,
Schizophoria pinguis, Unispirifer striatoconvolutus, U.
kozuboensis and Kitakamithyris hikoroitiensis.
KF180: Yokota area (39° 05′
23″N, 141° 33′22″E), middle Myojinsawa
Valley, light greenish grey coarse-grained tuffaceous
sandstone, lower AR2 Unit of the Arisu Formation,withTomiproductus elegantulus, Rhipidomella michelini,
Schizophoria pinguis and Acuminothyris triangularis.
KF181: Yokota area (39° 05′
25″N, 141° 33′
20″E), middle Myojinsawa
Valley, light greenish grey coarse-grained tuffaceous
sandstone, lower AR2 Unit of the Arisu Formation, with
Tolmatchoffia robusta, Tomiproductus elegantulus,
Orthotetes keokuk, Rhipidomella michelini,
Schizophoria pinguis, Grandispirifer mylkensis,
Unispirifer kozuboensis, Syringothyris texta andPseudosyrinx jumonjiensis.
KF207: Yokota area (39° 05′
04″N, 141° 33′56″E), road cutting
along the middle Otsubosawa Valley, 50 m W ofjunction of Otsubosawa and Suesawa valleys, blackshale, upper AR3 Unit of the Arisu Formation, with
Orbiculoidea sp., Marginatia burlingtonensis andBrachythyrina sp.
KF208: Yokota area (39° 05′
06″N, 141° 34′02″E), road cutting
along middle Otsubosawa Valley, 75 m E of junction of
Otsubosawa and Suesawa valleys, black shale, lower
AR3 Unit of the Arisu Formation, with Marginatia
burlingtonensis.
KF210: Yahagi area (39° 02′
31″N, 141° 35′08″E), road cutting
along the lower YukisawaValley, at 1 km W of junctionof the Yukisawa Valley and the Kesengawa River, grey
calcareous shale, upper OD3 Unit of the Odaira
Formation, with Marginicinctus marginicinctus,

Fluctuaria undata and Torynifer asiatica.
KYT1: Yokota area (39°05′
20″N, 141°33′
57″E), middle Suesawa
Valley, black shale, lower AR3 Unit of the Arisu
Formation, with Unispirifer minnewankensis andPunctospirifer sp.
KYT2: Yokota area (39° 05′
48″N, 141° 33′39″E), upper Suesawa
Valley, greenish grey fine-grained tuffaceous sandstone,
lower AR2 Unit of the Arisu Formation, with Martinia
georgei and Unispirifer kozuboensis.
KYT3: Yokota area (39° 05′
41″N, 141° 33′45″E), upper Suesawa
Valley, light greenish grey coarse-grained tuffaceous
sandstone, lower AR2 Unit of the Arisu Formation, with
Marginatia burlingtonensis, Acuminothyris triangularisand Asyrinxia nipponotrigonalis.
KYT4: Yokota area (39° 05′
41″N, 141° 33′39″E), upper Suesawa
Valley, light greenish grey coarse-grained tuffaceous
sandstone, lower AR2 Unit of the Arisu Formation, with
Acuminothyris triangularis.
YKA12: Shizu area (39°09′
44″N, 141°29′
13″E), near the mouthof the Konikurazawa Valley, greenish grey fine-grained
tuffaceous sandstone, upper OD2 Unit of the Odaira
Formation, with Levitusia elongata sp. nov.
YKA16: Shizu area (39° 09′
00″N, 141° 29′41″E), road cutting
along the Omatagawa River, at 1.3 km SW of junctionof the Omatagawa and Komatagawa rivers, greenish
grey fine-grained tuffaceous sandstone, middle OD2Unit of the Odaira Formation, with Rhipidomellamichelini and Schizophoria resupinata.
YKA18: Shizu area (39°08′
42″N, 141°29′
53″E), lower Suisawa
Valley, greenish grey fine-grained tuffaceous sandstone,
lower OD2 Unit of the Odaira Formation, with
Schizophoria resupinata.
YKA20: Shizu area (39°08′
41″N, 141°29′
55″E), lower Suisawa
Valley, greenish grey fine-grained tuffaceous sandstone,
lower OD2 Unit of the Odaira Formation, with
Absenticosta uldzejtuensis, Orthotetes keokuk,
Schizophoria resupinata and Rotaia hikoroichiensis.
YKA21: Shizu area (39°08′
30″N, 141°29′
43″E), middle Kominekurasawa
Valley, greenish grey fine-grained tuffaceous sandstone,
lower OD2 Unit of the Odaira Formation, with Levitusia
elongata sp. nov., Schizophoria resupinata and Spirifer
liangchowensis.
YUK1: Yahagi area (39°02′
26″N, 141°34′40″E), southern slope
facing the middle Yukisawa Valley, grey calcareous
shale, upper OD3 Unit of the Odaira Formation, with
Rugosochonetes extensus, Rugosochonetes sp.,
Plicatifera plicatilis, P. pseudoplicatilis,
Argentiproductus sp., Alitaria konincki, Orthotetes
keokuk, Rhipidomella michelini, Imbrexia forbesi,
Brachythyris chouteauensis and Kitakamithyris sp.