ARTICLE

Memoir of the Fukui Prefectural Dinosaur Museum 22: 13–141 (2023)© by the Fukui Prefectural Dinosaur Museum

MIDDLE PERMIAN (GUADALUPIAN) BRACHIOPODS FROM SETAMAI,KAMIYASSE–IMO AND MATSUKAWA IN THE SOUTH KITAKAMI BELT, JAPANJun-ichi TAZAWA1 and Yousuke IBARAKI2
Hamaura-cho 1-260-1, Chuo-ku, Niigata 951-8151, Japan
2Fossa Magna Museum, Itoigawa 941-0056, Japan

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ABSTRACTThis study describes 83 species of middle Permian (Guadalupian) brachiopods in 64 genera (including three new
species, Paramesolobus kitakamiensis sp. nov., Callispirina sheni sp. nov. and Whitspakia nipponica sp. nov.) fromthe Kanokura, Hosoo and Kamiyasse Formations in the central part (Setamai, Kamiyasse–Imo and Matsukawa areas)of the South Kitakami Belt (southern Kitakami Mountains), northeastern Japan. Most of the brachiopod species,
excluding two species Poikilosakos kamiyassensis and Martinia semiplana from the Hosoo Formation, occur in the
Wordian beds of the Kanokura and Kamiyasse Formations. The Wordian brachiopod fauna is characterised by a
mixture of Boreal and Tethyan elements, and has a close affinity with those of northern China (Inner Mongolia),
eastern Russia (South Primorye) and central Japan (the Hida Gaien Belt). Thus, the South Kitakami region belonged
to the Sino-Mongolian–Japanese Province between the Boreal and Tethyan realms in the Northern Hemisphere. Thisregion was probably a shallow sea bordering a microcontinent (the Proto-Japan Block) that was located near and to
the east of the North China Block at the eastern end of the Central Asian Orogenic Belt during the Wordian.
Key words: Brachiopoda, Japan, middle Permian, South Kitakami Belt, Wordian
田沢純一・茨木洋介 (2023) 南部北上帯世田米，上八瀬‒飯森，松川から産出する中期ペルム紀（グアダループ世）腕足類．福井県立恐竜博物館紀要 22：13‒141.
南部北上帯（南部北上山地）中央部の世田米・上八瀬‒飯森・松川地域に分布する叶倉層，細尾層，上八Paramesolobus kitakamiensis sp. nov., Callispirina sheni sp. nov.,
瀬 層 か ら 産 出 す る 64 属 83 種 (3 新 種，Paramesolobus
Whitspakia nipponica sp. nov. を含む ) の中期ペルム紀（グアダループ世）腕足類を記載した．これらの腕足類は，細尾層から産出する Poikilosakos
Poikilosakos kamiyassensis
kamiyassensis と Martinia semiplana の 2 種を除いてほかはすべて，叶倉層と上八瀬層の Wordian 相当層から産出する．古生物地理学的に，世田米・上八瀬‒飯森・松川
地域の Wordian の腕足類フォーナはボレアル区とテチス区の要素が混在するボレアル型‒テチス型混合
フォーナで，中国北部（内蒙古），ロシア東部（プリモリエ），中部日本（飛騨外縁帯）の同時代の腕足類フォーナに類似する．以上のことから，南部北上地域は中期ペルム紀（Wordian）に北半球中緯度地域のシノモン
ゴル‒日本区に属し，おそらく中央アジア造山帯の東端，北中国地塊の東方に位置したマイクロコンチネン
ト（原日本地塊）の縁辺浅海域であったと推定される．
せた まい

かみ やっ せINTRODUCTION

い

も

かのくら

ほそ お

South Kitakami Belt. Since the work of Endo (1924), manystudies have investigated the stratigraphy of the middle
Permian Kanokura Formation (named by Onuki, 1956) in the
Setamai area (e.g., Minato et al., 1954; Onuki, 1956; Saito,
1966; Choi, 1973; Tazawa and Ibaraki, 2001). In addition,
the Kamiyasse–Imo (i.e., Kamiyasse, Kesennuma City,
Miyagi Prefecture, and Imo, Yahagi-cho, Rikuzentakata City,
Iwate Prefecture; Figs. 1, 2, 3B and 12) and Matsukawa (i.e.,
Matsukawa, Kesennuma City, Miyagi Prefecture; Figs. 1 and13) areas, both of which are located in the central part of theSouth Kitakami Belt, are classical and renowned fossil
localities for middle Permian (Wordian) marine invertebrates.
The Wordian brachiopod fauna of the Kamiyasse–Imo area is

Permian marine sedimentary rocks are widely distributedin the South Kitakami Belt (southern Kitakami Mountains),
northeastern Japan. The Setamai area (i.e., Setamai,
Sumita-cho, Kesen-gun, Iwate Prefecture; Figs. 1, 3A and 11)
in the central part of the belt is a classical and important area
for the middle Permian stratigraphy of Japan as well as theReceived June 8, 2023. Accepted November 25, 2023.
Corresponding author―Jun-ichi Tazawa
E-mail: j1025-tazawa*memoad.jp
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JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 1. Maps showing the location and geology of the Setamai, Kamiyasse–Imo and Matsukawa areas, South Kitakami Belt, northeastern Japan. A,
Geotectonic map of the Japanese Islands, showing the distribution of the South Kitakami Belt (based on Tazawa, 2018); B, geotectonic map of thenortheastern Honshu, Japan, showing the distribution of the Permian rocks in the South Kitakami Belt (based on Kawamura et al., 2013).

the most diverse Japanese Permian brachiopod fauna. Since
Yabe (1900), many studies have investigated the taxonomy of
the brachiopod fauna (e.g., Hayasaka, 1922a, 1925a;
Hayasaka and Minato, 1956; Nakamura, 1959b, 1972;
Tazawa, 1974b, 2016b).
In this study, we describe middle Permian (Guadalupian;Roadian–Capitanian, mostly Wordian) brachiopods from the
Kanokura Formation in the Setamai area and from the Hosooand Kamiyasse Formations in the Kamiyasse–Imo
and Matsukawa areas and discuss the age andpalaeobiogeography of the Wordian brachiopod fauna. Mostof the materials were collected by the present authors: J.
Tazawa collected the brachiopod specimens from the
Kamiyasse–Imo and Matsukawa areas in 1970–1975 during
the course of graduate study at the Department of Geology
and Mineralogy, Faculty of Science, Hokkaido University,
under the supervision of M. Minato; and Y. Ibaraki collectedbrachiopod specimens from the Setamai area in 1996 during
the course of graduate study at the Department of Geology,
Faculty of Science, Niigata University, under the supervisionof J. Tazawa.

PREVIOUS WORK
Stratigraphy
Previous studies on the stratigraphy of the middle Permianrocks in the Setamai area are summarized in Fig. 4. In theearly studies, Endo (1924) named the lower part of thesandstone–limestone unit “the Alternation of Slate andSandstone” and the upper part “the Fusulina Limestone”.
Onuki (1938) named the middle Permian rocks the Kanokura
Stage (= upper part of the Yukizawa Series). Modern
stratigraphical studies on the Permian rocks of the Setamai
area began in the early 1940s. Minato (1942) and Minato etal. (1954) established the Permian stratigraphy in the Setamai
area. According to those studies, the Permian rocks in the
Setamai area can be divided into three series: theSakamotozawa Series (Sakmarian; Pseudoschwagerinaand Pseudofusulina zones), the Kanokura Series(Artinskian–Wordian; Lyttonia and Lepidolina zones) and the
Toyoma Series (upper Permian?), in ascending stratigraphic
order. Moreover, the middle Permian Kanokura Series was
subdivided into two stages: the lower part was named theKattisawa Stage and the upper part the Iwaizaki Stage. This
stratigraphic division was accepted and subsequently

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

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FIGURE 2. A view of central part of the Kamiyasse – Imo area, around the Kokitamukaisawa and Matashirosawa valleys (A), and outcrop of argillaceous
limestone of the KY1 Unit of the Kamiyasse Formation at locality KF67, upper Shigejisawa Valley (B).

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JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 3. Outcrop of sandstone of the KN2 Unit, Kanokura Formation at locality KS9 in the middle Kanokurasawa Valley, Setamai (A), and outcrop of
limestone of the KY2 Unit, Kamiyasse Formation at upper Minamizawa Valley, Kamiyasse (B).

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

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FIGURE 4. Comparison of stratigraphic schemes proposed for the middle Permian (Guadalupian) formations in the Setamai area, South Kitakami Belt.

followed by Onuki (1956, 1969), Saito (1966, 1968), Choi
(1973), Choi et al. (1979) and Tazawa and Ibaraki (2001).
In contrast, there have been few previous studies on the
Permian stratigraphy of the Kamiyasse–Imo and Matsukawa
areas (Fig. 5). Shiida (1940) named the Permian rocks the
Kesennuma Series. The lower part of the series was called
the Kamiyasse Formation; the upper part was named theFutatsumori Formation. In that study, the upper part of the
Kamiyasse Formation was assigned to the middle Permian.
Subsequently, Onuki (1956), Kambe and Shimazu (1961) andTazawa (1973, 1976) classified the Permian rocks into thesame three formations (or groups, series) as those of the
Setamai area; i.e., the lower Permian Sakamotozawa
Formation, the middle Permian Kanokura Formation and theupper Permian Toyoma Formation (Onuki, 1956). However,Misaki and Ehiro (2004) divided the Permian rocks into four
formations: the Nakadaira Formation, the Hosoo Formation,
the Kamiyasse Formation and the Kurosawa Formation, in
ascending stratigraphic order. In that study, both the Hosooand Kamiyasse Formations were assigned to the middle
Permian.

Palaeontology
Many taxonomic studies have been published on the
middle Permian biota of the Setamai, Kamiyasse–Imo andMatsukawa areas: fusulines (Morikawa, 1953; Fujimoto,
1956; Kanomata and Chisaka, 1967; Kanomata andMiyawaki, 1967; Choi, 1970, 1973; Kobayashi et al., 2009);
conulariids (Hayasaka, 1920, 1923, 1963a; Sugiyama, 1942;
Murata, 1967); bryozoans (Sakagami, 1961); brachiopods(Yabe, 1900; Hayasaka, 1917, 1922a, 1925a, 1937, 1953,
1960, 1963b, 1963d, 1966, 1967; Hayasaka and Minato,
1956; Minato, 1955; Minato and Nakamura, 1956;
Nakamura, 1959b, 1960, 1970, 1972; Nakamura et al., 1970;
Tazawa, 1974a, 1974b, 1979, 1999c, 2008c, 2014, 2016a,
2016b, 2017; Tazawa and Araki, 1984a, 1984b, 1999, 2013,2016, 2017, 2018; Tazawa and Ibaraki, 2001; Tazawa andKaneko, 2016; Tazawa and Takaizumi, 1987; Shen andTazawa, 1997, 2014; Afanasjeva and Tazawa, 2007, 2010;
Afanasjeva et al., 2015; Shiino, 2009; Shiino and Suzuki,
2007; Masunaga and Shiino, 2021); bivalves (Hayasaka,
1925b; Murata, 1964; Nakazawa and Newell, 1968);
cephalopods (Hayasaka, 1940; 1963c; Ehiro and Araki, 1997;

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JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 5. Comparison of stratigraphic schemes proposed for the
middle Permian (Guadalupian) formations in the Kamiyasse–Imo area,
South Kitakami Belt.

Ehiro and Misaki, 2005; Ehiro, 2015); trilobites (Araki,
1961; Jimbo, 1966; Araki and Koizumi, 1968; Koizumi and
Sasaki, 1978; Kobayashi and Hamada, 1984) andcartilaginous fish (Araki, 1980). The brachiopods of the area
have been studied extensively. Yabe (1900) described two
species (Lyttonia sp. and Lyttonia cf. nobilis) from
Matsukawa: this was the first study of Palaeozoic
brachiopods in Japan. Subsequently, Hayasaka (1917, 1922a,
1925a, 1937, 1953, 1960, 1963b, 1966, 1967) and Hayasaka
and Minato (1956) described 40 species in 24 genera; Minato(1955) and Minato and Nakamura (1956) described fivespecies in five genera; Nakamura (1959b, 1960, 1970, 1972)
and Nakamura et al. (1970) described 25 species in nine
genera; Tazawa and colleagues (Tazawa, 1974a, 1974b,
1979, 1999c, 2008c, 2014, 2016a, 2016b, 2017; Tazawa andAraki, 1984a, 1984b, 1999, 2013, 2016, 2017, 2018; Tazawa
and Ibaraki, 2001; Tazawa and Kaneko, 2016; Tazawa andTakaizumi, 1987) described 51 species in 41 genera; Shen
and Tazawa (1997, 2014) described four species in four
genera; Afanasjeva and colleagues (Afanasjeva and Tazawa,
2007, 2010; Afanasjeva et al., 2015) described three species
in three genera; Shiino (2009) and Shiino and Suzuki (2007)
described three species in two genera; and Masunaga andShiino (2021) described three species in two genera.
In terms of biostratigraphy, Choi (1973) proposed threefusuline zones in the Kanokura Series of the Setamai area:
the Monodiexodina matsubaishi and Colania kotsuboensis
zones in the Kattisawa Stage and the Lepidolina multiseptata
Zone in the Iwaizaki Stage. In addition, Choi (1973) assigned
the Kattisawa Stage to the Kubergandian–Murgabian and the

Iwaizaki Stage to the lower Lopingian. Subsequently, Minato
et al. (1978) proposed the Monodiexodina kofuganensis,
Leptodus nobilis and Lepidolina multiseptata–L. minatoi
zones, assigning the Kattisawa Stage to the
Kungurian–Kazanian and the Iwaizaki Stage to the lower
Tatarian. Ehiro and Misaki (2005), on the basis of theammonoid fauna, assigned the middle part of the Hosoo
Formation to the Roadian and both the upper part of the
Hosoo Formation and the Kamiyasse Formatio to the
Wordian–Capitanian. Kobayashi et al. (2009) proposed threefusuline zones in the Kamiyasse Formation of the
Kamiyasse–Imo area (the Monodiexodina sutchanica,
Parafusulina motoyoshiensis and Lepidolina shiraiwensis
zones) and correlated all three zones with the Midian (latest
Wordian–Capitanian). Tazawa (2016b, 2017, 2018) assigned
the lower part of the Kamiyasse Formation in the
Kamiyasse–Imo and Matsukawa areas to the Wordian, basedon the brachiopods.
In terms of palaeobiogeography, Tazawa (1987) andNakamura and Tazawa (1990) recognized middle Permian
mixed Boreal–Tethyan brachiopod faunas in northeasternJapan (the South Kitakami Belt), central Japan (the Hida
Gaien Belt), eastern Russia (South Primorye), northeastern
China (Jilin) and northern China (Inner Mongololia).
Subsequently, Tazawa (1991, 1992, 1998) proposed the
middle Permian geography of the South Kitakami region andadjacent areas in eastern Asia to have been as follows: theSouth Kitakami region was probably located at a middle
latitude of the Northern Hemisphere, near and to the east of
the North China Block, during the middle Permian. Thishypothesis was confirmed by Tazawa (2003, 2007, 2016b,
2017) and Tazawa and Araki (2017). In contrast, Ehiro(1997, 1998, 2001, 2010) and Ehiro and Misaki (2005)
emphasised that the middle Permian ammonoid faunas of theSouth Kitakami Belt consisted of Tethyan (tropical) andcosmopolitan genera and completely lacked Boreal(antitropical) genera; and suggested that the South Kitakami
region was probably located in equatorial Panthalassa near
the South China Block during the middle Permian.
STRATIGRAPHY
Permian rocks are exposed in the Setamai, Kamiyasse–Imo
and Matsukawa areas in the form of two subparallel synclinesthat trend N–S to NNW–SSE and plunge gently towards thesouth (Figs. 1B, 6 and 8). The two synclines are separated by
the left-lateral strike-slip Hizume–Kesennuma Fault(Hirokawa and Yoshida, 1954; Ehiro, 1977), which trends
NNW–SSE. The Permian successions of the areas consist
mostly of sandstone and shale with subordinate limestone andconglomerate. Both sandstone and limestone are predominant
in the eastern syncline (Setamai area), but shale is dominant
in the western syncline (Kamiyasse–Imo and Matsukawa
areas).

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

FIGURE 6. Geologic map of the Setamai area (after Tazawa and Ibaraki, 2001).

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JUN-ICHI TAZAWA AND YOUSUKE IBARAKI

ascending stratigraphic order (Fig. 7). Brachiopods occur inthree horizons: sandstone of the uppermost part of the KN1
Unit; sandstone of the upper part of the KN2 Unit; andargillaceous limestone of the uppermost part of the KN2
Unit.
Kamiyasse–Imo
The middle Permian rocks in the Kamiyasse–Imo area are
divided into two formations: the Hosoo Formation in thelower, and the Kamiyasse Formation in the upper (Tazawa,
2016b). The Hosoo Formation (400–500 m thick) consists
mainly of dark grey sandy shale with thin layers ofsandstone, conglomerate and limestone; the Kamiyasse
Formation (150–250 m thick) consists mainly of grey togreenish-grey sandstone with subordinate shale,
conglomerate and limestone (dark grey argillaceous
limestone in its lower part and light grey limestone in its
upper part). In this study, the Kamiyasse Formation is
subdivided into two units, the KY1 Unit (mainly sandstone
and argillaceous limestone with subordinate shale, 160 m
thick) and the KY2 Unit (mainly sandstone and limestone
with subordinate shale and conglomerate, 130 m thick), in
ascending stratigraphic order (Fig. 9). Brachiopods occur
mostly in the KY1 Unit but have also been found in the upper
part of the Hosoo Formation and the lower part of the KY2
Unit.
Matsukawa
FIGURE 7. Generalized columnar section of the Permian formations in
the Setamai area, showing the fossil horizons in the KN1 and KN2 units
in the Kanokura Formation (modified and adapted from Tazawa and
Ibaraki, 2001).

Setamai
According to Tazawa and Ibaraki (2001), the Permianrocks in the Setamai area are divided into three formations:
the Sakamotozawa, Kanokura and Toyoma Formations, inascending stratigraphic order. The middle Permian Kanokura
Formation (470 m thick) is subdivided into a lower part(mainly grey to greenish-grey sandstone, with subordinate
dark grey argillaceous limestone and conglomerate, 300 m
thick) and an upper part (light grey limestone, 170 m thick).
The boundary between the Kanokura Formation and theunderlying Sakamotozawa Formation is an unconformity.
In the present study, the Kanokura Formation is subdivided
into three units: the KN1 Unit (sandstone with a basal
conglomerate, 160 m thick), the KN2 Unit (mainly sandstone,
with subordinate conglomerate and argillaceous limestone,
140 m thick); and the KN3 Unit (limestone, 170 m thick), in

According to Tazawa and Araki (2017), the lower part of
the Kamiyasse Formation (mainly grey to greenish-greysandstone with subordinate dark grey argillaceous limestone
and dark grey shale, 215 m thick) is locally exposed in thewestern wing of the syncline in the Matsukawa area (Fig. 8).
Brachiopods occur in three horizons—two in the lower partand one in the middle part—of the lower Kamiyasse
Formation (KY1 Unit; Fig. 10).
MATERIALS
The brachiopods described herein were collected from 62
localities (stations) in the Setamai, Kamiyasse–Imo andMatsukawa areas of the South Kitakami Belt. The
topographic and stratigraphic locations, lithologies andbrachiopod species of the 62 fossil localities are indicated in
Figs. 7 and 9–13, and summarized in Table 1 and appendix.
These specimens are now registered and stored in the
Department of Geology, Niigata University, Niigata (NU-B
prefix); the Hokkaido University Museum. Sapporo (UHR
prefix); the Tohoku University Museum, Sendai (IGPS
prefix); the Kesennuma Board of Education, Kesennuma,
Miyagi Prefecture (KCG prefix); the Rikuzentakata City
Museum, Rikuzentakata, Iwate Prefecture (RCM-F prefix);
and the Palaeontological Institute, Russian Academy of

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

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FIGURE 8. Geological map of the Kamiyasse–Imo and Matsukawa areas (modified from Tazawa, 1976). Legend 1 – 3, Toyoma Formation (1, Shale; 2,
Sandstone; 3, Conglomerate); 4–6, Kamiyasse Formation (4, Shale; 5, Sandstone; 6, Conglomerate); 7–9, Hosoo Formation (7, Shale; 8, Sandstone; 9,
Conglomerate); 10–12, Nakadaira Formation (10, Shale; 11, Sandstone; 12, Conglomerate); 13, Limestone of the Nakadaira, Hosoo, Kamiyasse andToyoma Formations; 14, Carboniferous Nagaiwa Formation; 15, Cretaceous andesite; 16, Dyke rock.

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JUN-ICHI TAZAWA AND YOUSUKE IBARAKI

Sciences, Moscow (PIN prefix).
BRACHIOPOD FAUNAFIGURE 9. Generalized columnar section of the middle Permian(Guadalupian) rocks in the Kamiyasse–Imo area, showing the fossil
horizons in the upper part of the Hosoo Formation and KY1 and KY2
units in the Kamiyasse Formation (modified and adapted from Tazawa,
2017).

The brachiopods described herein are the following 83
species in 64 genera, including three new species(Paramesolobus kitakamiensis sp. nov., Callispirina sheni sp.
nov. and Whitspakia nippponica sp. nov.): Orbiculoidea
verum Masunaga and Shiino, 2021, Isogramma heritschi
Nakamura, 1970, I. nakamurai Tazawa, 2016a, Neochonetes(Huangichonetes) matsukawensis Tazawa and Araki, 2018,
Paramesolobus kitakamiensis Tazawa, sp. nov.,
Kanokurachonetes kanokurensis Afanasjava and Tazawa,
2010, Capillomesolobus heritschi Pečar, 1986, Chonetinetes
elongatus Afanasjeva, Tazawa and Shintani, 2015,
Kitakamichonetes multicapillatus Afanasjeva and Tazawa,
2007, Transennatia gratiosa (Waagen, 1884),
Spinomarginifera lopingensis (Kayser, 1883), S.
kueichowensis Huang, 1932, S. alpha Huang, 1932,
Paramarginifera sp., Hexiproductus echidniformis (Chao,
1925), Kunlunia sp., Costiferina spiralis (Waagen, 1884),
Tyloplecta yangtzeensis (Chao, 1927), Bathymyonia
neimongolica (Wang and Zhang, 2003), Vediproductus
punctatiformis (Chao, 1927), Waagenoconcha humboldti(d’Orbigny, 1842), W. irginae (Stuckenberg, 1898),
Edriosteges multispinosus Muir-Wood and Cooper, 1960,
Limbella sp., Chonosteges sp., Urushtenoidea crenulata(Ding in Yang et al., 1962), Scacchinella gigantea
Schellwien, 1900b, Linoproductus hayasakai Tazawa, 1979,
Globiella tschernyschewi (Netschajew, 1911), Asperlinus
japonicus (Tazawa, 2008c), Lamiproductus kamiyassensis(Tazawa, 2016a), Anidanthus mizukoshiensis Tazawa, 2008b,
Costatumulus cancriniformis (Tschernyschew, 1889),
Permundaria asiatica Nakamura, Kato and Choi, 1970, P.
tenuistriata Tazawa, 1974a, Grandaurispina kozlowskiana(Fredericks, 1925), Yakovlevia mammata (Keyserling, 1846),
Y. kaluzinensis Fredericks, 1925, Neorichthofenia mabutii(Tazawa and Araki, 1984b), Leptodus nobilis (Waagen,1883), Keyserlingina sp., Petasmaia expansa Cooper and
Grant, 1969, Pararigbyella doulingensis Shen and Zhang,2008, Poikilosakos kamiyassensis Tazawa and Takaizumi,
1987, Paralyttonia kesennumensis Tazawa and Araki, 1984a,
Permianella typica He and Zhu, 1979, Dicystoconcha
lapparenti Termier and Termier in Termier et al., 1974,
Laterispina parallela Shen, Fan, Zhang and Zhang, 1994,
Derbyia grandis Waagen, 1884, D. nipponica Nakamura,
1972, Meekella eximia (de Verneuil, 1845), M. nodosa
Nakamura, 1972, Orthothetina polita (Fliegel, 1901), O.
kayseri (Fliegel, 1901), O. kitakamiensis (Hayasaka, 1953),
O. hayasakai Nakamura, 1972, O. transversa Nakamura,
1972, Streptorhynchus pelargonatus (von Schlotheim, 1816),
S. cataclinus Zhang in Yang et al., 1962, Rhipidomellamagna Tazawa, 2016a, Acosarina dorsisulcata Cooper and
Grant, 1969, A. rectimarginata Cooper and Grant, 1976b,
Orthotichia sp., Stenoscisma margaritovi (Tschernyschew,GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

231888), S. mutabilis (Tschernyschew, 1902), S. sokolskajae
Koczyrkevicz, 1979b, Rhynchopora tchernyshae
Koczyrkevicz, 1979a, Permocryptospirifer omeishanensis(Huang, 1933), Hustedia remota (Eichwald, 1860), H.
ratburiensis Waterhouse and Piyasin, 1970, Martinia
semiplana Waagen, 1883, M. triquetra Gemmellaro, 1899,
Neospirifer moosakhailensis (Davidson, 1862), Gypospirifer
kobiyamai Tazawa and Araki, 2013, Alispiriferella lita
(Fredericks, 1924b), Phricodothyris catatona (Cooper and
Grant, 1976a), Permophricodothyris grandis (Chao, 1929), P.
squamularioides (Huang, 1933), Licharewina arakii(Hayasaka, 1963b), Callispirina sheni Tazawa, sp. nov.,
Spiriferellina fredericksi Tazawa, 2014, Dielasma sp. andWhitspakia nipponica Tazawa, sp. nov.
The above-listed brachiopods are classified into threefaunas: the Kanokura fauna from the Kanokura Formation in
the Setamai area; the Kamiyasse–Imo fauna from the
Kamiyasse Formation in the Kamiyasse–Imo area; and the
Matsukawa fauna from the Kamiyasse Formation in the
Matsukawa area. Moreover, the three faunas are subdivided
into five assemblages: the KN1 assemblage from the KN1Unit of the Kanokura Formation; the KN2 assemblage fromthe KN2 Unit of the Kanokura Formation; the HO
assemblage from the upper part of the Hosoo Formation; theKY1 (KI) assemblage from the KY1 Unit of the Kamiyasse
Formation in the Kamiyasse–Imo area; and the KY1 (MK)assemblage from the KY1 Unit of the Kamiyasse Formationin the Matsukawa area.
Kanokura fauna

FIGURE 10. Generalized columnar section of the middle Permian(Guadalupian) rocks in the Matsukawa area, showing the fossil horizonsin the Hosoo Formation and the KY1 Unit in the Kamiyasse Formation(modified and adapted from Tazawa and Araki, 2017).

KN1 assemblage
The KN1 assemblage, from the uppermost part of the KN1Unit of the Kanokura Formation in the Setamai area
(localities KS1, KS2, KS3, KS4 and KS8), includes six
species in five genera: Transennatia gratiosa, Tyloplecta
yangtzeensis, Linoproductus hayasakai, Leptodus nobilis,
Derbyia grandis and D. nipponica. Of these species,
Linoproductus hayasakai and Leptodus nobilis are abundant;
and the other species are rare (Table 1).
KN2 assemblage
The KN2 assemblage, from the upper part of the KN2 Unitof the Kanokura Formation in the Setamai area (localities
KS7, KS9 and KS10), includes 11 species in 10 genera:
Transennatia gratiosa, Waagenoconcha irginae,
Urushtenoidea crenulata, Permundaria asiatica,
Grandaurispina kozlowskiana, Leptodus nobilis, Laterispina
parallela, Derbyia grandis, D. nipponica, Orthothetina
kitakamiensis and Alispiriferella lita. Of these species,
Leptodus nobilis and Alispiriferella lita are abundant;
Transennatia gratiosa is common; and the other species arerare (Table 1).

24

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 11. Map showing the fossil localities KS1, KS2, KS3, KS4, KS5, KS7, KS8, KS9 and KS10 in the Setamai area (using the electronic
topographical map of GSI).

Kamiyasse–Imo fauna
HO assemblage
The HO assemblage, from the upper part of the Hosoo
Formation in the Kamiyasse–Imo area (locality KF212),
includes two species: Poikilosakos kamiyassensis andMartinia semiplana. Of these species, Poikilosakos
kamiyassensis is abundant; and Martinia semiplana iscommon (Table 1).
KY1 (KI) assemblage
The KY1 (KI) assemblage, from the KY1 Unit of the
Kamiyasse Formation in the Kamiyasse–Imo area (localities
AR1, AR2, AR3, KF3, KF6, KF7, KF8, KF11, KF13, KF17,
KF18, KF20, KF21, KF22, KF27, KF28, KF29, KF33, KF37,
KF39, KF46, KF50, KF55, KF61, KF62, KF63, KF64, KF65,
KF67, KF69, KF71, KF79, KF80, KF84, KF86, KF88, KF89,

KF90, KF91, KF94, KF96, KF99, KF121, KF217, KF218 andTY1), includes 64 species in 49 genera: Orbiculoidea verum,
Isogramma heritschi, I. nakamurai, Paramesolobus
kitakamiensis sp. nov., Kanokurachonetes kanokurensis,
Chonetinetes elongatus, Kitakamichonetes multicapillatus,
Transennatia gratiosa, Spinomarginifera lopingensis, S.
kueichowensis, S. alpha, Paramarginifera sp., Costiferina
spiralis, Tyloplecta yangtzeensis, Bathymyonia neimongolica,
Vediproductus punctatiformis, Waagenoconcha humboldti,
W. irginae, Edriosteges multispinosus, Limbella sp.,
Chonosteges sp., Urushtenoidea crenulata, Globiellatschernyschewi, Asperlinus japonicus, Lamiproductus
kamiyassensis, Anidanthus mizukoshiensis, Permundaria
asiatica, P. tenuistriata, Grandaurispina kozlowskiana,
Leptodus nobilis, Pararigbyella doulingensis, Permianella
typica, Dicystoconcha lapparenti, Derbyia nipponica,GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

25

FIGURE 12. Map showing the fossil localities AR1, AR2, AR3, KF3, KF6, KF7, KF8, KF9, KF11, KF13, KF17, KF18, KF20, KF21, KF22, KF27,
KF28, KF29, KF33, KF37, KF39, KF46, KF50, KF55, KF61, KF62, KF63, KF64, KF65, KF67, KF69, KF71, KF79, KF80, KF84, KF86, KF88, KF89,
KF91, KF94, KF96, KF99, KF121, KF212, KF217, KF218 and TY1 in the Kamiyasse–Imo area (using the electronic topographical map of GSI).

26

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 13. Map showing the fossil localities AR4, AR5 and KZ9 in the Matsukawa area (using the electronic topographical map of GSI).

Meekella eximia, M. nodosa, Orthothetina polita, O. kayseri,
O. hayasakai, O. transversa, Streptorhynchus pelargonatus,
S. cataclinus, Rhipidomella magna, Acosarina dorsisulcata,
A. rectimarginata, Orthotichia sp., Stenoscisma margaritovi,
S. mutabilis, S. sokolskajae, Rhynchopora tchernyshae,
Permocryptospirifer omeishanensis, Hustedia remota, H.
ratburiensis, Martinia triquetra, Neospirifer moosakhailensis,
Gypospirifer kobiyamai, Alispiriferella lita, Phricodothyris
catatona, Permophricodothyris grandis, P. squamularioides,
Licharewina arakii, Callispirina sheni sp. nov., Spiriferellina
fredericksi and Whitspakia nipponica sp. nov.
Of these species, Kitakamichonetes multicapillatus,
Spinomarginifera kueichowensis, Waagenoconcha irginae,
Urushtenoidea crenulata, Leptodus nobilis, Permianella
typica, Stenoscisma margaritovi and Spiriferellina fredericksi
are abundant; Paramesolobus kitakamiensis sp. nov.,
Transennatia gratiosa, Spinomarginifera lopingensis,
Tyloplecta yangtzeensis, Vediproductus punctatiformis,
Waagenoconcha humboldti, Edriosteges multispinosus,
Grandaurispina kozlowskiana, Dicystoconcha lapparenti,
Derbyia nipponica, Orthothetina kayseri, Rhipidomellamagna, Stenoscisma mutabilis, Permocryptospirifer
omeishanensis, Martinia triquetra, Permophricodothyris
grandis, P. squamularioides and Whitspakia nipponica sp.
nov. are common; and the other species are rare (Table 1).

Matsukawa fauna
KY1 (MT) assemblage
The KY1 (MT) assemblage, from the KY1 Unit of the
Kamiyasse Formation in the Matsukawa area (localities AR4,AR5 and KZ9), includes 26 species in 24 genera:
Neochonetes (Huangichonetes) matsukawensis,
Capillomesolobus heritschi, Transennatia gratiosa,Hexiproductus echidniformis, Kunlunia sp., Urushtenoidea
crenulata, Scacchinella gigantea, Linoproductus hayasakai,
Costatumulus cancriniformis, Permundaria asiatica, P.
tenuistriata, Yakovlevia mammata, Y. kaluzinensis,
Neorichthofenia mabutii, Leptodus nobilis, Keyserlingina sp.,Petasmaia expansa, Paralyttonia kesennumensis,
Dicystoconcha lapparenti, Meekella nodosa, Orthothetina
kayseri, Stenoscisma margaritovi, Martinia triquetra,
Alispiriferella lita, Licharewina arakii and Dielasma sp. Of
these species, Alispiriferella lita is abundant; Scacchinella
gigantea and Neorichthofenia mabutii are common; and theother species are rare (Table 1).
AGE AND CORRELATION
Kanokura Formation

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

27

FIGURE 14. Stratigraphic distributions of brachiopod species of the
KN1 assemblage in the Setamai area.

Lower part of the Kanokura Formation (KN1 Unit)
The stratigraphic distributions of the brachiopod species of
the KN1 assemblage are described in the “Systematic
descriptions” section of the present paper and summarised inFig. 14. Of the brachiopods listed above, Linoproductus
hayasakai is known only from the Wordian; Transennatia
gratiosa, Tyloplecta yangtzeensis and Derbyia nipponica
have a stratigraphic range of Roadian–Changhsingian;
Leptodus nobilis is known from the Kungurian–Changhsingian;and Derbyia grandis ranges from the Asselian to the
Wuchiapingian. To summarise, the age of the KN1
assemblage is identified as Wordian; thus, the uppermost partof the KN1 Unit is correlated with the Wordian. The age ofthe lower part of the KN1 Unit is unknown due to a lack offossil evidence in the Setamai area. However, Tazawa andShintani (2021) determined the age to be Kungurian on thebasis of a brachiopod fauna called “the Tashiroyama fauna”(which includes Transennatia insculpta, Dicystoconcha
lapparenti and Crenispirifer sagus), collected from near thebase of the KN1 Unit of the Kanokura Formation in the
Nagaiwa–Sakamotozawa area, about 10 km east of Setamai.
Therefore, the age of the lower part of the Kanokura
Formation (KN1 Unit) can be identified as
Kungurian–Wordian.
Middle part of the Kanokura Formation (KN2 Unit)
The stratigraphic distributions of the brachiopod species of
the KN2 assemblage are described in the “Systematic
descriptions” section of the present paper and summarised inFig. 15. Of the brachiopods listed above, Grandaurispina
kozlowskiana and Orthothetina kitakamiensis are known only
from the Wordian; Permundaria asiatica is known from theWordian–Capitanian; Urushtenoidea crenulata has a

FIGURE 15. Stratigraphic distributions of brachiopod species of the
KN2 assemblage in the Setamai fauna.

stratigraphic range of Kungurian–Capitanian; Transennatia
gratiosa is known from the Roadian–Changhsingian;
Laterispina parallela is known from theWordian–Changhsingian; and Transennatia gratiosa,
Derbyia nipponica and Alispiriferella lita have astratigraphic range of Roadian–Changhsingian.
Waagenoconcha irginae, Leptodus nobilis and Derbyia
grandis are long-ranging species. In summary, the age of the
KN2 assemblage is identified as Wordian; thus, the middle
part (KN2 Unit) of the Kanokura Formation is correlatedwith the Wordian.
Upper part of the Kanokura Formation (KN3 Unit)
In this study, no brachiopods are described from the upper
part of the Kanokura Formation (KN3 Unit) in the Setamai
area. However, Minato et al. (1978) correlated the Iwaizaki
Stage (= upper Kanokura Formation) to the upper Maokou
Limestone of South China based on fusulines (Lepidolina
multiseptata and L. minatoi) described by Choi (1973). The
upper Maokouan is correlated with the Capitanian (Jin et al.,1997); thus, the upper part of the Kanokura Formation (KN3
Unit) can be correlated with the Capitanian.
Hosoo Formation
The HO assemblage, from the upper part of the Hosoo
Formation, consists of only two species (Poikilosakos
kamiyassensis and Martinia semiplana) and is inadequate forage determination. Instead, Tazawa and Nakamura (2015)

28

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 16. Stratigraphic distributions of brachiopod species of the KY1 (KI) assemblage in the Kamiyasse–Imo area. Broken lines show those of genera.

determined the age of the lower part of the Hosoo Formation
as Kungurian on the basis of a brachiopod fauna, the“Nakadaira Fauna” (including Kunlunia grabaui, Chaoiellataiyuanfuensis, Juresania juresanensis, Orthothetina curvata
and Orthotichia jiangxiensis) from Nakadaira, about 4 km
north of Kamiyasse. Ehiro and Misaki (2005) correlated the
middle and upper parts of the Hosoo Formation with theRoadian and Wordian, respectively, on the basis ofammonoids. The Roadian–Wordian boundary lies slightly
above the brachiopod fossil horizon (KF212). According toEhiro and Misaki (2005), the middle part of the Hosoo

Formation yields Agathiceras sp., Agathiceras? sp.,
Adrianites sp., Demarezites sp. and Demarezites? sp., and theupper part of the Hosoo Formation yields Agathiceras sp.,
Cardiella sp., Tauroceras? sp., Parastacheoceras bidentatus,
Waagenoceras sp. and Paraceltites elegans. To summarise,
the Hosoo Formation can be correlated with the
Kungurian–Wordian; and the fossil horizon (KF212) is
assigned to the Roadian.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

29

Kamiyasse Formation
Lower part of the Kamiyasse Formation
The stratigraphic distributions of the brachiopod species of
the KY1 (KI) and KY1 (MT) assemblages are described in
the “Systematic descriptions” section of the present paperand summarised in Fig. 16. Of the brachiopods listed above,
Orbiculoidea verum, Isogramma heritschi, I. nakamurai,
Kanokurachonetes kanokurensis, Chonetinetes elongatus,
Globiella tschernyschewi, Asperlinus japonicus,
Lamiproductus kamiyassensis, Permundaria tenuistriata,
Grandaurispina kozlowskiana, Pararigbyella doulingensis,
Orthothetina transversa, Rhipidomella magna, Licharewina
arakii and Spiriferellina fredericksi are known only from theWordian; Kitakamichonetes multicapillatus, Bathymyonia
neimongolica, Anidanthus mizukoshiensis, Permundaria
asiatica, Streptorhynchus cataclinus, Stenoscisma
sokolskajae and Rhynchopora tchernyshae have astratigraphic range of Wordian–Capitanian; Permianella
typica, Stenoscisma margaritovi and Gypospirifer kobiyamai
are known from the Wordian–Wuchiapingian; andSpinomarginifera lopingensis, S. kueichowensis, S. alpha,Orthothetina hayasakai, Neospirifer moosakhailensis,
Permophricodothyris grandis and P. squamularioides range
from th e Wordian to the Changhs ingian. In contras t,
Permocryptospirifer omeishanensis is known from theRoadian–Wordian; Edriosteges multispinosus andPhricodothyris catatona have a stratigraphic range ofArtinskian–Wordian; Acosarina dorsisulcata is known fromthe Sakmarian–Wordian; and Meekella eximia, Orthothetina
polita, Acosarina rectimarginata and Martinia triquetra havebeen reported from Asselian–Wordian strata. In summary, theage of the KY1 (KI) assemblage is identified as Wordian;thus, the lower part of the Kamiyasse Formation (KY2 Unit)in the Kamiyasse–Imo area is correlated with the Wordian.
This conclusion is consistent with that of Tazawa (2016b).
The stratigraphic distributions of the brachiopod species of
the KY1 (MT) assemblage are described in the “Systematic
descriptions” section of the present paper and summarised inFig. 17. Of the brachiopods listed above, Neochonetes(Huangichonetes) matsukawensis, Linoproductus hayasakai,Paralyttonia kesennumensis and Licharewina arakii are
known only from the Wordian; Permundaria asiatica and P.
tenuistriata are known from the Wordian–Capitanian; andStenoscisma margaritovi ranges from the Wordian to the
Wuchiapingian. In contrast, Petasmaia expansa is known
from the Artinskian–Wordian; Capillomesolobus heritschi
has a stratigraphic range of Sakmarian–Wordian; andHexiproductus echidniformis, Costatumulus cancriniformis
and Martinia triquetra are known from theKasimovian–Wordian. Urushtenoidea crenulata is known
from the Kungurian–Capitanian; and two speecies(Yakovlevia kaluzinensis and Dicystoconcha lapparenti) are
known from the Kungurian–Wuchiapingian. To summarise,
the KY1 (MT) assemblage is identified as Wordian in age,

FIGURE 17. Stratigraphic distributions of brachiopod species of the
KY1 (MT) assemblage in the Matsukawa area. Broken lines show thoseof genera.

so, the lower part of the Kamiyasse Formation (KY2 Unit) in
the Matsukawa area is also correlated with the Wordian. Thisconclusion is consistent with the findings of Tazawa andAraki (2017, 2018).
Upper part of the Kamiyasse Formation
Only one species, Waagenoconcha irginae, a long-ranging(Asselian–Capitanian) species, was collected from the upper
part of the Kamiyasse Formation (KY2 Unit) of the
Kamiyasse–Imo and Matsukawa areas; thus, age
determination of the upper part of the Kamiyasse Formation(KY2 Unit) is difficult on the basis of brachiopods. However,Tazawa (1976) correlated the upper part of the Kamiyasse

FIGURE 18 (1). Geographic distributions of brachiopod species of the KY1 (KI) and KY1 (MT) assemblages excluding three new species (Paramesolobus kitakamiensis sp. nov., Callispirina sheni
sp. nov. and Whitspakia nipponica sp. nov.) and seven uncertain species (Paramarginifera sp., Kunlunia sp., Limbella sp., Chonosteges sp., Keyserlingina sp. Orthothetina sp. and Dielasma sp.).

30
JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 18 (2). Continue

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

31

32

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 19. Middle Permian (Wordian) reconstruction map of the world (adapted from Scotese, 2004), showing the geographic distribution ofbrachiopod species of the KY1 (KI) and KY1 (MT) assemblages excluding three new species (Paramesolobus kitakamiensis sp. nov., Callispirina sheni
sp. nov. and Whitspakia nipponica sp. nov.) and seven uncertain species (Paramarginifera sp., Kunlunia sp., Limbella sp., Chonosteges sp., Keyserlingina
sp. Orthothetina sp. and Dielasma sp.). Location numbers are same in Fig.18, and the numbers appended to the circles in the legend indicate the species
numbers, M: Mongolia, NC: North China, SC: South China, Ⅰ: Boreal Realm, Ⅱ : Tethyan Realm, Ⅲ : Panthalassa Realm, Ⅳ : Gondwanan Realm.

Formation with the Lepidolina multiseptata Zone (=Lepidolina multiseptata–L. minatoi Zone of Minato et al.,
1 9 7 9 ) f r o m t h e p r e s e n c e o f L e p i d o l i n a m u l t i s e p t a t a,
Verbeekina verbeeki and Codonofusiella explicata. Ehiro andAraki (1997) considered the uppermost Kamiyasse Formationand the lowest Toyoma Formation (= Kurosawa Formation of
Misaki and Ehiro, 2004) to be late Capitanian in age becauseof the occurrence of Tainoceras carinatum, Tainoceras sp.,
Pleuronautilus sp., Stearoceras sp., Jilingites kesennumensis,
Stacheoceras sp., Timorites takaizumii and Pseudagathiceras
ornatum. To summarise, the upper part of the Kamiyasse
Formation can be correlated with the Capitanian.
PALAEOBIOGEOGRAPHY
The geographic distributions of the Wordian brachiopod
species of the KN1, KN2, KY1 (KI) and KY1 (MT)assemblages are documented in the “Systematic descriptions”
section and are summarised in Figs. 18 and 19. Of the 72
species (excluding three new species and six uncertain
species) of the Wordian brachiopods, 19 species also occur ineastern Russia (South Primorye); 16 species also occur innorthern China (Inner Mongolia); 15 species also occur insouthwestern China (Sichuan); 13 species are found incentral-southern China (Hunan); and 12 species are found incentral Japan (Hida Gaien Belt), central-southern China(Hubei) and southwestern China (Guizhou). The Wordianbrachiopod faunas of the South Kitakami Belt, South
Primorye, Inner Mongolia and the Hida Gaien Belt include
both antitropical (Waagenoconcha, Costatumulus,
Grandaurispina, Yakovlevia, Rhynchopora, Neospirifer,
Gypospirifer and Alispiriferella) and tropical (Transennatia,
Spinomarginifera, Tyloplecta, Leptodus, Pararigbyella,Permianella, Dicystoconcha, Permocryptospirifer andPermophricodothyris) genera; in contrast, the Wordianbrachiopod faunas of Hubei, Hunan, Guizhou and Sichuanlack antitropical genera. To summarise, the Wordianbrachiopod fauna, consisting of the KN1, KN2, KY1 (KI)and KY1 (MT assemblages, is a mixed Boreal–Tethyan faunaand exhibits affinities with those of northern China (Inner
Mongolia), eastern Russia (South Primorye) and central
Japan (Hida Gaien Belt). This conclusion is consistent withthe previous works on the Wordian brachiopod faunas of
Setamai (Tazawa and Ibaraki, 2001), Kamiyasse–Imo(Tazawa, 2016b), Matsukawa (Tazawa and Araki, 2017,2018), Obama (Tazawa, 2003) and Yakejima (Tazawa et al.,2000) in the South Kitakami Belt.
From the information provided above, we can conclude

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

that the Wordian brachiopod fauna of the South Kitakami
Belt is characterised by a mixture of both Boreal and Tethyanelements and belonged to the Sino-Mongolian–Japanese
Province (Shi and Tazawa, 2001; Shen et al., 2009) between
the Boreal and Tethyan realms in mid-latitudes of theNorthern Hemisphere (Fig. 19). The South Kitakami region
was probably located near and to the east of the North China
Block at the eastern end of the Central Asian Orogenic Belt
during the Wordian. This conclusion is consistent with earlier
studies of the Wordian brachiopod biogeography of eastern
Asia (Tazawa, 1991, 1992, 1998, 2002, 2007; Shi, 2006) andglobally (Shen et al., 2009), and with the bipolar distributionof the Wordian antitropical fusulinoidean genus
Monodiexodina (Ueno, 2006). Excluding the Permianbrachiopod and fusuline palaeobiogeography, Li and
Takeuchi (2022) supported the above-mentioned Permianreconstruction of the South Kitakami region and east Asia,
including the North China Block, based on U-Pb dating ofdetrital zircon from Permian successions of the South
Kitakami Belt.
CONCLUSIONSIn this study, middle Permian (Guadalupian;Roadian–Capitanian, mostly Wordian) brachiopods aredescribed from the Kanokura, Hosoo and Kamiyasse
Formations in the Setamai, Kamiyasse–Imo and Matsukawa
areas of the central part of the South Kitakami Belt (southern
Kitakami Mountains), northerastern Japan. A total of 83
species in 64 genera are described, of which three are new(Paramesolobus kitakamiensis sp. nov., Callispirina sheni sp.
nov. and Whitspakia nippponica sp. nov.). In terms ofbiostratigraphy, the lower (KN1 Unit) and middle (KN2 Unit)
parts of the Kanokura Formation and the lower part (KY1
Unit) of the Kamiyasse Formation are correlated with the
Wordian. Palaeobiogeographically, the Wordian brachiopod
fauna, including the KN1, KN2, KY1 (KI) and KY1 (MK)assemblages, is a mixed Boreal–Tethyan fauna and exhibits
affinities with those of northern China (Inner Mongolia),
eastern Russia (South Primorye) and central Japan (the Hida
Gaien Belt). Thus, the South Kitakami region belonged to the
Sino-Mongolian–Japanese Province (Shi and Tazawa, 2001;
Shen et al., 2009) between the Boreal and Tethyan realms in
the Northern Hemisphere. This region was probably a
shallow sea bordering the Proto-Japan Block(microcontinent), which was located near and to the east of
the North China Block at the eastern end of the Central Asian
Orogenic Belt during the Wordian.
SYSTEMATIC DESCRIPTIONS(by J. Tazawa)
The suprageneric classification given herein mainly
follows that of “Treatise on Invertebrate Paleontology, Part H
Brachiopoda Revised, Volumes 2–6”, edited by Kaesler

33(2000a , 2000b, 2002, 2006) a nd Se lde n ( 2007) , wi t h
exception that classification 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 forbessi Davidson, 1848.
Orbiculoidea verum Masunaga and Shiino, 2021
(Fig. 20A)
Orbiculoidea sp. Hayasaka, 1963b, p. 479, fig. 1.
Orbiculoidea jangarensis Ustritsky. Tazawa and Araki, 2013,
p. 4, fig. 2.1.
Orbiculoidea verum Masunaga and Shiino, 2021, p. 262, figs.
4–8.
Material.— One specimen from locality AR3, externalmould of a ventral valve, KCG7.
Remarks.—This specimen was previously described by
Tazawa and Araki (2013, p. 4, fig. 2.1) as Orbiculoidea
jangarensis Ustritsky, 1960 from the lower part of the
Kanokura Formation (= Kamiyasse Formation) of Kamiyassein the Kamiyasse–Imo area. Subsequently, Masunaga andShiino (2021, p. 262, figs. 4–8) redescribed the specimen,
together with newly added 95 specimens from the upper partof the Hosoo Formation at Minamizawa and Shigejisawa
valleys in Kamiyasse, as Orbiculoidea verum Masunaga andShiino, 2021. This species resembles well Orbiculoidea
jangarensis Ustritsky, 1960 in size, shape and external
ornament of ventral valve, but differs in having a
posterior-sited apex in dorsal valve.
Occurrence.—KY1 Unit in Kamiyasse (locality AR3).
Distribution.—Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt).
Order DICTYONELLIDA Cooper, 1956
Superfamily EICHWALDIOIDEA Schuchert, 1893
Family ISOGRAMMIDAE Schuchert, 1929
Genus ISOGRAMMA Meek and Worthen, 1870
Type species.—Chonetes? millepunctatus Meek andWorthen, 1870.
Isogramma heritschi Nakamura, 1970
(Fig. 20B–D)
Isogramma heritschi Nakamura, 1970, p. 308, pl. 4, figs.
3–7; Minato et al., 1979, pl. 57, figs. 2–4, 6; Tazawa et al.,2014, p. 378, fig. 2.4; Tazawa, 2015, p. 63, fig. 6.4.
Material.—Five specimens from locality KF217: (1)34JUN-ICHI TAZAWA AND YOUSUKE IBARAKI

external moulds of two ventral valves, UHR19043, 19044;(2) external and internal moulds of a dorsal valve,
UHR19045 (holotype); (3) external mould of a dorsal valve,
UHR19047; and (4) internal mould of a dorsal valve,
UHR19046.
Remarks.—These specimens were described by Nakamura(1970, p. 308, pl. 4, figs. 3–7) as Isogramma heritschi
Nakamura, 1970. This species is a middle-sized Isogramma(length 25 mm, width 53 mm in the holotype), with external
ornament consisting of numerous fine concentric growthlines (numbering 5–7 in 1 mm in dorsal valve) and somerugae parallel to postero-lateral margins of shell. Isogrammamanchoukuoensis Hatai and Omura (1941, p. 7, pl. 2, figs.
5–11, text-fig. 1), from the Penchi Formation of Liaoning,northeastern China, differs from I. heritschi in its smaller
size and less-transverse outline. Isogramma paotechowensis(Grabau and Chao in Chao, 1928), originally described by
Chao (1928, p. 33, pl. 1, fig. 27; pl. 4, figs. 1–5) as
Aulacorhynchus paotechowensis Grabau and Chao in Chao,
1928 from the Taiyuan Formation of Shanxi, northern China,differs from the present species by its much larger
dimensions. Isogramma concavum Cooper and Grant (1974,
p. 252, pl. 25, figs. 16, 17), from the Neal Ranch Formationof Texas in the USA, resembles the present species in sizeand outline, but differs in having deeply concave dorsal
valve.
Occurrence.—KY1 Unit in Imo (locality KF217).
Distriburion.—Wordian: northeastern Japan(Kamiyasse–Imo and Hitachi in the South Kitakami Belt).

and dorsal valves. Isogramma renfrarum Cooper (1952, p.
114, pl. 21, figs. 1–3; pl. 22, fig 1; pl. 23, fig. 1), from thePennsylvanian Gonzales Shale of north-central Texas, is alsoa large-sized species of Isogramma, but the Texan species
differs from I. nakamurai in its more transverse outline andin having more regular and coarser growth lines on dorsal
valve. The type species, Isogramma millepunctata (Meek andWorthen, 1870, p. 566, pl. 25, fig. 3), from the Pennsylvanianof Illinois, is readily distinguished from the present species
by its much smaller size and more regular and coarser growthlines on both valves.
Occurrence.—KY1 Unit in Imo (locality KF217).
Distribution. — Wordian: northeastern Japan (Kamiyasse-Imo
in the South Kitakami Belt).

Isogramma nakamurai Tazawa, 2016a(Fig. 21A–C)

Neochonetes (Huangichonetes) matsukawensis Tazawa andAraki, 2018, p. 12, fig. 4.1.

Isogramma paotechowensis (Grabau and Chao). Minato,
1955, p. 29, text-fig. 1; Nakamura, 1970, p. 306, pl. 3, figs.
1, 2; pl. 4, figs. 1, 2; Minato et al., 1979, pl. 57, figs. 1, 5.
Isogramma aff. paotechowensis (Grabau and Chao). Chen
and Shi, 2006b, p. 416, fig. 3.
Isogramma nakamurai Tazawa, 2016a, p. 82, figs. 3, 4.

Material.—One specimen from locality AR4, external andinternal moulds of a ventral valve, KCG61 (holotype).
Remarks.—This specimen was described by Tazawa andAraki (2018, p. 12, fig. 4.1) as Neochonetes(Huangichonetes) matsukawensis Tazawa and Araki, 2018.
The specimen from the lower Kamiyasse Formation of
Matsukawa is large in size for genus, transverse in outline(length 8 mm, width 13 mm), and having numerous costellae
on ventral valve, numbering 56 near anterior margin ofventral valve. Neochonetes (Huangichonetes) matsukawensis
is most similar to the type species, Neochonetes(Hunagichonetes) substrophomenoides (Huang, 1932),
redescribed by Shen and Archbold (2002, p. 337, fig. 5E–M)
from the Lopingian of Hunan and Guizhou, southern China,
in shape and external ornament of ventral valve, but differsfrom the C hine s e s pe cie s in its large r s iz e a nd m uchtransverse outline.
Occurrence.—KY1 Unit in Matsukawa (locality AR4).
Distribution. —Wordian: northeastern Japan (Matsukawa
in the South Kitakami Belt).

Material.—Four specimens from locality KF217: (1)external and internal moulds of a dorsal valve, UHR19040(holotype); (2) external and internal moulds of two dorsal
valves, UHR19041, 19042; and (3) external mould of a dorsal
valve, NU-B1657.
Remarks.—These specimens were previously describedby Tazawa (2016a, p. 82, figs. 3, 4) as Isogramma nakamurai
Tazawa, 2016a. This species is characterized by its large size(length 58 mm, width about 130 mm in the holotype,
UHR19040) and external ornament consisting of numerous
fine elevated and slightly undulated concentric growth lines.
Isogramma paotechowensis (Grabau and Chao in Chao,
1928), from the Taiyuan Formation of Shanxi, northern
China, differs from I. nakamurai by its smaller size and inhaving more regular concentric growth lines on both ventral

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 NEOCHONETES Muir-Wood, 1962
Subgenus NEOCHONETES (HUANANGICHONETES) Shenand Archbold, 2002
Type species.—Chonetes substrophomenoides Huang, 1932.
Neochonetes (Huangichonetes) matsukawensis Tazawa andAraki, 2018
(Fig. 22A)GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

35

FIGURE 20. A, Orbiculoidea verum Masunaga and Shiino, external mould (A1, A2) of ventral valve, KCG7; B–D, Isogramma heritschi Nakamura; B,
external cast of ventral valve, UHR19043; C, external mould of dorsal valve, UHR19047; D, external mould (D1) and internal mould (D2) of dorsal valve,
UHR19045 (holotype). Scale bars are 1 cm.

Genus PARAMESOLOBUS Afanasjeva, 1975
Type species.—Paramesolobus ivanovae Afanasjeva, 1975.
Paramesolobus kitakamiensis Tazawa, sp. nov.
(Fig. 22E–H)
Chonetes sinuosa Schellwien. Hayasaka, 1925a, p. 93, pl. 5,
figs. 5, 6.
Mesolobus sinuosa (Schellwien). Tazawa, 1976, pl. 2, fig.
12; Minato et al., 1979, pl. 61, figs. 4–6.
Etymology.—Named after the fossil locality, Kitakami
Mountains.
Material.—Eight specimens from localities KF79 andKF217: (1) external and internal moulds of a ventral valve,
NU-B2457; (2) external mould of a ventral valve,
NU-B2458; (3) internal moulds of two ventral valves,
NU-B2459, 2460; (4) external and internal moulds of threedorsal valves, NU-B2461, 2462 (holotype), 2463; and (5)external mould of a dorsal valve, NU-B2464.
Diagnosis.—Large-sized Paramesolobus with relatively
fine capillae on both ventral and dorsal valves.
Description.—Shell large in size for genus, transversely
subrectangular in outline, with greatest width at slightly
anterior to hinge; length 16 mm, width 28 mm in the largest
dorsal valve specimen (NU-B2464). Ventral valve
moderately convex in lateral profile; hinge straight, with two
spines preserved on one side; umbo small; ears small, flat toslightly convex, not clearly demarcated from visceral region;
sulcus wide and shallow, with a narrow rounded median lobe.

Dorsal valve moderately concave in lateral profile; fold broad
and low, having a narrow median sulcus, External surface ofventral valve ornamented with numerous bifurcating capillae,
numbering 8 –9 in 2 mm at about midlenth of valve. External
ornament of dorsal valve similar to those of ventral valve.
Internally, ventral valve with short median septum
posteriorly one fourth valve length. Dorsal valve with shortdiverging lateral septa and a long median septum, extending
to two-thirds valve length; cardinal process ill preserved.
Internal shell surface with radial rows of papillae; anterior
margin capillate.
Remarks.—Of these specimens from Kamiyasse–Imo, thelargest one (NUB2464) was previously figured by Tazawa(1976, pl. 2, fig. 12) and Minato et al. (1979, pl. 61, fig. 4) asMesolobus sinuosa (Schellwien). However, the genus
Mesolobus Dunbar and Condra, 1932 differs from the genus
Paramesolobus Afanasjeva, 1975 in having lamellose
external ornament; and Capillomesolobus sinuosa(Schellwien, 1900b), from the Trogkofel Limestone of theCarnian Alps, differs from the Kitakami species in being
much smaller size and in having finer capillae on both valves.
All the specimens including the largest one can be safely
assigned to the genus Paramesolobus by large size, external
ornament consisting of numerous bifurcating and relatively
coarse capillae, and in having a ventral sulcus with a median
lobe and a dorsal fold with a median sulcus. Paramesolobus
kitakamiensis sp. nov. is distinguished from the type species,
Paramesolobus ivanovae Afanasjeva (1975, p. 102, fig. 4),
from the upper Carboniferous (Kasimovian) of the Russian
Platform, by its larger size and in having finer capillae on
ventral valve. Shells, described by Chao (1928, p. 22, pl. 1,

36

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 21. A–C, Isogramma nakamurai Tazawa; A, external mould (A1), internal latex cast (A2) and posterior view (A3) of internal latex cast of dorsal
valve, UHR19041; B, external mould (B1) and internal latex cast (B2) of dorsal valve, UHR19042; C, external mould (C1) and internal Mould (C2) of
dorsal valve, UHR19040 (holotype). Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

figs. 23–25; pl. 2, figs. 3–12; pl. 4, fig. 7) as Chonetes
latesinuata Schellwien, 1892, from the Taiyuan Series ofnorthern China, differs from Paramesolobus kitakamiensis in
having coarser capillae on both ventral and dorsal valves.
Occurrence.—KY1 Unit in Kamiyasse (locality KF79)
and Imo (locality KF217).
Genus KANOKURACHONETES Afanasjeva and Tazawa,
2010
Type species.—Kanokurachonetes kanokurensis
Afanasjava and Tazawa, 2010.
Kanokurachonetes kanokurensis Afanasjava and Tazawa,
2010(Fig. 22B–D)
Kanokurachonetes kanokurensis Afanasjava and Tazawa,
2010, p. 19, pl. 2, figs. 1–10.
Material.—Five specimens from localities KF39 andKF55: (1) internal mould of a ventral valve, PIN5213/1(holotype); and (2) external moulds of four dorsal valves,
PIN5213/10, 5213/11, NU-B2455, 2456.
Remarks.—Of the specimens listed above, three(PIN5213/1, PIN5213/10 and PIN5213/11) from the locality
KF55 were previously described by Afanasjeva and Tazawa( 2 0 1 0 , p . 1 9 , p l. 2 , fig s . 1 – 1 0 ) a s Ka n o kur ac hone te s
kanokurensis Afanasjava and Tazawa, 2010. The two(NU-B2455 and NU-B2456) from the locality KF39 are also
referred to this species in size, shape and external ornamentof dorsal valves. The diagnosis of the genus
Kanokurachonetes is as follows (after Afanasjeva andTazawa, 2010, p. 18–19): Shell medium in size, semicircularin outline and relatively concavoconvex; sulcus and fold welldeveloped; cardinal spines inclined to cardinal margins atangle of 45–50˚; Ventral interarea apsacline and dorsal areaorthocline; external radial ornamentation composed of a few
bifurcated costellae on entire surface of valves;
microornamentation represented by concentric rows ofdensely spaced ovate papillae. Inside ventral valve, median
septum spanning one-third of valve length; diductor scars
large, bifid; inside dorsal valve, cardinal process relatively
low, with basal alveolus; socket ridges massive; weak median
septum only developed in middle of valve; lateral septa
hardly visible; brachial ridges absent; thin accessory septa
extending from alveolus to anterior margin; beyond thesestructures, internal structures of both valves covered withsmall, rounded, irregularly arranged papillae. No otherspecies has been known besides the type species.
Occurrence.—KY1 Unit in Kamiyasse (localities KF39
and KF55).
Distribution.—Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt).

37

Subfamily CAPILLOMESOLOBINAE Pečar, 1986
Genus CAPILLOMESOLOBUS Pečar, 1986
Type species.—Capillomesolobus karavankensis Pečar, 1986.
Capillomesolobus heritschi Pečar, 1986
(Fig. 22I)
Chonetes sinuosa Schellwien, 1900b, p. 38, pl. 9, figs. 17,
18; Hayasaka, 1925a, p. 93, pl. 5, figs. 5, 6.
Chonetes sp. Heritsch, 1938, p. 103, pl. 7, figs. 6, 7.
Mesolobus mesolobus (Norwood and Pratten). Nakamura,
1959a, p. 205, pl. 2, figs. 2, 3; Minato et al., 1979, pl. 46,
fig. 11.
Mesolobus sp. Tazawa, 1979, p. 25, pl. 4, fig. 2.
Mesolobus sinuosus (Schellwien). Minato et al., 1979, pl. 61,
figs. 4–6.
Capillomesolobus heritschi Pečar, 1986, p. 28, pl. 3, figs.
1–9, text-fig. 11; Tazawa and Nakamura, 2015, p. 159, fig.
4.1–4.3; Tazawa and Araki, 2017, p. 269, fig. 6.1.
Material.—One specimen from locality AR5, external andinternal moulds of a ventral valve, IGPS96237.
Remarks.—This specimen was previously described by
Tazawa (1979, p. 25, pl. 4, fig. 2) as Mesolobus sp. Tazawa,
1979, and subsequently redescribed by Tazawa and Araki(2017, p. 269, fig. 6.1) as Capillomesolobus heritschi Pečar,
1986. The Matsukawa species can be referred toCapillomesolobus heritschi Pečar (1986, p. 28, pl. 3, figs.
1‒9, text-fig. 11), from the Trogkofel Limestone of the
Karavanke Mountains, Slovenia, in its small size (length
about 10 mm, width about 12 mm) and in having deep ventral
sulcus with a median lobe and numerous capillae (numbering
13‒14 in 2 mm at midlength) on external surface of ventral
valve. Comparison with the other species ofCapillomesolobus has been discussed by Tazawa andNakamura (2015, p. 161).
Occurrence.—KY1 Unit in Matsukawa (locality AR5).
Distribution. — Sakmarian‒Wordian: northeastern Japan(Nakadaira and Matsukawa in the South Kitakami Belt) andSlovenia (Karavanke Mountains).
Subfamily SVALBARDIINAE Archbold, 1982
Genus CHONETINETES Cooper and Grant, 1969
Type species.—Chonetinetes reversus Cooper and Grant,
1969.
Chonetinetes elongatus Afanasjeva, Tazawa and Shintani,
2015(Fig. 22J)
Chonetinetes elongatus Afanasjeva, Tazawa and Shintani,
2015, p. 24, pl. 4, figs. 1–3.

38

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 22. A, Neochonetes (Huangichonetes) matsukawensis Tazawa and Araki, external latex cast (A1, A2), internal latex cast (A3) and internal mould(A4) of ventral valve, KCG61 (holotype); B–D, Kanokurachonetes kanokurensis Afanasjeva and Tazawa; B, internal mould (B1, B2) of ventral valve,
PIN5313/1 (holotype); C, external mould of dorsal valve, PIN5313/10; D, external mould of dorsal valve, PIN5313/11: E–H, Paramesolobus
kitakamiensis sp. nov.; E, external latex cast (E1, E2) and internal mould (E3) of ventral valve, NU-B2457; F, external mould of dorsal valve, NU-B2464;
G, external latex cast (G 1 ) and internal mould (G 2 ) of dorsal valve, NU-B2462 (holotype); H, internal mould of ventral valve, NU-B2460; I,
Capillomesolobus heritschi Pečar, external latex cast (I1, I2) of ventral valve, IGPS96237; J, Chonetinetes elongatus Afanasjeva, Tazawa and Shintani,internal mould of ventral valve, NU-B1834 (holotype). Scale bars are 1 cm, except for that of J.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

39

FIGURE 23. A–E, Kitakamichonetes multicapillatus Afanasjeva and Tazawa; A, internal mould of dorsal valve, with partly preserved interior of ventral
valve, NU-B2438; B, external latex cast (B1, B2) and internal mould (B3) of ventral valve, NU-B2439; C, external latex cast of ventral valve, NU-B2442;
D, internal mould of ventral valve, NU-B2445; E, internal mould (E1, E2) of ventral valve, NU-B2444. Scale bars are 1 cm.

Material.—One specimen from locality KF39, external
and internal moulds of a ventral valve, NU-B1827.
Remarks.—This specimen was described by Afanasjeva etal. (2015, p. 24, pl. 4, figs. 1–3) as Chonetinetes elongatus
Afanasjeva, Tazawa and Shintani, 2015. The Kitakami
species is characterized by its longer outline (length 10 mm,
width 8 mm), and in having a narrow and deep sulcus and apair of flat and triangular ears, clearly demarcated fromvisceral portion by bend. Chonetinetes elongatus is mostly
similar to Chonetinetes angustisulcatus Cooper and Grant(1975, p. 1282, pl. 477, figs. 1–28), from the Road Canyon
Formation of Texas, in having a narrow sulcus and a pair offlattened, triangular ears well demarcated from the visceralportion of ventral valve, but differs from the Texan species in
its longer outline. The type species, Chonetinetes reversus
Cooper and Grant, 1969, redescribed by Cooper and Grant(1975, p. 1284, pl. 477, figs. 29–48) from the Road Canyon
Formation of Texas, is readily distinguished from the present
species in having fold and sulcus not reaching anterior
margin of shell.
Occurrence.—KY1 Unit in Kamiyasse (locality KF39).
Distribution.—Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt).
Subfamily CHALIMOCHONETINAE Afanasjeva, 1988
Genus KITAKAMICHONETES Afanasjeva and Tazawa, 2007

40

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 24. A–G, Transennatia gratiosa (Waagen); A, internal mould (A1, A2) of ventral valve, NU-B1736; B, ventral (B1, B2), anterior (B3), posterior(B4) and lateral (B5) views of internal mould of ventral valve, NU-B1735; C, external mould of dorsal valve, UHR17113; D, ventral (D1, D2), anterior(D3), posterior (D4) and lateral (D5) views of ventral valve, UHR30097; E, internal mould of ventral valve, NU-B2472; F, dorsal (F1) and lateral (F2) viewsof external mould of dorsal valve, NU-B1737; G, external mould of dorsal valve, UHR12126. Scale bars are 1 cm.

Type species.—Kitakamichonetes multicapillatus
Afanasjeva and Tazawa, 2007.
Kitakamichonetes multicapillatus Afanasjeva and Tazawa,
2007(Fig. 23A–E)
Chonetes blanfordi var. lita Hayasaka, 1925a, p. 92, pl. 5,
figs. 3, 4.
Neochonetes sp. Tazawa, 1979, p. 25, pl. 4, fig. 1.
Kitakamichonetes multicapillatus Afanasjeva and Tazawa,
2007, p. 73, pl. 11, figs. 1–12; Tazawa, 2008b, p. 42, fig.
6.3, 6.4; Tazawa et al., 2014, p. 378, fig. 2.1; Tazawa,
2015, p. 64, fig. 6.1.
Dyoros (Dyoros) sp. Tazawa and Araki, 2017, p. 269, fig. 6.2.
Material.—Seventeen specimens from localities KF61,
KF71, KF79, KF94 and KF218: (1) external mould of adorsal valve, with partly preserved interior of ventral valve,

NU-B2438: (2) external and internal moulds of two ventral
valves, NU-B2439, 2440; (3) external moulds of three ventral
valves, NU-B2441–2443; (4) internal moulds of eight ventral
valves, NU-B2444–2452; and (5) external mould of two
dorsal valves, NU-B2453, 2454.
Remarks.—The specimens from the lower Kamiyasse
Formation in Kamiyase are referred to Kitakamichonetes
multicapillatus Afanasjeva and Tazawa, 2007. This species ischaracterized by its large, transverse shell (length 31 mm,
width about 60 mm in the largest dorsal valve specimen,
NU-B2444) and external ornament consisting of numerous
and very fine capillae (numbering 8–9 in 2 mm at nearanterior margin of ventral valve). Kitakamichonetes
multicapillatus somewhat resembles Eolissochonetes
maximus Afanasjeva (1977, p. 17, pl. 2, figs. 1–6, 11–14),
from the middle and upper Carboniferous of theKolyma–Omolon Massif, northern Russia, but differs
primarily in its more prominent beak and longer median
septum on ventral valve, presence of brachial ridges, andabsence of an alveolus inside dorsal valve.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Occurrence.—KY1 Unit in Kamiyasse (localities KF61,
KF71, KF79, KF94) and Imo (locality KF218).
Distribution.—Wordian–Capitanian: northeastern Japan(Kamiyasse–Imo, Matsukawa and Hitachi in the South
Kitakam Belt) and southwestern Japan (Mizukoshi in Kyushu
Island).
Suborder PRODUCTIDINA Waagen, 1883
Superfamily MARGINIFEROIDEA Stehli, 1954
Family MARGINIFERIDAE Stehli, 1954
Subfamily MARGINIFERINAE Stehli, 1954
Genus TRANSENNATIA Waterhouse, 1975
Type species.—Productus gratiosus Waagen, 1884.
Transennatia gratiosa (Waagen, 1884)(Fig. 24A–G)
Productus gratiosus Waagen, 1884, p. 691, pl. 72, figs. 3–7;
Diener, 1897a, p. 23, pl. 3, figs. 3–7; Mansuy, 1913, p.
115, pl. 13, fig. 1; Diener, 1915, p. 70, pl. 7, fig. 4; Colani,
1919, p. 10, pl. 1, fig. 2; Chao, 1927, p. 44, pl. 4, figs.
6–10; Chi-Thuan, 1962, p. 491, pl. 2, figs. 5–7.
Productus (Dictyoclostus) gratiosus Waagen. Huang, 1933, p.
88, pl. 11, fig. 14; Hayasaka, 1960, p. 49, pl. 1, fig. 8.
Marginifera gratiosa (Waagen). Reed, 1944, p. 98, pl. 19,
figs. 6, 7.
Dictyoclostus gratiosus (Waagen). Zhang and Ching (Jin),
1961, p. 411, pl. 4, figs. 12–18; Wang et al., 1964, p. 291,
pl. 45, figs. 14–19; Leman, 1994, pl. 1, figs. 11–13.
Gratiosina gratiosa (Waagen). Grant, 1976, pl. 33, figs.
19–26; Licharew and Kotlyar, 1978, pl. 12, figs. 5, 6; pl.
20, fig. 1; Minato et al., 1979, pl. 61, figs. 11–13.
Asioproductus gratiosus (Waagen). Yang et al., 1977, p. 350,
pl. 140, fig. 5; Feng and Jiang, 1978, p. 254, pl. 90, figs. 1,
2; Tong, 1978, p. 228, pl. 80, fig. 7; Lee et al., 1980, p.
373, pl. 164, fig. 14; pl. 166, figs. 5, 6.
Asioproductus bellus Chan (Zhan), 1979, p. 85, pl. 6, figs.
7–13; pl. 9, figs. 8–10, text-fig. 18.
Gratiosina sp. Minato et al., 1979, pl. 61, fig. 14.
Dictyoclostus minor Lee and Gu in Lee et al., 1980, p. 372,
pl. 166, figs. 1–4.
Transennatia gratiosus (Waagen). Wang et al., 1982, p. 214,
pl. 92, figs. 6–8; pl. 102, figs. 4–9; Liu et al., 1982, p. 185,
pl. 132, fig. 9; Ding and Qi, 1983, p. 280, pl. 95, fig. 14;
Zeng et al., 1995, pl. 5, figs. 14, 15.
Transennatia gratiosa (Waagen). Yang, 1984, p. 219, pl. 33,
fig. 7; Jin, 1985, pl. 4, figs. 33, 34, 45, 46; Tazawa andMatsumoto, 1998, p. 6, pl. 1, figs. 4–8; Tazawa, 2000, fig.
3.6, 3.7; Tazawa et al., 2000, p. 7, pl. 1, figs. 3–5; Tazawa,
2001, p. 289, fig. 6.1–6.7; Tazawa and Ibaraki, 2001, p. 7,
pl. 1, figs. 1–3; Shen et al., 2002, p. 676, fig. 4.27–4.31;
Tazawa, 2002, fig. 10.2; Chen et al., 2005, p. 354, figs.
10E–H, 11; Tazawa, 2008b, p. 43, fig. 6.6, 6.7; Shen andZhang, 2008, fig. 4.20–4.22; Shen and Clapham, 2009, p.

41718, pl. 1, figs. 13–22; Shen and Shi, 2009, p. 157, fig.
3K–O; Tazawa et al., 2014, p. 378, fig. 2.2, 2.3; Tazawa,
2015, p. 65, fig. 6.2, 6.3; Tazawa, 2016b, p. 14, fig.
5.1–5.5; Tazawa and Araki, 2017, p. 209, fig. 6.3.
Material.—Nineteen specimens from localities KF217,
KF218, KS2, KS9 and KZ9: (1) external moulds of threeventral valves, NU-B302, 303, UHR17098; (2) internalmoulds of seven ventral valves, NU-B304, 305, 1735, 1736,
1738, 2472, UHR30097; and (3) external moulds of ten
dorsal valves, NU-B306, 1737, 1739, 1740, UHR12126,
17099. 17112, 17113, 17115.
Remarks. —Most of the specimens from Setamiai,
Kamiyasse–Imo and Matsukawa were described by Tazawa
and Ibaraki (2001, p. 7, pl. 1, figs. 1–3), Tazawa (2016b, p.
14, fig. 5.1–5.5) and Tazawa and Araki (2017, p. 209, fig.
6.3) as Transennatia gratiosa (Waagen, 1884). These
specimens can be referred to Transennatia gratiosa (Waagen,
1884, p. 691, pl. 72, figs. 3–7), from the Wargal and Chhidru
Formations of the Salt Range, Pakistan, on the basis of small
size (length 15 mm, width 21 mm in the largest dorsal valve
specimen, UHR12126), large triangular ears, strongly
geniculated dorsal valve with nearly flat visceral disc andmoderately long trail, and sharply reticulate ornament on
visceral discs of both ventral and dorsal valves, although the
Kitakami specimens are smaller than the type specimensfrom the Salt Range. Dictyoclostus minor Lee and Gu (in Lee
et al., 1980, p. 372, pl. 166, figs. 1–4.), from the Miaoling
Formation of Jilin, northeastern China, is probably a juniorsynonym of the present species. Transennatia insculpta(Grant, 1976, p. 135, pl. 32, figs. 1–37; pl. 33, figs. 1–6),
from the Rat Buri Formation of Ko Muk, southern Thailand,differs from T. gratiosa in being smaller size and in having
more extended ears.
Occurrence.— KN1 Unit (Locality KS2) and KN2 Unit(locality KS9) in Setamai; KY1 Unit in Imo (localities
KF217 and KF218) and Matsukawa (locality KZ9) .
Distribution. — Roadian‒Changhsingian: northeasternJapan (Setamai, Kamiyasse–Imo, Matsukawa, Ogatsu,
Takakurayama and Hitachi in the South Kitakami Belt),
central Japan (Moribu and Oguradani in the Hida Gaien
Belt), southwestern Japan (Mizukoshi in Kyushu Island),northern China (Shanxi), northeastern China (Heilongjiang
and Jilin), eastern Russia (South Primorye), eastern China(Anhui, Zhejiang and Jiangxi), central-southern China(Hubei, Hunan, Guangdong and Guangxi), southwestern
China (Guizhou, Sichuan), Greece (Hydra Island),southwestern China (Yunnan), Vietnam, Malaysia, Cambodia(Sisophon), northwestern China (Tibet), Nepal (Kumaon
Himalayas) and Pakistan (Salt Range).
Superfamily PRODUCTOIDEA Gray, 1840
Superfamily MARGINIFEROIDEA Stehli, 1954
Family COSTISPINIFERIDAE Muir-Wood and Cooper,
1960

42

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 25. A, B, Spinomarginifera lopingensis (Kayser); A, dorsal view of external latex cast (A1, A2), dorsal view of external mould (A3), dorsal (A4)and ventral view (A5) of internal mould, and ventral view (A6) of ventral external mould of conjoined shell, UHR12642; B, ventral view of external mould(B 1 , B 2 ), ventral view of internal mould (B 3 ), ventral view of external latex cast (B 4 ), and dorsal view of internal mould (B 5 ) of conjoined shell,
NU-B2021. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

43

FIGURE 26. A, B, Spinomarginifera kueichowensis Huang; A, ventral (A1, A2), anterior (A3) and lateral (A4) views of external latex cast of ventral valve,
UHR30074; B, external latex cast (B1, B2) and internal latex cast (B3) of dorsal valve, UHR30071; C, D, Spinomarginifera alpha Huang; C, ventral (C1,
C2), anterior (C3) and lateral (C4) views of external latex cast of ventral valve, and external mould (C5) of dorsal valve, UHR30086; D, external mould (D1,
D2) of dorsal valve, UHR12369. Scale bars are 1 cm.

44

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 27. A–C, Hexiproductus echidniformis (Chao); A, internal mould (A1, A2) of ventral valve, KCG50; B, external mould of dorsal valve, KCG53;
C, external mould of dorsal valve, KCG51; D, E, Spinomarginifera alpha Huang; D, external mould (D1, D2) and internal latex cast (D3) of dorsal valve,
UHR12371; E, ventral (E1, E2) and dorsal (E3) views of internal mould of conjoined shell, NU-B2474. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Subfamily SPINOMARGINIFERINAE Waterhouse, 2002
Genus SPINOMARGINIFERA Huang, 1932
Type species.—Spinomarginifera kueichowensis Huang, 1932.
Spinomarginifera lopingensis (Kayser, 1883)(Fig. 25A, B)
Productus nystianus var. lopingensis Kayser, 1883, p. 187,
pl. 28, figs. 1‒5.
Productus (Marginifera) helicus var. Frech, 1911, p. 130, pl.
19, figs. 1‒3.
Marginifera lopingensis (Kayser). Chao, 1927, p. 153, pl. 16,
figs. 8‒12.
Spinomarginifera kueichowensis Huang. Nakamura, 1959b,
p. 143, pl. 15, fig. 1 only; Minato et al., 1979, pl. 63, fig. 1
only.
Spinomarginifera lopingensis (Kayser). Zhang and Ching(Jin), 1961, p. 412, pl. 4, figs. 26–33; Wang et al., 1964, p.
312, pl. 49, figs. 21–23; Yang et al., 1977, p. 349, pl. 139,
fig. 5; Tong, 1978, p. 222, pl. 79, fig. 6; Licharew and
Kotlyar, 1978, pl. 15, figs. 9, 10; Zhan, 1979, p. 80, pl. 5,
figs. 17, 18; Liao, 1980, pl. 5, figs. 35–39; Wang et al.,
1982, p. 219, pl. 92, figs. 1, 2; Wang, 1984, p. 187, pl. 80,
fig. 16; Yang, 1984, p. 217, pl. 33, fig. 4; Liao, 1987, pl. 5,
figs. 5, 7–18; Zeng et al., 1995, pl. 9, fig. 1; Shen et al.,2002, p. 677, figs. 4.32, 4.33, 5.1–5.4; He et al., 2008, p.
812, fig. 4.1–4.10; Li and Shen, 2008, p. 315, figs.
4.17‒4.19, 6.1‒6.7; Shen and Zhang, 2008, fig. 4.13‒4.19;
Shen and Shi, 2009, p. 157, fig. 3P‒3X; Tazawa, 2012, p.
20, fig. 4.1–4.3; Tazawa, 2017, p. 41,fig. 6.1–6.3.
Spinomarginifera lopingensis (Chao). Jin et al., 1985, p. 194,
pl. 9, figs. 3–9; Jin, 1985, pl. 7, figs. 5, 16, 18, 20.
Spinomarginifera lopingensis Huang. Liao and Xu, 2002, pl.
1, figs. 28–33.
Spinomarginifera huangi Wang and Zhang, 2003, p. 73, pl.
21, fig. 12 only.
Material.—Seven specimens from localities KF7, KF90
and KF217: (1) external and internal moulds of two
conjoined shells, UHR12642, 30077; (2) internal moulds oftwo conjoined shells, with external moulds of the ventral
valves, NU-B2021, 2022; and (3) external moulds of threedorsal valves, NU-B2023, 2024, 2473.
Remarks.—Most of the specimens were previously
described by Tazawa (2017, p. 41, figs. 6.1–6.3) as
Spinomarginifera lopingensis (Kayser, 1883). The specimensfrom Kamiyasse–Imo can be referred to Spinomarginifera
lopingensis (Kayser, 1883), from the upper Permian of
Loping, Jiangxi Province, eastern China, in medium size(length about 19 mm, width about 32 mm in thebest-preserved conjoined shell, UHR12642) and in having
prominent costae on trails of both valves. Spinomarginifera
kueichowensis Huang (1932, p. 56, pl. 5, figs. 1‒11), fromthe upper Permian (Wuchiapingian) of Guizhou,

45

southwestern China, differs from S. lopingensis in lacking
radial costae on trails of both valves.
Occurrence.—KY1 Unit in Kamiyasse (localities KF7 andKF90) and Imo (KF217).
Distribution.—Wordian–Changhsingian: northeasternJapan (Kamiyasse‒Imo and Nabekoshiyama in the South
Kitakami Belt), northwestern China (Qinghai), northern
China (Inner Mongolia), eastern Russia (South Primorye),eastern China (Jiangsu, Anhui, Zhejiang and Jiangxi),
central-southern China (Hubei, Hunan, Guangdong andGuangxi), southwestern China (Guizhou, Sichuan andYunnan) and northwestern China (Tibet).
Spinomarginifera kueichowensis Huang, 1932
(Fig. 26A, B)
Spinomarginifera kueichowensis Huang, 1932, p. 56, pl. 5,
figs. 1‒11; Nakamura, 1959b, p. 143, pl. 15, figs. 2‒4
only; Muir-Wood and Cooper, 1960, p. 65, figs. 15‒22, 24;
Wang et al., 1964, p. 316, pl. 51, figs. 9‒11; Jin et al.,
1974, p. 312, pl. 164, fig. 13; Tazawa, 1976, pl. 2, fig. 1;
Feng and Jiang, 1978, p. 252, pl. 89, figs. 5, 6; Minato etal., 1979, pl. 63, fig. 2 only; Zhan, 1979, p. 80, pl. 11, figs.
14‒17, 20; Liao, 1980, pl. 4, fig. 29; Liu et al., 1982, p.
184, pl. 131, figs. 8‒10; Wang, 1984, p. 187, pl. 74, fig.
16; pl. 76, fig. 3; Zeng et al., 1995, pl. 5, fig. 10; Tazawa,
2002, fig. 10.11; Chen in Chen et al., 2006, p. 314, fig. 8,table 2; Shen and Shi, 2009, p. 158, figs. 3DD, 3EE, 4I;
Tazawa et al., 2014, p. 381, fig. 2.10; Tazawa, 2015, p. 67,
fig. 6.10; Tazawa, 2017, p. 43, fig. 7.1, 7.2.
Spinomarginifera nipponica Shimizu, 1961, p. 244, pl. 8,
figs. 1‒20; pl. 9, figs. 14–16.
Spinomarginifera cf. kueichowensis Huang. Lee et al., 1980,
p. 357, pl. 165, fig. 11; pl. 166, fig. 25; Gu, 1992, p. 224,
pl. 69, fig. 6.
Material.—Twelve specimens from localities KF63,
KF71, KF84, KF88, KF90 and KF217: (1) external andinternal moulds of a conjoined shell, NU-B2017; (2) internalmoulds of two conjoined shells, with external mould of theventral valves, UHR12584, 30074; (3) external and internalmoulds of four dorsal valves, NU-B2018, 2019, 2020,
UHR30071; and (4) external moulds of five dorsal valves,
UHR11554, 12368, 30068, 30080, 30082.
Remarks.—These specimens were described by Tazawa(2017, p. 43, fig. 7.1, 7.2) as Spinomarginifera kueichowensis
Huang, 1932. The specimens from Kamiyasse–Imo can bereferred to Spinomarginifera kueichowensis Huang, 1932,
from the upper Permian (Wuchiapingian) of Guizhou,
southwestern China, in medium size (length 18 mm, widthabout 38 mm in the largest specimen, UHR30074), transverse
outline, numerous spine bases on ventral valve and strongly
developed marginal ridge in dorsal valve. Spinomarginifera
nipponica Shimizu, 1961, from the Gujo Formation of
Kawahigashi in the Maizuru Belt, southwestern Japan, is46

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 28. A, B, Paramarginifera sp.; A, ventral (A1, A2), anterior (A3), posterior (A4) and lateral (A5) views of internal mould of ventral valve,
NU-B2589; B, ventral view (B1, B2) of internal mould of ventral valve, NU-B2590. Scale bars are 1 cm.

regarded as a junior synonym of S. kueichowensis. The
preceding species, Spinomarginifera lopingensis (Kayser,
1883), is distinguished from S. kueichowensis in having
costae on trails of both ventral and dorsal valves.
Spinomarginifera alpha Huang, 1932, originally described by
Huang (1932, p. 60, pl. 5, figs. 12, 13) as Spinomarginifera
kueichowensis mut. alpha Huang, 1932, from the upper
Permian of Guizhou, southwestern China, differs from thepresent species in its larger size and in having coarser andsparser spine bases on ventral valve.
Occurrence.—KY1 Unit in Kamiyasse (localities KF63,
KF71, KF84, KF88 and KF90) and Imo (locality KF217).
Distribution.—Wordian–Changhsingian: northeasternJapan (Kamiyasse–Imo and Hitachi in the South KitakamiBelt), southwestern Japan (Kawahigashi in the Maizuru Belt),
northeastern China (Jilin), eastern China (Zhejiang),
central-southern China (Hubei, Hunan, Guangdong andGuangxi) and southwestern China (Guizhou and Sichuan).
Spinomarginifera alpha Huang, 1932
(Figs. 26C, D; 27D, E)
Spinomarginifera kueichowensis mut. alpha Huang, 1932, p.

60, pl. 5, figs. 12, 13; Wang et al., 1964, p. 316, pl. 49,
figs. 31–33; Jin et al., 1974, p. 313, pl. 164, figs. 11, 12;
Yang et al., 1977, p. 349, pl. 139, fig. 9.
Spinomarginifera huangi Nakamura, 1959b, p. 145, pl. 15,
figs. 5–7; Minato et al., 1979, pl. 63, figs. 3–5.
Spinomarginifera alpha Huang. Liao, 1980, p. 259, pl. 5,
figs. 44–47; Zhao et al., 1981, pl. 8, figs. 28, 29; Wang etal., 1982, p. 219, pl. 96, fig. 26; Shen et al., 2003a, p. 231,
pl. 1, figs. 6–9; Chen et al., 2005, p. 355, fig. 10C, D;
Tazawa, 2017, fig. 8.1–8.4.
Material. — Eight specimens from localities KF89, KF90
and KF217: (1) internal mould of a conjoined shell, withexternal mould of the ventral valve, NU-B2025; (2) internalmoulds of two conjoined shells, with external moulds of thedorsal valves, NU-B2026, UHR12370; (3) internal mould of
a conjoined shell, NU-B2474; (3) external moulds of aconjoined shell, UHR30086; (4) external and internal moulds
of a dorsal valve, UHR12371; and (5) external mould of two
dorsal valves, UHR12369, NU-B2475.
Remarks. — The specimens from Kamiyasse–Imo can bereferred to Spinomarginifera alpha Huang, 1932, from the
Lopingian of Guizhou, southwestern China, in being large

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

47

FIGURE 29. A, Kunlunia sp., external mould (A1) and external latex cast (A2) of dorsal valve, KCG59; B–D, Costiferina spiralis (Waagen); B, externallatex cast (B1) and internal mould (B2) of ventral valve, UHR16004; C, ventral (C1) and dorsal (C2) views of internal mould, and dorsal (C3) and lateral(C4) views of external mould of conjoined shell, UHR12373; D, dorsal (D1), anterior (D2) and lateral (D3) views of external mould of dorsal valve,
UHR13555. Scale bars are 1 cm.

48

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 30. A–C, Tyloplecta yangtzeensis (Chao); A, ventral (A1), dorsal (A2) and lateral (A3) views of internal mould of conjoined shell, NU-B1845; B,
external latex cast (B1) and internal mould (B2) of ventral valve, UHR13553; C, ventral (C1), anterior (C2), posterior (C3), lateral (C4) and dorsal (C5) viewsof internal mould, and dorsal view (C6) of external mould of conjoined shell, UHR12602. Scale bars are 1 cm.

size (length 29 mm, width 35 mm in the best preserved
ventral valve specimen, UHR30086; length 30 mm, width 36
mm in the best-preserved dorsal valve specimen, UHR12371)
and in having coarse, widely-spaced spine bases on both
ventral and dorsal valves. Spinomarginifera huangi
Nakamura (1959b, p. 145, pl. 15, figs. 5‒7), from the lower
Kamiyasse Formation of Kamiyasse, is regarded as a juniorsynonym of the present species. The preceding species,

Spinomarginifera kueichowensis Huang, 1932, is readily
distinguished from S. alpha by its smaller size, muchtransverse outline and more numerous and finer spine bases
on both ventral and dorsal valves.
Occurrence.—KY1 Unit in Kamiyasse (localities KF89
and KF90) and Imo (locality KF217).
Distribution.—Wordian‒Changhsingian: northeasternJapan (Kamiyasse‒Imo in the South Kitakami Belt), easternGUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

China (Zhejiang), central-southern China (Hunan,
Guangdong and Guangxi), southwestern China (Guizhou andSichuan) and northwestern China (Tibet).
Family PAUCISPINIFERIDAE Muir-Wood and Cooper,
1960
Subfamily PAUCISPINIFERINAE Muir-Wood and Cooper,
1960
Genus PARAMARGINIFERA Fredericks, 1916
Type species.—Marginifera clarkei Tschernyschew, 1902.
Paramarginifera sp.
(Fig. 28A, B)
Material.—Three specimens from localities KF121 andKF217, internal moulds of three ventral valves,
NU-B2589–2591.
Remarks.—These specimens are safely assigned to thegenus Paramarginifera Fredericks, 1916 by its medium-sized(length about 25 mm, width about 30 mm in the best
preserved specimen, NU-B2589) and highly inflated ventral
valve with large trigonal-shaped ears, a narrow and deep
sulcus on trail and a conspicuous cincture around
anterolateral margins of corpse. The external ornament ofventral valve, which is faintly preserved, is consisted ofnumerous fine costellae. Internal structures of ventral valve
are not well preserved, except for a highly raised adductor
muscle scars in central part of the valve. The Kitakami
species somewhat resembles Paramarginifera gobiensis(Chao, 1927), redescribed by Wang and Zhang (2003, p. 69,
pl. 1, figs. 38–42; pl. 12, figs. 1, 2; pl. 21, figs. 17–21) fromthe Zhesi Formation (Wordian–Capitanian) of Zhesi, Inner
Mongolia, northern China, but accurate comparison isdifficult owing to ill-preservation of the present material.
Occurrence.—KY1 Unit in Imo (localities KF121 andKF217).
Genus HEXIPRODUCTUS Shi, Chen and Tong, 2008
Type species.—Productus echidniformis Chao, 1925.
Hexiproductus echidniformis (Chao, 1925)(Fig. 27A–C)
Productus echidniformis Grabau em. Chao, 1925, p. 239, pl.
2, figs. 7–9.
Avonia echidniformis (Grabau em. Chao). Chao, 1927, p.
120, pl. 14, figs. 17–27; Chao, 1928, p. 55, pl. 6, fig. 7;
Ozaki, 1931, p. 108, pl. 10, figs. 6–9; Nakamura, 1959a, p.
201, pl. 1, figs. 2–8; Volgin, 1960, p. 47, pl. 4, fig. 4;
Sergunkova and Zhizhilo, 1975, p. 60, pl. 9, fig. 7; Lee andGu, 1976, p. 240, pl. 141, fig. 1; Minato et al., 1979, pl.
46, figs. 15–17; Lee et al., 1980, p. 350, pl. 145, fig. 25;
Lee and Duan, 1985, p. 227, pl. 66, figs. 16–21; Zhan and

49

Wu, 1987, p. 203, pl. 47, figs. 23–25; He et al., 1995, pl.
56, figs. 51, 52, 61–64; Wang, 1995, pl. 1, fig. 4; Wang
and Yang, 1998, p. 67, pl. 3, figs. 21–25.
P r o d u c t u s (A v o n i a ) e c h i d n i f o r m i s G r a b a u a n d C h a o .
Licharew, 1939, p. 86, pl. 17, figs. 9, 10.
Avonia sp. Minato et al., 1979, pl. 46, figs. 13, 14.
“Avonia” echidniformis (Grabau em. Chao). Zhang et al.,1983, p. 286, pl. 131, fig. 1.
“Avonia”? echidniformis (Grabau em. Chao). Chen and Shi,
2002, p. 299, fig. 4J.
Breileenia echidniformis (Grabau in Chao). Chen and Shi,
2006a, p. 137, pl. 1, figs. 13, 14; text-fig. 10.
Hexiproductus echidniformis (Grabau in Chao). Shi et al.,2008, p. 290, fig. 6A–D.
Hexiproductus echidniformis (Chao). Tazawa and Nakamura,2015, p. 161, fig. 4.4–4.10; Tazawa and Araki, 2017, p.
271, fig. 6.4–6.6.
Material.—Four specimens from localities AR4 and KZ9:(1) internal mould of a ventral valve, KCG50; and (2)external moulds of three dorsal valves, KCG51–53.
Remarks.—These specimens were previously describedby Tazawa and Araki (2017, p. 271, fig. 6.4–6.6) as
Hexiproductus echidniformis (Chao, 1925). The Matsukawa
specimens can be referred to Hexiproductus echidniformis(Chao, 1925), originally described from the upper
Carboniferous‒lower Permian of Gansu, northwestern China
and Shanxi, northern China, in transversely subrectangularoutline and external ornament of both ventral and dorsal
valves, consisting of irregular fine concentric rugae andstrong costae with numerous elongate spine bases.
Occurrence.—KY1 Unit in Matsukawa (localities AR4
and KZ9).
Distribution.—Kasimovian‒Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt), northwestern
China (Xinjiang, Gansu and Ningxia), northern China (Inner
Mongolia, Shanxi and Hebei), northeastern China (Liaoning),
eastern China (Shandong) and Uzbekistan (Fergana).
Family DICTYOCLOSTIDAE Stehli, 1954
Subfamily DICTYOCLOSTINAE Stehli, 1954
Genus KUNLUNIA Wang in Zhang et al., 1983
Type species.—Kunlunia aspera Wang in Zhang et al.,1983.
Kunlunia sp.
(Fig. 29A)
Kunlunia sp. Tazawa and Araki, 2018, p. 13, fig. 4.4.
Material. ― One specimen from locality AR4, externalmould of a dorsal valve, KCG59.
Remarks. ― This specimen was previously described by
Tazawa and Araki (2018, p. 13, fig. 4.4) as Kunlunia sp. on

50

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 31. A–D, Tyloplecta yangtzeensis (Chao); A, external mould of dorsal valve, NU-B1841; B, external mould (B1) and external latex cast (B2) of
dorsal valve, NU-B1839; C, ventral (C1), dorsal (C2) and lateral (C3) views of internal mould, external mould (C4) and internal latex cast (C5) of dorsal
valve of conjoined shell, NU-B1835; D, external latex cast of ventral valve, NU-B1837. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

51

FIGURE 32. A–C, Bathymyonia neimongolica (Wang and Zhang); A, ventral (A1), dorsal (A2), anterior (A3), posterior (A4) and lateral (A5) views ofinternal mould of conjoined shell, NU-B19810; B, ventral (B1) and dorsal (B2) views of internal mould, and dorsal (B3) and lateral (B4) views of externalmould of conjoined shell, UHR13131; C, ventral (C1) and dorsal (C2) views of internal mould, and internal latex cast (C3, C4) of dorsal valve of conjoined
shell, UHR13132. Scale bars are 1 cm.

52

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 33. A–E, Vediproductus punctatiformis (Chao); A, ventral (A1), anterior (A2), posterior (A3) and lateral (A4) views of internal mould of ventral
valve, UHR12472; B, internal mould (B1) and external latex cast (B2) of ventral valve, NU-B1660; C, ventral (C1) and dorsal (C2) views of internal mould,
and dorsal external mould (C3) of conjoined shell, NU-B1662; D, external mould (D1, D2) of dorsal valve, NU-B1666; E, ventral (E1, E2), anterior (E3),posterior (E4) and lateral (E5) views of internal mould of ventral valve, NU-B1664. Scale bars are 1 cm.

the basis of a dictyoclostid shell with large extended ears on
dorsal valve. The Matsukawa species somewhat resembles
Kunlunia grabaui (Nakamura, 1960), redescribed by Tazawa
and Nakamura (2015, p. 162, fig. 5.1–5.3) from the lower
part of the Hosoo Formation (Kungurian) of Nakadaira,South Kitakami Belt, in shape and external ornament of
dorsal valve, but differs in its larger size (length 54 mm,
width 76 mm in the dorsal valve specimen, KCG59). Specificidentification is, however, difficult for the poorly preserved
specimen.
Occurrence.―KY1 Unit in Matsukawa (locality AR4).

Subfamily CALLYTHARRELLINAE Waterhouse, 2002
Genus COSTIFERINA Muir-Wood and Cooper, 1960
Type species.―Productus indicus Waagen, 1884.
Costiferina spiralis (Waagen, 1884)(Fig. 29B–D)
Productus spiralis Waagen, 1884, p. 681, pl. 67, fig. 6; pl.
68, fig. 3; pl. 69, figs. 1–3; Diener, 1915, p. 67, pl. 6, fig.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

16; pl. 7, figs. 1, 2.
Productus vishnu Waagen. Hayasaka, 1922a, p. 60, pl. 10,
figs. 3–8.
Dictyoclostus aff. spiralis (Waagen). Nakamura, 1960, p.
497, pl. 1, figs. 1, 2; pl. 2, figs. 1, 2.
Costiferina aff. spiralis (Waagen). Minato et al., 1979, pl.
64, fig. 2.
Material. ― Four specimens from localities KF217 andKanokurazawa (exact location unclear): (1) external andinternal moulds of a conjoined shell, UHR12373; (2) internalmould of a conjoined shell, with external mould of theventral valve, UHR16004; and (3) external moulds of two
dorsal valves, UHR13555, 13556.
Remarks. ― Most of the specimens, except for the largest
one (UHR16004), were previously described by Nakamura(1960, p. 497, pl. 1, figs. 1, 2; pl. 2, figs. 1, 2) as
Dictyoclostus aff. spiralis Waagen (1884). All the specimens
are, however, referred to Costiferina spiralis (Waagen, 1884,
p. 681, pl. 67, fig. 6; pl. 68, fig. 3; pl. 69, figs. 1–3), from the
Amb Formation of the Salt Range, Pakistan, by large,transverse shell (length about 55 mm, width about 115 mm in
the largest specimen, UHR16004), ornamented with coarse
costae (numbering 3–4 in 10 mm at about midlength ofventral valve) on both ventral and dorsal valves and strongly
geniculated dorsal valve. The type species, Costiferina indica(Waagen, 1884, p. 687, pl. 70, figs. 1–6; pl. 71, fig. 1), fromthe Wargal and Chhidru Formations of the Salt Range, differsfrom C. spiralis in its smaller size and less transverse outline.
Occurrence. ― Lower Kanokura Formation in Setamai(exact location unclear); and KY1 Unit in Imo (locality
KF217).
Distribution. ― Roadian–Changhsingian: northeasternJapan (Setamai and Kamiyasse–Imo in the South KitakamiBelt), Pakistan (Salt Range) and India (Kashmir).
Family BUXTONIIDAE Muir-Wood and Cooper, 1960
Subfamily TYLOPLECTINAE Termier and Termier, 1970
Genus TYLOPLECTA Muir-Wood and Cooper, 1960
Type species. ― Productus scabriculus mut. nankingensis
Frech, 1911.
Tyloplecta yangtzeensis (Chao, 1927)(Figs. 30A–C, 31A–D)
Productus semireticulatus Martin. Kayser, 1883, p. 181, pl.
25, figs. 1–4.
Productus costatus Sowerby. Kayser, 1883, p. 182, pl. 25,
figs. 5–7.
Productus sumatrensis Roemer. Fliegel, 1901, p. 128, pl. 6,
fig. 1.
Productus yangtzeensis Chao, 1927, p. 50, pl. 5, figs. 1–3; pl.
8, fig. 9; Simić, 1933, p. 31, 92, pl. 1, figs. 11‒14.

53

Productus (Dictyoclostus) yangtzeensis Chao. Huang, 1932,
p. 26, pl. 1, figs. 18–21.
Dictyoclostus yangtzeensis (Chao). Ramovs, 1958, p. 506, pl.
3, fig. 1.
Dictyoclostus sino-indicus (Frech). Nakamura, 1960, p. 499,
pl. 2, figs. 2–4; pl. 3, fig. 1; Minato et al., 1979, pl. 63, fig.
6; pl. 64, fig. 1.
Tyloplecta yangtzeensis (Chao). Schréter, 1963, p. 124, pl. 6,
figs. 1‒7; Yanagida, 1964, p. 3, pl. 1, figs. 1, 3, text-fig.
2; Ting, 1965, p. 268, pl. 3, figs. 1, 2; Sarytcheva in
Ruzhentsev and Sarytcheva, 1965, pl. 37, fig. 1; Fantini
Sestini and Glaus, 1966, p. 909, pl. 64, figs. 1, 4; pl. 65,
fig. 1; Pitakpaivan et al., 1969, p. 10, pl. 17, figs. 13‒15;
Termier and Termier, 1970, p. 458, pl. 31, fig. 4; Yang etal., 1977, p. 361, pl. 143, fig. 4; Feng and Jiang, 1978, p.
258, pl. 92, figs. 5‒8; Licharew and Kotlyar, 1978, p.
12, figs. 11, 12; pl. 19, figs. 8, 9; Tong, 1978, p. 230, pl.
81, fig. 4; Zhan, 1979, p. 88, pl. 6, figs. 20–22; Liao,
1980, pl. 4, figs. 21, 22; Nakamura et al., 1981, p. 44,
pl. 1, figs. 1–3; pl. 3, figs. 1, 2; Liu et al., 1982, p. 186, pl.
133, figs. 4, 5; Wang et al., 1982, p. 212, pl. 89, figs. 5, 6;
pl. 92, figs. 10, 11; Ding and Qi, 1983, p. 286, pl. 98,
fig. 4; Liao, 1987, p. 104, pl. 5, figs. 23, 24; Liang, 1990,
p. 192, pl. 33, figs. 1–9; Zhu, 1990, p. 70, pl. 13, figs. 1–9;
pl. 14, fig. 28; Zeng et al., 1995, pl. 7, figs. 2, 3, 7; He etal., 2008, p. 815, fig. 3.14, 3.15; Shen and Zhang, 2008,
fig. 5.1, 5.2; Shen and Shi, 2009, p. 159, fig. 4A–H;
Tazawa, 2016b, p. 16, figs. 5.6, 6.1–6.3.
Tyloplecta sino-indicus (Frech). Tazawa, 1976, pl. 2, fig. 13.
Tyloplecta cf. yangtzeensis (Chao). Tazawa and Ibaraki,
2001, p. 8, pl. 1, fig. 5.
Tyloplecta sp. Tazawa, 2002, fig. 10.6.
Material. ― Twenty-one specimens from localities KF96,
KF121, KF217 and KS8: (1) external and internal moulds oftwo conjoined shells, UHR12602, NU-B1842; (2) internalmould of a conjoined shell, with external mould of theventral valve, NU-B1893; (3) internal mould of a conjoined
shell, with external mould of the dorsal valve, NU-B1835,
UHR12603; (4) internal moulds of three conjoined shells,
NU-B1845–1847; (5) external cast of a ventral valve,
NU-B1836; (6) external mould of a ventral valve NU-B1837;(7) internal mould of two ventral valves, NU-B1838,
UHR13554; and (8) external moulds of eight dorsal valves,
NU-B335, 1839‒1841, 1849, 1848–1850.
Remarks. —Most of the specimens were previously
described by Tazawa (2016b, p. 16, figs. 5.6, 6.1–6.3) as
Tyloplecta yangtzeensis (Chao, 1927). The specimens from
Setamai and Kamiyasse–Imo can be referred to Tyloplecta
yangtzeensis (Chao, 1927, p. 50, pl. 5, figs. 1–3; pl. 8, fig. 9),from the Lungtan Formation of Zhejiang, eastern China, bylarge size (length 55 mm, width 66 mm in the largest
specimen, NU-B1837), subquadrate to subglobose outline
and external ornament of both ventral and dorsal valves. The
preceding species, Tyloplecta sinoindica (Frech, 1911), is

54

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 34. A–D, Waagenoconcha humboldti (d’Orbigny); A, external latex cast of ventral valve, UHR12596; B, external latex cast (B1, B2) andenlarged external ornament (B3) of ventral valve, UHR14847; C, external mould (C1) and external latex cast (C2) of dorsal valve, UHR19846; D, internallatex cast (D1) and enlarged posterior portion showing internal structures (D2) of dorsal valve, UHR19842. Scale bars are 1 cm, except for B3 and D2.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

55

FIGURE 35. A–E, Waagenoconcha irginae (Stuckenberg); A, ventral (A1) and dorsal (A2) views of internal mould, dorsal external mould (A3) and ventral
external latex cast (A4) of conjoined shell, UHR19841; B, internal mould of ventral valve, UHR19828; C, internal mould (C1), external latex cast (C2, C3)
and enlarged external latex cast, showing spine bases on external surface (C4) of ventral valve, UHR19818; D, external mould (D1), external latex cast(D2), internal mould (D3), internal latex cast (D4) and enlarged posterior portion, showing cardinal process, median septum, lateral septa and muscle scars(D5) of dorsal valve, UHR19821; E, external cast (E1) and internal mould (E2) of ventral valve, UHR19822. Scale bars are 1 cm, except for C4 and D5.

56

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 36. A–D, Waagenoconcha irginae (Stuckenberg); A, external latex cast (A 1, A2) and internal mould (A3) of ventral valve, UHR19838; B,
external latex cast (B1) and internal latex cast (B2) of dorsal valve, UHR19826; C, external mould of dorsal valve, UHR19831; D, ventral (D1), dorsal (D2),anterior (D3), posterior (D4) and lateral (D5) views of internal mould of conjoined shell, UHR19813; E, F, Scacchinella gigantea Schellwien; E, anteriorview of internal mould of ventral valve, NU-B197; F, anterior (F1) and posterior (F2) views of internal mould of ventral valve, NU-B198. Scale bars are 1
cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

readily distinguished from T. yangtzeensis by its moreelongate and strongly inflated ventral valve and trapezoidal
dorsal valve. Tyloplecta nankingensis (Frech, 1911, p. 163,
pl. 22, fig. 3), from the Lungtan Formation of Jiangsu,
eastern China, differs from the present species by its muchsmaller size and in having more strongly convex ventral
valve. The specimens, described by Nakamura (1960, p. 499,
pl. 2, figs. 2–4; pl. 3, fig. 1) and figured by Minato et al.
(1979, pl. 63, fig. 6; pl. 64, fig. 1) as Dictyoclostus
sinoindicus (Frech, 1911) from the lower Kanokura Series (=lower Kamiyasse Formation) of Imo in the South Kitakami
Belt, are referred to Tyloplecta yangtzeensis.
Occurrence. — KN1 Unit in Setamai (locality KS8); andKY1 Unit in Kamiyasse (locality KF96) and Imo (localities
KF121 and KF217).
Distribution. — Roadian–Changhsingian: northeasternJapan (Setamai and Kamiyasse–Imo in the South KitakamiBelt), Hungary, Slovenia, Serbia, northwestern China(Qinghai), northern China (Shanxi), eastern Russia (South
Primorye), eastern China (Jiangsu, Anhui, Zhejiang, Jiangxi
and Fujian), central-southern China (Hubei, Hunan,
Guangdong and Guangxi), southwestern China (Guizhou andSichuan), Armenia, Iran and north-central Thailand.
Superfamily ECHINOCONCHOIDEA Stehli, 1954
Family ECHINOCONCHIDAE Stehli, 1954
Subfamily JURESANIINAE Muir-Wood and Cooper, 1960
Genus BATHYMYONIA Muir-Wood and Cooper, 1960
Type species.—Productus nevadensis Meek, 1877.

57

by Tazawa (2014, p. 17, fig. 3.1–3.4) as Bathymyonia
neimongolica (Wang and Zhang, 2003). These specimens canbe referred to Bathymyonia neimongolica (Wang and Zhang,2003), originally described by Wang and Zhang (2003, p. 97,
pl. 8, figs. 1–8; pl. 9, figs. 1–4; pl. 14, figs. 4–7) from theZhesi Formation of Inner Mongolia, northern China, on
account of elongate subrectangular outline, flattened visceraldisc , steep lateral slopes of ventral valve and enormously
large cardinal process in dorsal valve. Bathymyonia
barabaschensis Kotlyar in Licharew and Kotlyar (1978, p.
67, pl. 16, fig. 1), from the upper Barabashevka Formation ofSouth Primorye, eastern Russia, differs from B. neimongolica
by its less elongate, more strongly and evenly convex ventral
valve. The type species, Bathymyonia nevadensis (Meek,
1877), refigured by Muir-Wood and Cooper (1960, pl. 82,
figs 1–7; pl. 109, fig. 1) on the specimens from the
Phosphoria Formation of Wyoming, Utah and Nevada in theUSA, is distinguished from B. neimongolica by its smaller,
less elongate and more evenly convex ventral valve.
Occurrence.—KY1 Unit in Kamiyasse (locality KF46)
and Imo (localities KF217 and KF218).
Distribution.— Wordian–Capitanian: northeastern Japan(Kamiyasse–Imo and Hitachi in the South Kitakami Belt) andnorthern China (Inner Mongolia).
Genus VEDIPRODUCTUS Sarytcheva in Sarytcheva and
Sokolskaya, 1965
Type species. — Vediproductus vediensis Sarytcheva in
Sarytcheva and Sokolskaya, 1965.

Bathymyonia neimongolica (Wang and Zhang, 2003)(Fig. 32A–C)

Vediproductus punctatiformis (Chao, 1927)(Fig. 33A–E)

Waagenoconcha imperfecta Prensergast. Tazawa, 1974b, p.
127, pl. 3, fig. 2 only; Minato et al., 1979, pl. 65, figs. 1, 2;
Shiino and Suzuki, 2007, figs. 3–6.
Waagenoconcha (Yazengoconcha) neimongolica Wang andZhang, 2003, p. 97, pl. 8, figs. 1–8; pl. 9, figs. 1–4; pl. 14,
figs. 4–7.
Bathymyonia neimongolica (Wang and Zhang, 2003).
Tazawa, 2014, p. 17, fig. 3.1–3.4; Tazawa et al., 2014, p.
381, fig. 3.1, 3.2; Tazawa, 2015, p. 69, fig. 7.1, 7.2 .

Echinoconchus punctatiformis Chao, 1927, p. 72, pl. 6, figs.
9–12; Zhan and Lee, 1962, p. 477, pl. 2, fig. 9.
Echinoconchus cf. fasciatus (Kutorga). Minato et al., 1979,
pl. 65, figs. 5–7.
Vediproductus punctatiformis (Chao). Wang et al., 1982, p.
208, pl. 80, figs. 10, 11; pl. 84, fig. 4; Ding and Qi, 1983,
p. 282, pl. 96, fig. 3; Wang, 1984, p. 190, pl. 76, fig. 7;
Chang, 1987, p. 759, pl. 2, fig. 4; Liang, 1990, p. 187, pl.
27, fig. 5; Shiino, 2009, p. 255, fig. 4A–H; Tazawa, 2016b,
p. 19, fig. 7.1–7.5.
Vediproductus mugenjin Shiino, 2009, p. 252, fig. 3.

Material.—Six specimens from localities KF46, KF217
and KF218; (1) internal moulds of two conjoined shells, withexternal moulds of the dorsal valves, UHR13131, 13132; (2)internal mould of a conjoined shell, with partly preserved
shell substance, UHR19810; (3) internal moulds of two
ventral valves, UHR12137, 19811; and (4) external mould ofa dorsal valve, NU-B1658.
Remarks. — The specimens from the lower Kamiyasse
Formation in the Kamiyasse–Imo area were first described by
Tazawa (1974b, p. 127, pl. 3, fig. 2 only) as Waagenoconcha
imperfecta Prendergast, 1935, and subsequently redescribed

Material.—Fourteen specimens from localities KF64,
KF217 and KF218: (1) external and internal moulds of two
conjoined shells, NU-B1660, 1661; (2) internal moulds oftwo conjoined shells, with external moulds of the dorsal
valves, NU-B1662, 1663; (3) external and internal moulds ofa ventral valve, NU-B1664; (4) internal moulds of six ventral
valves, NU-B1666, 1667, UHR12022, 12472, 12612, 12641;
and (5) external moulds of three dorsal valves, NU-B1665,
1668, UHR17083.

58

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 37. A– D, Edriosteges multispinosus Muir-Wood and Cooper; A, external mould (A 1 ) and external latex cast (A 2 , A 3 ) of ventral valve,
UHR12634; B, ventral external latex cast (B1), ventral internal mould (B2), dorsal external mould (B3) and dorsal internal latex cast (B4, B5) of conjoined
shell, UHR12631; C, external latex cast (C1) and external mould (C2) of dorsal valve, UHR12633; D, external latex cast (D1) and external mould (D2) ofventral valve, UHR12665. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

59

FIGURE 38. A, Limbella sp., external latex cast (A1, A2, A3) and internal latex cast (A4) of dorsal valve, NU-B2572. Scale bars are 1 cm.

Remarks.—These specimens were described by Tazawa(2016b, p. 19, fig. 7.1‒7.5) as Vediproductus punctatiformis(Chao, 1927). The specimens from Kamiyasse–Imo arereferred to Vediproductus punctatiformis (Chao, 1927, p. 72,
pl. 6, figs. 9‒12), from the Ksiaokiang Limestone of Jiangxi,
eastern China, in medium size (length 34 mm, width 26 mm
in the best-preserved specimen, NU-B1660), elongate oval
and strongly convex ventral valve and numerous regularspinose bands on both ventral and dorsal valves.
Vediproductus mugenjin Shiino (2009, p. 252, fig. 3), fromthe lower Kamiyasse Formation of Kamiyasse, may be asynonym of the present species. The type species,
Vediproductus vediensis Sarytcheva (in Sarytcheva and
Sokolskaya, 1965, p. 221, pl. 35, figs. 1‒3, text-fig. 33) fromthe Gnishik Horizon (Roadian) of the Transcaucasus, differsfrom V. punctatiformis in its less convex and wider ventral
valve and broader concentric bands on both ventral anddorsal valves.
Occurrence.—KY1 Unit in Kamiyasse (locality KF64)
and Imo (localities KF217 and KF218).
Distribution.—Asselian‒Capitanian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt), northwestern
China (Xinjiang and Gansu), eastern China (Anhui, Zhejiang
and Jiangxi) and central-southern China (Hubei).
Family WAAGENOCONCHIDAE Muir-Wood and Cooper,
1960
Subfamily WAAGENOCONCHINAE Muir-Wood andCooper, 1960
Genus WAAGENOCONCHA Chao, 1927

Type species.—Productus humboldti d’Orbigny, 1842.
Waagenoconcha humboldti (d’Orbigny, 1842)(Fig. 34A–D)
Productus humboldti d’Orbigny, 1842, p. 54, pl. 5, figs. 4–7;
Tschernyschew, 1902, p. 275, 620, pl. 53, figs. 1–3;
Kozlowski, 1914, p. 40, pl. 7, figs. 7–9; Fredericks, 1925,
p. 19, pl. 2, fig. 84.
Waagenoconcha humboldti (d’Orbigny). Chao, 1927, p. 86,
pl. 15, figs. 2, 3; Sarytcheva in Sarytcheva and
Sokolskaya, 1952, p. 98, pl. 15, fig. 109; Chronic, 1953, p.
86, pl. 15, figs. 4–7; Muir-Wood and Cooper, 1960, pl. 89,
figs. 6–10: Samtleben, 1971, p. 60, pl. 2, figs. 17–19;
Ifanova, 1972, p. 102, pl. 3, figs. 11–13; Tazawa, 1974b,
p. 125, pl. 1, figs. 2, 3; pl. 2, fig. 1; pl. 4, fig. 6; Ding and
Qi, 1983, p. 283, pl. 96, fig. 9; Duan and Li, 1985, p. 108,
pl. 35, figs. 2, 3; Wang and Zhang, 2003, p. 94, pl. 9, figs.
5‒7; pl. 15, figs. 8–10; Tazawa and Shintani, 2010, p. 56,
fig. 4.1–4.5; Tazawa and Shintani. 2015, p. 45, fig. 4.1 ;Tazawa, 2017, p. 47, fig. 9.1–9.4; Tazawa and Shintani,
2022, p. 24, fig. 20C, D.
Waagenoconcha imperfecta Prendergast. Tazawa, 1974b, p.
127, pl. 2, fig. 6 only.
Material.—Eight specimens from localities KF121 andKF217: (1) internal mould of a conjoined shell, UHR19819;(2) external and internal moulds of two ventral valves,
UHR12133, 19847; (3) external moulds of two ventral
valves, UHR12595, 12596; (4) external mould of a dorsal
valve, UHR19846; and (5) internal moulds of two dorsal

60

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 39. A–D, Urushtenoidea crenulata (Ding); A, anterior view of internal mould (A1, A2) and internal latex cast (A3, A4) and dorsal view of
external mould (A5) of conjoined shell, UHR30386; B, external mould (B1) and external latex cast (B2) of dorsal valve, NU-B2150; C, ventral (C1, C2),
dorsal (C3), anterior (C4), posterior (C5) and lateral (C6) views of internal mould of conjoined shell, NU-B2148; D, ventral (D1, D2), dorsal (D3), anterior(D4), posterior (D5) and lateral (D6) views of internal mould of conjoined shell, NU-B2479. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

valves, UHR19842, 19843.
Remarks.—These specimens were previously describedby Tazawa (1974b, p. 125, pl. 1, figs. 2, 3; pl. 2, fig. 1; pl. 4,
fig. 6) and Tazawa (2017, p. 47, fig. 9.1–9.4) as
Waagenoconcha humboldti (d’Orbigny, 1842). The
specimens from Kamiyasse–Imo are referred to
Waagenoconcha humboldti (d’Orbigny, 1842), originallydescribed from the lower Permian (Asselian) of Yarbichambi,
Bolivia, on account of medium size (length about 33 mm,
width about 45 mm in the largest specimen, UHR12596),
transversely subrectangular outline and external ornament ofboth valves, particularly, in having elongate, relatively coarse
spine bases (numbering 4–5 spine bases in 5 mm width atmidlength) and strong rugae on ventral valve.
Waagenoconcha abichi (Waagen, 1884, p. 697, pl. 74, figs.
1–7), from the Wargal and Chhidru Formations of the Salt
Range, Pakistan, is also a medium-sized species, but the
Pakistani species differs from W. humboldti in having coarser
and sparser spine bases on ventral valve.
Occurrence.—KY1 Unit in Imo (localities KF121 andKF217).
Distribution. — Gzhelian–Capitanian: northeastern Japan(Nagaiwa–Sakamotozawa and Kamiyasse–Imo in the South
Kitakami Belt), northern Russia (Timan and Pechora Basin),western Russia (Moscow Basin), northwestern China(Gansu), northern China (Inner Mongolia), eastern Russia(South Primorye) and Bolivia.
Waagenoconcha irginae (Stuckenberg, 1898)(Figs. 35A–E, 36A–D)
Productus irginae Stuckenberg, 1898, p. 220, pl. 2, fig. 16;
Tschernyschew, 1902, p. 273, 618, pl. 30, figs. 3, 4; pl. 52,
figs. 1‒4; Miloradovich, 1935, p. 67, 133, pl. 5, figs. 1, 2.
Productus cf. humboldti irginae Stuckenberg. Fredericks,
1925, p. 19, pl. 4, fig. 117.
Waagenoconcha humboldti var. irginae (Stuckenberg).
Solomina, 1960, p. 31, pl. 2, figs. 1–4.
Waagenoconcha irginae (Stuckenberg). Muir-Wood andCooper, 1960, pl. 89, figs. 15, 16; Gobbett, 1963, p. 76, pl.
5, fig. 7; pl. 6, figs. 1–5; Zavodowsky and Stepanov, 1970,
p. 89, pl. 3, figs. 3, 4; Ifanova, 1972, p. 103, pl. 3, figs.
14‒16; Lee and Gu, 1976, p. 252, pl. 155, figs. 3, 4; pl.
170, fig. 3; Kalashnikov, 1986, pl. 118, figs. 2, 3;
Kalashnikov, 1993, p. 70, pl. 36, figs. 3–5; Tazawa andAraki, 2016, p. 157, fig. 4.1–4.6; Tazawa, 2016b, p. 22,
fig. 7.6–7.11; Tazawa, 2017, p. 49, fig. 10.1–10.4.
Waagenoconcha imperfecta Prendergast. Tazawa, 1974b, p.
127, pl. 1, figs. 4‒6; pl. 2, figs. 2–7; pl. 3, figs. 1–3; pl. 4,
figs. 1–4, 7 (excluding pl. 2, fig. 6; pl. 3, fig. 2); Tazawa,
1976, pl. 2, fig. 6; Minato et al., 1979, pl. 65, figs. 1, 2;
Manankov, 1991, p. 112, pl. 23, figs. 4–7; Tazawa. 2002,
figs. 10.12; Tazawa, 2007, fig. 4.12.
Waagenoconcha sp. Tazawa and Ibaraki, 2001, p. 9, pl. 1,
fig. 4.

61

Waagenoconcha cf. imperfecta Prendergast. Tazawa, 2001, p.
293, fig. 7.24.
Material.—Fifty-six specimens from localities KF9,
KF11, KF39, KS9 and TY1: (1) external and internal mouldsof two conjoined shells, UHR19813, 19841; (2) internalmoulds of three conjoined shells, with external moulds of thedorsal valves, UHR19816, 19817, 19827; (3) internal mouldsof eleven conjoined shells, KCG24–29, UHR19834, 19839;(4) external and internal moulds of six ventral valves,
UHR19812, 19818, 19822, 19828, 19838, 19840; (5) externalmould of a ventral valve, UHR19823; (6) internal moulds of
tw e nty-thre e ve ntra l va lve s , K C G 30–44 , NU- B1659,
UHR19815, 19825, 19830, 19832, 19835; (7) external andinternal moulds of eight dorsal valves, UHR19814, 19821,
19824, 19826, 19829, 19831, 19836, 19837; and (8) externalmoulds of seven dorsal valves, KCG45–49, NU-B336,
UHR19820.
Remarks.—Most of the specimens were described by
Tazawa and Araki (2016, p. 157, fig. 4.1–4.6) and Tazawa(2016b, p. 22, fig. 7.6 7.11; 2017, p. 49, fig. 10.1–10.4.) as
Waagenoconcha irginae (Stuckenberg, 1898). These
specimens can be referred to Waagenoconcha irginae(Stuckenberg, 1898), redescribed and refigured byTschernyschew (1902, p. 273, 618, pl. 30, figs. 3, 4; pl. 52,
figs. 1–4) from the lower Permian (Cora–Schwagerinahorizons) of Ufa, southern Urals, central Russia, in being
medium size (length 47 mm, width 40 mm in the largest
specimen, UHR19813; length 34 mm, width 34 mm inaverage-sized specimen, UHR19835), equidimensional toslightly longer subrectangular in outline and external
ornament of both ventral and dorsal valves, in particular, finequincunxially arranged spine bases becoming finer anteriorly(numbering 8 spine bases in 5 mm width at midlength, 8–9
spine bases in 5 mm width near anterior margin).
Waagenoconcha imperfecta Prendergast, 1935, redescribedby Archbold (1993, p. 20, figs. 11–13) from the upper
Permian (Wuchiapingian) of the Canning and Bonaparte Gulfbasins of western Australia, differs from the present species
in its much larger size and in having finer spine bases onventral valve. The preceding species, Waagenoconcha
humboldti (d’Orbigny, 1842), is readily distinguished fromW. irginae by coarser spine bases on ventral valve.
Occurrence.—NK2 Unit in Setamai (locality KS9); KY1
Unit in Kamiyasse (localities KF11 and KF39) and Imo(locality TY1); and KY2 Unit in Kamiyasse (locality KF9).
Distribution. — Asselian–Capitanian: northeastern Japan(Setamai and Kamiyasse–Imo in the South Kitakami Belt),
central Japan (Moribu in the Hida Gaien Belt), Spitsbergen,
northern Russia (Kanin Peninsula, Timan, northern Urals,
Pechora Basin and Kolyma Massif), central Russia (southern
Urals), southern Mongolia, northern China (Inner Mongolia)
and eastern Russia (South Primorye).

62

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 40. A, Chonosteges sp., dorsal (A1), anterior (A2) and lateral (A3) views of external mould, and internal mould (A4) of dorsal valve, NU-B2519;
B, C, Anidanthus mizukoshiensis Tazawa; B, external latex cast (B1) and internal mould (B2) of ventral valve, NU-B2543; C, external mould (C1), externallatex cast (C2) and internal mould (C3) of dorsal valve, NU-B2542. Scale bars are 1 cm.

Superfamily AULOSTEGOIDEA Muir-Wood and Cooper,
1960
Family ECHINOSTEGIDAE Muir-Wood and Cooper, 1960
Subfamily ECHINOSTEGINAE Muir-Wood and Cooper,
1960
Genus EDRIOSTEGES Muir-Wood and Cooper, 1960
Type species.—Edriosteges multispinosus Muir-Wood andCooper, 1960.
Edriosteges multispinosus Muir-Wood and Cooper, 1960
(Fig. 37A–D)
Aulosteges medlicottianus (Waagen). King, 1931, p. 93, pl.
26, figs. 7–11; pl. 27, figs. 8, 9.
Edriosteges multispinosus Muir-Wood and Cooper, 1960, p.
104, pl. 17, figs. 1–10; Cooper and Grant, 1975, p. 840, pl.
218, figs. 1–11; pl. 219, figs. 1–27; pl. 220, figs. 1–32; pl.

221, figs. 1–23; pl. 222, figs. 6–13; pl. 223, figs. 17, 18;Sun, 1991, p. 226, pl. 5, fig. 17.
Edriosteges poyangensis (Kayser). Minato et al., 1979, pl.
61, figs. 7–10.
Material.—Eight specimens from localities KF89 andKF218: (1) external and internal moulds of a conjoined shell,
UHR12631; (2) external mould of a conjoined shell,
UHR12632; (3) external and internal moulds of two ventral
valves, NU-B1276, UHR12634; (4) external mould of aventral valve, UHR12665; (5) internal mould of a ventral
valve, UHR12635; (6) external and internal moulds of adorsal valve, NU-B2569; and (7) external mould of a dorsal
valve, UHR12633, 12635.
Description.—Shell medium in size for genus, slightly
longer subquadrate in outline, with greatest width at hinge;
length 37 mm, width 34 mm in the largest specimen,
UHR12634. Ventral valve strongly and unevenly convex inGUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

63

FIGURE 41. A–I, Linoproductus hayasakai Tazawa; A, external latex cast (A1) and internal mould (A2) of ventral valve, IGPS96239; B, external latex
cast of ventral valve, IGPS96241; C, external latex cast of dorsal valve, IGPS96249; D, external latex cast (D 1, D2) of ventral valve, IGPS96240; E,
internal mould (E1) and external latex cast (E2) of ventral valve, NU-B337; F, ventral (F1), anterior (F2), posterior (F3) and lateral (F4) views of internalmould of ventral vale, NU-B344; G, exterior of ventral valve, NUB2482; H, external mould of dorsal valve, NU-B2483; I, internal mould of dorsal valve,
NU-B341. Scale bars are 1 cm.

64

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 42. A, Globiella tschernyschewi (Netschajew), external latex cast (A 1 , A 2 ) and internal mould (A 3 ) of ventral valve, NU-B6197; B–F,
Lamiproductus kamiyassensis (Tazawa); B, external latex cast (B1, B2) and external mould (B3) of ventral valve, NU-B1978 (holotype); C, external latex
cast of ventral valve, NU-B1983; D, external mould of ventral valve, NU-B1979; E, external latex cast (E 1) and external mould (E2) of dorsal valve,
NU-B1990; F, external mould of dorsal valve, NU-B1991; G, H, Costatumulus cancriniformis (Tschernyschew); G, ventral (G1, G2), anterior (G3) andlateral (G4) views of internal mould of ventral valve, IGPS96217; H, external latex cast of ventral valve, IGPS96218. Scale bars are 1 cm.

lateral profile, most convex at posterior portion, flattened
medially; ears large, triangular; sulcus shallow on anterior
two-thirds. Dorsal valve slightly concave, geniculated withshort trail. External surface of ventral valve ornamented withnumerous spines, dense on ears and sporadically on venter;dorsal valve ornamented with irregular fine concentric rugae.
Interior of ventral valve not well preserved. Interior of dorsal
valve with prominent trilobate cardinal process on shaft; thin,long median breviseptum extending to two-thirds valve
length; strong, short lateral ridges; a pair of large dendritic
adductor scars on posterior of valve. .
Remarks.—These specimens are referred to Edriosteges
multispinosus MuirWood and Cooper (1960, p. 104, pl. 17,
figs. 1–10), from the upper Leonardian of the Glass
Mountains, Texas, by medium size and slightly longer
subquadrate outline. Minato et al. (1979, pl. 61, figs. 7–10)

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

figured some of the present specimens (UHR12631–12633)
as Edriosteges poyangensis (Kayser). But Edriosteges
poyangensis (Kayser, 1883, p. 190, pl. 28. figs. 8–10), fromthe upper Permian of Loping, Jiangxi, eastern China, differsfrom E. multispinosus in its smaller size, transverse outline,
and in having numerous fine concentric lines on both ventral
and dorsal valves.
Occurrence. — KY1 Unit in Kamiyasse (locality KF89)
and Imo (locality KF218).
Distribution.—Artinskian–Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt), USA (Texas)
and northwestern China (Tibet).
Genus LIMBELLA Stehli, 1954
Type species.—Aulosteges wolfcampensis King, 1931.
Limbella sp.
(Fig. 38A)
Material.—One specimen from locality KF37, external
and internal moulds of a dorsal valve, NU-B2572.
Remarks.—The single specimen from Imo can be
assigned to the genus Limbella Stehli, 1954 by medium size(length 20 mm, width 23 mm), subquadrate outline,
moderately concave dorsal valve with flattened ears, external
ornament consisting of numerous capillae, fine costellae anddimples scattered over valve, and in having bilobed cardinal
process and short median septum in dorsal valve. The Imo
species most resembles Limbella wolfcampensis (King,
1931), redescribed and refigured by Cooper and Grant (1975,
p. 834, pl. 213, figs. 1–29; pl. 214, figs. 1–21; pl. 215, figs.
1–3; pl. 216, figs. 11–13, 15) from the lower Wolfcamian ofTexas, in size, shape and external ornament of dorsal valve,but, exact comparison is difficult for the poor material.
Occurrence.—KY1 Unit in Kamiyasse (locality KF37).
Subfamily CHONOSTEGINAE Muir-Wood and Cooper,
1960
Genus CHONOSTEGES Muir-Wood and Cooper, 1960
Type species.—Aulosteges magnicostatus Girty, 1909.
Chonosteges sp.
(Fig. 40A)
Material.—One specimen from locality KF62, external
and internal moulds of a dorsal valve, NU-B2519.
Remarks.—The single specimen from Kamiyasse can be
assigned to the genus Chonosteges Muir-Wood and Cooper,
1960 by its small size (length 14 mm, width 14 mm), flat andsmooth visceral disc which is strongly geniculated with ashort trail, and numerous strong costae on dorsal trail. But
the specific identification is difficult for the poorly preserved
specimen.

65

Occurrence.—KY1 Unit in Kamiyase (locality KF62).
Genus URUSHTENOIDEA Jin and Hu, 1978
Type species.—Urushtenia chaoi Jin, 1963.
Urushtenoidea crenulata (Ding in Yang et al., 1962)(Fig. 39A–D)
Eomarginifera crenulata Ding in Yang et al., 1962, p. 85, pl.
37, figs. 6–8.
Urushtenia crenulata (Ding). Jin, 1963, p. 20, 29, pl. 1, figs.
17–24; pl. 2, figs. 9, 10, 18–20, text-fig. 5; Jin et al., 1974,
p. 309, pl. 162, figs. 1–3; Yang et al., 1977, p. 335, pl.
136, fig. 11; Tong, 1978, p. 218, pl. 78, fig. 17; Yang andGao, 1996, pl. 34, figs. 7, 8.
Urushtenoidea crenulata (Ding). Nakamura, 1979, p. 228, pl.
1, figs. 5–9; pl. 3, figs. 1, 2; Yang, 1984, p. 213, pl. 31,
fig. 19; Jin, 1985, pl. 6, fig. 41; Tazawa, 2000, fig. 3.10,
3.11; Tazawa, 2001, p. 296, fig. 7.1–7.9; Shen et al.,2003c, p. 1131, fig. 4.11–4.13; Tazawa, 2008b, p. 50, fig.
7.15, 7.16; Shen and Shi, 2009, p. 155, fig. 3B–I; Tazawa,
2016b, p. 24, fig. 8.1–8.4; Tazawa and Araki, 2017, p. 272,
fig. 6.8, 6.9.
Urushtenoidea maceus (Jin). Nakamura, 1979, p. 227, pl. 1,
figs. 1–4; pl. 2, figs. 1–3; Minato et al., 1979, pl. 65, figs.
8–11; Tazawa, 2002, fig. 10.8.
Uncisteges crenulata (Ding). Liu et al., 1982, p. 178, pl. 129,
fig. 1; Zhu, 1990, p. 74, pl. 14, figs. 4–14; pl. 17, fig. 12.
Material. —Twenty specimens from localities AR4,
KF39, KF90, KF217, KF218, KS9 and KZ9: (1) externalmould of a conjoined shell, UHR30386; (2) internal mouldsof two conjoined shells, with external moulds of the ventral
valves, NU-B2476, 2477; (3) internal moulds of two
conjoined shells, with external moulds of the dorsal valves,
NU-B2142, 2143; (4) internal moulds of six conjoined shells,
NU-B2144–2148, 2478; (5) internal moulds of four ventral
valves, NU-B2149, 2479, UHR30387, 30388; (6) external
and internal moulds of a dorsal valve, NU-B2480; (7)external moulds of three dorsal valves, NU-B2150, 2151,
2481; and (8) internal mould of a dorsal valve, UHR17068.
Remarks.—These specimens were previously describedby Tazawa (2016b, p. 24, fig. 8.1–8.4) and Tazawa and Araki(2017, p. 272, fig. 6.8, 6.9) as Urushtenoidea crenulata (Dingin Yang et al., 1962). The specimens from Kamiyasse–Imo
and Matsukawa are referred to Urushtenoidea crenulata(Ding in Yang et al., 1962), from the Maokouan of Qinghai,
northwestern China, in size (length about 17 mm, widthabout 25 mm in the largest specimen, NU-B2148),transversely subquadrate outline, and external ornament ofboth ventral and dorsal valves, especially in having relatively
fine costae (6–7 in 5 mm) on ventral trail. Urushtenoidea
m a c e u s J i n ( 1 9 6 3 , p . 1 9 , p l . 2 , f i g s . 1 –6 ) , f r o m t h e
Chihsian‒Maokouan of eastern China (Jiangsu, Anhui and66

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 43. A–E, Asperlinus japonicus (Tazawa); A, ventral internal mould (A 1), dorsal internal mould (A2) and dorsal external mould (A3, A4) ofconjoined shell, NU-B702 (holotype); B, ventral internal mould (B1), dorsal external mould (B2) and dorsal internal mould (B3) of conjoined shell,
NU-B704; C, ventral internal mould (C1) and dorsal internal mould (C2) of conjoined shell, NU-B705; D, internal mould (D1) and external mould (D2, D3)
of ventral valve, NU-B703; E, external latex cast (E1, E2) and internal mould (E3) of ventral valve, NU-B2526. Scale bars are 1 cm.

Zhejiang) and central-southern China (Hubei), differs fromU. crenulata in having finer costae (8‒9 in 5 mm) on ventral
valve. Urushtenoidea chaoi Jin (1963, p. 15, 28, pl. 1, figs.
1 –4 , 9 –1 2 ; p l . 2 , f i g s . 7 , 8 , 1 3 –1 7 ) , f r o m t h e u p p e r
Chihsian‒lower Maokouan of Jiangxi and Anhui, eastern
China, is readily distinguished from the present species inhaving coarser costae (3–4 in 5 mm) on ventral valve.
Occurrence. — KN2 Unit in Setamai (locality KS9); andKY1 Unit in Kamiyasse (localities KF39 and KF90), Imo(localities KF217 and KF218) and Matsukawa (locality
AR4).
Distribution. — Kungurian‒Capitanian: northeasternJapan (Kamiyasse‒Imo and Matukawa in the South KitakamiBelt), central Japan (Moribu in the Hida Gaien Belt),southwestern Japan (Mizukoshi in Kyushu Island),northwestern China (Qinghai and Gansu), eastern China(Jiangsu and Fujian), central-southern China (Hubei, Hunan,
Guangdong and Guangxi), southwestern China (Sichuan),Laos, Cambodia and northwestern China (Tibet).
Family SCACCHINELLIDAE Licharew, 1928
Subfamily SCACCHINELLINAE Licharew, 1928
Genus SCACCHINELLA Gemmellaro, 1891
Type species.—Scacchinella variabilis Gemmelaro, 1891.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

67

FIGURE 44. A, B, Permundaria asiatica Nakamura, Kato and Choi; A, external latex cast of ventral valve, KCG54; B, external latex cast of ventral valve,
UHR19015 (holotype); C, D, Permundaria tenuistriata Tazawa; C, external mould (C1) and external latex cast (C2) of dorsal valve, KCG14; D, internalmould of ventral valve, UHR19700 (holotype); E, Yakovlevia kaluzinensis Fredericks, internal mould of ventral valve, KCG8; F, Yakovlevia mammata(Keyserling), internal latex cast (F1, F2) and internal mould (F3) of ventral valve, KCG15. Scale bars are 1 cm.

Scacchinella gigantea Schellwien, 1900b(Fig. 36E, F)
Scacchinella gigantea Schellwien, 1900b, p. 35, pl. 4, figs.
1–3; pl. 5, figs. 1–8, text-figs. 5, 6, 8; Heritsch, 1938, p.
101, pl. 5, figs. 1, 2, 9: Licharew, 1939, p. 96, pl. 23, fig.
2; Ramovs, 1965, p. 357, pl. 13, figs. 3–6; Tazawa andAraki, 1999, p. 453, fig. 2.1–2.4; Tazawa and Araki, 2017,p. 272, fig. 7.8, 7.9.
Material.—Five specimens from locality KZ9: (1)external mould of a ventral valve, NU-B197; and (2) internalmoulds of four ventral valves, NU-B198‒201.
Remarks.—These specimens were previously describedby Tazawa and Araki (1999, 2017) as Scacchinella gigantea
Schellwien, 1900b. The Matsukawa specimens are strongly

68

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 45. A–D, Grandaurispina kozlowskiana (Fredericks); A, external mould (A1, A2) and internal mould (A3) of dorsal valve, NU-B2001; B,
external latex cast (B1) and external mould (B2) of ventral valve, NU-B2006; C, external mould of dorsal valve, NU-B2002; D, external latex cast (D1, D2)
and external mould (D3) of ventral valve, NU-B1997. Scale bars are 1 cm.

deformed and imperfect, but they can be referred to
Scacchinella gigantea Schellwien (1900b), from the
Trogkofel Formation of the Carnian Alps, by medium tolarge, transversely subelliptical ventral valve with a broadand very shallow depression on the middle of the anteriorside of the valve. Scacchinella exasperata Cooper and Grant(1975, p. 921, pl. 271, figs. 14–24; pl. 273, figs. 26–28), fromthe lower Wolfcampian of Texas in the USA, differs from S.
gigantea in its more rounded and cyrindrical outline and inhaving a broader ventral interarea. Scacchinella titan Cooper
and Grant (1975, p. 923, pl. 270, figs. 12–16; pl. 272, figs.
1–6; pl. 273, figs. 1–25; pl. 274, figs. 1–6; pl. 275, figs. 1–4;

pl. 276, figs. 1–3; pl. 277, figs. 1–4; pl. 278, figs. 1–19; pl.
279, figs. 1–9; pl. 280, figs. 1–8; pl. 281, figs. 1–18; pl. 282,
figs. 1–19; pl. 283, figs. 1–22; pl. 284, figs. 16–30), from theupper Wolfcampian of Texas, differs from the present species
in its larger dimensions, more rounded, nearly circularanterior profile and much broader ventral interarea.
Occurrence.—KY1 Unit in Matsukawa (locality KZ9).
Distribution.— Asselian‒Capitanian: northeastern Japan(Matsukawa in the South Kitakami Belt), Slovenia (CarnianAlps), central Russia (southern Urals) and Uzbekistan(Fergana).

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Superfamily LINOPRODUCTOIDEA Stehli, 1954
Family LINOPRODUCTIDAE Stehli, 1954
Subfamily LINOPRODUCTINAE Stehli, 1954
Genus LINOPRODUCTUS Chao, 1927
Type species.—Productus cora d’Orbigny, 1842.
Linoproductus hayasakai Tazawa, 1979
(Fig. 41A–I)
Productus cora d’Orbigny. Hayasaka, 1925a, p. 94, pl. 5,
figs. 7–9.
Linoproductus cora (d’Orbigny). Hayasaka and Minato,
1956, p. 145, pl. 23, figs. 9, 10; Tazawa, 1976, pl. 2, fig.
11; Minato et al., 1979, pl. 62, figs. 1, 2; Tazawa and
Ibaraki, 2001, p. 10, pl. 1, figs. 11–13; pl. 2, figs. 1–8.
Linoproductus sp. Minato et al., 1979, pl. 62, figs. 3, 4.
Linoproductus hayasakai Tazawa, 1979, p. 26, pl. 4, figs.
5–11; Tazawa and Araki, 2017, p. 274, fig. 7.2–7.4.
Material.— Forty specimens from localities AR5, KS1,
KS3 and KS4: (1) external and internal moulds of eightventral valves, IGPS96239, 96240–96242, NU-B337–340;(2) exterior of a ventral valve, NU-B2482; (3) external mouldof a ventral valve, IGPS96243 (holotype); (4) internal mouldsof twenty-seven ventral valves, IGPS96244–96248,
NU-B341–359; and (5) external mould of six dorsal valves,
IGPS17042, 96249, NU-B360–362, 2483.
Remarks.—Part of the specimens were previously
described by Tazawa (1979, p. 26, pl. 4, figs. 5–11) as
Linoproductus hayasakai Tazawa, 1979; and the others were
described by Tazawa and Ibaraki (2001, p. 10, pl. 1, figs.
11–13; pl. 2, figs. 1–8) as Linoproductus cora (d’Orbigny,
1842). All the specimens are, however, referred to
Linoproductus hayasakai Tazawa, 1979, which ischaracterized by its medium size (length 41 mm, width 42
mm in the holotype, IGPS96243), fine costellae (numbering
15–18 in 10 mm at midlength of ventral valve) and no spines
on ventral valve. The type species, Linoproductus cora(d’Orbigny, 1842), redescribed by Tschernyschew (1902, p.
279, 621, pl. 33, figs. 2, 3; pl. 35, fig. 1; pl. 54, figs. 1–5,
text-figs. 69–71), from the lower Permian of Timan, northern
Russia, differs from L. hayasakai in its larger size and inhaving coarser costellae and sporadically distributed spines
on ventral valve. Linoproductus kaseti Grant (1976, p. 154,
pl. 41, figs. 8–28), from the Rat Buri Formation of Phangnga,
southern Thailand, is also a medium-sized Linoproductus
species, but the Thailand species differs from the present
species in its elongate outline and in having coarser costellae
and some spines on ventral valve.
Occurrence.—KN1 Unit in Setamai (localities KS1, KS3
and KS4) and KY1 Unit in Matsukawa (locality AR5).
Distribution. —Wordian: northeastern Japan (Setamai,
Kamiyasse–Imo and Matsukawa in the South Kitakami Belt).

69

Genus GLOBIELLA Muir-Wood and Cooper, 1960
Type species.—Productus hemispharium Kutorga, 1844.
Globiella tschernyschewi (Netschajew, 1911)(Fig. 42A)
Productus tschernyschewi Netschajew, 1911, p. 31, 141, pl.
1, figs. 5, 7; pl. 2, figs. 5–11, 16.
Stepanoviella tschernyschewi (Netschajew). Grigorjewa,
1962, p. 45, pl. 10, figs. 1–3; pl. 13, fig. 5; pl. 14, fig. 3.
Globiella tschernyschewi (Netschajew). Grigorjewa et al. in
Sarytcheva, 1977, p. 164, pl. 27, figs. 10, 11; Tazawa,
2016b, p. 26, fig. 9.1.
Material.—One specimen from locality KF21, external
and internal moulds of a ventral valve, NU-B1977.
Remarks.—This specimen was previously described by
Tazawa (2016b, p. 26, fig. 9.1) as Globiella tschernyschewi(Netschajew, 1911). The specimen from Kamiyasse can bereferred to Globiella tschernyschewi (Netschajew, 1911),from the Kazanian beds of the Pinega River region, northern
Russia, by its small (length 17 mm, width 22 mm), slightly
transverse and hemisphaerical ventral valve ornamented withnumerous capillae (numbering 16–17 in 5 mm at about
midlength) and strong, irregular concentric rugae. The typespecies, Globiella hemisphaerium (Kutorga, 1844),
redescribed by Grigorjewa (1962, p. 42, pl. 9, figs. 1–9; pl.
13, figs. 6–9; text-figs. 3, 5, 14, 15) from the Kazanian bedsof the Pinega River and the Kama River regions, northern
Russia, is distinguished from G. tschernyschewi by its longer
ventral valve.
Occurrence.—KY1 Unit in Kamiyasse (locality KF21).
Distribution.— Wordian: northeastern Japan(Kamiyasse‒Imo in the South Kitakami Belt) and northern
Russia (Pinega River).
Subfamily LAMIPRODUCTINAE Liang, 1990
Genus ASPERLINUS Waterhouse and Piyasin, 1970
Type species.—Productus asperulus Waagen, 1884.
Asperlinus japonicus (Tazawa, 2008c)(Fig. 43A–E)
Terrakea japonica Tazawa, 2008c, p. 336, fig. 3A–J.
Material.—Thirty-three specimens from localities KF17,
KF28, KF79, KF86, KF90, KF217 and KF218: (1) external
and internal moulds of two conjoined shells, NU-B2520,
2521; (2) internal moulds of six conjoined shells, withexternal moulds of the dorsal valves, NU-B702 (holotype),704, 2522–2525; (3) internal moulds of three conjoined
shells, NU-B705, 706, 1786; (4) external and internal mouldsof three ventral valves, NU-B703, 2526, 2527; (5) external

70

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 46. A–D, Neorichthofenia mabutii (Tazawa and Araki); A, dorsal (A1, A2), ventral (A3), anterior (A4), posterior (A5) and lateral (A6) views ofinternal mould of conjoined shell, IGPS98874; B, dorsal (B 1), ventral (B2), anterior (B3), posterior (B4) and lateral (B5) views of internal mould ofconjoined shell, IGPS98870 (holotype); C, external mould of dorsal valve, IGPS98877; D, dorsal (D 1, D2) and ventral (D3) views of internal mould ofconjoined shell, IGPS98872. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

71

FIGURE 47. A–K, Leptodus nobilis (Waagen); A, internal mould of ventral valve, UHR12110; B, internal mould of ventral valve, UHR12114; C, internalmould of ventral valve, NU-B1691; D, internal mould of ventral valve, NU-B1701; E, internal mould of ventral valve, NU-B1697; F, internal mould ofventral valve, UHR12111; G, internal latex cast (G1) and internal mould (G2) of ventral valve, UHR12117; H, internal mould of ventral valve, NU-B1722;
I, internal mould of ventral valve, UHR12109; J, internal mould (J1) and internal latex cast (J2) of ventral valve, NU-B1681; K, internal mould of ventral
valve, NU-B1693.Scale bars are 1 cm.

72

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 48. Longitudinal section of internal latex cast of ventral valve, showing lateral septa, a: anterior, p: posterior. A, Leptodus nobilis (Waagen),
NU-B1681; B, Petasmaia expansa Cooper and Grant, KCG55.

moulds of two ventral valves, NU-B2528, 2529; and (6)internal moulds of seventeen ventral valves, NU-B707–710,
1785, 2530–2541.
Remarks.—Of the above-listed specimens, nine from Imo
were previously described by Tazawa (2008c, p. 336, fig.
3A–J) as Terrakea japonica Tazawa, 2008c. This species is,however, assigned to the genus Asperlinus Waterhouse andPiyasin, 1970 in having somewhat irregular costellae (7–8 in
5 mm at about midvalve) and numerous spine bases onventral valve. The type species, Asperlinus asperulus(Waagen, 1884), redescribed by Waterhouse and Piyasin(1970, p. 132, pl. 23, figs. 2–5) from the Wordian beds of
Khao Phrik, southern Thailand, differs from the Kitakami
species in its much smaller in size. The Kitakami species
most resembles the shells, figured by Licharew and Kotlyar(1978, pl. 14, figs. 11–12) as Asperlinus asperulus (Waagen)from the upper Barabashevka Formation (Capitanian) of
Primorye, eastern Russia, but accurate comparison is difficult
owing to lack of description on the Russian specimens.
Occurrence.—KY1 Unit in Kamiyasse (localities KF17,
KF28, KF79, KF86 and KF90) and Imo (localities KF217 andKF218).
Distribution.—Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt).

Genus LAMIPRODUCTUS Liang, 1990
Type species.—Lamiproductus typica Liang, 1990.
Lamiproductus kamiyassensis (Tazawa, 2016a)(Fig. 42B–F)
Globiella kamiyassensis Tazawa, 2016a, p. 84, fig. 5.
Material.—Nineteen specimens from locality KF99: (1)external moulds of six ventral valves, NU-B1978 (holotype),
1979–1983; (2) internal moulds of six ventral valves,
NU-B1984–1989; and (3) external moulds of seven dorsal
valves, NU-B1990–1996.
Remarks.—These specimens were previously describedby Tazawa (2016a, p. 84, fig. 5) as Globiella kamiyassensis
Tazawa, 2016a. However, the genus of this species is not
Globiella but Lamiproductus Liang, 1990 in having strong
costae on both ventral and dorsal valves. This species ischaracterized by its medium size (length 25 mm, width 21
mm in the holotype, NU-B1978), narrow, tapering umbo andexternal ornament of ventral valve consisting of numerous
strong costae (numbering 10–11 in 5 mm at midlength ofventral valve). The type species, Lamiproductus typica Liang(1990, p. 205, pl. 35, figs. 1–12, 14–18, text-fig. 26), fromthe Lengwu Formation (Capitanian) of Zhejiang, easternGUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

China, is distinguished from the Kitakami species in having
irregular costae on both valves.
Occurrence.—KY1 Unit in Kamiyasse (locality KF99).
Distribution.— Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt).
Subfamily ANIDANTHINAE Waterhouse, 1968b
Genus ANIDANTHUS Hill, 1950
Type species.—Linoproductus springsurensis Booker, 1932.
Anidanthus mizukoshiensis Tazawa, 2008b(Fig. 40B, C)
Anidanthus mizukoshiensis Tazawa, 2008b, fig. 7.6–7.8.
Material.—Three specimens from localities KF3 andKF39: (1) external and internal moulds of two ventral valves,
NU-B2543, 2544; and (2) external and internal moulds of adorsal valve, NU-B2542.
Remarks.—The specimens from Kamiyasse can bereferred to Anidanthus mizukoshiensis Tazawa (2008b, p. 47,
fig. 7.6–7.8), from the upper part of the Mizukoshi Formation(correlated with the Capitanian, by Ishida et al., 2010) of
Mizukoshi, central Kyushu, southwestern Japan, in being
small and transverse outline (length 12 mm, width 25 mm in
the largest specimen, NU-B2543) and in having external
ornament consisting of numerous strong lamellae and finecostellae on both ventral and dorsal valves. This species like
Anidanthus minor Cooper (1957, p. 38, pl. 6A, figs. 1–17),
from the Coyote Butte Formation of central Oregon, USA, in
its small size, but the American species is less transverse inoutline. The type species Anidanthus springsurensis (Booker,
1932), redescribed by Briggs (1998, p. 204, fig. 97A–K) fromthe Kungurian bed of the Sydney–Bowen Basin, eastern
Australia, differs from A. mizukoshiensis in its larger size andless transverse outline.
Occurrence.—KY1 Unit in Kamiyasse (localities KF3 andKF39).
Distribution.—Wordian–Capitanian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt) andsouthwestern Japan (Mizukoshi in Kyushu Island).
Family KANSUELLIDAE Muir-Wood and Cooper, 1960
Subfamily AURICULISPININAE Waterhouse,1986
Genus COSTATUMULUS Waterhouse in Waterhouse andBriggs, 1986
Type species.—Auriculispina tumida Waterhouse in
Waterhouse, Briggs and Parfrey, 1983.
Costatumulus cancriniformis (Tschernyschew, 1889)(Fig. 42G, H)

73

Productus cancriniformis Tschernyschew, 1889, p. 283, 373,
pl. 7, figs. 32, 33; Tschernyschew, 1902, p. 292, 629, pl.
52, figs. 5, 6; Fredericks, 1925, p. 27, pl. 4, figs. 115, 116.
Cancrinella cancriniformis (Tschernyschew). Kaschirzew,
1959, p. 39, pl. 15, figs. 4, 5; Solomina, 1960, p. 49, pl. 8,
figs. 3–7; Ustritsky and Tschernjak, 1963, p. 84, pl. 13,
figs. 6–8; pl. 14, figs. 1–5; Abramov, 1970, p. 124, pl. 5,
figs. 9–12; Solomina, 1970, p. 85, pl. 5, fig. 9;
Zavodowsky and Stepanov, 1970, p. 101, pl. 24, fig. 8; pl.
27, fig. 10; pl. 35, figs. 4–7; Grigorjeva et al. in
Sarytcheva, 1977, p. 134, pl. 20, fig. 1; Alexandrov andEinor, 1979, pl. 24, fig. 5; Tazawa, 1979, p. 27, pl. 4, figs.
3, 4; Lee and Duan, 1985, p. 239, pl. 75, figs. 2–5; Pavlova
and Lazarev in Tatarinov et al., 1991, p. 114, pl. 26, figs.
5, 6, 8, 14; Kalashnikov, 1993, p. 81, pl. 33, fig. 10.
Costatumulus cancriniformis (Tschernyschew). Tazawa andAraki, 2017, p. 274, figs. 6.10, 7.1.
Material.—Two specimens from locality AR5, external
and internal moulds of two ventral valves, IGPS96217,
96218.
Remarks.—These specimens were first described by
Tazawa (1979, p. 27, pl. 4, figs. 3, 4) as Cancrinella
cancriniformis (Tschernyschew, 1889); and subsequently
redescribed by Tazawa and Araki (2017, p. 274, figs. 6.20,
7.1) as Costatumulus cancriniformis (Tschernyschew, 1889).
The specimens from Matsukawa can be referred to
Costatumulus cancriniformis (Tschernyschew, 1889), fromthe lower Permian (Artinskian) of the northern Urals, by its
strongly inflated ventral valve which is ornamented withnumerous undulate concentric rugae. Costatumulus tazawai
Shen, Archbold, Shi and Chen (2000, p. 743, figs. 12.1–12.8,
11–14), from the Selong Group (Wuchiapingian) of Xizang(Tibet), differs from C. cancriniformis in having more
numerous, finer costellae on ventral valve. The type species,
Costatumulus tumida (Waterhouse in Waterhouse et al.,1983, p. 133, pl. 3, figs. 2–4, 6, 7), from the Tiverton
Formation of the Bowen Basin in Queensland, eastern
Australia, is readily distinguished from the present species inhaving less strong concentric rugae on ventral valve.
Occurrence.—KY1 Unit in Matsukawa (locality AR5).
Distribution.—Moscovian‒Wordian; northeastern Japan(Matsukawa in the South Kitakami Belt), northern Russia(Timan, northern Urals, Pechora Basin, Taimyr Peninsula,
Verkhoyansk Range and Kolyma Massif), central Russia(southern Urals), southern Mongolia, northern China(Shanxi) and eastern Russia (South Primorye).
Genus PERMUNDARIA Nakamura, Kato and Choi, 1970
Type species.—Permundaria asiatica Nakamura, Katoand Choi, 1970.
Permundaria asiatica Nakamura, Kato and Choi, 1970
(Fig. 44A, B)

74

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 49. A, Keyserlingina sp., internal latex cast (A1, A2) and internal mould (A3) of ventral valve, KCG19; B, Petasmaia expansa Cooper and Grant,internal mould (B1) and internal latex cast (B2, B3) of ventral valve, KCG55; C, Pararigbyella doulingensis Shen and Zhang, internal mould (C1, C2) ofventral valve, NU-B1756. Scale bars are 1 cm.

Striatifera? sp. Hayasaka and Minato, 1956, p. 144, pl. 23,
figs. 6, 7.
Permundaria asiatica Nakamura, Kato and Choi, 1970, p.
296, pl. 2, figs. 1, 2; Tazawa, 1976, pl. 2, fig. 7; Minato etal., 1979, pl. 62, figs. 12, 13; Tazawa, 2000, fig. 3.4;
Tazawa, 2001, p. 296, fig. 7.17‒7.19; Tazawa and Araki,
2018, p. 15, fig. 4.5.
Material. ― Four specimens from localities AR4, KF80,

KF99 and KS10: (1) external and internal mould of a ventral
valve, UHR19015 (holotype); (2) external mould of a ventral
valve, KCG54; (3) internal moulds of two ventral valves,
UHR 19698, 19699.
Remarks. ― These specimens were previously describedby Nakamura et al. (1970, p. 296, pl. 2, figs. 1, 2), Tazawa(1974a, p. 315, pl. 43, figs. 3, 4) and Tazawa and Araki(2018, p. 15, fig. 4.5) as Permundaria asiatica Nakamura,
Kato and Choi, 1970 on the basis of its large, subquadrate
and flattened ventral valve (length 47 mm, width about 37

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

75

FIGURE 50. A–F, Poikilosakos kamiyassensis Tazawa and Takaizumi; A, internal mould (A1, A2) of ventral valve, IGPS99009; B, internal mould ofventral valve, IGPS99012; C, internal mould of ventral valve, IGPS99015; D, internal mould of ventral valve, IGPS99016; E, internal mould of ventral
valve, IGPS99011; F, internal mould of ventral valve, IGPS99010 (holotype). Scale bars are 1 cm.

mm in the holotype; length more than 65 mm, width morethan 64 mm in a ventral valve specimen, KCG54),
ornamented with numerous fine costellae (numbering 5 in 2mm at about midlength) and numerous regular but slightly
undulate concentric rugae. As noted by Nakamura et al.
(1970), Striatifera? sp. Hayasaka and Minato, 1956, from thelower part of the Kanokura Series (= Kamiyasse Formation)of Imo, South Kitakami Belt, is a synonym of the present
species. Permundaria tenuistriata Tazawa (1974a, p. 317, pl.
43, figs. 1, 2), from the lower Kamiyasse Formation of
Kamiyasse, South Kitakami Belt, is distinguished from P.
asiatica in having finer capillae on both ventral and dorsal
valves.
Occurrence. ― KN2 Unit in Setamai (locality KS10); KY1
Unit in Kamiyasse (localities KF80 and KF99) andMatsukawa (AR4).
Distribution. ― Wordian—Capitanian: northeastern Japan(Setamai, Kamiyase–Imo and Matsukawa in the South KitakamiBelt) and central Japan (Moribu in the Hida Gaien Belt).
Permundaria tenuistriata Tazawa, 1974a(Fig. 44C, D)

Permundaria tenuistriata Tazawa, 1974a, p. 317, pl. 43, figs.
1, 2; Tazawa and Araki, 2017, p. 275, fig. 6.11.
Material.—Three specimens from localities AR4 andKF99: (1) internal mould of a ventral valve, UHR19700(holotype); and (2) external moulds of two dorsal valves,
KCG14 and UHR19701.
Remarks.—These specimens were described by Tazawa(1974a, p. 317, pl. 43, figs. 1, 2) and Tazawa and Araki(2017, p. 275, fig. 6.11) as Permundaria tenuistriata Tazawa,
1974a, on account of large (length about 30 mm, width 61
mm in the holotype, UHR19700; length 45 mm, width about
50 mm in a dorsal valve specimen, KCG14), semicircular andalmost flat dorsal valve, ornamented by numerous regularconcentric rugae and numerous capillae (11–12 capillae in 2mm at about midlength). The preceding species,
Permundaria asiatica Nakamura, Kato and Choim 1970, isreadily distinguished from Permundaria tenuistriata inhaving coarser capillae (6–8 in 3 mm at 10 mm from umbo)
on ventral valve.
Occurrence.—KY1 Unit in Kamiyasse (locality KF99)

76

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 51. A, Paralyttonia kesennumensis Tazawa and Araki, internal mould (A1, A2) and internal latex cast (A3) of ventral valve, IGPS98393; B–G,
Permianella typica He and Zhu; B, external latex cast (B1, B2) and internal mould (B3) of ventral valve, NU-B58; C, external mould of dorsal valve,
NU-B60; D, internal mould of dorsal valve, NU-B55; E, internal mould (E1, E2) and external latex cast (E3) of ventral valve, NU-B63; F, external mouldof dorsal valve, NU-B57; G, internal mould of ventral valve, NU-B65. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

and Matsukawa (locality AR4).
Distribution.—Wordian: northeastern Japan(Kamiyasse–Imo and Matsukawa in the South KitakamiBelt).
Subfamily PAUCISPINAURIINAE Waterhouse, 1986
Genus GRANDAURISPINA Muir-Wood and Cooper, 1960
Type species.—Grandaurispina kingorum Muir-Wood andCooper, 1960.
Grandaurispina kozlowskiana (Fredericks, 1925)(Fig. 45A–D)
Productus villiersi kozlowskianus Fredericks, 1925, p. 18, pl.
1, figs. 36‒40; pl. 2, figs. 86, 87; Hayasaka, 1925a, p. 96,
pl. 5, figs. 10, 11.
Cancrinella villiersi kozlowskiana (Fredericks). Hayasaka
and Minato, 1956, p. 144, pl. 23, fig. 5.
Cancrinella cancriniformis spinosa Hayasaka and Minato,
1956, p. 144, pl. 23, fig. 4.
Cancrinella spinosa Hayasaka and Minato. Minato et al.,
1979, pl. 62, figs. 5‒8, 11; Tazawa, 1976, pl. 2, fig. 5;
Tazawa, 2002, fig. 10.3.
Cancrinella kozlowskiana (Fredericks). Minato et al., 1979,
pl. 62, figs. 9, 10.
Cancrinella truncata (Chao). Lee et al., 1980, p. 380, pl. 165,
figs. 13, 14, 24; Gu, 1992, p. 237, pl. 68, figs. 18, 20.
Cancrinella sp. Tazawa and Ibaraki, 2001, p. 11, pl. 1, fig. 6.
Cancrinella cf. spinosa Hayasaka and Minato. Tazawa, 2001,
p. 295, fig. 6.17.
Grandaurispina kozlowskiana (Fredericks). Tazawa, 2016b,
p. 27, fig. 9.2‒9.5.
Material.— Eighteen specimens from localities KF21,
KF22, KF46, KF50, KF218 and KS9: (1) external andinternal moulds of nine ventral valves, NU-B1997‒2000,
2003–2005, 2484, 2485; (2) external mould of two ventral
valves, NU-B307, 2006; (3) internal moulds of three ventral
valves, NU-B 2007‒2009; (4) external and internal moulds oftwo dorsal valves, NU-B2001, 2002; and (5) external mouldsof two dorsal valves, NU-B2010, 2011.
Remarks.—Most of the specimens listed above werepreviously described by Tazawa (2016b, p. 27, fig. 9.2‒9.5)
as Grandaurispina kozlowskiana (Fredericks, 1925). The
specimens from the lower Kamiyasse Formation of
Kamiyasse and Imo in the Kamiyasse–Imo area can bereferred to Grandaurispina kozlowskiana (Fredericks, 1925),
originally described by Fredericks (1925, p. 18, pl. 1, figs.
36‒40; pl. 2, figs. 86, 87) from the Chandalaz Formation ofSouth Primorye, eastern Russia, on account of small size(length 23 mm, width 22 mm in the largest specimen,
NU-B2006) and in having numerous, quincunxially arranged,
elongate spine bases on venter and dense strong spine baseson ears and adjacent lateral slopes of ventral valve .

77

Grandaurispina bella Cooper and Grant (1975, p. 1162, pl.
442, figs. 1–38), from the Word Formation of Texas in theUSA, is also a small-sized Grandaurispina with numerous
spine bases on ventral valve, but differs from G.
kozlowskiana in its more rounded outline. The type species,
Grandaurispina kingorum Muir-Wood and Cooper (1960, p.
306, pl. 121, figs. 1–13), from the Word Formation of Texas,
differs from the present species in its larger size and inhaving more distant spine bases on ventral valve.
Occurrence. —KN2 Unit in Setamai (locality KS9); andKY1 Unit in Kamiyasse (localities KF21, KF22 and KF46)
and Imo (locality KF218).
Distribution.—Wordian: northeastern Japan (Setamai,
Kamiyasse–Imo and Iwaizaki in the South Kitakami Belt),
central Japan (Moribu in the Hida Gaien Belt), northern
China (Inner Mongolia), northeastern China (Jilin) andeastern Russia (South Primorye),.
Family YAKOVLEVIIDAE Waterhouse, 1975
Genus YAKOVLEVIA Fredericks, 1925
Type species.—Yakovlevia kaluzinensis Fredericks, 1925.
Yakovlevia mammata (Keyserling, 1846)(Fig. 44F)
Productus mammatus Keyserling, 1846, p. 206, pl. 4, fig. 5;
de Koninck, 1847, p. 49, pl. 7, fig. 4; Tschernyschew,
1902, p. 295, pl. 35, figs. 4–6; Keidel, 1906, p. 367, pl. 12,
fig. 5.
Linoproductus? mammatus (Keyserling). Chao, 1927, p. 146,
pl. 15, figs. 10–14.
Productus (Linoproductus?) mammatus Keyserling. Grabau,
1931, p. 288, pl. 29, figs. 10–14.
Productus (Thomasina) mammatus Keyserling. Stepanov,1937a, p. 127, 177, pl. 2, figs. 5–7.
Muirwoodia mammata (Keyserling). Muir-Wood and Cooper,
1960, pl. 120, figs. 9‒11; Harker, 1960, p. 58, pl. 16, figs.
1–5; Gobbett, 1963, p. 112, pl. 13, figs. 23–28; Lee andGu, 1976, p. 263, pl. 159, figs. 7–9; pl. 163, fig. 2; pl. 164,
figs. 3, 4; pl. 170, figs. 6, 7; Licharew and Kotlyar, 1978,
pl. 14, figs. 3–5; Liu and Waterhouse, 1985, p. 17, pl. 4,
figs. 4–6; Nakamura et al., 1992, pl. 1, fig. 4; Kalashnikov,
1993, p. 63, pl. 19, figs. 1–3.
Yakovlevia mammatus (Keyserling). Kotlyar, 1961, text-figs.
4‒6.
Yakovlevia mammata (Keyserling). Brabb and Grant, 1971, p.
16, pl. 1, figs. 9–12, 33–36; Ifanova, 1972, p. 121, pl. 7,
figs. 4, 5; Kalashnikov, 1986, pl. 121, figs. 5, 6;
Malkowski, 1988, p. 40, pl. 5, fig. 6; Zhang, 1990, pl. 2,
figs. 4, 7, 9; Tazawa, 1999b, fig. 2.1, 2.2; Tazawa, 1999c,p. 90, fig. 3.1–3.5; Wang and Zhang, 2003, p. 85, pl. 6,
figs. 1–8; pl. 7, figs. 1–10; Klets, 2005, pl. 11, figs. 1–7 ;Tazawa and Araki, 2017, p. 275, fig. 6.7.
Yakovlevia paramammata Lee and Gu in Lee et al., 1980, p.

78

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 52. A–E, Dicystoconcha lapparenti Termier and Termier; A, internal mould (A1, A2) of ventral valve, NU-B2493; B, internal mould of ventral
valve, NU-B2494; C, internal mould of ventral valve, NU-B2491; D, internal mould of ventral valve, NU-B2492; E, internal mould of ventral valve,
NU-B1680; F, Laterispina parallela Shen, Fan, Zhang and Zhang, internal mould (F1, F2) of ventral valve, NU-B70. Scale bars are 1 cm.

382, pl. 171, figs. 4, 15.
Muirwoodia sp. Tazawa, 1987, fig. 1.6.
Material.—One specimen from locality AR4, internalmould of a ventral valve, KCG15.
Remarks.— This specimen was previously described by
Tazawa and Araki (2017, p. 275, fig. 6.7) as Yakovlevia
mammata (Keyserling, 1846). The Matsukawa specimen canbe referred to Yakovlevia mammata (Keyserling, 1846), fromthe lower Permian (Sakmarian?) of the Pechora Basin,

northern Russia, in its small size (length 19 mm, width 33mm), and in having large, acute ears and a narrow,moderately deep sulcus on ventral trail. Yakovlevia
greenlandica (Dunbar, 1955, p. 103, pl. 16, figs. 1–17), fromthe Guadalupian of eastern Greenland, is also a small-sized
Yakovlevia species, but the Greenlandic species isdistinguished from Y. mammata by its less transverse outline.
Yakovlevia mammatiformis (Fredericks, 1926, p. 87, pl. 3,
figs. 4–6), from the lower Permian (Artinskian) of thePechora Basin, northern Russia, is clearly distinguished from

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

79

FIGURE 53. A, B, Derbyia grandis Waagen; A, external latex cast (A1), internal mould (A2) and internal latex cast (A3) of ventral valve, NU-B363; B,
external latex cast (B1) and internal mould (B2) of ventral valve, NU-B364. Scale bars are 1 cm.

the present species by its larger size and more transverse
outline.
Occurrence.—KY1 Unit in Matsukawa (locality AR4).
Distribution.—Kasimovian–Capitanian: northeasternJapan (Kamiyasse–Imo and Matsukawa in the South
Kitakami Belt), northern USA (Alaska), northern Canada(Devon Island), Spitsbergen, northern Russia (Timan,
Pechora Basin and Verkhoyansk Range), northwestern China(Xinjiang), northern China (Inner Mongolia), northeastern

China (Heilongjiang) and eastern Russia (South Primorye).
Yakovlevia kaluzinensis Fredericks, 1925
(Fig. 44E)
Chonetes (Yakovlevia) kaluzinensis Fredericks, 1925, p. 7, pl.
2, figs. 64–66.
Yakovlevia kaluzinensis Fredericks. Kotlyar, 1961, text-figs.
1–3; Licharew and Kotlyar, 1978, pl. 14, figs. 1, 2;

80JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 54. A–F, Derbyia nipponica Nakamura; A, external latex cast (A1) and internal mould (A2) of ventral valve, UHR12470: B, external latex cast(B1) and internal mould (B2) of ventral valve, NU-B365; C, internal mould of dorsal valve, NU-B369; D, internal mould of ventral valve, NU-B2485; E,internal mould of dorsal valve, NU-B2486; F, ventral internal mould (F1), dorsal internal mould (F2) and ventral external latex cast (F3) of conjoined shell,
UHR17035. Scale bars are 1 cm.

Manankov, 1998, pl. 8, figs. 18, 19; Tazawa, 1999b, fig
2.4–2.6; Tazawa, 1999c, p. 90, fig. 3.7–3.15; Tazawa,
2000, fig. 3.18; Tazawa, 2001, p. 291, fig. 6.20–6.25;
Tazawa, 2008b, p. 49, fig. 7.14; Tazawa and Araki, 2013,
p. 5, fig. 2.2; Tazawa and Araki, 2017, p. 276, fig. 6.12.
Material.—One specimen from locality AR5, internalmould of a ventral valve, KCG8.

Remarks.— This specimen was previously described by
Tazawa and Araki (2013, p. 5, fig. 2.2) and Tazawa andAraki (2017, p. 276, fig. 6.12) as Yakovlevia kaluzinensis
Fredericks, 1925. The Matsukawa specimen can be referred
to Yakovlevia kaluzinensis Fredericks, 1925, originallydescribed by Fredericks (1925, p. 7, pl. 2, figs. 64–66) fromthe Chandalaz Formation of Vladivostok, South Primorye,eastern Russia, by its large size (length 49 mm, width 58

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWAmm), transversely rectangular outline and in having internal
structures, consisting of a pair of small, elongate subtrigonal
and dendritic adductor scars and two large, flabellate andradially striated diductor scars in ventral valve. Yakovlevia
impressa (Toula, 1875, p. 236, pl. 5, fig. 11), from the middle
Permian of Spitsbergen, differs from Y. kaluzinensis in
having larger and more prominent ears. The preceding
species, Yakovlevia mammata (Keyserling, 1846), is readily
distinguished from the present species by its much smaller
size and larger and more acute ears.
Occurrence.—KY1 Unit in Matsukawa (locality AR5).
Distribution.—Kungurian–Wuchiapingian: northeasternJapan (Matsukawa in the South Kitakami Belt), central Japan(Moribu in the Hida Gaien Belt), southwestern Japan(Mizukoshi in Kyushu Island), southern Mongolia andeastern Russia (South Primorye).
Superfamily RICHTHOFENIOIDEA Waagen, 1885
Family HERCOSIIDAE Cooper and Grant, 1975
Genus NEORICHTHOFENIA Shen, He and Zhu, 1992
Type species.—Richthofenia mabutii Tazawa and Araki,
1984b.
Neorichthofenia mabutii (Tazawa and Araki, 1984b)(Fig. 46A–D)
Richthofenia mabutii Tazawa and Araki, 1984b, p. 3, pl. 1,
figs. 1–7.
Neorichthofenia mabutii (Tazawa and Araki). Shen et al.,
1992, p. 180, pl. 3, figs. 13–22.
Material.—Eight specimens from locality AR4: (1)internal moulds of five conjoined shells, IGPS98870(holotype), 98871–98874; (2) internal moulds of two ventral
valves, IGPS98875. 98876; and (3) external mould of adorsal valve, IGPS98877.
Remarks.—The specimens from Matsukawa were firstdescribed by Tazawa and Araki (1984b, p. 3, pl. 1, figs. 1‒7)
as Richthofenia mabutii Tazawa and Araki, 1984b.
Subsequently, Shen et al. (1992, p. 180) proposed the genus
Neorichthofenia with the Matsukawa species as type species.
Wardlaw et al. (2000, p. 614) classified the Matsukawa
species as the type species of Neorichthofenia Shen, He andZhu, 1992, which belonged to the family Hercosiidae Cooper
and Grant, 1975. The genus Neorichthofenia is characterizedby having a median ridge in ventral valve. Neorichthofenia
mabutii was described by Shen et al. (1992) from also the
Changhsing Formation of Sichuan, southwestern China. No
other species assigned to the genus has been known.
Occurrence.—KY1 Unit in Matsukawa (locality AR4).
Distribution.— Wordian‒Changhsingian: northeasternJapan (Matsukawa in the South Kitakami Belt) andsouthwestern China (Sichuan).

81

Suborder LYTTONIIDINA Williams, Harper and Grant,
2000
Superfamily LYTTONIOIDEA Waagen, 1883
Family LYTTONIIDAE Waagen, 1883
Subfamily LYTTONIINAE Waagen, 1883
Genus LEPTODUS Kayser, 1883
Type species.—Leptodus richthofeni Kayser, 1883.
Leptodus nobilis (Waagen, 1883)(Figs. 47A–K, 48A)
Lyttonia nobilis Waagen, 1883, p. 398, pl. 29, figs. 1–3; pl.
30, figs. 1, 2, 5, 6, 8, 10, 11; Noetling, 1904, p. 112,
text-figs. 4–7; Noetling, 1905, p. 140, pl. 17, figs. 1, 2; pl.
18, figs. 1–11, text-fig. 2; Mansuy, 1913, p. 123, pl. 13,
fig. 10; Mansuy, 1914, p. 32, pl. 6, fig. 7; pl. 7, fig. 1;
Albrecht, 1924, p. 289, fig. 1; Huang, 1932, p. 89, pl. 7,
figs. 9, 10; pl. 8, figs. 8, 9; pl. 9, figs. 1–8, text-figs. 8–11.
Lyttonia sp. Yabe, 1900, p. 2, text-figs. 1, 2.
Oldhamina (Lyttonia) richthofeni var. nobilis Waagen.
Fredericks, 1916, p. 76, pl. 4, fig. 2, text-fig. 22.
Lyttonia richthofeni Kayser. Hayasaka, 1917, p. 43, pl. 18,
figs. 1–8; Hayasaka, 1922a, p. 62, pl. 11, figs. 1–6;
Hayasaka, 1922b, p. 103, pl. 4, figs. 12, 13; Mashiko,
1934, p. 182, text-fig. on p. 182.
Lyttonia (Leptodus) richthofeni Kayser. Hamlet, 1928, p. 31,
pl. 6, figs. 1–4.
Lyttonia richthofeni forma nobilis Waagen: Licharew, 1932b,
p. 69, 96, pl. 2, figs. 13, 14; pl. 5, figs. 1–4, 6, text-fig. 3.
Lyttonia cf. nobilis Waagen. Huang, 1936, p. 493, pl. 1, fig. 5.
Leptodus nobilis (Waagen). Termier and Termier, 1960, p.
241, text-pl. 3, figs. 1–10; Chi-Thuan, 1961, p. 274, pl. 1,
fig. 1; Ding in Yang et al., 1962, p. 90, pl. 37, fig. 4;
Schréter, 1963, p. 107, pl. 3, figs. 5–8; Cooper and Grant,
1974, pl. 191, figs. 8, 9; Grant, 1976, pl. 43, figs. 18, 19;
Lee and Gu, 1976, p. 267, pl. 162, figs. 1, 2; Tazawa,
1976, pl. 2, fig. 8; Yang et al., 1977, p. 371, pl. 147, fig. 5;
Feng and Jiang, 1978, p. 269, pl. 100, fig. 2; Licharew and
Kotlyar, 1978, pl. 14, figs. 13–15; Jin et al., 1979, p. 82,
pl. 23, fig. 15; Minato et al., 1979, pl. 66, figs. 1, 4, 5;Zhan, 1979, p. 93, pl. 9, fig. 12; Lee et al., 1980, p. 389,
pl. 172, figs. 15, 16; Liao, 1980, pl. 6, figs. 42, 43; Wanget al., 1982, p. 229, pl. 95, fig. 20; Gu and Zhu, 1985, pl. 1,
figs. 31, 33, 34; Liao and Meng, 1986, p. 81, pl. 2, figs. 24,
25; Sremac, 1986, p. 30, pl. 10, figs. 1, 2; Liang, 1990, p.
225, pl. 40, figs. 1, 5; Leman, 1994, pl. 1, figs. 3, 4; Zenget al., 1995 , pl. 11, fig. 3; Tazawa et al., 1998, p. 241, figs.
2.1, 2.2, 4; Tazawa and Matsumoto, 1998, p. 7, pl. 2, figs.
7–12; Kato et al., 1999, p. 47, fig. 4; Tazawa, 2000, figs.
3.14, 3.15, 7.1; Tazawa, 2001, p. 297, fig. 7.13–7.16;
Tazawa and Ibaraki, 2001, p. 11, pl. 1, figs. 7–10; Shen etal., 2002, p. 678, fig. 5.28; Tazawa, 2002, fig. 10.14;
Tazawa, 2003, p. 31, fig. 4.1, 4.2; Wang and Zhang, 2003,
p. 118, pl. 22, figs. 13–18; Tazawa, 2009, p. 71, fig. 4.7;

82

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 55. A, B, Meekella eximia (de Verneuil); A, external latex cast of dorsal valve, UHR12576: B, internal mould of ventral valve, UHR13574; C–F,
Meekella nodosa Nakamura; C, ventral view (C1) and posterior view (C2) of internal mould of ventral valve, UHR12578 (holotype); D, external latex castof ventral valve, KCG60; E, external latex cast (E1) and internal mould (E2) of ventral valve, UHR12577; ventral internal mould (F1, F2) and dorsalinternal mould (F3) of conjoined shell, UHR19533.Scale bars are 1 cm.

Tazawa, 2016b, p. 29, figs. 10.1–10.5, 11.5–11.7; Tazawa
and Araki, 2017, p. 276, fig. 8.1, 8.2.
Gubleria armenica Sarytcheva, 1964, p. 68, pl. 8, figs. 1–3;
Sarytcheva and Sokolskaya, 1965, p. 39, figs. 9, 10.
Gubleria sp. Licharew and Kotlyar, 1978, pl. 15, figs. 5, 6.
Leptodus ivanovi Fredericks. Minato et al., 1979, pl. 66, fig. 3.
Leptodus sp. Minato et al., 1979, pl. 66, fig. 2.
Leptodus elongatus Ching and Hu: Wang et al., 1982, p. 229,
pl. 91, figs. 16, 17; pl. 93, fig. 4.

Gubleria sp. Zhu, 1990, p. 80, pl. 16, fig. 24.
Leptodus sp. Yanagida et al., 1993, p. 5, pl. 1, figs. 8, 9.
Leptodus sp. Yanagida, 1996, fig. 2.14.
Leptodus sp. Tazawa, 1999a, p. 5, pl. 1, fig. 1; Tazawa et al.,1999, fig. 2.1.
Gubleria sp. Sone et al., 2001, p. 185, fig. 6.9–6.12.
Leptodus sp. Shen and Zhang, 2008, fig. 5.4.
Material.—Twenty-three specimens from localities AR4,GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

KF13, KF67, KF84, KF217, KF218, KS1 and KS7: (1)internal mould of a ventral valve, with external and internalmoulds of the dorsal valve (internal plate), UHR12111; (2)external and internal moulds of a ventral valve, NU-B1681;(3) external moulds of two ventral valves, NU-B1682, 1683;
and (4) internal moulds of fifty-seven ventral valves,
KCG16–18, NU-B308–314, 1684–1694, 1696–1704,
1706–1726, 2486, UHR11450, 12109, 12110, 12114, 12117.
Remarks.—These specimens were previously describedby Tazawa and Ibaraki (2001, p. 11, pl. 1, figs. 7–10),
Tazawa (2016b, p. 29, figs. 10.1–10.5, 11.5–11.7) andTazawa and Araki (2017, p. 276, fig. 8.1, 8.2) as Leptodus
nobilis (Waagen, 1883). The specimens from Setamai,
Kamiyasse–Imo and Matsukawa are referred to Leptodus
nobilis (Waagen, 1883), originally described from the Wargal
and Chhidru Formations of the Salt Range, by flat ventral
valve with numerous, regularly and symmetrically disposed
broad and solid lateral septa on both sides of median septum.
Leptodus richthofeni Kayser (1883 p. 161, pl. 21, figs. 9–11),
from the upper Permian of Loping, Jiangxi Province, eastern
China, and refigured by Cooper and Grant (1974, pl. 191,
figs. 11–15) on the lectotype, is readily distinguished from L.
nobilis by its more highly convex ventral valve, sharp lateral
septa and wider interseptal spaces.
Occurrence. — KN1 Unit (locality KS1) and KN2 Unit(locality KS7) in Setamai; and KY1 Unit in Kamiyasse(localities KF13, KF67 and KF84), Imo (localities KF217
and KF218) and Matsukawa (locality AR4).
Distribution. — Kungurian–Changhsingian; northeasternJapan (Setamai, Kamiyasse–Imo, Matsukawa, Ogatsu andIchinoseki in the South Kitakami Belt), central Japan(Moribu and Oguradani in the Hida Gaien Belt, and Akasaka
in the Mino Belt), southwestern Japan (Yakuno and Yachiyo
in the Maizuru Belt, and Tsunemori in the Akiyoshi Belt),
Hungary, Croatia, Serbia, northwestern China (Qinghai),
northern China (Inner Mongolia), northeastern China(Heilongjiang and Jilin), eastern Russia (South Primorye),eastern China (Zhejiang, Jiangxi and Fujian),central-southern China (Hubei, Hunan, Guangdong andGuangxi), southwestern China (Guizhou and Sichuan),
Armenia (Transcaucasia), southwestern China (Yunnan),
Malaysia, Cambodia, Indonesia (Timor) and Pakistan (Salt
Range).
Genus KEYSERLINGINA Tschernyschew, 1902
Type species.—Keyserlingina schellwieni Tschernyschew,
1902.
Keyserlingina sp.
(Fig. 49A)
Keyserlingina sp. Tazawa and Araki, 2017, p. 277, fig. 8.3.
Material.—One specimen from locality KZ9, internal

83

mould of a ventral valve, KCG19.
Remarks.— This specimen was previously described by
Tazawa and Araki (2017, p. 277, fig. 8.3) as Keyserlingina
sp. The specimen from Matsukawa is safely assigned to thegenus Keyserlingina on the basis of its small (length 10 mm,
width 8 mm), nearly flat internal plate, with a broad medianridge and symmetrically arranged, broad and deeply grooved
lateral ridges numbering 2 on each side of median ridge. The
Matsukawa species resembles Keyserlngina filicis(Keyserling, 1853), redescribed by Tschernyschew (1902, p.
56, 474, pl. 42, figs. 16, 17) from the lower Permian
Schwagerina Limestone of the Urals, in having lateral ridges
slightly inclined towards anterior. Accurate comparison ofthese two species is, however, difficult for the poorly
preserved material of the Kitakami species.
Occurrence.—KY1 Unit in Matsukawa (locality KZ9).
Genus PETASMAIA Cooper and Grant, 1969
Type species.—Petasmaia expansa Cooper and Grant,
1969.
Petasmaia expansa Cooper and Grant, 1969
(Figs. 48B, 49B)
Petasmaia expansa Cooper and Grant, 1969, p. 10, pl. 2, figs.
15–18; Cooper and Grant, 1974, p. 430, pl. 163, figs. 1–8;
pl. 164, figs. 1–16; pl. 165, figs. 1–23; pl. 169, figs.
11–16; Tazawa and Ono, 2013, p. 51, figs. 2, 3; Tazawa etal., 2016, p. 374, fig. 7.6; Tazawa and Araki, 2018, p. 15,
figs. 5, 6.4.
Material.—One specimen from locality KZ9, internalmould of a ventral valve, KCG55.
Remarks.—This specimen was described by Tazawa andAraki (2018, p. 15, figs. 5, 6.4) as Petasmaia expansa Cooper
and Grant, 1969 on account of transversely wider ventral
valve (length about 45 mm, w idth about 58 mm) andregularly arranged thin lateral septa (numbering 11 pairs),
being gently convex anteriorly and inclined posteriorly. The
Matsukawa specimen, smaller than the type specimens ofWest Texas, may be a young individual. Petasmaia ehiroi
Tazawa and Miyake (2011, p. 8, figs. 3.10, 3.11, 4), from the
Toyoma Formation of Maeda, South Kitakami Belt, differsfrom P. expansa in having lateral septa with narrowerinterspaces.
Occurrence.—KY1 Unit in Matsukawa (locality KZ9).
Distribution.— Artinskian–Wordian: northeastern Japan(Matsukawa in the South Kitakami Belt), central Japan(Akasaka in the Mino Belt) and USA (Texas).
Genus PARARIGBYELLA Shen and Zhang, 2008
Type species.—Pararigbyella quadrilobata Shen andZhang, 2008.

84

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 56. A–C, Orthothetina polita (Fliegel); A, ventral (A1, A2) and dorsal (A3) views of internal mould of conjoined shell, UHR13663; B, ventral(B1) and dorsal (B2) views of internal mould of conjoined shell, UHR13664; C, internal mould of ventral valve, UHR13668; D–G, Orthothetina kayseri(Fliegel); D, ventral (D1) and dorsal (D2) views of internal mould of conjoined shell, UHR13693; E, ventral (E1) and dorsal (E2) views of internal mould ofconjoined shell, UHR13686; F, ventral (F1, F2) and dorsal (F3) views of internal mould of conjoined shell, NU-B2499; G, ventral internal mould (G1),
dorsal internal mould (G2) and ventral external latex cast (G3) of conjoined shell, NU-B2502. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Pararigbyella doulingensis Shen and Zhang, 2008
(Fig. 49C)
Pararigbyella doulingensis Shen and Zhang, 2008, p. 933,
fig. 5.12–5.14; Shen and Tazawa, 2014, p. 247, fig. 3.6–3.8.
Material.—One specimen from locality KF218, internalmould of a ventral valve, NU-B1756.
Remarks.—This specimen was previously described by
Shen and Tazawa (2014, p. 247, fig. 3.6–3.8) as
Pararigbyella doulingensis Shen and Zhang, 2008. The
specimen from Imo is not well preserved, but large size(length 25 mm, width about 16 mm), elongate triangularoutline, bifurcated lobes and distinct median septum which isbounded by two sinuses in posterior one-third, arecomparable with those of Pararigbyella doulingensis Shenand Zhang, 2008, from the uppermost Douling Formation(lower Wuchiapingian) in Chenxian, Hunan, central-southern
China. The type species, Pararigbyella quadrilobata Shenand Zhang (2008, p. 933, fig. 5.5–5.11), from the Douling
Formation of Hunan, is readily distinguished from the present
species by its much smaller dimensions and fewer sinuses.
Occurrence.—KY1 Unit in Imo (locality KF218).
Distribution. —Wordian; northeastern Japan (Kamiyasse–Imoin the South Kitakami Belt) and central-southern China(Hunan).
Subfamily POIKILOSAKINAE Williams, 1953
Genus POIKILOSAKOS Watson, 1917
Type species.—Poikilosakos petaloides Watson, 1917.
Poikilosakos kamiyassensis Tazawa and Takaizumi, 1987
(Fig. 50A–F)
Poikilosakos kamiyassensis Tazawa and Takaizumi, 1987, p.
12, fig. 3.1–3.7.
Material.—Eight specimens from locality KF212, internalmoulds of eight ventral valves, IGPS99009, 99010(holotype), 99011–99016.
Remarks.—These specimens were described by Tazawa
and Takaizumi (1987, p. 12, fig. 3.1–3.7) as Poikilosakos
kamiyassensis Tazawa and Takaizumi, 1987. The Kitakami
species is large, transverse Poikilosakos (length about 21mm, width about 34 mm in the holotype, IGPS99010), having
a pair of long, narrow, often irregular brachial lobes.
Poikilosakos kamiyassensis most resembles Poikilosakos
tschernyschewi Fredericks (1926, p. 83, fig. 1), from thelower Permian (Sakmarian) of the central Ural Mountains,
central Russia, in size, shape and in having long, narrow
lobes, but the Russian species differs in having lessfrequently branched and more symmetrically developed
lobes. The type species, Poikilosakos petaloides Watson

85(1917, p. 213, pl. 14, figs. 1–5), from the upper
Pennsylvanian of north-central Texas, is readily distinguishedfrom the Kitakami species by its smaller size and in having
massive lobes.
Occurrence.—Upper part of the Hosoo Formation in
Kamiyasse (locality KF212).
Distribution.—Roadian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt).
Family RIGBYELLIDAE Williams, Harper and Grant, 2000
Genus PARALYTTONIA Wanner in Wanner and Sieverts,
1935
Type species.—Paralyttonia permica Wanner in Wannerand Sieverts, 1935.
Paralyttonia kesennumensis Tazawa and Araki, 1984a(Fig. 51A)
Paralyttonia kesennumensis Tazawa and Araki, 1984a, p.
122, fig. 2.1, 2.2; Tazawa and Araki, 2017, p. 279, fig. 8.4.
Material.—Two specimens from locality AR4, internalmoulds of two ventral valves, IGPS98393, 98394.
Remarks.—These specimens were previously describedby Tazawa and Araki (1984a, p. 122, fig. 2.1, 2.2) andTazawa and Araki (2017, p. 279, fig. 8.4) as Paralyttonia
kesennumensis Tazawa and Araki, 1984a, on the basis ofsmall, transverselly elliptical shell (length about 7 mm, widthabout 11 mm in the holotype) with 5 septa in ventral valve
and 6 lobes in dorsal valve. The Kitakami species resembles
Paralyttonia tenax Grant (1976, p. 168, pl. 44, figs. 4–36; pl.
45, figs. 32–42, text-fig. 15), from the Rat Buri Formation ofKo Muk, southern Thailand, but differs from the Thailand
species in its smaller size and in having longer and moreregular septa.
Occurrence.—KY1 Unit in Matsukawa (locality AR4).
Distribution.—Wordian: northeastern Japan (Matsukawa
in the South Kitakami Belt).
Superfamily PERMIANELLOIDEA He and Zhu, 1979
Family PERMIANELLIDAE He and Zhu, 1979
Genus PERMIANELLA He and Zhu, 1979
Type species.—Permianella typica He and Zhu, 1979.
Permianella typica He and Zhu, 1979
(Fig. 51B–G)
Permianella typica He and Zhu, 1979, p. 132, 137, pl. 1, fig.
1; pl. 2, figs. 1–3; pl. 3, figs. 1–3; Wang and Jin, 1991, p.
496, pl. 2, figs. 1–3; Zeng et al., 1995, pl. 21, fig. 16; Shen
and Tazawa, 1997, p. 288, figs. 2–4, 5.1–5.14; Campi etal., 2000, p. 41, figs. 4A –C, 5A, B; Campi et al., 2002,
fig. 6P; Tazawa, 2002, fig. 10.13; Campi et al., 2005, p.

86JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 57. A, B, Orthothetina transversa Nakamura; A, ventral (A1) and dorsal (A2) views of internal mould of conjoined shell, UHR19532; B, externallatex cast (B1) and internal mould (B2) of ventral valve, NU-B2498; C, D, Orthothetina hayasakai Nakamura; C, ventral internal mould (C1, C2), dorsalinternal Mould (C3) and ventral external latex cast (C4) of conjoined shell, UHR13572 (holotype); D, ventral (D1) and dorsal (D2) views of internal mouldof conjoined shell, NU-B2505; E, Orthothetina kitakamiensis (Hayasaka), ventral (E1, E2) and dorsal (E3) views of internal mould of conjoined shell, solespecimen (not registered) figured by Hayasaka (1953, pl. 9, figs. 1, 2). Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

127, pl. 4, figs. I, J; Tazawa, 2008b, p. 50, fig. 8.1;
Tazawa et al., 2014, p. 383, fig. 2.7; Tazawa, 2015, p. 72,
fig. 6.7; Shen et al., 2017, pl. 20, figs. 3–5.
Material.—Thirty-five specimens from localities KF46,
KF88, KF91, KF94, KF217 and KF218: (1) two conjoined
shells, NU-B51, 59: (2) a ventral valve, NU-B52; (3) external
and internal moulds of three ventral valves, NU-B58, 62, 63;(4) internal moulds of twenty-six ventral valves, NU-B54–56,
61, 64–66, 68, 69, 1669–1678, 2027, 2028, 2487–2490; and(5) external moulds of four dorsal valves, NU-B50, 57, 60,
67.
Remarks.—Most of the specimens were described by
Shen and Tazawa (1997, p. 288, figs. 2–4, 5.1–5.14) as
Permianella typica He and Zhu, 1979. The specimens from
Setamai and Kamiyasse–Imo can be referred to Permianella
typica He and Zhu (1979), originally described from theLungtan Formation of Jiangxi, eastern China and Sichuan,southwestern China, by large size (length 49 mm, width 17
mm in the largest specimen, NU-B64), elongate ovate
outline, deep incision, irregular marginal brim and nearlyparallel anterolateral sides. Permianella sp. He and Zhu(1979, p. 133, 139, pl. 1, figs. 2, 3), from the Lungtan
Formation of Jiangxi, eastern China, differs from
Permianella typica in its nearly plano-convex shell and inhaving a very low dorsal fold.
Occurrence.—KY1 Unit in Kamiyasse (localities KF46,
KF88, KF91 and KF94) and Imo (localities KF217 andKF218).
Distribution.—Wordian–Wuchiapingian: northeasternJapan (Setamai, Kamiyasse–Imo and Hitachi in the South
Kitakami Belt), southwestern Japan (Mizukoshi in Kyushu
Island), eastern China (Zhejiang and Jiangxi), southwestern
China (Sichuan) and Malaysia.
Genus DICYSTOCONCHA Termier and Termier in Termieret al., 1974
Type species.—Dicystoconcha lapparenti Termier andTermier in Termier et al., 1974.
Dicystoconcha lapparenti Termier and Termier in Termier etal., 1974(Fig. 52A–E)
Dicystoconcha lapparenti Termier and Termier in Termier etal., 1974, p. 123, pl. 22, figs. 1, 2, text-fig. 22; Wang andJin, 1991, p. 495, pl. 1, figs. 1‒9; pl. 3, figs. 1‒7; Shen andTazawa, 2014, p. 248, fig. 3.1‒3.5; Tazawa et al., 2014, p.
383, fig. 2.6; Tazawa, 2015, p. 73, fig. 6.6; Tazawa andAraki, 2018, p. 16, fig. 4.2; Tazawa and Shintani, 2021, p.
11, fig. 4I.
Dipunctella contracta Liang in Wang et al., 1982, p. 229, pl.
102, fig. 3.
Guangjiayanella guangjiayanensis Yang, 1984, p. 212, pl.

87

31, figs. 11‒16, text-fig. 5.9.
Guangdongina xiamaoensis Mou and Liu, 1989, p. 458, pl. 1,
figs. 1‒9; pl. 2, figs. 1‒7, text-fig. 5.
Guangdongina leguminiformis Mou and Liu, 1989, p. 458, pl.
3, figs. 4‒8.
Guangdongina perforatus Mou and Liu, 1989, p. 459, pl. 2,
fig. 8; pl. 3, figs. 1‒3.
Guangdongina sp. Mou and Liu, 1989, p. 459, pl. 2, fig. 9.
Paritisteges latesulcata Liang, 1990, p. 380, pl. 42, figs. 1, 2.
Febulasteges planata Liang, 1990, p. 381, pl. 42, figs. 3, 4.
Material. ― Thirteen specimens from localities AR4,
KF217 and KF218, internal moulds of ventral valves,
KCG58. NU-B1679, 1680, 2491–2494, RCMF10001–10006.
Remarks.—These specimens were previously describedby Shen and Tazawa (2014, p. 248, fig. 3.1‒3.5) and Tazawa
and Araki (2018, p. 16, fig. 4.2) as Dicystoconcha lapparenti
Termier and Termier in Termier et al., 1974. The specimensfrom Kamiyasse–Imo and Matsukawa are referred toDicystoconcha lapparenti Termier and Termier in Termier etal., 1974 in small size (length 25 mm, width 27 m in thelargest specimen, RCM-F10001; length 19 mm, width 22 mm
in an average-sized specimen, RCM-F10003), slightly
triangular ovate outline, and strongly convex and bilobate
ventral valve. Shen and Tazawa (2014, p. 248) treated thefollowing six species from the Permian of South China as
junior synonyms of Dicystoconcha lapparenti:
Guangjiayanella guangjiayanensis Yang, 1984,
Guangdongina xiamaoensis Mou and Liu, 1989,
Guangdongina leguminiformis Mou and Liu, 1989,
Guangdongina perforatus Mou and Liu, 1989,
Guangdongina sp. Mou and Liu, 1989 and Fabulasteges
planata Liang, 1990. Moreover, Paritisteges latesulcata
Liang, 1990, from the lower Lengwu Formation of Lengwu,
Zhejiang, eastern China, is also considered to be a juniorsynonym of the present species.
Occurrence.—KY1 Unit in Imo (localities KF217 andKF218) and Matsukawa (locality AR4).
Distribution.—Kungurian–Wuchiapingian: northeasternJapan (Nagaiwa–Sakamotozawa, Kamiyasse–Imo,
Matsukawa and Hitachi in the South Kitakami Belt), northern
China (Inner Mongolia), eastern China (Zhejiang),
central-southern China (Hubei, Hunan and Guangdong) andAfghanistan.
Genus LATERISPINA Wang and Jin, 1991
Type species.—Laterispina liaoi Wang and Jin, 1991.
Laterispina parallela Shen, Fan, Zhang and Zhang, 1994
(Fig. 52F)
Laterispina parallela Shen, Fan, Zhang and Zhang, 1994, p.
478, pl. 1, figs. 1–12; pl. 2, figs. 1–11, 14, text-figs. 1–5;
Shen and Tazawa, 1997, p. 289, fig. 5.15.

88

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 58. A, Orthothetina transversa Nakamura, ventral internal mould (A1, A2), dorsal internal mould (A3) and ventral external latex cast (A4) ofconjoined shell, UHR13573 (holotype); B, C, Streptorhynchus pelargonatus (Schlotheim); B, dorsal internal mould (B1 , B 2 ) of conjoined shell,
UHR12367; C, ventral (C1, C2), dorsal (C3) and posterior (C4) views of internal mould and dorsal external latex cast (C5) of conjoined shell, UHR13584;
D, Streptorhynchus cataclinus Zhang, ventral external latex cast (D 1, D2) and ventral (D3), dorsal (D4) and posterior (D5) views of internal mould ofconjoined shell, NU-B2508. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Material.—One specimen from locality KS9, internalmoulds of a ventral valve, NU-B70.
Remarks.—This specimen was previously described by
Shen and Tazawa (1997, p. 289, fig. 5.15) as Laterispina
parallela Shen, Fan, Zhang and Zhang, 1994. The Setamai
specimen can be referred to Laterisma parallela Shen, Fan,
Zhang and Zhang (1994, p. 478, pl. 1, figs. 1–12; pl. 2, figs.
1–11, 14, text-figs. 1–5), from the Changhsing Formation of
Sichuan, southwestern China, by large, very long shell(length 55 mm, width 14 mm), with nearly parallel sides andin having marginal brim along lateral commissure. The typespecies, Laterispina liaoi Wang and Jin (1991, p. 497. pl. 2,
figs. 4–12), from the Changhsing Formation of Guangxi,
central-southern China, differs from L. parallela in its lesselongate, subtriangular outline.
Occurrence.—KN2 Unit in Setamai (locality KS9).
Distribution.—Wordian–Changhsingian: northeasternJapan (Setamai in the South Kitakami Belt) and southwestern
China (Sichuan).
Order ORTHOTETIDA Waagen, 1884
Suborder ORTHOTETIDINA Waagen, 1884
Superfamily ORTHOTETOIDEA Waagen, 1884
Family DERBYIIDAE Stehli, 1954
Genus DERBYIA Waagen, 1884
Type species.—Derbyia regularis Wagen, 1884.
Derbyia grandis Waagen, 1884
(Fig. 53A, B)
Derbyia grandis Waagen, 1884, p. 597, pl. 51, fig. 1–1c; pl.
52, figs. 1, 3; pl. 53, figs. 3, 5; Tschernyschew, 1902, p.
207, 580, pl. 24, figs. 1, 2; pl. 24, figs. 1, 2; pl. 26, fig. 5,
text-figs. 59, 60; Broili, 1916, p. 7, pl. 115, fig. 9; Frebold,
1950, p. 41, pl. 1, fig. 5: Grunt, 1973, p. 84, pl. 3, figs.
1–4; Manankov, 1973, pl. 8, figs. 4, 5; Kulikov, 1974, p.
82, pl. 1, fig. 6; Lee and Gu, 1976, p. 236, pl. 161, fig. 12;
Licharew and Kotlyar, 1978, pl. 12, fig. 1; Lee et al., 1980,
p. 336, pl. 159, fig. 13; Kalashnikov, 1986, pl. 114, fig. 1;
Nakamura et al., 1992, pl. 1, fig. 1; Gu, 1992, p. 215, pl.
67, fig. 17; Kalashnikov, 1993, p. 23, pl. 6, figs. 1, 2; pl. 7,
fig. 7; Fang and Fan, 1994, p. 73, pl. 27, figs. 5–8; pl. 28,
figs. 1, 2; Angiolini, 1996, p. 9, pl. 1, figs. 7–9; Tazawa
and Ibaraki, 2001, p. 13, pl. 2, fig. 9; pl. 3, fig. 3a –c;
Wang and Zhang, 2003, p. 120, pl. 24, figs. 1–3; pl. 25,
fig. 4; pl. 27, figs. 13–15; Shen et al., 2003b, p. 60, pl. 1,
figs. 1–3, text-fig. 4.
Derbyia cf. grandis Waagen. Frech, 1911, p. 125, pl. 18, fig.
4a–d; Stepanov, 1937a, p. 110, 174, pl. 1, fig. 5; Harker,
1960, p. 52, pl. 16, figs. 9, 10; Bamber and Waterhouse,
1971, pl. 18, fig. 1; Manankov, 1979, p. 57, pl. 4, fig. 1,
text-fig. 25.
Derbyia aff. grandis Waagen. Gobbett, 1963, p. 54, pl. 2,

89

figs. 1, 2.
Wardakia grandis (Waagen). Termier et al., 1974, p. 95, pl. 9,
figs. 2–5; pl. 10, figs. 1–3, text-fig. 14.
Material.—Two specimens from localities KS1 and KS9,external and internal moulds of two ventral valves, NU-B363,
364.
Remarks.—These specimens were previously describedby Tazawa and Ibaraki (2001, p. 13, pl. 2, fig. 9; pl. 3, fig. 3)
as Derbyia grandis Waagen, 1884. The specimens from
Setamai can be referred to Derbyia grandis Waagen (1884, p.
597, pl. 51, fig. 1; pl. 52, figs. 1, 3; pl. 53, figs. 3, 5), fromthe Wargal and Chhidru Formations of the Salt Range,
Pakistan, on account of large, transverse shell (length about
63 mm, w idth 115 mm in better pres erved specimen,
NU-B363) and external ornament of ventral valve, consistingof strong, irregular rugae and numerous costellae with broadinterspaces. The Setamai specimens closely resemble to theshells, described by Harker (1960, p. 52, pl. 16, figs. 9, 10) as
Derbyia cf. grandis Waagen, from the Assistance Formationof Devon Island, northern Canada, in having slightly concave
to nearly flat ventral valve.
Occurrence.—KN1 Unit (locality KS1) and KN2 Unit in
Setamai (locality KS9).
Distribution.—Asselian–Wuchiapingian: northeasternJapan (Setamai in the South Kitakami Belt), northern Canada(Devon Island and northern Yukon Territory), northeastern
Greenland, Spitsbergen, northern Russia (Timan, northern
Urals and Kolyma Massif), northern China (Inner Mongolia),
northeastern China (Jilin), eastern Russia (South Primorye),eastern China (Jiangsu), Afghanistan (Wardak), Tajikistan(Pamir), southwestern China (Yunnan), Indonesia (Timor),
northwestern China (Tibet) and Pakistan (Karakoram and Salt
Range).
Derbyia nipponica Nakamura, 1972
（Fig. 54A–F）
Derbyia magnifica Licharew. Hayasaka and Minato, 1956, p.
141, pl. 23, fig. 1; Hayasaka, 1960, p. 45, pl. 2, figs. 5, 6.
Derbyia nipponica Nakamura, 1972, p. 399, pl. 7, figs. 1, 4,
9; Minato et al., 1979, pl. 60, figs. 1–4; Tazawa and
Ibaraki, 2001, p. 14, pl. 3, figs. 1, 2, 4; Tazawa, 2008b, p.
50, fig. 8.12–8.16.
Derbyia cf. buchi d’Orbigny. Tazawa and Matsumoto, 1998,
p. 5, pl. 1, figs. 1, 2.
Material.—Nine specimens from localities KF22, KF121,
KS1 and KS9: (1) internal mould of a conjoined shell, withexternal mould of the ventral valve, UHR17035; (2) external
and internal moulds of four ventral valves, NU-B365–367,
UHR12470; (3) internal mould of two ventral valves,
NU-B38, 2485; and (4) internal mould of two dorsal valves,
NU-B369, 2486.
Remarks.—These specimens were previously described

90

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 59. A–F, Rhipidomella magna Tazawa; A, ventral (A1, A2), dorsal (A3), anterior (A4), posterior (A5) and lateral A6) views of conjoined shell,
NU-B1757; B, ventral (B1) and dorsal (B2) views of conjoined shell, NU-B1759; C, internal mould of ventral valve, NU-B1974; D, ventral valve,
NU-B1760; E, dorsal valve, NU-B1762; F, ventral (F1) and dorsal (F2) views of conjoined shell, NU-B1762. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

by Tazawa and Ibaraki (2001, p. 14, pl. 3, figs. 1, 2, 4) as
Derbyia nipponica Nakamura, 1972. The specimens from
Setamai are referred to Derbyia nipponica Nakamura, 1972,
originally described from the lower Kanokura Formation of
Imo in the South Kitakami Belt, by medium-sized (length 41mm, width 49 mm in the best-preserved ventral valve
specimen, NU-B365; length 51 mm, width about 73 mm in
the largest dorsal valve specimen, NU-B369), weakly
biconvex and transverse shell with short hinge and regularly
developed fine costellae with narrow interspaces. Derbyia
magnifica Licharew (1932a, p. 18, 40, pl. 1, figs. 1, 2, 6, 7,
13; pl. 2, fig. 1; pl. 3, fig. 1), from the lower Permian of thenorthern Caucasus, differs from D. nipponica in having long
hinge marking the greatest width, ventral fold and dorsal
sulcus. Derbyia regularis Waagen (1884, p. 504, pl. 53, figs.
1, 2, 4), from the Amb and Wargal Formations of the Salt
Range, is also medium to large, transverse Derbyia species,
but differs from the present species in having less dense
costellae.
Occurrence.—KN1 Unit (locality KS1) and KN2 Unit in
Setamai (locality KS9); KY1 Unit in Kamiyasse (locality
KF22) and Imo (locality KF121).
Distribution.—Roadian–Wuchiapingian: northeasternJapan (Setamai and Kamiyasse–Imo in the South KitakamiBelt), central Japan (Oguradani in the Hida Gaien Belt) andsouthwestern Japan (Mizukoshi in Kyushu Island).
Family MEEKELLIDAE Stehli, 1954
Subfamily MEEKELLINAE Stehli, 1954
Genus MEEKELLA White and St. John, 1867
Type species.―Plicatula striatocostata Cox, 1857.
Meekella eximia (de Verneuil, 1845)(Fig. 55A, B)
Orthis eximia de Verneuil, 1845, p. 192, pl. 11, fig. 2.
Meekella striatocostata (Cox). Tschernyschew, 1902, p. 211,
582, pl. 24, figs. 7, 8; pl. 26, fig. 4; pl. 51, fig. 3; Dunbar
and Condra, 1932, p. 125, pl. 16, figs. 1 –10; pl. 17, fig. 3.
Meekella eximia (Eichwald). Tschernyschew, 1902, p. 213,
582, pl. 24, fig. 5; pl. 51, fig. 13; Stepanov, 1937a, p. 111,
174, pl. 1, fig. 7; Licharew and Einor, 1939, p. 21, 203, pl.
2, fig. 1; Lapina, 1957, p. 23, pl. 1, fig. 6; Sokolskaya in
Sarytcheva and Sokolskaya, 1952, p. 58, pl. 9, fig. 46;
Sokolskaya, 1954, p. 157, pl. 17, figs. 4–8; Zavodowsky
and Stepanov, 1970, p. 72, pl. 42, figs. 3–5; Kalashnikov,
1980. p. 26, pl. 1, fig. 9; Zhang et al., 1983, p. 277, pl.
125, figs. 14, 15; Wang, 1995, pl. 1, fig. 10; Wang andYang, 1998, p. 63, pl. 2, fig. 7.
Meekella striatocostata (Cox). Hayasaka, 1937, p. 268,
text-figs. 1, 2; Hayasaka, 1963d, fig. 5.
Meekella cf. eximia (de Verneuil). Czarniecki, 1969, p. 262,
pl. 2, fig. 1.
Meekella cf. striatocostata (Cox). Nakamura, 1972, p. 385,

91

pl. 5, fig. 1; Minato et al., 1979, pl. 59, figs. 3, 4.
Meekella cf. eximia (Eichwald). Nakamura, 1972, p. 386, pl.
6, figs. 5, 6; Minato et al., 1979, pl. 59, fig. 5.
Mekella eximia (de Verneuil). Manankov, 1979, p. 39, pl. 1,
figs. 2–4, text-figs. 2, 15.
Material.—Two specimens from localities KF64 andKF121: (1) internal mould of a ventral valve, UHR13574;
and (2) external mould of a dorsal valve, UHR12576.
Remarks.—Among two specimens from Kamiyasse–Imo,the larger one (UHR12576) was previously described byNakamura (1972, p. 386, pl. 6, figs. 5, 6) as Meekella cf.
eximia (von Eichwald, 1840); and the smaller one(UHR13574) was described by Nakamura (1972, p. 385, pl.
5, fig. 1) as Meekella cf. striatocostata (Cox, 1857).
Subsequently, Manankov (1979, p. 39) treated the Kitakami
specimens as synonyms of Mekella eximia (de Verneuil,1845). Czarniecki (1969, p. 262) commented about the authorof Meekella eximia as follows: According to the Code ofZoological Nomenclature, the author of this species is deVerneuil, who first published its description and figures it.
The Kitakami specimens can be referred to Meekella eximia(de Verneuil, 1845), redescribed by Sokolskaya (1954, p.
157, pl. 17, figs. 4–8), from the Moscovian–Gzhelian of theRussian Platform, in medium, transverse shell (length about
28 mm, width about 45 mm in the larger specimen,
UHR12576) and in having thick and relatively few costae onboth ventral and dorsal valves. Meekella uralica
Tschernyschew (1902, p. 215, 583, pl. 51, figs. 1, 2), fromthe lower Permian (Schwagerina Limestone) of the Urals,differs from M. eximia in having thick but shorter costae
which occur on posterior half of both ventral and dorsal
valves.
Occurrence.—KY1 Unit in Kamiyasse (locality KF64)
and Imo (locality KF121).
Distribution.—Moscovian–Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt), USA (Iowa,
Missouri, Nebraska and Kansas), Spitsbergen, northern
Russia (Pechora Basin, Novaya Zemlya and Omolon Massif),
western Russia (Moscow Basin), central Russia (southern
Urals) and northwestern China (Xinjiang).
Meekella nodosa Nakamura, 1972
(Fig. 55C–F)
Meekella sp. Huang, 1933, p. 31, pl. 4, figs. 8, 9.
Meekella garnieri Bayan. Nakamura, 1972, p. 387, pl. 5, figs.
5–7; Minato et al., 1979, pl. 59, figs. 1, 2.
Meekella nodosa Nakamura, 1972, p. 388, pl. 6, figs. 3, 4;
Minato et al., 1979, pl. 59, fig. 8; Tazawa and Araki, 2018,p. 17, fig. 4.3.
Material.—Four specimens from localities KF64, KF121
and KZ9: (1) internal mould of a conjoined shell,
UHR19533; (2) external moulds of two ventral valves,

92

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 60. A, B, Acosarina rectimarginata Cooper and Grant; A, ventral (A 1, A2), dorsal (A3), anterior (A4), posterior (A5) and lateral (A6) views ofinternal mould and ventral (A7) and dorsal (A8) views of external cast of conjoined shell, NU-B2467; B, ventral (B1) and dorsal (B2) views of internalmould of conjoined shell, NU-B2468; C, D, Acosarina dorsisulcata Cooper and Grant; C, ventral (C1, C2), dorsal (C3), anterior (C4), posterior (C5) andlateral (C6) views of internal mould of conjoined shell, NU-B2465; D, internal mould of ventral valve, NU-B2466; E, F, Orthotichia sp.; E, internal mould(E1, E2) of ventral valve, NU-B2497; F, internal mould of ventral valve, UHR12624. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

KCG60, UHR12577; and (3) internal mould of a ventral
valve, UHR12578 (holotype).
Remarks. —One of the specimens (UHR12578) from Imo
was previously described by Nakamura (1972) as theholotype of Meekella nodosa Nakamura, 1972. The
specimens from Kamiyasse, Imo and Matsukawa can bereferred to Meekella nodosa Nakamura, 1972, by medium tolarge size (length about 44 mm, width about 60 mm in theholotype, UHR12578) and external ornament consisting of
numerous nodose costae and irregularly developed strong
rugae. As noted by Nakamura (1972, p. 389), Meekella sp.
Huang, 1933, from the Wuchiapingian of Guizhou,
southwestern China, is a synonym of Meekella nodosa.
Furthermore, two specimens described by Nakamura (1972,
p. 388, pl. 6, figs. 3, 4) as Meekella garnieri Bayan arer e f e r r e d t o t h e p re s e n t s p ec ie s . Me e k e lla ir r e gular is
Schellwien, 1900b (p. 20, pl. 2, figs. 8, 9), from the
Trogkofel Limestone of the Karavanke Mountains, Slovenia,
also has nodose ornament on ventral valve, but the
Karavanke species differs from M. nodosa in its muchelongate outline. Mekella magnifica Cooper and Grant (1974,
p. 365, pl. 100, figs. 1–33; pl. 116, figs. 9–18), from theWolfcampian of Texas, differs from M. nodosa in having
much longer interarea of the ventral valve.
Occurrence.—KY1 Unit in Kamiyasse (locality KF64),
Imo (locality KF121) and Matsukawa (locality KZ9).
Distribution.—Roadian–Wuchiapingian: northeasternJapan (Kamiyasse–Imo and Matsukawa in the South
Kitakami Belt) and southwestern China (Guizhou).
Genus ORTHOTHETINA Schellwien, 1900a
T y pe spe cie s . —O r th o th e te s p e r s ic u s S c huc he rt in
Schuchert and LeVene, 1929.
Orthothetina polita (Fliegel, 1901)(Fig. 56A–C)
Orthothetes politus Fliegel, 1901, p. 97, pl. 6, fig. 8.
Orthotetina polita Fliegel. Nakamura, 1972, p. 380, pl. 3,
figs. 4; pl. 4, figs. 1, 2 .
Orthothetina polita Fliegel. Minato et al., 1979, pl. 58, fig. 5.
Material.—Four specimens from localities KF121and
KF217: (1) internal mould of two conjoined shells,
UHR13663, 13664; (2) external and internal moulds of aventral valve, UHR13668; and (3) internal mould of a ventral
valve, NU-B2501.
Remarks.—These specimens were previously describedby Nakamura (1972, p. 380, pl. 3, fig. 4; pl. 4, figs. 1, 2) as
Orthotetina polita Fliegel, and refigured by Minato et al.
(1979, pl. 58, fig. 5) as Orthothetina polita Fliegel. The
specimens from Imo can be referred to Orthothetina polita(Fliegel, 1901) on the basis of medium size (length about 31
mm, width about 40 mm in the largest specimen,

93

UHR13663), subtriangular outline and in having a broad anddeep sulcus on dorsal valve and somewhat twisted ventral
beak. Orthothetina planoconvexa Hayasaka (1933, p. 32, pl.
9, fig. 2; text-fig. 5), from the Nabeyama Limestone of the
Ashio Mountains, central Japan, resembles O. polita inoutline and shape of shell, but larger in size. Orthothetina
azarjani Kotlyar (1989, p. 118, pl. 21, fig. 7; pl. 22, figs.
2–5), from the Midian of the Caucasus, western Russia,
appears to be conspecific with the present species, althoughcomparison of the two species is difficult owing to ill
preservation of the Kitakami specimens.
Occurrence.—KY1 Unit in Imo (localities KF121 andKF217).
Distribution.—Upper Carboniferous–Wordian:northeastern Japan (Kamiyasse–Imo in the South KitakamiBelt) and Indonesia (Sumatra).
Orthothetina kayseri (Fliegel, 1901)(Fig. 56D–G)
Orthothetes kayseri Jäkel. Fliegel, 1901, p. 127, pl. 6, fig. 9.
Orthotetina kayseri (Jäkel) Fliegel. Minato and Nakamura,
1956, p. 152, pl. 24, fig. 4.
Orthotetina kayseri (Jaekel). Hayasaka, 1963d, p. 753, figs.
1, 2 only.
Orthotetina kayseri (Jäkel). Nakamura, 1972, p. 380, pl. 2,
figs. 2–5.
Orhtothetina kayseri (Jäkel). Minato et al., 1979, pl. 58, figs.
3, 4.
Material.—Seven specimens from localities KF91,
KF121, KF217 and KZ9: (1) external and internal moulds ofa conjoined shell, NU-B2499; (2) internal mould of aconjoined shell, with external mould of the ventral valve,
NU-B2502; (3) internal mould of a conjoined shell, withexternal mould of the dorsal valve, UHR17071; (4) internalmoulds of two conjoined shells, UHR13686, 13693; and ( 5)internal mould of two ventral valves, NU-B2500, 2503.
Remarks.—Most of the specimens (UHR13686, 13693,
17071) were previously described by Minato and Nakamura(1956, p. 152, pl. 24, fig. 4) and Nakamura (1972, p. 380, pl.
2, figs. 2–5) as Orthothetina kayseri (Flieger, 1901). This
species is medium in size for the genus, slightly longer
subrectangular in outline; ventral valve flat to slightly convex
in lateral profile, interarea moderately high; dorsal valeslightly convex; length about 36 mm, width about 33 mm,height of ventral interarea about 9 mm in the best-preserved
specimen (NU-B2499). The preceding species, Orthothetina
polita (Fliegel, 1901), is distinguished from the present
species in having somewhat twisted ventral beak.
Orthothetina ruber (Frech, 1911, p. 124, 174, pl. 26, fig. 4),
from the Lopingian of Jiangxi, eastern China is also
subrectangular in outline, but differs from O. kayseri in its
much larger size.
Occurrence.—KY1 Unit in Kamiyasse (locality KF91),

94

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 61. A–D, Stenoscisma margaritovi (Tschernyschew); A, ventral (A1, A2), dorsal (A3), anterior (A4), posterior (A5) and lateral (A6) views ofinternal mould of conjoined shell, NU-B2556; B, ventral valve, NU-B2547; C, ventral valve, NU-B2550; D, ventral (D1, D2), dorsal (D3), anterior (D4),posterior (D5) and lateral (D6) views of conjoined shell, NU-B2555. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Imo (localities KF121 and KF217) and Matsukawa (locality
KZ9).
Distribution.—Roadian–Changhsingian: northeasternJapan (Kamiyasse–Imo and Matsukawa in the South
Kitakami Belt) and eastern China (Jiangxi).
Orthothetina kitakamiensis (Hayasaka, 1953)(Fig. 57E)
Hamletella kitakamiensis Hayasaka, 1953, p. 93, pl. 9, figs.
1, 2.
Orthotetina kitakamiensis (Hayasaka). Nakamura, 1972, p.
375, text-fig. 6.
Material.—One specimen from locality KS10, internalmould of a conjoined shell (figured by Hayasaka, 1953, pl. 9,
figs. 1, 2).
Remarks.—The single specimen from Setamai were firstdescribed by Hayasaka (1953, p. 93, pl. 9, figs. 1, 2) as
Hamletella kitakamiensis Hayasaka, 1953. Subsequently, thegenus Hamletella was treated as a synonym of the genus
Orthothetina by Muir-Wood and Williams (1965, p. 406).
Orthothetina kitakamiensis (Hayasaka, 1953) has a pair oflong, thin and subparallel dental pates in the ventral valve,and is characterized by its elongate outline (length 44 mm,
width about 21 mm in the sole specimen), with high ventral
interarea (height 23 mm, width of hinge line 17 mm). This
species is the most elongate Orthothetina species from theSouth Kitakami region, as noted and illustrated by Nakamura(1972, text-fig. 6).
Occurrence.—KN2 Unit in Setamai (locality KS10).
Distribution.—Wordian: northeastern Japan (Setamai in
the South Kitakami Belt).
Orthothetina hayasakai Nakamura, 1972
(Fig. 57C, D)
Orthotetes rugosa Fredericks. Hayasaka, 1925a, p. 92, pl. 9,
fig. 1.
Orthotetina kayseri (Jaekel). Hayasaka, 1963b, p. 753, fig. 3
only.
Orthotetina hayasakai Nakamura, 1972, p. 376, pl. 1, figs. 1, 2.
Orthotetina elongata Nakamura, 1972, p. 379, pl. 1, figs.
3–6; pl. 2, fig. 1.
Orthothetina hayasakai Nakamura. Minato et al., 1979, pl.
58, fig. 1.
Orthothetina elongata Nakamura. Minato et al., 1979, pl. 58,
fig. 2; Shen et al., 1992, p. 176, pl. 2, figs. 7–10; Shen andShi, 2007, p. 23, pl. 7, figs. 21–24.
Orthothetina rara Shen, He and Zhu, 1992, p. 176, pl. 2, figs.
7–10.
Material.—Seven specimens from localities KF121 andKF217: (1) external and internal moulds of a conjoined shell,
UHR13572 (holotype); (2) internal mould of a conjoined95

shell, with external mould of the dorsal valve, UHR13567;(3) external and internal moulds of a ventral valve,
UHR13689; and (4) internal moulds of four ventral valves,
NU-B2504, 2505, UHR13690, 13692.
Remarks.—Five specimens from Imo consists of theholotype of Orthothetina hayasakai Nakamura, 1972
(UHR13572) and the holotype (UHR13567) and theparatypes (UHR13689, 13690, 13692) of Orthothetina
elongata Nakamura, 1972. In the present study, Orthothetina
elongata is treated as a synonym of Orthothetina hayasakai
because of both species are characterized by medium size andelongate oval outline. As noted by Shen and Shi (2007, p.
23), Orthothetina rara Shen, He and Zhu, 1992 (p. 176, pl. 2,
figs. 7–10), from the Changhsingian Formation ofBeifengjing, Sichan, southwestern China, is a synonym of thepresent species. Orthothetina zhejiangensis Liang (1990, p.
135, pl. 14, figs. 5–14), from the Lengwu Formation ofZhejiang, eastern China, is also elongate in outline, butdiffers from O. hayasakai in its much larger size.
Occurrence.—KY1 Unit in Imo (localities KF121 andKF217).
Distribution.—Wordian–Changhsingian: northeasternJapan (Kamiyasse–Imo in the South Kitakami Belt) andsouthwestern China (Sichuan).
Orthothetina transversa Nakamura, 1972
(Figs. 57A, B, 58A)
Orthotetina transversa Nakamura, 1972, p. 382, pl. 3, figs.
1–3
Orthothetina transversa Nakamura. Minato et al., 1979, pl.
58, fig. 6.
Material.—Three specimens from locality KF217: (1)external and internal moulds of a conjoined shell, UHR13573(holotype); (2) internal mould of a conjoined shell, withexternal mould of the ventral valve, UHR19532; and (3)external and internal moulds of a ventral valve, NU-B2498.
Remarks.—The specimens from Imo, except for one
specimen (NU-B2498), were previously described byNakamura (1972, p. 382, pl. 3, figs. 1–3) as Orthotetinatransversa Nakamura, 1972. Subsequently Minato et al.
(1979, pl. 58, fig. 6) figured the type specimen (UHR13573)
as Orthothetina transversa Nakamura. This species ischaracterized by its transverse outline (length about 25 mm,
width about 43 mm in the holotype) and in having a broad
and large delthyrium on ventral valve. The preceding species,
Orthothetina hayasakai Nakamura, 1972, is readily
distinguished from O, transversa by its much elongate
outline. Orthothetina ruber (Frech, 1911, p. 124, 174, pl. 26,
fig. 4), from the Lopingian of Jiangxi, eastern China, differsfrom the present species in having less transverse ventral
valve.
Occurrence.—KY1 Unit in Imo (locality KF217).
Distribution.—Wordian: northeastern Japan (Kamiyasse–Imo

96

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 62. A–D, Stenoscisma margaritovi (Tschernuschew), A, ventral (A1, A2) and dorsal (A3) views of internal mould and ventral external latex castof conjoined shell, NU-B2554; B, ventral valve, NU-B2548; C, ventral valve, NU-B2549; D, ventral (D1, D2), dorsal (D3), anterior (D4), posterior (D5) andlateral (D6) views of internal mould of conjoined shell, NU-B2546. Scale bars are 1 cm.

in the South Kitakami Belt).
Family SCHUCHERTELLIDAE Williams, 1953
Subfamily STREPTORHYNCHINAE Stehli, 1954
Genus STREPTORHYNCHUS King, 1850
Type species.—Terebratulites pelargonatus vonSchlotheim, 1816.

Streptorhynchus pelargonatus (von Schlotheim, 1816)(Fig. 58B, C)
Terebratulites pelargonatus von Schlotheim, 1816, p. 28, pl.
8, figs. 21–24.
Streptorhynchus pelargonatus (von Schlotheim). Davidson,
1858, p. 32, pl. 2, figs. 32–42; Waagen, 1884, p. 579, pl.
50, figs. 3–5, 7; Licharew, 1932a, p. 11, 36, pl. 4, figs. 2,4; Huang, 1933, p. 17, pl. 2, fig. 13; Minato andGUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Nakamura, 1956, p. 149, pl. 24, fig. 1; Nakamura, 1972, p.
397, pl. 9, fig. 3; Minato et al., 1979, pl. 60, figs. 6, 7; Leeet al., 1980, p. 332, pl. 158, figs. 8, 20; Yang, 1984, p. 207,
pl. 30, fig. 12; Campi et al., 2005, p. 132, pl. 5, figs. Q–S.
Streptorhynchus pseudopelargonatus Broili, 1916, p. 5, pl.
115, figs. 4, 5.
Streptorhynchus broili Grabau, 1931, p. 245, pl. 24, figs. 3, 4.
Streptorhynchus amygdalis Lee and Gu, 1976, p. 234, pl.
159, fig. 1.
Material.—Four specimens from localities KF27 andKF121: (1) internal mould of a conjoined shell, with externalmould of the dorsal valve, UHR13584; (2) internal mould of
a conjoined shell, UHR 12367; (3) external and internalmoulds of a ventral valve, NU-B2506; and (4) external andinternal moulds of a dorsal valve, NU-B2507.
Remarks.—One of the specimens (UHR12367) was
previously described by Nakamura (1972, p. 397, pl. 9, fig.
3) and figured by Minato et al. (1979, pl. 60, fig. 7) as
Streptorhynchus pelargonatus (von Schlotheim, 1816). This
species is characterized by its elongate, oval and small shell(length 15 mm, width 15 mm, height of ventral interarea 6mm, width of hinge line 11 mm in a conjoined shell
specimen, UHR12367), high interarea with narrow andconvex pseudodeltidium in ventral valve, and a pair of strong
socket plates in dorsal valve. Shells described by Frech(1911, p. 122, pl. 18, fig. 7) as Streptorhynchus pelargonatus(von Schlotheim, 1816), from Loping, Jiangxi Province,
eastern China, differs from the lectotype figured by Williams
and Brunton (2000, fig. 481.4) in much larger size. Chinese
specimens, described by Grabau (1931, p. 243, pl. 24, fig. 2)
and Shen and Shi (2007, p. 33, pl. 12, figs. 15–24) as
Streptorhynchus pelargonatus (von Schlotheim, 1816), fromthe Jisu Honguer Formation of Inner Mongolia, and from the
Lopingian of Guizhou and Sichuan, respectively, differ fromthe type specimen in transverse wider outline and in having
lower ventral interarea. Streptorhynchus sulculatum Grant(1976, p. 50, pl. 8, figs. 1–56), from the Ratburi Formation ofKo Muk, southern Thailand, differs from S. pelargonatus inhaving shallow sulcus in ventral valve.
Occurrence.—KY1 Unit in Kamiyasse (locality KF27)
and Imo (locality KF121).
Distribution.—Artinskian–Changhsingian: northeasternJapan (Kamiyasse–Imo in the South Kitakami Belt), UK(England), Germany, western Russia (Caucasus), northern
China (Inner Mongolia), northeastern China (Jilin),
central-southern China (Hubei), southwestern China(Guizhou), Malaysia, Indonesia (Timor) and Pakistan (Salt
Range).
Streptorhynchus cataclinus Zhang in Yang et al., 1962
(Fig. 58D)
Streptorhynchus cataclinus Zhang in Yang et al., 1962, p. 36,
pl. 10, figs. 7–9, text-fig. 17; Lee and Gu, 1976, p. 234, pl.

97

161, fig. 5.
Streptorhynchus pelargonatus (von Schlotheim). Tazawa,
1976, pl. 2, fig. 2.
Material.—Three specimens from locality KF20: (1)internal mould of a conjoined shell, with external mould of
the ventral valve, NU-B2508; (2) external and internalmoulds of ventral valve, NU-B2509; and (3) internal mouldof a ventral valve, NU-B2510.
Remarks.—One of the specimens from Kamiyasse waspreviously figured by Tazawa (1976, pl. 2, fig. 2) as
Streptorhynchus pelargonatus (von Schlotheim, 1816). But
the Kamiyasse specimens can be referred to Streptorhynchus
cataclinus Zhang (in Yang et al., 1962, p. 36, pl. 10, figs.
7–9, text-fig. 17), from the Maokouan of Qinghai,
northwestern China, in being medium size (length about 20
mm, width about 26 mm in the best-preserved specimen,
NU-B2508) and transverse outline and in having acomparatively low ventral interarea (height of ventral
interarea about 5 mm in the same specimen). The preceding
species, Streptorhynchus pelargonatus (von Schlotheim,
1816), differs from S. cataclinus in its smaller and longer
shell, with higher ventral interarea.
Occurrence.—KY1 Unit in Kamiyasse (locality KF20).
Distribution.—Wordian–Capitanian: northeastern Japan(Kamiyasse–Imo in the South KItakami Belt), northwestern
China (Qinghai) and northern China (Inner Mongolia).
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 Léveilé, 1835.
Rhipidomella magna Tazawa, 2016a(Fig. 59A–F)
Rhipidomella sp. Tazawa, 2002, fig. 10.7.
Rhipidomella magna Tazawa, 2016a, p. 85, fig. 6.
Material.—Eight specimens from localities KF8, KF94
and KF217: (1) two conjoined shells, NU-B1757 (holotype),
1758; (2) internal mould of a conjoined shell, with externalmould of the ventral valve, NU-B2570; (3) internal mould of
a conjoined shell, NU-B1759; (4) external cast and mould ofa ventral valve, NU-B1760; (5) external and internal moulds
of a ventral valve, NU-B1974; (6) external mould of a ventral
valve, NU-B1761; and (7) external cast of a dorsal valve,
NU-B1762.
Remarks.—These specimens were previously describedby Tazawa (2016a, p. 85, fig. 6) as Rhipidomella magna
Tazawa, 2016a. This species is characterized by its large size

98

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 63. A–D, Stenoscisma mutabilis (Tschernyschew); A, ventral (A1, A2), dorsal (A3), anterior (A4), posterior (A5) and lateral (A6) views of internalmould of conjoined shell, NU-B2574; B, ventral external latex cast (B1) and ventral (B2) and dorsal (B3) views of internal mould of conjoined shell,
NU-B2557; C, ventral (C1) and dorsal (C2) views of internal mould of conjoined shell, NU-B2573; D, ventral (D1) and dorsal (D2) views of internal mouldof conjoined shell, NU-B2560; E, F, Stenoscisma sokolskajae Koczyrkevicz; E, ventral (E1) and dorsal (E2) views of internal mould of conjoined shell,
NU-B2563; F, ventral (F1, F2), dorsal (F3), anterior (F4), posterior (F5) and lateral (F6) views of internal mould of conjoined shell, NU-B2564. Scale bars
are 1 cm.

(length 22 mm, width 25 mm in the holotype, NU-B1757;
length 24 mm, width 30 mm in the largest dorsal valve
specimen, NU-B1762) and slightly transverse outline.
Rhipidomella corallina (Waagen, 1884, p. 572, pl. 56, fig. 1),
from the Wargal Formation of the Salt Range, Pakistan,
resembles the Kitakami species in size and outline of shell,
but differs in having more strongly concave ventral valve.
The type species, Rhipidomella michelini (Léveilé, 1835),

from the Lower Carboniferous of Europe and Asia, is readily
distinguished from R. magna by its smaller size and morerounded posterior portion of the shell.
Occurrence.—KY1 Unit in Kamiyasse (localities KF8 andKF94) and Imo (locality KF217).
Distribution.—Wordian: northeastern Japan (Kamiyasse–Imoin the South Kitakami Belt).

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Superfamily ENTELETOIDEA Waagen, 1884
Family SCHIZOPHORIIDAE Schuchert and LeVene, 1929
Genus ACOSARINA Cooper and Grant, 1969
Type species.—Acosarina dorsisulcata Cooper and Grant,
1969.
Acosarina dorsisulcata Cooper and Grant, 1969
(Fig. 60C, D)
Acosarina dorsisulcata Cooper and Grant, 1969, p. 2, pl. 5,
figs. 19–23; Cooper and Grant, 1976b, p. 2621, pl. 667,
figs. 1–26; pl. 673, figs. 1–6.
Material.—Two specimens from locality KF217: (1)internal mould of a conjoined shell, NU-B2465; and (2)internal mould of a ventral valve, NU-B2466.
Description.—Shell small in size for genus, roundly
subquadrate in outline, wider than long, widest at about
midlength, slightly biconvex and having weakly sulcate
anterior commissure; length 10 mm, width 12 mm in thelarger specimen (NU-B2465). Internally, ventral valve having
a pair of short dental plates divergent anteriorly and a long
median septum attaining to midlength of valve. Dorsal valve
interior not well preserved.
Remarks.—The specimens from Imo can be referred to
Acosarina dorsisulcata Cooper and Grant (1969, p. 2, pl. 5,
figs. 19–23), from the Bone Spring Formation of Texas, inbeing small size and transversely subeliptical outline, and inhaving slightly sulcate anterior commissure and relatively
short median septum in ventral valve. Acosarina indica(Waagen, 1884, p. 568, pl. 56, figs. 7, 8, 14–16), from the
Amb and Wargal Formations of the Salt Range, Pakistan,differs from A, dorsisulcata in its longer outline. Acosarina
strophiria Xu and Grant (1994, p. 18, figs. 4, 5.1–17, 5.19,5.20), from the Changhsing Formation of Zhejiang, eastern
China, is also having slightly emarginate anterior
commissure, but distinguished from A. dorsisulcata in its
longer median septum extending near anterior margin ofventral valve.
Occurrence.—KY1 Unit in Imo (locality KF217).
Distribution.—Sakmarian–Wordian: northeastern Japan(Kamiyasse–Imo in the South KItakami Belt) and USA(Texas).
Acosarina rectimarginata Cooper and Grant, 1976b(Fig. 60A, B)
Acosarina rectimarginata Cooper and Grant, 1976b, p. 2624,
pl. 674, figs. 1–46; Shen et al., 2011, p. 564, fig. 6.1–6.18;
Tazawa in Tazawa et al., 2012, p. 63, fig. 4.9; Tazawa andOyagi, 2019, p. 34, fig. 3E, F.
Material.—Four specimens from localities KF7 andKF217: (1) external and internal moulds of a conjoined shell,

99

NU-B2467; (2) internal mould of a conjoined shell,
NU-B2468; and (3) internal moulds of two ventral valves,
NU-B2469, 2571.
Description.—Shell small in size for genus, transversely
subquadrate in outline, with greatest width at about
midlength, dorsibiconvex and reticulate anterior commissure;length 9 mm, width 11 mm in the largest specimen(NU-B2571); length 7 mm, width 10 mm in thebest-preserved specimen (NU-B2467). Ventral valve gently
convex, flattened anteriorly. Dorsal valve moderately andunevenly convex in lateral profile, most convex at posterior
one-third of valve; no sulcus. External surface of both valvesornamented with numerous fine costellae, numbering 4 in 1mm at anterior margin of dorsal valve and one or two
irregular rugae near anterior margin. Ventral valve interior
with a pair of short dental plates slightly diverging anteriorly
and a long median septum extending at about three-quarters
valve length. Interior of dorsal valve with two strongly
divergent brachiophore plates and a large muscle scar.
Remarks.—These specimens are referred to Acosarina
rectimarginata Cooper and Grant (1976b, p. 2624, pl. 674,
figs. 1–46), from the Neal Ranch Formation of the Glass
Mountains, Texas, in size and outline of shell, particularly inhaving reticulate anterior commissure. Acosarina kanmerai
Yanagida and Nakornsri (1999, p. 111, pl. 26, figs. 1–7),
from the Middle–lower Upper Permian of Khao Hin Kling,north-central Thailand, resembles A rectimarginata in its
small size and transversely subquadrate outline, but differsfrom the latter in having slightly sulcate anterior commissure.
Acosarina dunbari Cooper and Grant (1976b, p. 2622, pl. 67,
figs. 1–8), from the lower Wolfcampian of Nebraska in theUSA, is also having reticulate to faintly sulcate anterior
commissure, but differs from A. rectimarginata in its rounder
outline and in having coarser costellae and numerous
concentric rugae on both valves.
Occurrence.—KY1 Unit in Kamiyasse (locality KF7) andImo (locality KF217).
Distribution.—Asselian–Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt), central Japan(Hatahoko, Kuzu and Ryozensan in the Mino Belt) and USA(Texas).
Genus ORTHOTICHIA Hall and Clarke, 1892
Type species.—Orthis? morganiana Derby, 1874.
Orthotichia sp.
(Fig. 60E, F)
Material.—Two specimens from localities KF6 and
KF121, internal moulds of two ventral valves, NU-B2497,
UHR12624.
Remarks.—These specimens can be assigned to the genus
Orthotichia by medium-sized, slightly convex andtransversely subelliptical shell (length 29 mm, width 35 mm

100

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 64. A, Rhynchopora tchernyshae Koczyrkevicz, ventral (A1, A2), dorsal (A3), anterior (A4), posterior (A5) and lateral (A6) views of internalmould of conjoined shell, NU-B2588: B–G, Permocryptospirifer omeishanensis (Huang); B, internal mould of ventral valve, NU-B2514; C, internalmould of ventral valve, NU-B2513; D, ventral valve, NU-B2014; E, ventral valve, NU-B2013; F, abraded ventral valve, NU-B2015; G, internal mould of
dorsal valve, NU-B2515. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

in the larger specimen, NU-B2497), and in having a pair ofstrong subparallel dental plates and a long median septum inventral valve. But the specific identification is difficult for
the ill-preserved specimens.
Occurrence.—KY1 Unit in Kamiyasse (locality KF6) andImo (locality KF121).
Order RHYNCHONELLIDA Kuhn, 1949
Superfamily STENOSCISMATOIDEA Oehlert, 1887
Family STENOSCISMATIDAE Oehlert, 1887
Subfamily STENOSCISMATINAE Oehlert, 1887
Genus STENOSCISMA Conrad, 1839
Type species.—Terebratula schlottheimii von Buch, 1834.
Stenoscisma margaritovi (Tschernyschew, 1888)(Figs. 61A–D, 62A–D)
Camarophoria margaritovi Tschernyschew, 1888, p. 355, pl.
1, figs. 1–3; Fredericks, 1924b, p. 48, pl. 1, figs. 32–42,
text-fig. 4.
Camarophoria humbletonensis Howse. Hayasaka, 1922a, p.
62, pl. 9, figs. 10–12; pl. 10, fig. 9; Hayasaka, 1966, p.
1226, text-figs. 6‒8.
Stenoscisma humbletonensis (Howse). Tazawa, 1976, pl. 2,
figs. 9, 10; Minato et al., 1979, pl. 66, figs. 6–8.
Stenoscisma gigantea (Diener). Lee and Gu, 1976, p. 272, pl.
176, fig. 3; pl. 177, fig. 18; Lee et al., 1980, p. 395, pl.
173, figs. 6, 8.
Stenoscisma margaritovi (Tschernyschew). Licharew and
Kotlyar, 1978, pl. 17, fig. 7; Koczyrkevicz, 1979b, p. 50,
pl. 11, figs. 5, 6; Duan and Li, 1985, p. 120, pl. 43, figs.
5–8; Tazawa and Matsumoto, 1998, p. 9, pl. 2, figs. 1–5;
Tazawa et al., 2000, p. 10, pl. 1, figs. 7‒11; Tazawa, 2001,
p. 298, fig. 8.1–8.4; Tazawa, 2002, fig. 10.5; Wang andZhang, 2003, p. 130, pl. 33, figs. 6, 7, 12–16; pl. 50, fig.
19; Tazawa and Chen, 2006, p. 333, fig. 5.7, 5.8; Tazawa
and Araki, 2018, p. 19, fig. 6.3.
Stenoscisma purdoni (non Davidson). Lee et al., 1980, p.
395, pl. 173, figs. 4, 5, 7.
Material.—Thirteen specimens from localities AR4,
KF46, KF86, KF89, KF94, KF217 and KF218: (1) external
and internal moulds of two conjoined shells, NU-B2554,
2555; (2) internal moulds of four conjoined shells, KCG56,
NU-B2445, 2446, 2556; (3) external and internal moulds of aventral valve, NU-B2547; (4) external casts of two ventral
valves, NU-B2548, 2549; (5) external mould of a ventral
valve, NU-B2550; (6) internal mould of a ventral valve,
NU-B2551; and (7) internal moulds of two dorsal valves,
NU-B2552, 2553.
Remarks.—Part of the specimens from Kamiyasse, Imo
and Matsukawa were previously described by Hayasaka(1922a, p. 62, pl. 9, figs. 10‒12; pl. 10, fig. 9) as
Camarophoria humbletonensis (Howse, 1848) and figured by

101

Tazawa (1976, pl. 2, figs. 9, 10) and Minato et al. (1979, pl.
66, figs. 6‒8) as Stenoscisma humbletonensis (Howse, 1848).
However, the Kitakami specimens are referred to
Stenoscisma margaritovi (Tschernyschew, 1888), originallydescribed by Tschernyschew (1888, p. 355, pl. 1, figs. 1–3)
as Camarophoria margaritovi Tschernyschew, 1888, from
the Middle Permian of Vladivostok, eastern Russia, in having
large elongate shell (length about 34 mm, width about 28 mm
in the largest specimen, NU-B2551) with shallow ventral
sulcus and ornamented with numerous costae over the valve.
Stenoscisma humbletonensis (Howse, 1848) differs from S.
margaritovi in its transverse outline and in having a deeper
sulcus on ventral valve. Stenoscisma gigantea (Diener,
1897a, p. 72, pl. 12, figs. 5, 7, 10), from the
Capitanian–Wuchiapingian of the Chitichun No. 1, Himalaya,is also large in size, but differs from S. margaritovii in its
transverse outline.
Occurrence.—KY1 Unit in Kamiyasse (localities KF46,
KF86, KF89 and KF94), Imo (localities KF217 and KF218)
and Matsukawa (locality AR4).
Distribution.—Wordian–Wuchiapingian: northeasternJapan (Kamiyasse–Imo, Matsukawa, Ogatsu andTakakurayama in the South Kitakami Belt), central Japan(Moribu and Oguradani in the Hida Gaien Belt); northern
China (Inner Mongolia), northeastern China (Heilongjiang
and Jilin) and eastern Russia (South Primorye).
Stenoscisma mutabilis (Tschernyschew, 1902)(Fig. 63A–D)
Camarophoria mutabilis Tschernyschew, 1902, p. 81, 491,
pl. 22, fig. 18; pl. 23, fig. 10; pl. 45, figs. 1–15; pl. 46, fig.
14, text-fig. 18; Grabau, 1931, p. 211, pl. 4, fig.7; pl. 5,
figs. 1, 2; Grabau, 1934, p. 16, pl. 2, figs. 1, 2; Stepanov,1937 b, p. 5, pl. 6, figs. 9, 10, text-fig. 1.
Stenoscisma mutabilis (Tschernyschew). Zhao, 1965, p. 429,
pl. 2, fig. 9; Mironova, 1967, p. 38, pl. 3, fig. 6;
Kalashnikov, 1980, p. 71, pl. 20, figs. 9 –11; Lee and Gu,
1980, p. 486, pl. 1, fig. 5 only: Kalashnikov, 1986, pl. 112,
fig. 9; Gu, 1992, p. 243, pl. 74, figs. 11, 12; He et al.,
1995, pl. 70, figs. 8–11; Shi and Waterhouse, 1996, p. 114,
pl. 21, figs. 4–14; Shen and Shi, 2007, p. 59, pl. 23, figs.
25–35; pl. 24, figs. 1–31, text-fig. 20.
Material.—Eight specimens from localities KF8, KF33,
KF121, KF217 and KF218: (1) external and internal moulds
of a conjoined shell, NU-B2573; (2) internal moulds of three
conjoined shells, with external moulds of the ventral valves,
NU-B2557, 2558, 2574; (3) internal moulds of two conjoined
shells, NU-B2559, 2560; and (4) external and internal mouldsof two ventral valves, NU-B2561, 2562.
Remarks.—The present specimens from Kamiyasse–Imo
are small-sized and transverse-shaped Stenoscisma (length
about 16 mm, width about 22 mm in the largest specimen,
NU-B2560), with relatively small spondylium in ventral

102

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 65. A–D, Hustedia remota (von Eichwald); A, internal mould (A1 , A 2 ) of ventral valve, NU-B2016; B, internal mould of ventral valve,
NU-B2017; C, internal mould of ventral valve, NU-B2018; D, internal mould of dorsal valve, NU-B2019; E, F, Hustedia ratburiensis Waterhouse andPiyasin; E, external latex cast (E1, E2) of ventral valve, NU-B2512; F, ventral (F1, F2), dorsal (F3), anterior (F4), posterior (F5) and lateral (F6) views ofinternal mould of conjoined shell, NU-B2511; G, H, Martinia semiplana Waagen; G, internal mould of ventral valve, NU-B2585; H, ventral (H1, H2) anddorsal (H3) views of internal mould of conjoined shell, NU-B2583; I, Neospirifer moosakhailensis (Davidson), ventral (I1, I2) and dorsal (I3) views ofinternal mould of conjoined shell, KCG3. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

valve and small camarophorium in dorsal valve, and external
ornament consisting of simple costae on both valves (4–5 insulcus and 5 in each flank of the ventral valve). These
specimens can be referred to Stenoscisma mutabilis(Tschernyschew, 1902), from the Schwagerina Horizon of
the Urals and Timan, in being medium size and transverse
outline, and in having moderately developed ventral sulcus
and dorsal fold. Stenoscisma purdoni (Davidson, 1861),
redescribed by Waagen (1883, p., p. 90, pl. 2, fig. 4) from theWargal Formation of the Salt Range, Pakistan, differs from S.
mutabilis in its less transverse and nearly triangular outline.
The preceding species, Stenoscisma margaritovi(Tschernyschew, 1888), is readily distinguished from S.
mutabilis by its much larger size and longer outline.
Occurrence.—KY1 Unit in Kamiyasse (localities KF8 andKF33) and Imo (localities KF121, KF217 and KF218).
Distribution.—Kasimovian–Changhsingian: northeasternJapan (Kamiyasse–Imo in the South Kitakami Belt), northern
Canada (northern Yukon Territory), northern Russia (Timan,
Vaygach Island and northern Urals), central Russia (southern
Urals), northern China (Inner Mongolia and Shanxi),
northeastern China (Jilin), central-southern China (Hunan)and southwestern China (Guizhou and Sichan).
Stenoscisma sokolskajae Koczyrkevicz, 1979b(Fig. 63E, F)
Stenoscisma tenuistriata (Hamlet). Licharew and Kotlyar,
1978, pl. 21, figs. 11, 12.
Stenoscisma sokolskajae Koczyrkevicz, 1979b, p. 53, pl. 12,
figs. 7–9.
Material.—Three specimens from locality KF217: (1)internal mould of a conjoined shell, with external mould of
the ventral valve, NU-B2563; and (2) internal moulds of two
conjoined shells, NU-B2564, 2565.
Remarks.—These specimens can be referred to
Stenoscisma sokolskajae Koczyrkevicz (1979b, p. 53, pl. 12,
figs. 7–9), from the middle part of the Chandalaz Formation(Parafusulina stricta Zone) of South Primorye, eastern
Russia, in being small size and longer outline (length about
20 mm, width about 15 mm in the largest specimen,
NU-B2563), and in having external ornament consisting of
simple costae numbering 4–5 in sulcus and 4 in each flank of
the ventral valve. Stenoscisma tenuistriata Hamlet (1928, p.
61, pl. 10, figs. 1, 2), from the Wordian–Wuchiapingian ofTimor, differs from S. sokolskajae in its smaller size and lesselongate outline and in having more numerous fine costae onboth valves. The preceding species Stenoscisma mutabilis(Tschernyschew, 1902) is readily distinguished from thepresent species in its larger size and transverse outline.
Occurrence.—KY1 Unit in Imo (locality KF217).
Distribution.—Wordian–Capitanian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt) and eastern
Russia (South Primorye).

103

Superfamily RHYNCHOPOROIDEA Muir-Wood, 1955
Family RHYNCHOPORIDAE Muir-Wood, 1955
Subfamily RHYNCHOPORINAE Muir-Wood, 1955
Genus RHYNCHOPORA King, 1865
Type species.—Terebratula geinitziana de Verneuil, 1845.
Rhynchopora tchernyshae Koczyrkevicz, 1979a(Fig. 64A)
Rhynchopora tchernyshae Koczyrkevicz, 1979a, p. 47, pl. 11,
figs. 1–4, text-fig. 4; Tazawa et al., 2000, p. 10, pl. 1, fig. 6.
Material.—One specimen from locality KF217, internalmould of a conjoined shell, NU-B2588.
Description.—Shell small in size for genus, bluntly
subpentagonal in outline, with greatest width at about
midlength; length 9 mm, width 9 mm in the sole specimen.
Ventral valve slightly concave, nearly flat in lateral profile;umbo small; sulcus shallow and broad. Dorsal valve gently
convex in lateral profile, with low and broad fold. Externalsurface of both valves ornamented with numerous simple
costae; 4 in ventral sulcus, 4–5 on each lateral flank ofventral valve; 5 on dorsal fold, 5 on each flank of dorsal
valve. Internally, ventral valve with rather long dental plates
although they are not well preserved; dorsal valve with ashort but high and sharp median septum extending to about
one sixth of valve length.
Remarks.—This specimen can be referred to Rhynchopora
tchernyshae Koczyrkevicz (1979a, p. 47, pl. 11, figs. 1–4,
text-fig. 4), from the lower Barabash Formation(Monodiexodina sutschanica Zone) of Barabash, South
Primorye, eastern Russia, by its size, outline and number of
costae on both valves, particularly in its nearly flattened
shell. Rhynchopora tenuicostata Koczyrkevicz (1979a, p. 42,
pl. 10, figs. 1–9, text-fig. 1) from the Chandalaz andBalabash Formations of South Primorye, is also a flattened( w e a k l y b i c o n v e x ) s h e l l s p e c i e s , b u t d i f f e r s f r o m R.
tchernyshae in having more numerous finer costae on both
valves.
Occurrence.—KY1 Unit in Imo (locality KF217).
Distribution.—Wordian–Capitanian: northeastern Japan(Kamiyasse–Imo and Ogatsu in the South Kitakami Belt) andeastern Russia (South Primorye).
Order ATHYRIDIDA Boucot, Johnson and Staton, 1964
Suborder ATHYRIDIDINA Boucot, Johnson and Staton,
1964
Superfamily ATHYRYDOIDEA Davidson, 1881
Family ATHYRIDIDAE Davidson, 1881
Subfamily LOCHENGINAE Ching (Jin) and Yang in Yang etal., 1977
Genus PERMOCRYPTOSPIRIFER Shen and Grunt in Shen etal., 2017

104

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 66. A–E, Martinia triquetra Gemmellaro; A, ventral (A1) and posterior (A2) views of internal mould of ventral valve, NU-B2577; B, internalmould of dorsal valve, NU-B2578; C, internal mould of dorsal valve, NU-B2579: D, internal mould of ventral valve, NU-B2575; E, internal mould ofventral valve, NU-B2012; F, G, Gypospirifer kobiyamai Tazawa and Araki; F, external latex cast of ventral valve, KCG2; G, dorsal external latex cast(G1) and ventral (G2) and dorsal (G3) views of internal mould of conjoined shell, KCG1 (holotype). Scale bars are 1 cm.

Type species.—Cryptospirifer omeishanensis Huang, 1933.
Permocryptospirifer omeishanensis (Huang, 1933)(Fig. 64B–G)
Cryptospirifer omeishanensis Huang, 1933, p. 44, pl. 6, fig.
4; pl. 8, fig. 1; Wang et al., 1964, p. 512, pl. 95, figs. 4, 7;
Jin et al., 1974, p. 310, pl. 163, fig. 17; Yang et al., 1977,
p. 413, pl. 163, fig. 5; Tong, 1978, p. 253, pl. 89, fig. 1;
Wang, 1984, p. 220, pl. 88, fig. 12; Zeng, 1992, pl. 2, figs.

1–4; Shi and Shen, 2001, p. 250, pl. 3, figs. 1–7, text-figs.
4–6; Jin and Zhan, 2008, p. 9, figs. 8, 9E; Tazawa, 2016b,
p. 32, fig. 11.2–11.4.
Cryptospirifer sp. Minato et al., 1979, pl. 67, fig. 9.
Cryptospirifer sp. Hu, 1983, pl. 1, figs. 1–3.
Cryptospirifer minor Yang, 1984, p. 233, pl. 38, fig. 1.
“Cryptospirifer” omeishanensis Huang. Shen et al., 2016, p.
531, fig. 7A, B.
Permocryptospirifer omeishanensis (Huang). Shen and Gruntin Shen et al., 2017, p. 766, pl. 27, figs. 2–5, text-figs. 29,
30.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Material.—Seven specimens from localities KF217 andKF218: (1) an abraded conjoined shell, NU-B2518; (2) threeabraded ventral valves, NU-B2013–2015; (3) internal mouldsof two ventral valves, NU-B2513, 2514; and (4) internalmould of a dorsal valve, NU-B2515.
Remarks.—Some of the specimens from Imo in the
Kamiyasse–Imo area (NU-B2013–2015) were previously
described by Tazawa (2016b, p. 32, fig. 11.2–11.4) as
Cryptospirifer omeishanensis Huang, 1933. These specimens
are represented by imperfect abraded ventral valves, but they
can be referred to Cryptospirifer omeishanensis Huang, 1933,
from the Maokou Formation of Omeishan, Sichuan Province,
southwestern China, on account of large size (length about 66
mm, width more than 100 mm in the largest specimen,
NU-B2015), transversely elliptical outline, and gently
convex, smooth ventral valves. Newly added specimens from
Imo (NU-B2013–2015), represented by internal moulds ofventral or dorsal valves, are also referred to Cryptospirifer
omeishanensis in large, transversely elliptical shell (length
about 63 mm, width about 73 mm in the largest specimen,
NU-B2513) and in having long and parallel crural bases in
dorsal valve. The generic name Cryptospirifer was recently
replaced by Shen et al. (2017, p. 766) with
Permocryptospirifer Shen and Grunt in Shen et al., 2017.
Occurrence.—KY1 Unit in Imo (localities KF217 andKF218).
Distribution.—Roadian–Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt), eastern China(Jiangxi), central-southern China (Hubei), southwestern
China (Sichuan and Yunnan) and northwestern China (Tibet).
Suborder RETZIIDINA Boucot, Johnson and Staton, 1964
Superfamily RETZIOIDEA Waagen, 1883
Family NEORETZIIDAE Dagys, 1972
Subfamily HUSTEDIINAE Grunt, 1986
Genus HUSTEDIA Hall and Clarke, 1893
Type species.—Terebratula mormoni Marcou, 1858.
Hustedia remota (von Eichwald, 1860)(Fig. 65A–D)
Rhynchonella remota von Eichwald, 1860, p. 768, pl. 35, fig. 10.
Hustedia remota (Eichwald). Tschernyschew, 1902, p. 107,
pl. 47, figs. 8–11; Huang, 1933, p. 79, pl. 11, figs. 4, 5;
Licharew, 1939, p. 111, pl. 28, fig.6; Campi et al., 2002,
fig. 7A–C; Shen et al., 2003a, p. 250, pl. 5, figs. 22, 23;
Shen and Zhang, 2008, fig. 6.16–6.21; He et al., 2008, p.
819, fig. 5.15–5.18.
Retzia (Hustedia) radialis var. grandicosta Davidson. Broili,
1916, p. 11, pl. 124, figs. 14–22; pl. 125, figs. 1–3.
Material.—Four specimens from localities KF22 andKF218: (1) internal moulds of three ventral valves,

105

NU-B2016–2018; and (2) internal mould of a dorsal valve,
NU-B2019.
Remarks.—These specimens can be referred to Hustedia
remota (von Eichwald, 1860), redescribed by Tschernyschew(1902, p. 107, pl. 47, figs. 8–11) from the Schwagerina
Limestone of Ufa, southern Urals, central Russia, in having
relatively few strong costae (eight in ventral valve, nine indorsal valve), with wide and flat-bottomed intercostal
troughs. Huang (1933, p. 79) treated Retzia radialis var,
grandicosta Davidson, 1862 [=Hustedia grandicosta(Davidson, 1862)], from the Salt Range, Pakistan, as asynonym of Hustedia remota (von Eichwald). The Pakistanispecies is, however, distinguished from H. remota in having
more numerous costae on both ventral and dorsal valves, andthe intercostal troughs are narrower and having roundbottom. The type species, Hustedia mormoni (Marcou, 1858),
refigured by Alvarez and Rong (2002, fig. 1087.1), from thePennsylvanian of Nebraska in the USA, differs from H.
remota in having more numerous costae on both valves.
Occurrence.—KY1 Unit in Kamiyasse (locality KF22)
and Imo (locality KF218).
Distribution.—Asselian–Changhsingan: northeasternJapan (Kamiyasse–Imo in the South Kitakami Belt), central
Russia (southern Urals), northwestern China (Qinghai),central-southern China (Hunan), southwestern China(Guizhou), Malaysia, Indonesia (Timor) and northwestern
China (Tibet),
Hustedia ratburiensis Waterhouse and Piyasin, 1970
(Fig. 65E, F)
Hustedia ratburiensis Waterhouse and Piyasin, 1970, p. 138,
pl. 23, figs. 15–30; Grant, 1976, p. 241, pl. 66, figs. 1–69;
pl. 67, figs. 51–58; Sun, 1991, p. 254, pl. 6, figs. 5–8;
Yanagida and Nakornsri, 1999, p. 118, pl. 32, figs. 11–16;
Archbold, 1999, fig. 5E–H; Tazawa, 2001, p. 299, fig. 8.6;
Tazawa, 2008b, p. 53, fig. 8.2–8.6; Tazawa in Tazawa etal., 2015, p. 44, fig. 6.7; Tazawa and Nakamura, 2015, p.
169, fig. 7.1–7.7; Tazawa and Shintani, 2021, p. 13, fig.
5C, D.
Hustedia nakornsrii Yanagida, 1970, p. 79, pl. 14, fig. 9.
Hustedia indica (non Waagen). Koizumi, 1979, pl. 1, fig. 5.
Material.—Two specimens from locality KF218: (1)internal mould of a conjoined shell, NU-B2511; and (2)external mould of a dorsal valve, NU-B2512.
Remarks.—These specimens can be referred to Hustedia
ratburiensis Waterhouse and Piyasin (1970), originallydescribed from the Middle Permian (Kazanian) limestones of
Khao Phrik, southern Thailand, in being small to medium
size (length about 8 mm, width about 5 mm in the smaller
specimen, NU-B2511; length 9 mm, width 10 mm in thelarger specimen, NU-B2512) and in having rotund dorsal
valve, and surface ornament of both ventral and dorsal valvesconsisting of subangular costae (8 in ventral valve, and 9 in

106

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 67. A–F, Alispiriferella lita (Fredericks); A, ventral (A1) and dorsal (A2) views of internal mould of conjoined shell, KCG20; B, ventral (B1) anddorsal (B2) views of internal mould of conjoined shell, KCG21; C, ventral (C1) and dorsal (C2) views of external latex cast of conjoined shell, IGPS96219;
D, external latex cast (D1, D2) of dorsal valve, IGPS96220; E, internal mould of ventral valve, NU-B326; F, external latex cast (F1, F2) and internal mould(F3) of ventral valve, NU-B315. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

dorsal valve), of which the middle one of the dorsal valve isslightly depressed. Hustedia ratburiensis is clearly
distinguished from the preceding species, Hustedia remota(von Eichwald, 1860), by its rotund outline and subangularcostae with narrow intercostal troughs.
Occurrence.—KY1 Unit in Imo (locality KF218).
Distribution.—Artinskian–Wuchiapingian: northeasternJapan (Nagaiwa–Sakamotozawa, Kamiyasse–Imo and
Takakurayama in the South Kitakami Belt), central Japan(Moribu in the Hida Gaien Belt), southwestern Japan(Mizukoshi in Kyushu Island), north-central Thailand (KhaoNong Ta On and Khao Hin King), southern Thailand (KhaoPhrik, Khao Tok Nam and Ko Muk) and northwestern China(Tibet).
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 semiplana Waagen, 1883
(Fig. 65G, H)
Martinia semiplana Waagen, 1883, p. 536, pl. 43, fig. 4;
Hayasaka, 1922c, p. 130, pl. 6, fig. 18; Grabau, 1936, p.
237, pl. 24, figs. 1–4; Hayasaka and Minato, 1956, p. 146,
pl. 23, fig. 2; Zhang and Ching (Jin), 1961, p. 406, pl. 2,
figs. 16–19; He and Fan, 1990, pl. figs. 19–23; Zeng, 1990,
p. 228, pl. 7, figs. 6, 7; pl. 8, fig. 11; Shen and Zhang,2008, fig. 6.6, 6.7; Tazawa, 2008a, p. 37, fig. 5.3, 5.4;
Martinia semiplana var. lata Grabau, 1936, p. 239, pl. 21,
figs. 1–3; Hayasaka and Minato, 1956, p. 146, pl. 23, fig. 3.
Martinia lata Grabau. Tazawa, 2008a, p. 38, fig. 5.7–5.14;
Tazawa and Shintani, 2021, p. 14, fig. 5E–G.
Material.—Four specimens from locality KF212: (1)internal mould of a conjoined shell, with external mould of
the ventral valve, NU-B2583; and (2) internal moulds ofthree ventral valves, NU-B2584–2586.
Remarks.—These specimens can be referred to Martinia
semiplana Waagen, 1883, originally described by Waagen(1883, p. 536, pl. 43, fig. 4) from the Middle Productus
Limestone (Wargal Formation) of the Salt Range, Pakistan,by medium-sized (length about 20 mm, width about 25 mm
in the largest specimen, NU-B2585), transversely subelipticaland gently biconvex shell with radial vascular scars on
ventral valve. The shells, described by Tschernyschew (1902,
p. 182, 565, pl. 60, figs. 15–17) as Martinia semiplana
Waagen, 1883 from the Schwagerina Limestone of the Urals,
are deemed to be a different species in its less-transverse
outline and much stronger convex ventral valve. Martinia

107

nealranchensis Cooper and Grant (1976a, p. 2270, pl. 649,
figs. 1–14), from the Neal Ranch Formation of Texas,
resembles M. semiplana, but differs in its subpentagonal
outline and in having shorter hinge.
Occurrence.—Upper part of the Hosoo Formation in
Kamiyasse (locality KF212).
Distribution.—Kasimovian–Wuchiapingian: northeasternJapan (Nagaiwa–Sakamotozawa, Kamiyasse–Imo and
Takakurayama in the South Kitakami Belt), northern China(Inner Mongolia and Shanxi), northeastern China (Liaoning),
eastern China (Anhui), central-southern China (Hunan andGuangxi), southwestern China (Guizhou) and Pakistan (Salt
Range).
Martinia triquetra Gemmellaro, 1899
(Fig. 66A–E)
Martinia triquetra Gemmellaro, 1899, p. 17, pl. 32, figs.
23–28; Tschernyschew, 1902, p. 178, 562, pl. 16, figs.
1–6; Heritsche, 1935, p. 363, pl. 1, figs. 1, 10; Grunt,
1973, p. 147, pl. 12, figs. 3, 4; pl. 16, fig. 2, text-fig. 48;
Zhang et al., 1983, p. 371, pl. 143, fig. 1; Zhan and Wu,
1987, p. 228, pl. 62, figs. 1–7; Wang, 1995, pl. 4, fig. 9;
Wang and Yang, 1998, p. 130, pl. 22, figs. 16–19, 23.
Martinia spp. (Martinia sp. A, Martinia sp. B and Martinia
sp. C) Hayasaka, 1960, p. 52, pl. 3, figs. 2–4.
Martinia cf. triquetra Gemmellaro. Yang et al., 1962, p. 114,
pl. 43, fig. 4.
Martinia sp. Minato et al., 1979, pl. 67, fig. 8.
Martinia sp. Tazawa and Araki, 2017, p. 279, fig. 8.7.
Material.—Nine specimens from localities AR4, KF121
and KF218: (1) internal moulds of four ventral valves,
NU-B2012, 2575–2577; and (2) internal moulds of fivedorsal valves, NU-B2578–2582.
Remarks.—One of the specimens (NU-B2012) from thelower Kamiyasse Formation of Kamiyasse–Imo andMatsukawa was previously described by Tazawa and Araki(2017, p. 279, fig. 8.7) as Martinia sp. However, these
specimens can be referred to Martinia triquetra Gemmellaro,
1899, redescribed by Tschernyschew (1902, p. 178, 562, pl.
16, figs. 1–6) from the Schwagerina Limestone of the Urals,
in being large size (length about 30 mm, width about 46 mm
in the largest specimen, NU-B2577), rounded subrectangularoutline and in having shallow ventral sulcus. Martinia
uralica Tschernyschew (1902, p. 183, 566, pl. 18, figs. 1–4)
from the Schwagerina Limestone of the Urals and Timan, isalso large-sized Martinia species, but differs from M.
triquetra in its longer outline. Martinia karawanica Volgin(1959, p. 121, pl. 6, figs. 8–10), from the Upper
Carboniferous of Fergana, differs from M. triquetra in having
a conspicuous but very narrow ventral sulcus. Martinia
miranda Cooper and Grant (1976a, p. 2268, pl. 643, figs.
1–39), from the Cathedral Mountains Formation of Texas, isdistinguished from the present species in its longer

108

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 68. A–E, Alispiriferella lita (Fredericks); A, external latex cast of dorsal valve, NU-B321; B, internal mould (B1) and internal latex cast (B2) of
dorsal valve, NU-B332; C, external latex cast of ventral valve, NU-B318; D, ventral (D1) and dorsal (D2) views of internal mould of conjoined shell,
NU-B319; E, external latex cast of ventral valve, NU-B317; F, Licharewina arakii (Hayasaka), dorsal external latex cast (F1, F2) and ventral (F3) anddorsal (F4) views of internal mould of conjoined shell, KCG4. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

pentagonal outline. The preceding species, Martinia
semiplana Waagen, 1883, is readily distinguished from thepresent species by its much smaller size and transversely
subelliptical outline.
Occurrence.—KY1 Unit in Imo (localities KF121 andKF218) and Matsukawa (locality AR4).
Distribution.—Kasimovian–Wordian: northeastern Japan(Kamiyasse–Imo and Matsukawa in the South KitakamiBelt), Austria (Carnian Alps), central Russia (southern
Urals), northwestern China (Xinjiang and Qinghai), Italy(Sicily) and Tajikistan (Pamir).
Superfamily SPIRIFEROIDEA King, 1846
Family TRIGONOTRETIDAE Schuchert, 1893
Subfamily NEOSPIRIFERINAE Waterhouse, 1968a
Genus NEOSPIRIFER Fredericks 1924aType species.—Spirifer fasciger Keyserling, 1846.
Neospirifer moosakhailensis (Davidson, 1862)(Fig. 65I)
Spirifer moosakhailensis Davidson, 1862, p. 28, pl. 2, fig. 2;
Beyrich, 1865, p. 77, pl. 1, fig. 7; Grabau, 1931, p. 168, pl.
23, figs. 5–7 only; Noda, 1951, p. 46, pl. 1, figs. 1–4.
Spirifer musakheylensis Davidson. Waagen, 1883, p. 512, pl.
45, figs. 1–6; Diener, 1897b, p. 35, pl. 3, figs. 3, 4; pl. 4,
figs. 1, 2; pl. 5, fig. 1; Diener, 1899, p. 63, pl. 5, figs. 3–7.
Spirifer fasciger Keyserling. Broili, 1916, p. 37, pl. 120, figs.
10–15; pl. 121, figs. 1–3.
Neospirifer fasciger ambiensis Waagen. Fredericks, 1925, p.
28, pl. 4, fig. 113.
Spirifer fasciger var. musakheylensis Davidson. Reed, 1925,
p. 42, pl. 6, fig. 2.
Neospirifer moosakhailensis (Davidson). Muir-Wood andOakley, 1941, p. 30, p. 2, figs. 12, 13; Lee and Gu, 1976,
p. 286, pl. 177, fig. 13 only; Lee et al., 1980, p. 412, pl.
177, figs. 7, 10; Tazawa, 2014, p. 19, fig. 3.8.
Spirifer cf. moosakhailensis Davidson. Noda, 1956, p. 17, pl.
6, fig. 10 only.
Neospirifer fasciger (Keyserling). Hayasaka, 1960, p. 42, pl.
1, figs. 1–7 only; Koizumi, 1979, pl. 1, fig. 17 only.
Material.—One specimen from locality AR1, internalmould of a conjoined shell, KCG3.
Remarks.—This specimen was previously described by
Tazawa (2014, p. 19, fig. 3.8) as Neospirifer moosakhailensis(Davidson, 1862). The Kamiyasse specimen is poorly
preserved, but can be referred to Neospirifer moosakhailensis(Davidson, 1862, p. 28, pl. 2, fig. 2), from the middle andupper Permian of the Salt Range, Pakistan, in its large,transverse shell (length about 40 mm, width about 70 mm),with a deep ventral sulcus and a high dorsal fold andnumerous fasciculate costae on both ventral and dorsal
valves. Shells, described by Hayasaka (1960, p. 42) as

109

Neospirifer fasciger (Keyserling, 1846) from the lower
Kamiyasse Formation of Tagara and Takayashiki in the
Kesennuma area, South Kitakami Belt, include two different
species: Neospirifer moosakhailensis (Davidson, 1862), atransverse spiriferid having distinct fasciculate costae(Hayasaka, 1960, pl. 1, figs. 1–7) and Gypospirifer
kobiyamai Tazawa and Araki, 2013, a broader and larger
spiriferid with less fasciculate costae (Hayasaka, 1960, pl. 2,
figs. 1, 2). Neospirifer fasciger (Keyserling, 1846, p. 231, pl.
8, fig. 3), from the lower Permian of Timan, northern Russia,
is readily distinguished from N. moosakhailensis by its muchsmaller size.
Occurrence.—KY1 Unit in Kamiyasse (locality AR1).
Distribution.—Wordian–Changhsingian: northeasternJapan (Kamiyasse–Imo in the South Kitakami Belt), northern
China (Inner Mongolia), northeastern China (Heilongjiang
and Jilin), eastern Russia (South Primorye), Indonesia(Timor), Pakistan (Salt Range) and northern India (Sikkim,
Garhwal, Spiti and Kashmir).
Genus GYPOSPIRIFER Cooper and Grant, 1976aType species.—Gypospirifer nelsoni Cooper and Grant,
1976a.
Gypospirifer kobiyamai Tazawa and Araki, 2013
(Fig. 66F, G)
Spirifer fasciger var. simplex Grabau. Kobiyama, 1956, fig. 4.
Neospirifer fasciger (Keyserling). Hayasaka, 1960, p. 42, pl.
2, figs. 1, 2 only; Yanagida, 1963, p. 71, pl. 8, figs. 3, 6;
pl. 9, fig. 3 only; Koizumi, 1979, pl. 1, fig 16 only.
Neospirifer aff. cameratus Morton. Yanagisawa, 1967, p. 91,
pl. 2, fig. 11.
Neospirifer striato-paradoxus (Toura). Licharew and
Kotlyar, 1978, pl. 18, fig. 1; Lee et al., 1980, p. 412, pl.
177, figs. 3, 6, 9.
Gypospirifer kobiyamai Tazawa and Araki, 2013, p. 7, fig.
2.3, 2.4.
Material.—Two specimens from locality AR1: (1) internalmould of a conjoined shell, with external mould of the dorsal
valve, KCG1; and (2) external mould of a ventral valve,
KCG2.
Remarks.—These specimens were previously describedby Tazawa and Araki (2013, p. 7, fig. 2.3, 2.4) as
Gypospirifer kobiyamai Tazawa and Araki, 2013. This
species is characterized by its large size (length 49 mm,
width about 65 mm in the better-preserved specimen, KCG1)
and in having a narrow deep ventral sulcus and a narrow highdorsal fold and numerous fine costae on the both valves(10–11 in 10 mm at midlength of ventral valve). Gypospirifer
kobiyamai most resembles Gypospirifer gryphus Cooper and
Grant 1976a, p. 2211, pl. 591, figs. 1–5), from the Graham
Formation of Texas, in size and shape of shell, but the Texan

110

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 69. A, B, Phricodothyris catatona (Cooper and Grant); A, ventral (A1), dorsal (A4), anterior (A5), posterior (A6) and lateral (A7) views ofconjoined shell, and enlarged concentric lamellae and rows of very fine spinules on the ventral valve, NU-B2516; B, dorsal valve (B 1, B2) and enlarged
concentric lamellae and rows of very fine spinules on the dorsal valve, NU-B2517. Scale bars are 1 cm, except for A3 and B3.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

species differs from the present species in having a shallower
sulcus and coarser costae on ventral valve. Gypospirifer
volatilis Duan and Li (1985, p. 127, 207, pl. 48, figs. 1, 2; pl.
49, figs. 1, 2) from the Zhesi (Jisu) Formation of Zhesi, Inner
Mongolia, north China, differs from G. kobiyamai in having
much broader and higher fold on dorsal valve.
Occurrence.—KY1 Unit in Kamiyasse (locality AR1).
Distribution.—Wordian–Wuchiapingian: northeasternJapan (Kamiyasse–Imo, Kesennuma and Takakurayama in
the South Kitakami Belt), central Japan (Moribu in the Hida
Gaien Belt), southwestern Japan (Mizukoshi in Kyushu
Island), northeastern China (Heilongjiang) and eastern Russia(South Primorye).
Family SPIRIFERELLIDAE Waterhouse, 1968a
Genus ALISPIRIFERELLA Waterhouse and Waddington,
1982
Type species.—Spirifer (Spiriferella) keilhavii var.
ordinaria Einor in Licharew and Einor, 1939.
Alispiriferella lita (Fredericks, 1924b)(Figs. 67A–F, 68A–E)
Spiriferella saranae mut. lita Fredericks, 1924b, p. 36, pl. 1,
figs. 16–27; text-fig. 2.
Spirifer cf. saranae mut. lita Fredericks. Hayasaka, 1925a, p.
98, pl. 5, fig. 14.
Spiriferella cf. saranae mut. lita Fredericks. Nonaka, 1944,
p. 86, pl. 7, figs. 12–14.
Spiriferella keilhavii (von Buch). Yanagida, 1963, p. 72, pl.
9, figs. 4–9; pl. 10, figs. 1–7; Minato et al., 1979, pl. 67,
figs. 1–3: Tazawa and Gunji, 1982, p. 70, pl. 4, figs. 4–7;
Tazawa, 1999a, p. 5, pl. 1, figs. 2–6; Tazawa and Ibaraki,
2001, p. 16, pl. 4, figs. 1–10; Tazawa, 2002, fig. 10.4;
Tazawa and Chen, 2006, p. 335, fig. 6.1, 6.2, 6.8.
Alispirifer aff. laminosus transversa Maxwell. Yanagisawa,
1967, p. 90, pl. 2, fig. 3.
Cancellospirifer? maxwelli Campbell. Yanagisawa, 1967, p.
92, pl. 3, fig. 16.
Timaniella harkeri Waterhouse. Licharew and Kotlyar, 1978,
pl. 18, figs. 2, 3.
Spiriferella grandis Kotlyar in Licharew and Kotlyar, 1978,
p. 73, pl. 18, figs. 7, 8.
Spiriferella lita (Fredericks). Tazawa, 1979, p. 28, pl. 4, figs.
12, 13; pl. 5, figs. 1–4, 6; Tazawa, 2001, p. 302, fig.
8.19–8.22; Tazawa and Chen, 2006, p. 336, fig. 6.4.
Alispiriferella sp. Yanagida, 1996, fig. 2.2, 2.4.
Spiriferella cf. lita (Fredericks). Tazawa et al., 2000, p. 12,
pl. 1, figs. 16, 17.
Alispiriferella ordinaria (Einor). Tazawa, 2001, p. 302, fig.
8.14.
Alispiriferella japonica Tazawa, 2001, p. 303, fig. 8.15‒8.18.
Alispiriferella neimongolensis Wang and Zhang, 2003, p.
154, pl. 46, figs. 9–18; pl. 50, figs. 5, 9; Tazawa and Chen,

111

2006, p. 336, fig. 6.3.
Alispiriferella lita (Fredericks). Tazawa and Hasegawa, 2007,
p. 9, fig. 5.3–5.11; Tazawa, 2008b, p. 55, fig. 9.8–9.14;
Tazawa, 2009, p. 74, fig. 5.4–5.9; Tazawa, 2016b, p. 34,
fig. 11.1; Tazawa and Araki, 2017, p. 279, fig. 8.8–8.11.
Spiriferella sp. Tazawa, Kikuchi, Nikaido, Adachi andOkumura, 2014, p. 385, fig. 4.3.
Spiriferella sp. Tazawa, 2015, p. 79, fig. 8.3.
Material.—Forty specimens from localities AR4, AR5,KF217 and KS9: (1) external and internal moulds of aconjoined shell, IGPS96219: (2) internal mould of aconjoined shell, with external mould of the ventral valve,
IGPS96221; (3) internal mould of two conjoined shells, withexternal mould of the dorsal valves, IGPS96220, NU-B319;(4) internal moulds of three conjoined shells, KCG20, 21,
NU-B2016, NU-B325; (5) external and internal moulds offive ventral valves, IGPS96222–96224, NU-B315, 316; (6)external moulds of two ventral valves, NU-B317, 318; (7)internal moulds of nine ventral valves, IGPS96225–96227,
NU-B326–331; (8) external and internal moulds of four
dorsal valves, IGPS96228–96230, NU-B320; (9) externalmould of a dorsal valve, IGPS96231; and (9) internal moulds
of seven dorsal valves, IGPS96232–96235, NU-B332–334.
Remarks.—Most of the specimens from Setamai andMatsukawa were previously described by Tazawa (1979, p.
28, pl. 4, figs. 12, 13; pl. 5, figs. 1–4, 6) as Spiriferella lita
(Fredericks, 1924b) and by Tazawa and Ibaraki (2001, p. 16,
pl. 4, figs. 1–10) as Spiriferella keilhavii (von Buch, 1846).
Subsequently, Tazawa and Araki (2017, p. 279, fig. 8.8–8.11)
redescribed the specimens as Alispiriferella lita (Fredericks,
1924b). These specimens can be referred to Alispiriferellalita (Fredericks, 1924b), from the middle Permian (Wordian)of South Primorye, eastern Russia, by large transverse shells
and deep ventral sulcus with smooth V-shaped bottom andstrong simple costae on ventral valve. The type species,
Alispiriferella ordinaria (Einor in Licharew and Einor, 1939,
p. 140, 217, pl. 23, figs. 6, 7; pl. 24, fig. 1), from the lower
Permian of Novaya Zemlya, northern Russia, differs from A.
lita by its smaller and less transverse shell with ventral sulcus
bearing two prominent sulcal costae. Alispiriferella keilhavii(von Buch, 1846), redescribed by Dunbar (1955, p. 139, pl.
25, figs. 1–9; pl. 26, figs. 1–11; pl. 27, figs. 1–44), from the
middle Permian of eastern Greenland, differs from thepresent species in having weakly fasciculate costae on both
valves.
Occurrence.—KN2 Unit in Setamai (locality KS9); andKY1 Unit in Imo (locality KF217) and Matsukawa (localities
AR4 and AR5).
Distribution.—Roadian–Changhsingian: northeasternJapan (Setamai, Kamiyasse–Imo, Matsukawa, Ogatsu,
Ichinoseki, Takakurayama, Soma and Hitachi in the South
Kitakami Belt), central Japan (Moribu in the Hida Gaien
Belt), southwestern Japan (Mizukoshi in Kyushu Island, andTsunemori in the Akiyoshi Belt), northern China (Inner

112

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 70. A–C, Permophricodothyris grandis (Chao); A, ventral external mould (A1) enlarged rows of spine bases on the external mould of ventral
valve (A2 , A 3 ), external latex cast (A4 ) and ventral (A5 ) and dorsal (A6 ) views of internal mould of conjoined shell, NU-B1961; B, ventral valve,
NU-B1965; C, ventral (C1) and dorsal (C2, C3) views of internal mould of conjoined shell, NU-B1962. Scale bars are 1 cm, except for A2 and A3.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Mongolia), northeastern China (Heilongjiang) and eastern
Russia (South Primorye).
Suborder DELTHYRIDINA Ivanova, 1972
Superfamily RETICULARIOIDEA Waagen, 1883
Family ELYTHIDAE Fredericks, 1924a
Subfamily PHRICODOTHYRIDINAE Caster, 1939
Genus PHRICODOTHYRIS George, 1932
Type species.—Phricodothyris lucerna George, 1932.
Phricodothyris catatona (Cooper and Grant, 1976a)(Fig. 69A, B)
Neophricodothyris catatona Cooper and Grant, 1976a, p.
2250, pl. 636, figs. 1–28.
Phricodothyris sp. Minato et al., 1979, pl. 67, fig. 6.
Phricodothyris catatona (Cooper and Grant). Tazawa et al.,2016, p. 381, fig. 9.4.
Material.—Two specimens from locality KF84: (1) an
abraded conjoined shell, NU-B2516; and (2) an abradeddorsal valve, NU-B2517.
Description.—Shell large in size for genus, transversely
suboval in outline, moderately biconvex, anterior commissure
slightly uniplicate, with greatest width at about midlength;
length 28 mm, width 59 mm in the larger specimen(NU-B2517; length 32 mm, width 46 mm in the smatter
specimen (NU-B2516). Ventral valve moderately and evenly
convex in lateral profile; sulcus weak near anterior margin.
Brachial valve slightly less strongly convex than opposite
valve; fold absent or very low near anterior margin. Externalsurface of both valves ornamented with numerous regularconcentric lamellae bearing dense very fine spinules;
concentric lamellae numbering 9–15 in 5 mm at near anterior
margin of ventral valve. Interior of both valves not wellpreserved.
Remarks.—These specimens can be referred toPhricodothyris catatona (Cooper and Grant, 1976a),
originally described by Cooper and Grant (1976a, p. 2250, pl.
636, figs. 1–28) as Neophricodothyris catatona Cooper and
Grant, 1976a, in large, transverse shell with slightly sulcate
anterior commissure and in having numerous regularconcentric lamellae. Phricodothyris transversa (Cooper and
Grant, 1976a, p. 2255, pl. 639, figs. 1–28), from the lower
Wolfcampian of Texas, is also a large, transverse
Phricodothyris species, but differs from P. catatona inhaving strong uniplicate anterior commissure.
Occurrence.—KY1 Unit in Kamiyasse (locality KF84).
Distribution.—Artinskian–Wordian: northeastern Japan(Kamiyasse–Imo in the South Kitakami Belt), central Japan(Kuzu in the Mino Belt) and USA (Texas).
Genus PERMOPHRICODOTHYRIS Pavlova, 1965

113

Type species.—Permophricodothyris ovata Pavlova, 1965.
Permophricodothyris grandis (Chao, 1929)(Fig. 70A–C)
Squamularia grandis Chao, 1929, p. 97, pl. 11, figs. 1–3; Jinet al., 1974, p. 312, pl. 164, figs. 17–19; Yang et al., 1977,
p. 452, pl. 179, fig. 11; Feng and Jiang, 1978, p. 296, pl.
105, figs. 2, 3; Tong, 1978, p. 261, pl. 91, fig. 2; Liao,
1980, pl. 8, figs. 50, 51; Liu et al., 1982, p. 214, pl. 158,
fig. 1; Wang et al., 1982, p. 248, pl. 94, fig. 10; Ding and
Qi, 1983, p. 412, pl. 139, fig. 10; Yang, 1984, p. 238, pl.
38, fig. 23; Xu, 1987, p. 235, pl. 16, figs. 4–9; Kotlyar,
1989, pl. 24, figs. 6–8; Zeng et al., 1995, pl. 13, figs. 1, 2.
Permophricodothyris grandis (Chao). Licharew and Kotlyar,
1978, pl. 22, fig. 3; Zhan, 1979, p. 97, pl. 8, figs. 1, 2, 4–6,
text-fig. 19; Shi et al., 2002, p. 378, figs. 3B, C, 4B,
5.1–5.13, 6.1–6.9; Tazawa and Kaneko, 2016, p. 26, fig. 3.
Material.—Six specimens from localities KF62, KF79,
KF217 and KF218: (1) internal moulds of two conjoined
shells, with external moulds of the ventral valves,
NU-B1961, 1962; (2) external and internal moulds of two
ventral valves, NU-B1963, 1964; and (3) external casts oftwo ventral valves, NU-B1965, 1966.
Remarks.—These specimens were previously describedby Tazawa and Kaneko (2016, p. 26, fig. 3) as
Permophricodothyris grandis (Chao, 1929). The
Kamiyasse–Imo specimens can be referred toPermophricodothyris grandis (Chao, 1929), from theLungtan Formation of Zhejiang, eastern China, in large size(length about 61 mm, width about 48 mm in the largest
specimen, NU-B1965), slightly elongate oval outline, and inhaving broad and shallow sulcus on ventral valve. Besides,
the Kitakami species is ornamented with numerous regularconcentric lamellae with a row of very fine biramous spine
bases on anterior margin, and has posterior directed spiralia.
Permophricodothyris indica (Waagen, 1883, p. 542, pl. 43,
fig. 6; pl. 44, fig. 2), from the Wargal Formation of the Salt
Range, Pakistan, is distinguished from P. grandis by its
nearly equidimentional shell and shallower ventral sulcus.
The type species, Permophricodothyris ovata Pavlova (1965,
p. 135, figs. 1e, 2–4), from the upper Permian (Dzulfian) of
Transcaucasia, is clearly distinguished from P. grandis by its
smaller size, more elongate outline and shallower ventral
sulcus.
Occurrence.—KY1 Unit in Kamiyasse (localities KF62
and KF79) and Imo (localities KF217 and KF218).
Distribution.—Wordian‒Changhsingian: northeasternJapan (Kamiyasse–Imo in the South Kitakami Belt),
northwestern China (Shaanxi), eastern Russia (South
Primorye), eastern China (Zhejiang and Jiangxi),
central-southern China (Hubei, Hunan, Guangdong andGuangxi) and southwestern China (Guizhou and Sichuan).

114

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 71. A–D, Permophricodothyris squamularioides (Huang); A, ventral external mould (A1), ventral external latex cast (A2) and ventral (A3), dorsal(A4), anterior (A5), posterior (A6) and lateral (A7) views of internal mould of conjoined shell, UHR12582; B, ventral (B1, B2) and dorsal (B3) views ofabraded conjoined shell, NU-B1969; C, ventral external mould (C1), enlarged rows of spine bases and interspinous pustues on the external mould (C2, C3)
of ventral valve, NU-B1970; D, external mould (D1) and external latex cast (D2) of ventral valve, NU-B1972. Scale bars are 1 cm, except for C2 and C3.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

115

Permophricodothyris squamularioides (Huang, 1933)(Fig. 71A–D)

Type species.—Licharewina praetriassica Kotlyar,
Zakharov and Polubotko, 2004.

Martinia squamularioides Huang, 1933, p. 50, pl. 7, fig. 2;
pl. 8, fig. 8.
Squamularia squamularioides (Huang). Feng and Jiang,
1978, p. 295, pl. 105, fig. 1; Tong, 1978, p. 261, pl. 90, fig.
11; Liao, 1979, pl. 1, fig. 23; Liao, 1980, p. 266, pl. 7, figs.
36, 37; Yang, 1984, p. 238, pl. 38, fig. 20; Zeng et al.,
1995, pl. 14, figs. 6, 8.
Phricodothyris cf. caroli (Gemmellaro). Minato et al., 1979,
pl. 67, fig. 7.
Permophricodothyris squamularioides (Huang). Shen andZhang, 2008, fig. 5.24–5.30; Tazawa and Kaneko, 2016, p.
28, fig. 4.

Licharewina arakii (Hayasaka, 1963b)(Fig. 68F)

Material.—Eight specimens from localities KF18, KF28,
KF94 and KF217: (1) external and internal moulds of three
conjoined shells, NU-B1967–1969; (2) internal moulds ofthree conjoined shells, with external moulds of the ventral
valves, NU-B1970, 1971, UHR12582; (3) external andinternal moulds of a ventral valve, NU-B1972; and (4)external cast of a ventral valve, NU-B1973.
Remarks.—These specimens were previously describedby Tazawa and Kaneko (2016, p. 28, fig. 4) as
Permophricodothyris squamularioides (Huang, 1933), on thebasis of ventral valve of medium size (length 36 mm, width
30 mm in the largest specimen, UHR12582), elongate oval
outline, and in having a distinct ventral sulcus, besides
regularly spaced growth lamellae with a row of numerous
dense spine bases and very fine interspinous pustules atfrontal margin. Permophricodothyris calori (Gemmellaro,1899), redescribed by Pavlova (1969, p. 103, pl. 10, fig. 1) on
the specimens from the upper Permian of the Caucasus
Mountains, differs from P. squamularioides in its larger sizeand shallower ventral sulcus. The preceding species,
Permophricodothyris grandis (Chao, 1929), is readily
distinguished from the present species by its larger size, lesselongate outline, and in having a wider and shallower ventral
sulcus.
Occurrence.—KY1 Unit in Kamiyasse (localities KF18,
KF28 and KF94) and Imo (locality KF217).
Distribution.—Wordian‒Changhsingian: northeasternJapan (Kamiyasse–Imo in the South Kitakami Belt),
central-southern China (Hubei and Hunan) and southwestern
China (Guizhou and Sichuan).
Order SPIRIFERINIDA Ivanova, 1972
Suborder CYRTINIDINA Carter and Johnson in Carter et al.,
1994
Superfamily CYRTINOIDEA Fredericks, 1911
Family CYRTINIDAE Fredericks, 1911
Genus LICHAREWINA Kotlyar, Zakharov and Polubotko,
2004

Geyerella arakii Hayasaka, 1963b, p. 481, figs. 2, 3.
Licharewina arakii (Hayasaka). Tazawa and Araki, 2013, p.
9, fig. 3.1, 3.2; Tazawa and Araki, 2017, p. 280, fig. 8.5.
Material.—Two specimens from localities AR2 and AR4:(1) internal mould of a conjoined shell, with external mouldof the dorsal valve and interarea of the ventral valve, KCG4;and (2) internal mould of a conjoined valve, KCG6.
Remarks.—These specimens were first described by
Hayasaka (1963b, p. 481, figs. 2, 3) as Geyerella arakii
Hayasaka, 1963b. Subsequently, Tazawa and Araki (2013, p.
9, fig. 3.1, 3.2) redescribed the specimens as Licharewina
arakii (Hayasaka, 1963b). This species is characterized by its
large size (length about 27 mm, width 19 mm in the
better-preserved specimen, KCG4) and the highly pyramidal
ventral valve. Licharewina josephinae (Gemmellaro, 1899),
redescribed by Shen and Clapham (2009, p. 732, pl. 6, figs.
1–15) from the Episkopi Formation (Wuchiapingian) of
Hidra island, Greece, differs from L. arakii in its smaller sizeand less pyramidal outline. The type species, Licharewina
praetriassica Kotlyar, Zakharov and Polubotko (2004, p.
522, fig. 6.13–6.20), from the upper Permian(Changhsingian) of the Caucasus Mountains, is readily
distinguished from the present species by its much smallersize.
Occurrence.—KY1 Unit in Kamiyasse (locality AR2) andMatsukawa (locality AR4).
Distribution.—Wordian: northeastern Japan (Kamiyasse–Imo
and Matsukawa in the South Kitakami Belt).
Suborder SPIRIFERINIDINA Ivanova, 1972
Superfamily PENNOSPIRIRINOIDEA Dagys, 1972
Family PARASPIRIFERINIDAE Cooper and Grant, 1976b
Genus CALLISPIRINA Cooper and Muir-wood, 1951
Type species.—Spiriferina ornata Waagen, 1883.
Callispirina sheni Tazawa, sp. nov.
(Fig. 72A, B)
Etymology.—Named for Shen Shuzhong, who contributed
to the Permian brachiopod taxonomy and biogeography, andstudied with the present author (J. Tazawa) on the Permanbrachiopod faunas of Japan.
Material.—Three specimens from localities KF65 andKF69: (1) external and internal moulds of a dorsal valve,
NU-B2566 (holotype); (2) external mould of a ventral valve,
NU-B2567; and (3) external mould of a dorsal valve,
NU-B2568.

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JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 72. A, B, Callispirina sheni sp. nov.; A, external latex cast (A1, A2) of ventral valve, NU-B2567; B, external latex cast (B1, B2) and internalmould (B3) of dorsal valve, NU-B2566 (holotype); C–E, Spiriferellina fredericksi Tazawa, C, ventral external latex cast (C1, C2, C3) and dorsal externallatex cast (C4) of conjoined shell, NU-B1763 (holotype); D, external latex cast (D1, D2) and internal mould (D3) of ventral valve, NU-B1767; E, ventral(E1, E2) and dorsal (E3) views of internal mould of conjoined shell, NU-B1768. Scale bars are 1 cm.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

Diagnosis.—Medium-sized Callispirina having a ventral
sulcus with flat bottom.
Description.—Shell medium in size for genus, transversely
subelliptical in outline, widest at midlength; length 9 mm,
width 14 mm in the largest specimen (NU-B2566). Ventral
valve strongly convex in lateral profile; sulcus originating at
beak, widening anteriorly, with flat bottom. Dorsal valve
unevenly convex in lateral profile, most convex at umbonal
region; fold originating at beak, widening anteriorly. Externalsurface of both valves ornamented with some coarse rounded
costae and microornament consisting of numerous regularconcentric lamellae; 4–5 costae on each flank of both ventral
and dorsal valves; 3–4 lamellae in 1 mm at near anterior
margin of both valves. Internal structure of dorsal valve notwell preserved.
Remarks.—These specimens are safely assigned to thegenus Callispirina Cooper and Muir-Wood, 1951 in being
small size and in having microornament consisting of
numerous dense concentric lamellae over both ventral anddorsal valves. The Kamiyasse specimens are seemed to be a
new species in having flat-bottomed ventral sulcus.
Callispirina rotunda Cooper and Grant (1976b, p. 2743, pl.
705, figs. 66–82), from the Bell Canyon Formation of Texas,
differs from Callispirina sheni by its larger size and inhaving ventral sulcus with narrow depression bearing a low
costa. Callispirina austrina Grant (1976, p. 231, pl. 63, figs.
1–37), from the Ratburi Formation of Ko Muk, southern
Thailand, differs from C. sheni by its smaller size and inhaving ventral sulcus with V-shaped bottom. The typespecies, Callispirina ornata (Waagen, 1883), originallydescribed by Waagen (1883, p. 505, pl. 50, figs. 1, 2) fromthe Chhidru Formation of the Salt Range, Pakistan, differsfrom the present new species in having V-shaped bottom inventral valve.
Occurrence.—KY1 Unit in Kamiyasse (localities KF65
and KF69).
Family SPIRIFERELLINIDAE Ivanova, 1972
Genus SPIRIFERELLINA Fredericks, 1924aType species.—Terebratulites cristatus von Schlotheim, 1816.
Spiriferellina fredericksi Tazawa, 2014
（Figs. 72C–E, 73A, B）
Spiriferina cristata (Schellwien). Hayasaka, 1922a, p. 66, pl.
9, figs. 5–9.
Spiriferina cristata (von Schlotheim). Hayasaka, 1960, p. 53,
pl. 1, fig. 10.
Spiriferellina cristata biplicata (Davidson). Fredericks,
1924b, p. 35, pl. 1, fig. 15.
Spiriferellina cristata (von Schlotheim). Tazawa, 1976, pl. 2,
fig. 2, fig. 3; Minato et al., 1979, pl. 67, figs. 4, 5.
Spiriferellina fredericksi Tazawa, 2014, p. 19, fig. 3.5-3.7.

117

Material.—Sixteen specimens from localities KF29,
KF88, KF217 and KF218: (1) external mould of a conjoined
shell, NU-B1763 (holotype); (2) internal moulds of two
conjoined shells, with external moulds of the ventral valves,
NU-B1764, 1765; (3) internal mould of a conjoined shell,
with external mould of the dorsal valve, NU-B2592; (4)internal moulds of three conjoined shells, NU-B1768–1770;(5) external and internal moulds of two ventral valves,
NU-B1766, 1767; (6) internal moulds of four ventral valves,
NU-B1771–1774; (7) external cast of a dorsal valve,
NU-B1775; and (8) internal moulds of two dorsal valves,
NU-B1776, 1777.
Remarks.—The specimens from Kamiyasse–Imo werepreviously described by Tazawa (2014, p. 19, fig. 3.5–3.7) as
Spiriferellina fredericksi Tazawa, 2014. This species ischaracterized by its large, transverse shell (length about 21mm, width about 50 mm in the holotype, NU-B1763; length
about 21 mm, width about 54 mm in the largest ventral valve
specimen, NU-B1766), with 5–6 costae on each lateral flankof both valves. The type species, Spiriferellina cristata (vonSchlotheim, 1816), redescribed and refigured by Campbell(1959, p. 358, pl. 59, figs. 1–9; pl. 60, fig. 3) on thespecimens from the Zechstein of Thuringia, Germany, differsfrom S. fredericksi by its smaller size, less transverse outline
and in having fewer costae on both ventral and dorsal valves.
Spiriferellina cristata biplicata (non Davidson, 1863),
described by Fredericks (1924b, p. 35, pl. 1, fig. 15) from the
Chandalaz Formation of South Primorye, eastern Russia, may
be a synonym of the present species.
Occurrence.—KY1 Unit in Kamiyasse (localities KF29
and KF88) and Imo (localities KF217 and KF218).
Distribution.—Wordian: northeastern Japan (Kamiyasse–Imoin the South Kitakami Belt) and eastern Russia (South
Primorye).
Order TEREBRATULIDA Waagen, 1883
Suborder TEREBRATULIDINA Waagen, 1883
Superfamily DIELASMATOIDEA Schuchert, 1913
Family DIELASMATIDAE Schuchert, 1913
Subfamily DIELASMATINAE Schuchert, 1913
Genus DIELASMA King, 1859
Type species.—Terebratulites elongatus von Schlotheim, 1816.
Dielasma sp.
(Fig. 74A)
Dielasma sp. Tazawa, 1979, p. 30, pl. 5, fig. 5; Tazawa andAraki, 2017, p. 280, fig. 8.6.
Material.—One specimen from locality AR5, external andinternal moulds of a dorsal valve, IGPS96236.
Remarks.—This specimen was previously described by
Tazawa (1979, p. 30, pl. 5, fig. 5) and Tazawa and Araki(2017, p. 280, fig. 8.6) as Dielasma sp. Tazawa, 1979. The

118

JUN-ICHI TAZAWA AND YOUSUKE IBARAKIFIGURE 73. A, B, Spiriferellina fredericksi Tazawa; A, ventral external latex cast (A1, A2) and ventral (A3), dorsal (A4), anterior (A5), posterior (A6) andlateral (A7) views of internal mould of conjoined shell, NU-B1764; B, ventral (B1, B2), dorsal (B3), anterior (B4), posterior (B5) and lateral (B6) views ofconjoined shell, NU-B1770. Scale bars are 1 cm.

Matsukawa species is large in size for genus (length 29 mm,
width 20 mm in the dorsal valve, IGPS96236), andcharacterized by presence of a sharp fold on anterior portionof dorsal valve. The Matsukawa species resembles Dielasma
plica (Kutorga, 1842), redescribed by Diener (1903, p. 44, pl.
2, fig. 2) from the middle Permian (Capitanian) of the
Chitichun No. 1, southern Tibet, in having a dorsal fold. But
accurate comparison is difficult for the poorly preserved
specimen. Dielasma sp. Tazawa (2001, p. 303, fig. 8.27,8.28), from the Moribu Formation of Moribu, Hida Gaien
Belt, central Japan, is readily distinguished from the present
species in having a distinct median fold on ventral valve.
Occurrence.—KY1 Unit in Matsukawa (locality AR5).
Genus WHITSPAKIA Stehli, 1964
Type species.—Dielasma biplex Waagen, 1882.

Whitspakia nipponica Tazawa, sp. nov.
(Fig. 74B–D)
Etymology.—Named after the country, Nippon.
Matrial.—Seven specimens from localities KF217 andKF218: (1) internal and external moulds of a conjoined shell,
NU-B1864; (2) internal moulds of three conjoined shells,
NU-B1865 (holotype), 2471, 2587; (3) internal mould of aventral valve, NU-B2470; (4) a dorsal valve, NU-B1866; and(5) external and internal moulds of a dorsal valve,
NU-B1975.
Diagnosis.—Large, elongate subpentagonal and gently
biconvex shell, with biplicate anterior commissure.
Description.—Shell medium in size for genus, elongate
subpentagonal in outline, with greatest width slightly anterior
to midlength of valve; length about 42 mm, width about 24
mm in the holotype (NU-B1865). Ventral valve gently andunevenly convex in lateral profile, rather strongly convex at

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

119

FIGURE 74. A, Dielasma sp., external latex cast (A1, A2), internal latex cast (A3) and internal mould (A4) of dorsal valve, IGPS96236; B–D, Whitspakia
nipponica sp. nov., B, ventral (B1, B2), dorsal (B3), anterior (B4), posterior (B5) and lateral (B6) views of internal mould of conjoined shell, NU-B1865(holotype); C, dorsal view (C1, C2) of dorsal valve, NU-B1866; D, internal mould of dorsal valve, NU-B1975. Scale bars are 1 cm.

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JUN-ICHI TAZAWA AND YOUSUKE IBARAKI

umbonal region but nearly flattened on venter; foramen large,
permesothirid; sulcus broad and shallow, with two shortdiverging costae on anterior one third valve length. Dorsal
valve slightly convex in lateral profile; fold wide and low,with a median costa and two diverging costae on anterior halfof valve. External surface of both valves ornamented withsome irregular concentric rugae. Internal structures of both
ventral and dorsal valves not well preserved.
Remarks.—These specimens can be assigned to the genus
Whitspakia by its large, elongate subpentagonal and slightly
biconvex shell, with a large permesothirid foramen. The
present species resembles Whitspakia biplex (Waagen, 1882),
originally described by Waagen (1882, p. 349, pl. 25, figs.
3–5) as Dielasma biplex Waagen from the Wargal Formationof the Salt Range, in size and outline of shell, but differsfrom the Pakistani species in having valves with biplicate
anterior commissure. Whitspakia introplicata Yang (1984, p.
239, pl. 38, fig. 24) from the Maokouan of Hubei,
central-southern China has also biplicate anterior
commissure, but differs from W. nipponica sp. nov. by its
much smaller size and less elongate outline.
Occurrence.—KY1 Unit in Imo (localities KF217 andKF218).
ACKNOWLEDGEMENTS
We sincerely thank Kenshiro Ogasawara (Professor
Emeritus of the University of Tsukuba, Tsukuba) andSoichiro Kawabe (Fukui Prefectural Dinosaur Museum,
Katsuyama) for editing the manuscript; Shuzhong Shen
(Nanjing University, Nanjing) and an anonymous reviewer
for their constructive comments and suggestions on the
manuscript; Yoshitsugu Kobayashi and Hiroki Echizenya(Hokkaido University Museum, Sapporo), Yasufumi Iryu and
Jun Nemoto (Tohoku University Museum, Sendai), andMasaru Kumagai (Rikuzentakata City Museum,Rikuzentakata, Iwate Prefecture) for loan of the typespecimens; Tomoharu Ikeda (Fuji City, Shizuoka Prefecture)for help in the field (Imo); and Hideo Araki (Kesennuma
City, Miyagi Prefecture) for providing part of the brachiopod
specimens.
REFERENCES
Abramov, B. S. 1970. Biostratigraphy of the Carboniferous of
Sette -Davana (Southern Verkhoyansk). Nauka, Moskva, 176
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Afanasjeva, G. A. 1975. Middle and Late Carboniferous
Chonetacea (Brachiopoda) of the Russian Platform.
Paleontologicheskiy Zhurnal 1975 (2): 96–113.***Afanasjeva, 1977, G. A. 1977. Systematic descriptions;
Suborder Chonetidina; pp. 5–41 in T. G. Sarytcheva (ed.),Late Palaeozoic Productida of Siberia and Arctic Region.
Trudy Paleontologicheskogo Instituta, Akademiya NaukSSSR 161.***Afanasjeva, G. A. 1988. Brachiopoda of the order Chonetida.
Trudy Paleontologicheskogo Instituta, Akademiya NaukSSSR 228: 1–122.***Afanasjeva, G. A., and J. Tazawa. 2007. New large chonetids(Brachiopoda) from the Middle Permian of the southern
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穴淵
Anabuchi
葡萄沢
Budosawa Valley
茶屋沢
Chayazawa Valley
洞屋沢Doyasawa Valley
二ツ森層
Futatsumori Formarion
Hizume–Kesennuma Fault
日詰‒気仙沼断層
Hosoo Formation
細尾層
Imo
飯森
Imosawa Valley
飯森沢
Iwaizaki Stage
岩井埼階合地沢
Kacchizawa (Kattizawa, Kattisawa) Valley
上八瀬
Kamiyasse
叶倉層
Kanokura Formation

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地名・地層名Kesen-gun
Kiritoshi
Kokitamukaisawa Valley
Kurosawa Formation
Matashirosawa Valley
Matsukawa
Minamizawa Valley
Minokerazawa Valley
Mt. Kurosawayama
Nagaiwa Formation
Nakadaira Formation
Obama
Omotematsukawa

＞
気仙郡
切通
小北向沢
黒沢層
又四郎沢
松川
南沢
ミノケラ沢
黒沢山
長岩層
中平層
小浜
表松川

Onimarusawa Valley
Rikuzentakata
Sakamotozawa Series
Setamai
Shigejisawa Valley
Sumita-cho
Tateishizawa Valley
Toyazawa Valley
Toyoma Formation
Yahigi-cho
Yakejima
Yukizawa Series

鬼丸沢
陸前高田
坂本沢統
世田米
茂路沢
住田町
立石沢
戸屋沢
登米層
矢作町
八景島
雪沢統

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

APPENDIX
AR1: Kamiyasse in the Kamiyasse – Imo area (38° 59′13″N,141° 31′44″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, upper KY1 Unit of the
Kamiyasse Formation, with Neospirifer moosakhailensis
and Gypospirifer kobiyamai.
AR2: Kamiyasse in the Kamiyasse – Imo area (38°59′04″N,
141° 31′33″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, upper KY1 Unit of the
Kamiyasse Formation, with Licharewina arakii.
AR3: Kamiyasse in the Kamiyasse – Imo area (38°58′53″N,141° 31′19″E), lower Kokitamukaisawa Valley, a tributaryof Shigejisawa Valley, greenish grey fine-grainedsandstone, middle KY1 Unit of the Kamiyasse
Formation, with Orbiculoidea verum and Anidanthus
mizukoshiensis.
AR4: Anabuchi in the Matsukawa area (38° 55′12″N, 141° 32′32″E),
cliff along the Matsukawa River, dark grey argillaceous
limestone, middle KY1 Unit of the Kamiyasse
Formation, with Neochonetes (Huangichonetes)
matsukawensis, Hexiproductus echidniformis, Kunluniasp., Urushtenoidea crenulata, Permundaria asiatica,
Permundaria tenuistriata, Yakovlevia mammata,
Neorichthofenia mabutii, Leptodus nobilis, Paralyttonia
kesennumensis, Dicystoconcha lapparenti, Stenoscisma
margaritovi, Martinia triquetra, Alispiriferella lita andLicharewina arakii.
AR5: Kiritoshi in the Matsukawa area (38°54′55″N, 141°32′33″E),
road cutting, greenish grey fine-grained sandstone,
lower KY1 Unit of the Kamiyasse Formation, with
Capillomesolobus heritschi, Linoproductus hayasakai,
Costatumulus cancriniformis, Yakovlevia kaluzinensis,
Alispiriferella lita and Dielasma sp.
KF3: Kamiyasse in the Kamiyasse – Imo area (38 ° 59′03″N,
141°31′41″E), upper Shigejisawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Anidanthus mizukoshiensis.
KF6: Kamiyasse in the Kamiyasse – Imo area (38 ° 59′23″N,141° 31′08″E), upper Minamizawa Valley, float, dark grey
argillaceous limestone, upper KY1 Unit of the
Kamiyasse Formation, with Orthotichia sp.
KF7: Kamiyasse in the Kamiyasse – Imo area (38 ° 59′23″N,141° 31′08″E), upper Minamizawa Valley, float, dark grey
argillaceous limestone, upper KY1 Unit of theK a m i y a s s e F o r m a t i o n , w i t h S p i n o m a rg i n i f e r alopingensis and Acosarina rectimarginata.
KF8: Kamiyasse in the Kamiyasse – Imo area (38° 59′31″N,
141°30′53″E), upper Minokerazawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Kitakamichonetes
multicallatus, Rhipidomella magna and Stenoscisma
mutabilis.
KF9: Kamiyasse in the Kamiyasse – Imo area (38° 59′02″N,141° 31′44″E), upper Shigejisawa Valley, greenish grey137

fine-grained sandstone, lower KY2 Unit of the
Kamiyasse Formation, with Waagenoconcha irginae.
KF11: Kamiyasse in the Kamiyasse – Imo area (38° 58′59″N,
141° 30′31″E), upper Shigejisawa Valley, dark grey argillaceous
limestone, middle KY1 Unit of the Kamiyasse
Formation, with Waagenoconcha irginae.
KF13: Kamiyasse in the Kamiyasse – Imo area (38° 59′11″N,141° 31′22″E), lower Kokitamukaisawa Valley, a tributaryof Shigejisawa Valley, dark grey argillaceous limestone,
middle KY1 Unit of the Kamiyasse Formation, with
Leptodus nobilis.
KF17: Kamiyasse in the Kamiyasse –Imo area (38° 59′07″N,
141°31′57″E), northern ridge of Mt. Kurosawayama, float,
greenish grey fine-grained sandstone, upper KY1 Unitof the Kamiyasse Formation, with Asperlinus japonicus.
KF18: Kamiyasse in the Kamiyasse – Imo area (38° 59′06″N,
141° 31′49″E), upper Shigejisawa Valley, float, dark grey
argillaceous limestone, lower KY1 Unit of the
Kamiyasse Formation, with Permophricodothyris
squamularioides.
KF20: Kamiyasse in the Kamiyasse – Imo area (38° 58′59″N,
141°31′29″E), upper Shigejisawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Streptorhynchus cataclinus.
KF21: Kamiyasse in the Kamiyasse – Imo area (38° 59′04″N,
141° 31′22″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, upper KY1 Unit of the
Kamiyasse Formation, with Globiella tschernyschewi
and Grandaurispina kozlowskiana.
KF22: Kamiyasse in the Kamiyasse – Imo area (38° 59′06″N,
141° 31′35″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, upper KY1 Unit of the
Kamiyasse Formation, with Grandaurispina
kozlowskiana, Derbyia nipponica and Hustedia remota.
KF27: Kamiyasse in the Kamiyasse – Imo area (38° 59′16″N,141°31′07″E), upper Minamizawa Valley, greenish grey
fine-grained sandstone, lower KY1 Unit of theK a m i y a s s e F o r m a t i o n , w i t h S t re p t o r h y n c h u spelargonatus.
KF28: Kamiyasse in the Kamiyasse – Imo area (38° 59′18″N,141° 31′08″E), upper Minamizawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Asperlinus japonicus andPermophricodothyris squamularioides.
KF29: Kamiyasse in the Kamiyasse – Imo area (38° 59′18″N,141° 31′08″E), upper Minamizawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Spiriferelina fredericksi.
KF33: Kamiyasse in the Kamiyasse – Imo area (38° 59′42″N,141° 30′55″E), upper Tateishizawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Stenoscisma mutabilis.
KF37: Kamiyasse in the Kamiyasse – Imo area (38° 59′12″N,141°31′17″E), upper Shigejisawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the

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JUN-ICHI TAZAWA AND YOUSUKE IBARAKI

Kamiyasse Formation, with Limbella sp.
KF39: Kamiyasse in the Kamiyasse – Imo area (38° 58′35″N,141° 31′10″E), upper Chayazawa Valley, greenish grey
fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Kanokurachonetes
kanokurensis, Chonetinetes elongatus, Waagenoconcha
irginae, Urushtenoidea crenulata and Anidanthus
mizukoshiensis.
KF46: Kamiyasse in the Kamiyasse – Imo area (38° 59′11″N,141° 31′39″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, upper KY1 Unit of the
Kamiyasse Formation, with Bathymyonia neimongolica,
Grandaurispina kozlowskiana, Permianella typica andStenoscisma margaritovi.
KF50: Kamiyasse in the Kamiyasse – Imo area (38° 59′21″N,
141°31′02″E), upper Minamizawa Valley, greenish grey
fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Grandaurispina kozlowskiana.
KF55: Kamiyasse in the Kamiyasse – Imo area (38° 58′19″N,141° 31′1″E), upper Doyasawa Valley, dark grey shale,
upper KY1 Unit of the Kamiyasse Formation, with
Kanokurachonetes kanokurensis.
KF61: Kamiyasse in the Kamiyasse – Imo area (38° 59′09″N,
141°31′41″E), upper Shigejisawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of
the Kamiyasse Formation, with Kitakamichonetes
multicapillatus.
KF62: Kamiyasse in the Kamiyasse – Imo area (38° 59′21″N,
141° 31′01″E), upper Minamizawa Valley, float, greenish
grey fine-grained sandstone, upper KY1 Unit of
the Kamiyasse Formation, with Chonosteges sp. andPermophricodothyris grandis.
KF63: Kamiyasse in the Kamiyasse – Imo area (38° 59′22″N,141°31′02″E), upper Minamizawa Valley, greenish grey
fine-grained sandstone, middle KY1 Unit of
the Kamiyasse Formation, with Spinomarginifera
kueichowensis.
KF64: Kamiyasse in the Kamiyasse – Imo area (38° 59′26″N,141°31′04″E), upper Minamizawa Valley, greenish grey
fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Vediproductus punctatiformis,
Meekella eximia and M. nodosa.
KF65: Kamiyasse in the Kamiyasse – Imo area (38° 58′51″N,
141 ° 31′22″E), lower Kokitamukaisawa Valley, atributary of Shigejisawa Valley, float, dark grey
argillaceous limestone, middle KY1 Unit of the
Kamiyasse Formation, with Callispirina sheni sp. nov.
KF67: Kamiyasse in the Kamiyasse – Imo area (38° 59′02″N,141° 31′39″E), upper Shigejisawa Valley, float, dark grey
argillaceous limestone, middle KY1 Unit of the
Kamiyasse Formation, with Leptodus nobilis.
KF69: Kamiyasse in the Kamiyasse – Imo area (38° 59′01″N,141° 31′31″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Callispirina sheni sp. nov.

KF71: Kamiyasse in the Kamiyasse – Imo area (38° 59′13″N,141°31′10″E), upper Shigejisawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Kitakamichonetes
multicapillatus and Spinomarginifera kueichowensis.
KF79: Kamiyasse in the Kamiyasse – Imo area (38° 59′07″N,
141°31′40″E), upper Shigejisawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Paramesolobus kitakamiensis
sp. nov., Kitakamichonetes multicapillatus, Asperlinus
japonicus and Permophricodothyris grandis.
KF80: Kamiyasse in the Kamiyasse – Imo area (38° 59′08″N,
141° 31′47″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, upper KY1 Unit of the
Kamiyasse Formation, with Permundaria asiatica.
KF84: Kamiyasse in the Kamiyasse – Imo area (38° 59′26″N,141° 31′04″E), upper Minamizawa Valley, float, dark grey
argillaceous limestone, lower KY1 Unit of theK a m i y a s s e F o r m a t i o n , w i t h S p i n o m a rg i n i f e r akueichowensis, Leptodus nobilis and Phricodothyris
catatona.
KF86: Kamiyasse in the Kamiyasse – Imo area (38° 59′20″N,141° 31′26″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, upper KY1 Unit of the
Kamiyasse Formation, with Asperlinus japonicus,
Permianella typica and Stenoscisma margaritovi.
KF88: Kamiyasse in the Kamiyasse – Imo area (38° 59′15″N,141° 31′22″E), upper Shigejisawa Valley, float, dark grey
argillaceous limestone, middle KY1 Unit of the
Kamiyasse Formation, with Kitakamichonetes
multicallatus, Spinomarginifera kueichowensis,
Permianella typica, Stenoscisma margaritovi andSpiriferellina fredericksi.
KF89: Kamiyasse in the Kamiyasse – Imo area (38° 59′16″N,141° 31′22″E), upper Shigejisawa Valley float, greenish
grey fine-grained sandstone, middle KY1 Unit of the
Kamiyasse Formation, with Spinomarginifera alpha,
Edriosteges multispinosus and Stenoscisma margaritovi.
KF90: Kamiyasse in the Kamiyasse – Imo area (38° 59′14″N,141° 31′23″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, upper KY1 Unit of theK a m i y a s s e F o r m a t i o n , w i t h S p i n o m a rg i n i f e r alopingensis, S. kueichowensis, S. alpha, Urushtenoidea
crenulata and Asperlinus japonicus.
KF91: Kamiyasse in the Kamiyasse – Imo area (38° 59′05″N,
141° 31′45″E), upper Shigejisawa Valley, greenish grey
fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Permianella typica andOrthothetina kayseri.
KF94: Kamiyasse in the Kamiyasse – Imo area (38° 59′30″N,141° 31′08″E), upper Minamizawa Valley, float, dark grey
argillaceous limestone, middle KY1 Unit of the
Kamiyase Formation, with Kitakamichonetes
multicapillatus, Permianella typica, Rhipidomellamagna, Stenoscisma margaritovi and Permophricodothyris

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWA

squamularioides.
KF96: Kamiyasse in the Kamiyasse – Imo area (38° 59′28″N,141° 31′08″E), upper Minamizawa Valley, float, greenish
grey fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Tyloplecta yangtzeensis.
KF99: Kamiyasse in the Kamiyasse – Imo area (38° 59′07″N,
141 ° 31′43″E), upper Shigejisawa Valley, dark grey
argillaceous limestone, upper KY1 Unit of the
Kamiyasse Formation, with Lamiproductus kamiyassensis,
Permundaria asiatica and P. tenuistriata.
KF121: Imo in the Kamiyasse–Imo area (38 ° 59′55″N,
141 ° 31′18″E), upper Imosawa Valley, greenish grey
fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Paramarginifera sp.,
Tyloplecta yangtzeensis, Waagenoconcha humboldti,
Derbyia nipponica, Meekella eximia, M. nodosa,
Orthothetina polita, O. kayseri, O. hayasakai,
Streptorhynchus pelargonatus, Orthotichia sp.,
Stenoscisma mutabilis and Martinia triquetra.
KF212: Kamiyasse in the Kamiyasse –Imo area (38° 58′49″N,
141° 31′11″E), middle Matashirosawa Vallley, a tributaryof Shigejisawa Valley, dark grey shale, upper Hosoo
Formation, with Poikilosakos kamiyassensis andMartinia semiplana.
KF217: Imo in the Kamiyasse–Imo area (38 ° 59′55″N,
141 ° 31′14″E), upper Imosawa Valley, greenish grey
fine-grained sandstone, lower KY1 Unit of the
Kamiyasse Formation, with Isogramma heritschi, I.
nakamurai, Paramesolobus kitakamiensis sp. nov.,
Transennatia gratiosa, Spinomarginifera lopingensis, S.
kueichowensis, S. alpha, Paramarginifera sp., Costiferina
spiralis, Tyloplecta yangtzeensis, Bathymyonian e i m o n g o l i c a , Ve d i p ro d u c t u s p u n c t a t i f o r m i s ,Waagenoconcha humboldti, Urushtenoidea crenulata,
Asperlinus japonicus, Leptodus nobilis, Permianella
typica, Dicystoconcha lapparenti, Orthothetina polita,
O. kayseri, O. hayasakai, O. transversa, Rhipidomellamagna, Acosarina dorsisulcata, A. rectimarginata,
Stenoscisma margaritovi, S. mutabilis, S. sokolskajae,
Rhynchopora tchernyshae, Permocryptospirifer
omeishanensis, Alispiriferella lita, Permophricodothyris
grandis, P. squamularioides, Spiriferellina fredericksi
and Whitspakia nipponica sp. nov.
KF218: Imo in the Kamiyasse–Imo area (38°59′55″N, 141°31′16″E),
upper Imosawa Valley, dark grey argillaceous limestone,
lower KY1 Unit of the Kamiyasse Formation, with
Kitakamichonetes multicapillatus, Transennatia
gratiosa, Bathymyonia neimongolica, Vediproductus
punctatiformis, Edriosteges multispinosus, Urushtenoidea
crenulata, Grandaurispina kozlowskiana, Asperlinus
japonicus, Leptodus nobilis, Pararigbyella doulingensis,
Permianella typica, Dicystoconcha lapparenti, Stenoscisma
margaritovi, S. mutabilis, Permocryptospirifer

139

omeishanensis, Hustedia remota, H. ratburiensis,
Martinia triquetra, Permophricodothyris grandis,
Spiriferellina fredericksi and Whitspakia nipponica sp.
nov.
KS1: Setamai area (39 ° 07′53″N, 141 ° 31′33″E), middle
Kanokurasawa Valley, greenish grey fine-grainedsandstone, upper KN1 Unit of the Kanokura Formation,with Linoproductus hayasakai, Leptodus nobilis,
Derbyia grandis and D. nipponica.
KS2: Setamai area (39 ° 06′44″N, 141 ° 31′25″E), upper
Onimarusawa Valley, a tributary of Kacchizawa Valley,greenish grey fine-grained sandstone, upper KN1 Unitof the Kanokura Formation, with Transennatia gratiosa.
KS3: Setamai area (39 ° 07′52″N, 141 ° 30′51″E), upper
Kanokurasawa Valley, greenish grey fine-grainedsandstone, upper KN1 Unit of the Kanokura Formation,with Linoproductus hayasakai.
KS4: Setamai area (39 ° 07′51″N, 141 ° 31′18″E), upper
Kanokurasawa Valley, greenish grey fine-grainedsandstone, upper KN1 Unit of the Kanokura Formation,with Linoproductus hayasakai.
KS7: Setamai area (39°07′42″N, 141°32′10″E), a tributary of
Kanokurasawa Valley, greenish grey fine-grainedsandstone, upper KN2 Unit of the Kanokura Formation,with Leptodus nobilis.
KS8: Setamai area (39°07′33″N, 141°32′26″E), a tributary of
Kacchizawa Valley, greenish grey fine-grainedsandstone, upper KN1 Unit of the Kanokura Formation,with Tyloplecta yangtzeensis.
KS9: Setamai area (39° 08′01″N, 141° 32′09″E), middle
Kanokurasawa Valley, greenish grey fine-grainedsandstone, upper KN2 Unit of the Kanokura Formation,with Transennatia gratiosa, Waagenoconcha irginae,
Urushtenoidea crenulata, Grandaurispina kozlowskiana,
Laterisma parallela, Derbyia grandis, Derbyia
nipponica and Alispiriferella lita.
KS10: Setamai area (39° 07′43″N, 141° 32′18″E), upper
Budosawa Valley, a tributary of Kacchizawa Valley, darkgrey impure limestone, upper KN2 Unit of the Kanokura
Formation, with Permundaria asiatica and Orthothetina
kitakamiensis.
KZ9: Omotematsukawa in the Matsukawa area (38° 55′09″N,
141° 32′33″E), sandstone quarry, greenish grey fine-grainedsandstone, lower KY1 Unit of the Kamiyasse
Formation, with Transennatia gratiosa, Hexiproductus
echidniformis, Scacchinella gigantea, Keyserlingina sp.,Petasmaia expansa, Meekella nodosa and Orthothetina
kayseri.
TY1: Imo in the Kamiyasse-Imo area (38 ° 59′22″N,141° 31′59″E), upper Toyazawa Valley, greenish grey
fine-grained sandstone, middle KY1 Unit of the
Kamiyasse Formation, with Waagenoconcha irginae.

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JUN-ICHI TAZAWA AND YOUSUKE IBARAKITABLE 1 (1). Occurrence of brachiopod species from the Kanokura, Hosoo and Kamiyasse Formations in the Setamai, Kamiyasse–Imo and Matsukawa
areas, South Kitakami Belt.

GUADALUPIAN BRACHIOPODS FROM SETAMAI, KAMIYASSE–IMO AND MATSUKAWATABLE 1 (2). Continue

141