ARTICLE

Memoir of the Fukui Prefectural Dinosaur Museum 17: 27–87 (2018)© by the Fukui Prefectural Dinosaur Museum

EARLY CARBONIFEROUS (MISSISSIPPIAN) BRACHIOPODS FROM THE HIKOROICHIFORMATION, SOUTH KITAKAMI BELT, JAPANJun-ichi TAZAWA
Hamaura-cho 1-260-1, Chuo-ku, Niigata 951-8151, Japan
ABSTRACTThis study describes an early Carboniferous brachiopod fauna (the Hikoroichi fauna), consisting of 51 species in 36
genera, including two new species (Leiochonetes onimarensis sp. nov. and Punctospirifer iwatensis sp. nov.), from thelower part (HK1 Unit), the middle part (HK2 Unit) and the upper part (HK4 Unit) of the Hikoroichi Formation in the
Hikoroichi area, South Kitakami Belt, northeastern Japan. The HK1 assemblage (18 species in 16 genera) indicates a late
Tournaisian age, the HK2 assemblage (28 species in 23 genera) indicates an early Visean age, and the HK4 assemblage (15species in 15 genera) indicates a late Visean age. Thus, the Hikoroichi Formation is correlated with the upper Tournaisian–
upper Visean. In terms of palaeobiogeography, the three assemblages have close afﬁnities with the early Carboniferous
brachiopod faunas of North America (Missouri and Iowa), western Europe (England), northern Russia (Verkhoyansk
Range), central Russia (southern Urals and Kuznetsk Basin), Kazakhstan and northwestern China (Xinjiang). In otherwords, the Hikoroichi fauna is allied with those of the North China Province in northwestern‒northeastern China. Thus,
the South Kitakami region was probably located at the eastern extension of the province, namely east to the North China(Sino-Korea) block in the early Carboniferous.
Key words : Brachiopoda, early Carboniferous, Hikoroichi fauna, Japan, South Kitakami Belt
田沢純一（2018）南部北上帯の日頃市層から産出する前期石炭紀（ミシシッピー亜紀）腕足類．福井県立恐竜博物館紀要 17: 27‒87.
南部北上帯日頃市地域に分布する日頃市層の下部（HK1 ユニット）, 中部（HK2 ユニット）, 上部（HK4 ユニット）から産出する 36 属 51 種の前期石炭紀腕足類（日頃市フォーナ）を記載した．これらのなかには 2 種の新種（Leiochonetes onimarensis sp. nov., Punctospirifer iwatensis sp. nov.）が含まれる．HK1 群集（16 属 18 種）
は後期Tournaisian，HK2 群集（23 属 28 種）は前期Visean，HK4 群集（15 属 15 種）は後期Viseanを示す．したがって，日頃市層は上部Tournaisian〜上部Viseanに対比される．古生物地理学的に，HK1，HK2，HK4 群
集はいずれも北アメリカ（ミズーリ州，アイオワ州），西ヨーロッパ（イングランド），ロシア北部（ベルホヤ
ンスク山脈），ロシア中央部（ウラル山脈南部，クズネツク盆地），カザフスタン，中国西北部（新疆）などの
前期石炭紀腕足類フォーナに近縁である．すなわち日頃市フォーナは中国西北部〜東北部に分布する中国北方区の前期石炭紀腕足類フォーナに種属構成が類似する．このことから，前期石炭紀に南部北上帯は中国北方区の東方延長部に位置し，北中国（中朝地塊）の東方に存在したと推定される．

INTRODUCTION(i.e., Hikoroichi-cho, Ofunato City, Iwate Prefecture; Figs. 1 and
2) is a classical and important area for the lower Carboniferous
stratigraphy of Japan as well as the South Kitakami Belt. Since
the work of Onuki (1937) and Yabe and Sugiyama (1937),
many studies have investigated the stratigraphy of the lower
Carboniferous rocks in the Hikoroichi area (e.g., Minato, 1950b;
Okubo, 1951; Minato et al., 1953, 1979b, 1979c; Onuki, 1969;
Mori and Tazawa, 1980; Tazawa, 1981a, 1984b; Kawamura,1983).
The lower Carboniferous rocks of the Hikoroichi area are

The South Kitakami Belt of northeastern Japan contains
excellent sections of lower Carboniferous (Mississippian) rocks.
The Hikoroichi area in the eastern part of the South Kitakami Belt
Received August 29, 2017. Accepted May 8, 2018.
Hamaura-cho 1-260-1, Chuo-ku, Niigata 951-8151, Japan
E-mail: j1025-tazawa＊memoad.jp27

28

JUN-ICHI TAZAWA

FIGURE 1. Maps showing the location and geology of the Hikoroichi area, South Kitakami Belt. (A) Geotectonic map of the Japanese Islands,
showing the distribution of the South Kitakami Belt (based on Tazawa, 2004); (B) geotectonic map of the northeastern part of Honshu, Japan, showing
the distribution of the Carboniferous rocks in the South Kitakami Belt (based on Kawamura et al., 2013); and (C) topographical map showing the
mapped area and fossil localities in the Hikoroichi area, South Kitakami Belt (using the electronic topographical map of GSI).

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

29

FIGURE 2. A distant view (A) and a close view (B) of the outcrop of sandstone and shale of the HK2 Unit of the Hikoroichi Formation in the Onimaru quarry,
Hikoroichi-cho, Ofunato City, Iwate Prefecture. Arrows show the fossil locality KF202. Mt. Goyosan in the background is composed of Cretaceous granitic
rocks.

divided into two formations: the Tournaisian‒upper Visean,
volcaniclastic-dominated Hikoroichi Formation and the upper
Visean, limestone-dominated Onimaru Formation (Minato, 1950b;
Kawamura, 1983; Tazawa, 1984b, 1985). However, the age ofthe lower part of the Hikoroichi Formation is poorly constrained
because of a lack of palaeontological data. Brachiopods are themost common megafossils in the Hikoroichi Formation of the
Hikoroichi area, and 23 species in 20 genera have been described

30

JUN-ICHI TAZAWA

FIGURE 3. Comparison of stratigraphic schemes proposed for the lower Carboniferous formations in the Hikoroichi area, South Kitakami Belt.

by Minato (1951, 1952, 1953), Minato and Kato (1977), Tazawa(1984a, 1984b, 1985, 1989, 1996, 2006) and Tazawa and Ibaraki(2009). However, these brachiopods were obtained and described
mostly from the middle and upper parts of the Hikoroichi
Formation. Thus, the brachiopod species from the lower part of
the Hikoroichi Formation are poorly known.
In this study, I describe brachiopods from the lower part (HK1
Unit), the middle part (HK2 Unit) and the upper part (HK4 Unit)of the Hikoroichi Formation in the Hikoroichi area, and discuss
the age and palaeobiogeography of the Hikoroichi fauna (HK1,HK2 and HK4 assemblages). Most of the materials were collectedby the present author; the rest were provided by M. Harashinai,
T. Kawamura and A. Kaneko. The specimens described herein
are registered and stored in the Department of Geology, Niigata
University, Niigata (NU-B prefix) and the Tohoku University
Museum, Sendai (IGPS preﬁx).
PREVIOUS WORK
Stratigraphy
Previous studies on the stratigraphy of the lower Carboniferous
rocks in the Hikoroichi area are summarized in Fig. 3. In the earlystudies of the area, Onuki (1937) and Yabe and Sugiyama (1937)
named the lower Carboniferous rocks the Kanenokura Stageand the Tyoanzi (Choanji) and Onimaru Groups, respectively.
Minato (1950b) established the Hikoroiti (Hikoroichi) Series(Etroeungtian‒lower Tournaisian) for the lower strata and the
Onimaru Series (upper Visean) for the upper strata. Furthermore,
Minato (1950a) emphasized the pre-Onimaru hiatus as evidenceof the Shizu Folding , one of the Palaeozoic orogenic events
recorded in the South Kitakami region. This proposal was
accepted by many workers (e.g., Okubo, 1951; Minato et al.,1953, 1979b, 1979c; Minato, 1955, 1966; Onuki, 1956, 1969).
Subsequently, Mori and Tazawa (1980) revealed that the Onimaru
Formation conformably overlies the Hikoroichi Formation in the
Hikoroichi area. This new ﬁnding was supported and strengthenedby stratigraphical studies (Tazawa, 1981a, 1984b: Kawamura,1983) and palaeontological studies (Kawamura, 1983; Tazawa,
1984a, b, 1985, 2006).
Recently, Tazawa (2017a) and Tazawa and Niikawa (2018)
described the Late Devonian brachiopods, Cyrtospirifer
choanjiensis, Cyrtospirifer sp. 1, Cyrtospirifer sp. 2 andDesquamatia (Seratrypa) sp., from sandstone and shale that
had been previously assigned to the Hikoroichi Formation at
Choanji, just east to the Hikoroichi area (Fig. 1C). Tazawa andNiikawa (2018) redeﬁned the fossil-bearing sandstone and shaleof Choanji as the Upper Devonian (Frasnian‒Famennian) Choanji
Formation. Thus, the Hikoroichi Formation is now restricted to
the Hikoroichi area and does not occur in the Choanji area.
Palaeontology
Many studies have examined on the fossils of the Hikoroichi

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

Formation, including rugose corals (Kawamura, 1983; Kato etal., 1989), tabulate corals (Kawamura, 1983; Kato et al., 1989),
bryozoans (Sakagami, 1962, 1989), brachiopods (Minato, 1951,
1952, 1953; Minato and Kato, 1977; Tazawa, 1984a, b, 1985,
1989, 1996, 2006; Tazawa and Ibaraki, 2009), gastropods (Kase,
1988, 1989), cephalopods (Niko, 1989), trilobites (Kobayashi and
Hamada, 1980; Kaneko, 1983, 1986, 1989), crinoids (Minato,
1951) and land plants (Asama et al., 1985, 1989). However, mostof these studies focussed on materials from the middle and upperparts of the Hikoroichi Formation.
In terms of biostratigraphy, the upper part of the Hikoroichi
Formation is correlated with the upper Visean on the basisof rugose corals such as Amygdalophyllum etheridgei,
Dibunophyllum sp., Koninckophyllum sp. and Siphonodendron
sp. (Mori and Tazawa, 1980; Kawamura, 1983) and brachiopods
such as Schizophoria resupinata, Orthotetes sp., Pustula cf.
pustulosa, Karavankina sp. and Choristites bisulcatus (Mori and
Tazawa, 1980; Tazawa, 1984b). The middle part of the formationis correlated with the lower Visean on the basis of rugose corals
such as Amygdalophyllum etheridgei, Lophophyllidium sp.
and Multithecopora sp. (Kawamura, 1983) and brachiopods such
as Marginatia burlingtonensis, Syringothyris transversa, Rotaia
hikoroichiensis, Linoprotonia ashfellensis and Gigantoproductus
cf. crassiventer (Tazawa, 1985, 2006; Tazawa and Ibaraki, 2009).
However, the age of the lower part of the Hikoroichi Formationis unclear. Kawamura (1983) suggested a late Tournaisian age
based on rugose corals, Caninia cornucopiae, Palaeosmiliasp., Amygdalophyllum sp. and Cyathaxonia sp., although thecorals remain undescribed. Minato (1950b) considered thatthe Hikoroichi Series is correlated with the Etroeungtian‒lower Tournaisian based on brachiopods, mostly from Choanji.
However, as mentioned above, the fossil-bearing beds of Choanji
are assigned to the Upper Devonian Choanji Formation, not to the
Hikoroichi Formation.
In terms of palaeobiogeography, Tazawa (2002, 2006)
concluded that the South Kitakami region was the easternextension of the Tianshan‒Jilin Province of Yang (1994), and was
located on the continental shelf bordering the eastern margin ofNorth China (Sino-Korea) in the early Visean, as inferred from
the Marginatia‒Syringothyris‒Rotaia brachiopod assemblage
from the middle part of the Hikoroichi Formation and equivalentfaunas in the Shimoarisu and Yokota areas in the South Kitakami
Belt.
STRATIGRAPHY AND MATERIALS
Carboniferous rocks crop out in the Hikoroichi area, forming
a syncline that plunges gently to the north (Fig. 4). The strata are
subdivided lithologically into three formations: the Hikoroichi
Formation (upper Tournaisian‒upper Visean; this study), theOnimaru Formation (upper Visean; Minato and Kato, 1979)
and the Nagaiwa Formation (Serpukhovian‒Moscovian; Katoand Nakamura, 1979), in ascending order (Fig. 5). According
to Tazawa (1984b), the Hikoroichi Formation consists mostlyof sandstone, with subordinate shale, conglomerate, tuff and31

arenaceous limestone, and has a total thickness of 440‒560 m.
The formation overlies the Middle Devonian Nakazato Formation
across an angular unconformity, and is in turn conformably
overlain by the Onimaru Formation.
In this paper, I propose three parts (the lower, middle and upperparts) and four lithostratigraphic units (the HK1, HK2, HK3 andHK4 units) in the Hikoroichi Formation. The lower part consistsof the HK1 Unit (mainly sandstone with subordinate shale, felsic
tuff and arenaceous limestone, 133‒207 m in total thickness); themiddle part consists of both the HK2 Unit (mainly shale, withsubordinate sandstone and arenaceous limestone, 67‒100 m intotal thickness) and the HK3 Unit (mainly andesitic to basaltic
tuff, with subordinate sandstone, shale and arenaceous limestone,
40‒87 m in total thickness); and the upper part consists ofthe HK4 Unit (mainly sandstone, with subordinate shale andarenaceous limestone, 147‒240 m in total thickness). Brachiopods
described herein were collected from 16 localities (stations):
three localities (KAC3, KF200 and KF201) in the HK1 Unit, ﬁve
localities (KF171, KF172, KF174, KF188 and KF202) in the HK2
Unit, and eight localities (KF156, KF167, KAC1, KF148, KF163,
KF152, KF151 and KF153) in the HK4 Unit. The topographic
and stratigraphic locations, lithologies and brachiopod speciesof the 16 fossil localities are indicated in Figs. 1, 5, 6 andsummarized in Appendix.
HIKOROICHI FAUNA
The brachiopods described herein are the following 51
species in 36 genera, including two new species (Leiochonetes
onimarensis sp. nov. and Punctospirifer iwatensis sp. nov.):
Leptagonia analoga (Phillips, 1836), Rugosochonetes cf.
hardrensis (Phillips, 1841), Plicochonetes sp., Leiochonetes
onimarensis sp. nov., Argentiproductus margaritaceus(Phillips, 1836), Tolmatchoffia robusta (Tolmatchoff, 1924),
Marginatia burlingtonensis (Hall, 1858), M. magna Carter,
1968, Echinoconchus punctatus (Sowerby, 1822), Karavankina
jizodoensis Tazawa, 1980, Pustula pustulosa (Phillips, 1836),
Ovatia laevicosta (White, 1860), O. cf. prolata Carter, 1987,
Ovatia sp., Linoprotonia ashfellensis Ferguson, 1971, L.
hikoroichiensis Tazawa and Ibaraki, 2009, Gigantoproductus
crassiventer (Prentice, 1949), G. latiexpansus Sarytcheva in
Sarytcheva and Sokolskaya, 1952, Orthotetes keokuk (Hall, 1858),
Rhipidomella michelini (Léveillé, 1835), Schizophoria resupinata(Martin, 1809), Rhynchopora sp., Rotaia hikoroichiensis Tazawa,
2006, Actinoconchus expansus (Phillips, 1836), A. paradoxus
MʼCoy, 1844, Lamellosathyris qaidamensis Chen, Shi andZhan, 2003, Cleiothyridina harkeri Carter, 1987, Cleiothyridinasp., Hustedia sp., Spirifer liangchowensis Chao, 1929, S.
karagai (Litvinovich, 1962), Unispirifer striatoconvolutus(Dun and Benson, 1920), U. mediocris (Tolmatchoff, 1924),
U. kozuboensis (Minato, 1952), Angiospirifer bisulcatus(Sowerby, 1825), Tegulocrea incerta (Hall, 1858), Acuminothyris
triangularis Roberts, 1963, A. keokuk Carter, 1990, Brachythyris
suborbicularis (Hall, 1858), Tylothyris laminosa (MʼCoy, 1844),
Kitakamithyris tyoanjiensis Minato, 1951, K. hikoroitiensis

32

JUN-ICHI TAZAWA

FIGURE 4. Geologic map of the Hikoroichi area (after Tazawa, 1984b). 1, 2, Lower Permian Sakamotozawa Formation (1, limestone; 2, sandstone); 3, 4,
upper Carboniferous Nagaiwa Formation (3, limestone; 4, sandstone); 5, limestone of the lower Carboniferous Onimaru Formation; 6‒8, lower Carboniferous
Hikoroichi Formation (6, sandstone and shale; 7, tuff of the HK3 Unit; 8, limestone); 9, Middle Devonian Nakazato Formation; 10, Lower Devonian Ono
Formation; 11, Silurian Kawauchi Formation; 12, conglomerate of the Hikoroichi, Nagaiwa and Sakamotozawa Formations; 13, shale of the Onimaru, Nagaiwa
and Sakamotozawa Formations; 14, tuff of the HK1 and HK2 units of the Hikoroichi Formation and the Nagaiwa Formation; 15, granitic rocks of the Hikami
Granite; 16, dyke rocks.

Minato, 1951, Torynifer asiatica Besnossova in Besnossova
et al., 1962, Syringothyris sibirica Sokolskaya in Sarytcheva
et al., 1963, Pseudosyrinx cf. missouriensis Weller, 1914,
Dimegelasma sp., Punctospirifer plicatosulcatus Glenister, 1955,

P. gigantiformis Lee and Gu in Lee et al., 1980, P. iwatensis sp.
nov., Punctospirifer sp. and Beecheria khalfini Besnossova in
Sarytcheva et al., 1963.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

33

FIGURE 5. Generalized columnar section of the Carboniferous formations in the Hikoroichi area, showing the fossil horizons in the HK1, HK2 and HK4
units in the Hikoroichi Formation (modiﬁed and adapted from Tazawa, 1984b). 1, Shale; 2, sandstone; 3, conglomerate; 4, limestone (Nagaiwa Formation); 5,
limestone (Onimaru Formation); 6, limestone (Hikoroichi Formation); 7, tuff.

HK1 assemblage (late Tournaisian)
The HK1 assemblage includes 18 species in 16 genera:
Leptagonia analoga, Plicochonetes sp., Tolmatchoffia robusta,
Marginatia burlingtonensis, M. magna, Ovatia sp., Schizophoria
resupinata, Rhynchopora sp., Cleiothyridina harkeri, Unispirifer
mediocris, Acuminothyris keokuk, Brachythyris suborbicularis,

Tylothyris laminosa, Kitakamithyris tyoanjiensis, K. hikoroitiensis,
Syringothyris sibirica, Punctospirifer sp. and Beecheria khalfini.
Of these species, Leptagonia analoga, Tolmatchoffia robusta
and Acuminothyris keokuk are abundant; Marginatia magna,
Unispirifer mediocris and Syringothyris sibirica are common; andthe others are rare (Fig. 6).
The stratigraphic distributions of the brachiopod species

34

JUN-ICHI TAZAWA

FIGURE 7. Stratigraphic distributions of brachiopod species of the HK1
assemblage in the Hikoroichi fauna. Broken lines show those of the genera.

FIGURE 6. Occurrence of brachiopod species from the Hikoroichi
Formation in the Hikoroichi area, South Kitakami Belt. A, abundant; C,
common; R, rare.

of the HK1 assemblage are summarized in Fig. 7. Of thebrachiopods listed above, Marginatia magna is known only
from the upper Tournaisian, Cleiothyridina harkeri is known
from the lower‒upper Tournaisian, and Kitakamithyris
tyoanjiensis has a range of upper Famennian‒upper Tournaisian.
In contrast, Acuminothyris keokuk and Syringothyris sibirica
are known from the upper Tournaisian‒lower Visean, four
species (Marginatia burlingtonensis, Schizophoria resupinata,
Brachythyris suborbicularis and Tylothyris laminosa) occur inthe upper Tournaisian‒upper Visean, three species (Tolmatchoffia
robusta, Unispirifer mediocris and Beecheria khalfini) range
from the lower Tournaisian‒lower Visean, and Kitakamithyris
hikoroitiensis has a range of upper Fammenian‒lower Visean.
Leptagonia analoga is a long-ranging species, known from thelower Tournaisian to the upper Visean. At the generic level,
Plicochonetes is known from the lower Tournaisian‒Serpukhovian(Racheboeuf, 2000), Ovatia ranges from the lower Tournaisianto the Serpukhovian (Brunton et al., 2000), Rhynchopora is
known from the upper Tournaisian‒Gzhelian (Savage, 2002),
and Punctospirifer is known from the upper Famennian to theupper Visean (Carter, 2006b). In summary, the age of the HK1
assemblage is identiﬁed as late Tournaisian. Thus the lower partof the Hikoroichi Formation (HK1 Unit) is correlated with theupper Tournaisian.
The geographic distributions of the brachiopod species ofthe HK1 assemblgae are summarized in Figs. 8 and 9. Amongthe 18 species of the HK1 assemblage, seven species also occur
in central Russia (Kuznetsk Basin); six species also occur in
Kazakhstan; ﬁve species also occur in the USA (Missouri), central
Russia (southern Urals) and northwestern China (Xinjiang); four
species also occur in northern Russia (Verkhoyansk Range);
and three species also occur in southwestern Japan (Akiyoshi

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

35

FIGURE 8. Geographic distributions of brachiopod species of the HK1 assemblage in the Hikoroichi fauna.

FIGURE 9. Late Tournaisian reconstruction map of the world (adapted from Qiao and Shen, 2014), showing the geographic distributions of brachiopod speciesof the HK1 assemblage in the Hikoroichi fauna. Solid circles indicate numbers of brachiopod species listed in the HK1 assemblage. Station numbers are sameas in Fig. 8.

36

JUN-ICHI TAZAWA

Belt), the USA (Iowa), Canada (Alberta), the UK (England)
and Belgium. In summary, the HK1 assemblage has a close
afﬁnity with the early Carboniferous (Mississippian) brachiopod
faunas of North America, western Europe, northern and central
Russia, Kazakhstan and northwestern China. Thus, the HK1
assemblage is allied with those of the North China Province ofYang (1983), which covered northwestern‒northeastern China inearly Carboniferous, and is characterized by the Boreal elements
such as Syringothyris, Pseudosyrinx, Rotaia, Grandispirifer andTolmatchoffia, but lacks the South China (Tethyan) elements suchas Yanguania, Eochoristites and Martiniella. According to Yang
(1983), the faunas of the North China Province are essentiallysimilar to those of northeastern Siberia and North America.
HK2 assemblage (early Visean)
The HK2 assemblage includes 28 species in 23 genera:
Leptagonia analoga, Rugosochonetes cf. hardrensis, Leiochonetes
onimarensis, Tolmatchoffia robusta, Marginatia burlingtonensis,
Echinoconchus punctatus, Ovatia laevicosta, Linoprotonia
ashfellensis, L. hikoroichiensis, Gigantoproductus crassiventer,
G. latiexpansus, Orthotetes keokuk, Rhipidomella michelini,
Schizophoria resupinata, Rotaia hikoroichiensis, Actinoconchus
expansus, A. paradoxus, Lamellosathyris qaidamensis,
Hustedia sp., Spirifer karagai, Unispirifer striatoconvolutus, U.
kozuboensis, Tegulocrea incerta, Acuminothyris triangularis,
Kitakamithyris hikoroitiensis, Pseudosyrinx cf. missouriensis,
Punctospirifer plicatosulcatus and P. gigantiformis. Of these,
Leiochonetes onimarensis, Tolmatchoffia robusta, Marginatia
burlingtonensis, Linoprotonia ashfellensis, L. hikoroichiensis,
Lamellosathyris qaidamensis, Unispirifer striatoconvolutus, U.
kozuboensis and Tegulocrea incerta are abundant; Echinoconchus
punctatus, Gigantoproductus crassiventer, G. latiexpansus,
Orthotetes keokuk, Rhipidomella michelini, Schizophoria
repupinata, Actinoconchus expansus and Spirifer karagai arecommon; and the others are rare (Fig. 6).
The stratigraphic distributions of the brachiopod speciesof the HK2 assemblage are summarized in Fig. 10. Of thebrachiopods listed above, Linoprotonia hikoroichiensis andActinoconchus paradoxus are known only from the lower Visean,
five species (Ovatia laevicosta, Lamellosathyris qaidamensis,
Unispirifer kozuboensis, Tegulocrea incerta and Acuminothyris
triangularis) are known from the upper Tournaisian‒lower Visean, Tolmatchoffia robusta and Punctospirifer
plicatosulcatus are known from the lower Tournaisian‒lower
Visean, and Kitakamithyris hikoroitiensis is known fromthe upper Famennian‒lower Visean. In contrast, five species(Linoprotonia ashfellensis, Gigantoproductus crassiventer, Rotaia
hikoroichiensis, Spirifer karagai and Punctospirifer gigantiformis)
are known from the lower‒upper Visean, and Gigantoproductus
latiexpansus and Actinoconchus expansus are known from thelower Visean‒Serpukhovian. Three species are long-ranging
species: Leptagonia analoga ranges from the lower Tournaisianto the upper Visean, Echinoconchus punctatus ranges from theupper Tournaisian to the Gzhelian, and Rhipidomela michelini

FIGURE 10. Stratigraphic distributions of brachiopod species of the HK2
assemblage in the Hikoroichi fauna. Broken lines show those of the genera.

ranges from the upper Tournaisian to the Bashkirian. At thegeneric level, Rugosochonetes has a long range from Tournaisianto Gzhelian (Racheboeuf, 2000), Leiochonetes is known from theupper Tournaisian‒upper Visean (Carter, 1967; Roberts, 1976),
Hustedia is known from the Tournaisian‒Changhsingian (Alvarez
and Rong, 2002b), and Pseudosyrinx is known from the upper
Tournaisian‒lower Visean (Carter, 2006a). In summary, the ageof the HK2 assemblage is identified as early Visean. Thus, themiddle part of the Hikoroichi Formation (HK2 Unit) is correlatedwith the lower Visean.
The geographic distributions of the brachiopod species of theHK2 assemblage are summarized in Figs. 11 and 12. Amongthe 28 species of the HK2 assemblage, seven species also occur
in the UK (England) and Kazakhstan; six species also occur inMISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

37

FIGURE 11. Geographic distributions of brachiopod species of the HK2 assemblage in the Hikoroichi fauna.

FIGURE 12. Early Visean reconstruction map of the world (adapted from Qiao and Shen, 2014), showing the geographic distributions of brachiopod species ofthe HK2 assemblage in the Hikoroichi fauna. Solid circles indicate numbers of brachiopod species listed in the HK2 assemblage. Station numbers are same as
in Fig. 11.

38

JUN-ICHI TAZAWA

northwestern China (Xinjiang); five species also occur in the
USA (Iowa) and central Russia (southern Urals and Kuznetsk
Basin); four species also occur in the USA (Oklahoma), Belgium,
western Russia (Moscow Basin and Donetz Basin), northwestern
China (Qinghai) and southwestern China (Yunnan); and three
species also occur in southwestern Japan (Akiyoshi Belt), theUSA (Missouri), northern Russia (Taimyr Peninsula and Pechora
Basin), the UK (Wales), Spain, Kyrgyzstan and eastern Australia.
In summary, the HK2 assemblage has a close affinity with theearly Carboniferous (Mississippian) brachiopod faunas of North
America, western Europe (England and Belgium), northern,western and central Russia, Kazakhstan and northwestern China(Xinjiang and Qinghai). Thus, the HK2 assemblage also is alliedwith those of the North China Province of Yang (1983).
HK4 assemblage (late Visean)
The HK4 assemblage includes 15 species in 15 genera:
Leptagonia analoga, Argentiproductus margaritaceus, Karavankina
jizodoensis, Pustula pustulosa, Ovatia cf. prolata, Orthotetes
keokuk, Rhipidomella michelini, Schizophoria resupinata, Rotaia
hikoroichiensis, Cleiothyridina sp., Spirifer liangchowensis,
Angiospirifer bisulcatus, Torynifer asiatica, Dimegelasma sp. andPunctospirifer iwatensis. Of these species, Schizophoria resupinata
is abundant; Rhipidomella michilini and Torynifer asiatica arecommon; and the other species are rare (Fig. 6).
The stratigraphic distributions of the brachiopod species of theHK4 assemblage are summarized in Fig. 13. Of the brachiopods
listed above, Orthotetes keokuk and Schizophoria resupinata
are known from the upper Tournaisian‒upper Visean, Rotaia
hikoroichiensis and Spirifer liangchowensis are known from thelower‒upper Visean, and Karavankina jizodoensis is restricted tothe upper Visean. In contrast, Angiospirifer bisulcatus is known
from the upper Visean‒Bashkirian, four species (Argentiproductus
margaritaceus, Pustula pustulosa, Rhipidomella michelini andTorynifer asiatica) are long-ranging species, though within thelower Tournaisian‒Bashkirian. At the generic level, Ovatia is
known from the lower Tournaisian‒Serpukhovian (Brunton
et al., 2000), Cleiothyridina ranges from the Famennian to
the Wuchiapingian (Alvarez and Rong, 2002a), Dimegelasma
is known from the lower‒upper Visean (Carter, 2006a), andPunctospirifer is known from the upper Famennian‒upper
Visean (Carter, 2006b). In summary, the age of the HK4 fauna is
identiﬁed as late Visean. Thus, the upper part of the Hikoroichi
Formation (HK4 Unit) is correlated with the upper Visean.
The geographic distributions of the brachiopod species of theHK4 assemblage are summarized in Figs. 14 and 15. Among the
15 species of the HK4 assemblage, six species also occur in the
UK (England), Kazakhstan and northwestern China (Xinjiang);
five species also occur in northern Russia (Pechora Basin), theUK (northern Ireland), Belgium, western Russia (Donetz Basin),central Russia (southern Urals) and Uzbekistan; four species
also occur in northern Russia (Verkhoyansk Range), Germany
and Poland; and three species also occur in northern Russia(Taimyr Peninsula), the UK (Wales and Isle of Man), western

FIGURE 13. Stratigraphic distributions of brachiopod species of the HK4
assemblage in the Hikoroichi fauna. Broken lines show those of the genera.

Russia (Moscow Basin), central Russia (Kuznetsk Basin),
Kyrgyzstan and northwestern China (Gansu). In summary, theHK4 assemblage has a close afﬁnity with the early Carboniferous(Mississippian) brachiopod faunas of the UK, northern, westernand central Russia, Kazakhstan, Uzbekistan and northwestern
China. Thus, the HK4 assemblage also is allied with those of theNorth China Province of Yang (1983).
CONCLUSIONSIn this study, an early Carboniferous (Mississippian)brachiopod fauna (Hikoroichi fauna: HK1, HK2 and HK4
assemblages), consisting of 51 species in 36 genera (including
two new species, Leiochonetes onimarensis sp. nov. andPunctospirifer iwatensis sp. nov.) are described from the lower
part (HK1 Unit), the middle part (HK2 Unit) and the upper part
(HK4 Unit) of the Hikoroichi Formation in the Hikoroichi area,
South Kitakami Belt, northeastern Japan. The HK1 assemblage,
consisting of 18 species in 16 genera, indicates a late Tournaisianage; the HK2 assemblage, consisting of 28 species in 23 genera,
indicates an early Visean age; and the HK4 assemblage, consistingof 15 species in 15 genera, indicates a late Visean age. Thus, theHikoroichi Formation is correlated with the upper Tournaisian–
upper Visean. In terms of palaeobiogeography, the HK1, HK2
and HK4 assemblages have close affinities with the early
Carboniferous brachiopod faunas of North America (Missouri andIowa), western Europe (England), northern Russia (Verkhoyansk
Range), central Russia (southern Urals and Kuznetsk Basin),MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

39

FIGURE 14. Geographic distributions of brachiopod species of the HK4 assemblage in the Hikoroichi fauna.

FIGURE 15. Late Visean reconstruction map of the world (adapted from Qiao and Shen, 2014), showing the geographic distributions of brachiopod species ofthe HK4 assemblage in the Hikoroichi fauna. Solid circles indicate numbers of brachiopod species listed in the HK4 assemblage. Station numbers are same as
in Fig. 14.

Kazakhstan and northwestern China (Xinjiang). In other words,
these assemblages are allied with those of the North China
Province of Yang (1983). Thus, the South Kitakami region wasprobably the eastern extension of the province, and located east to
the North China (Sino-Korea) block in the early Carboniferous.

SYSTEMATIC DESCRIPTIONSThe suprageneric classiﬁcation given herein mainly follows thatof Treatise on Invertebrate Paleontology, Part H Brachiopoda
Revised, Volumes 2‒6 , edited by Kaesler (2000a, b, 2002,
2006) and Selden (2007), with exception that classiﬁcation of the

40

JUN-ICHI TAZAWA

Productidina follows Waterhouse (2002).
Order STROPHOMENIDA Öpik, 1934
Superfamily STROPHOMENOIDEA King, 1846
Family RAFINESQUINIDAE Schuchert, 1893
Subfamily LEPTAENINAE Hall and Clarke, 1894
Genus LEPTAGONIA MʼCoy, 1844
Type species.―Producta analoga Phillips, 1836.
Leptagonia analoga (Phillips, 1836)(Figs. 16B‒F, 22A, B, 30A)
Producta analoga Phillips, 1836, p. 116, pl. 7, ﬁg. 10.
Strophomena rhomboidalis var. analoga Phillips. Davidson, 1861,
p. 119, pl. 28, ﬁgs. 1‒6, 9‒13; Etheridge, 1872, p. 333, pl. 18,
ﬁg. 1.
Leptaena analoga (Phillips). Weller, 1914, p. 49, pl. 2, ﬁgs. 1‒10;
Frech, 1916, p. 237, pl. 2, ﬁgs. 2, 3d; Girty, 1920, pl. 54, ﬁg. 3;
Tolmatchoff, 1924, p. 209, 569, pl. 13, ﬁg. 8; Girty, 1927, pl.
22, ﬁgs. 6‒8; Demanet, 1934, p. 61, pl. 5, ﬁgs. 1‒14, text-ﬁgs.
1‒14; Branson, 1938, p. 24, pl. 5, ﬁg. 31; Nelson, 1961, pl. 4,
ﬁg. 26; Sokolskaya in Sarytcheva et al., 1963, p. 80, pl. 4, ﬁgs.
9‒14.
Leptaena rhomboidalis Wilckens. Sommer, 1909, p. 626, pl. 29,
ﬁg. 14.
Leptagonia cf. analoga (Phillips). Cvancara, 1958, p. 860, pl.
110, ﬁgs. 6‒13, text-ﬁgs. 3, 4.
Leptaenella analoga (Phillips). Yang, 1964, p. 61, pl. 1, fig. 5;
Grechishnikova, 1966, p. 94, pl. 1, figs. 19, 20; pl. 2, figs.
1‒6; Abramov, 1970, p. 108, pl. 1, ﬁgs. 11, 12; Aisenverg and
Poletaev, 1971, pl. 60, ﬁgs. 2, 3; Nalivkin and Fotieva, 1973, p.
20, pl. 1, ﬁgs. 9‒13.
Leptagonia analoga (Phillips). Brunton, 1968, p. 29, pl. 3, ﬁgs.
26‒31; pl. 4, ﬁgs. 1‒9; text-ﬁgs. 6‒17; Gaetani, 1968, p. 688,
pl. 47, ﬁg. 3; Thomas, 1971, p. 30, pl. 18, ﬁgs. 1‒8, text-ﬁg.
11; Bublichenko, 1971, p. 37, pl. 3, figs. 1‒5; Brand, 1972,
p. 59, pl. 8, figs. 1‒6, text-figs. 1a, 3; Kalashnikov, 1974, p.
23, pl. 3, ﬁg. 5; Litvinovich et al., 1975, p. 53, pl. 16, ﬁg. 11;
Bublichenko, 1976, p. 22, pl. 1, ﬁg. 10; Yang et al., 1977, p.
316, pl. 131, ﬁg. 2; Nalivkin, 1979, p. 18, pl. 3, ﬁgs. 1‒3, 5,
6; Ding and Qi, 1983, p. 251, pl. 89, figs. 9, 10, 12; Zhang
et al., 1983, p. 271, pl. 107, ﬁg. 13; pl. 106, ﬁg. 3; Tazawa etal., 1984, p. 350, pl. 67, ﬁgs. 2‒4; Yang, 1984, p. 205, pl. 29,
ﬁg. 11; Xu and Yao, 1988, p. 274, pl. 67, ﬁgs. 4, 6‒10; Carter,
1999, p. 96, figs. 1A‒E; Shi et al., 2005, p. 39, figs. 3A, E;
Tazawa et al., 2016, p. 51, ﬁg. 5.5; Tazawa, 2017b, p. 333, ﬁg.
6.1.
Material.―Fourteen specimens from localities KAC1, KF172,
KF201 and KF202: (1) external and internal mould of a ventral
valve, NU-B2122; (2) external moulds of two ventral valves, NUB2081, 2082, (3) internal mould of a ventral valve, NU-B2123;(4) external and internal moulds of two dorsal valves, NU-B2125,
2126; (5) external moulds of six dorsal valves, NU-B2083, 2124,

2127‒2129, 2239; and (6) internal moulds of two dorsal valves,
NU-B2130, 2131.
Description.―Shell large in size for genus, transversely
trapezoidal in outline, hinge equal to greatest width; length 34
mm, width 66 mm in the largest dorsal valve specimen (NUB2127). Ventral valve gently convex in visceral disc, strongly
geniculated, and followed by short trail; umbo small; sulcus
absent. Dorsal valve flat to slightly concave in visceral disc,strongly geniculated, and followed by a short trail; fold absent;
marginal ridge around visceral disc often developed in adult shell.
External surface of both valves ornamented with fine rounded
costellae and regular, slightly ﬂexuous rugae; numbering 10‒11
costellae in 5 mm at about midlength, and 16 ‒17 rugae invisceral disc of ventral valve. Ventral interior with strong teeth
and large subcircular pseudospondylium, about one-third as longas length of disc. Dorsal interior with a bilobate cardinal process
supported by low divergent socket ridges; adductor scars raised,
pear-shaped, and divided by a long median septum extending forabout two-ﬁfths length of disc.
Remarks.―These specimens can be referred to Leptagonia
analoga (Phillips, 1836), redescribed by Thomas (1971, p.
30, pl. 18, figs. 1‒8, text-fig. 11) from the upper Tournaisianof the Bonaparte Gulf Basin, western Australia, in generalshape, external ornament and internal structure of both valves,particularly, the large transversely trapezoidal shell, ornamented
with numerous costellae and regular concentric rugae, and inhaving a prominent marginal ridge around visceral disc of dorsal
valve and raised muscle ﬁeld on both ventral and dorsal valves.
Comparison with the other species of Leptagonia was fully
discussed by Brand (1972).
Occurrence.―HK1, HK2 and HK4 units.
Distribution.―Lower Tournaisian‒upper Visean: northeasternJapan (Hikoroichi, Yokota and Soma in the South Kitakami Belt),southwestern Japan (Hina in the Akiyoshi Belt), USA (Illinois,
Iowa, Missouri, Oklahoma, Utah and New Mexico), western
Canada (Alberta), northern Russia (Verkhoyansk Range andPechora Basin), UK (England, lsle of Man and northern Ireland),Germany, Belgium, western Russia (Donetz Basin), Turkey(Taurus Mountains), Iran (Elburz Range), central Russia (southern
Urals and Kuznetsk Basin), Kazakhstan, northwestern China(Xinjiang and Gansu), central-southern China (Hubei, Guangdong
and Guangxi), southwestern China (Yunnan), western Australia(Bonaparte Gulf Basin) and eastern Australia (Queensland and
New South Wales).
Order PRODUCTIDA Sarytcheva and Sokolskaya, 1959
Suborder CHONETIDINA Muir-Wood, 1955
Superfamily CHONETOIDEA Bronn, 1862
Family RUGOSOCHONETIDAE Muir-Wood, 1962
Subfamily RUGOSOCHONETINAE Muir-Wood, 1962
Genus RUGOSOCHONETES Sokolskaya, 1950
Type species.―Orthis hardrensis Phillips, 1841.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

41

FIGURE 16. Brachiopods of the HK1 assemblage (1). A, Plicochonetes sp., external latex cast (A1, A2) and external mould (A3) of ventral valve, NU-B2173;
B‒F, Leptagonia analoga (Phillips); B, external latex cast (B1) and internal latex cast (B2, B3) of ventral valve, NU-B2122; C, internal mould of dorsal valve,
NU-B2130; D, internal mould of ventral valve, NU-B2123; E, external latex cast of dorsal valve, NU-B2127; F, external mould (F1), external latex cast (F2),internal mould (F3) and internal latex cast (F4, F5) of dorsal valve, NU-B2126. Scale bars represent 1 cm.

Rugosochonetes cf. hardrensis (Phillips, 1841)(Fig. 22C)
cf. Rugosochonetes hardrensis (Phillips). Muir-Wood, 1962, p.
66‒68, pl. 8, ﬁgs. 7‒10, 13.
Material.―One specimen from locality KF202, external andinternal moulds of a dorsal valve, NU-B2042.
Remarks.―The single dorsal valve specimen from the
Onimaru quarry, Hikoroichi is safely assigned to the genus
Rugosochonetes by the small (length 8 mm, width 8 mm),subquadrate and ﬂat to slightly concave dorsal valve, ornamented
with numerous capillae (numbering 6 in 1 mm at midlengthof valve) and in having strong diverging socket ridges. The
Hikoroichi species somewhat resembles Rugosochonetes
hardrensis (Phillps, 1841), redescribed by Muir-Wood (1962,
p. 66, pl. 8, ﬁgs. 7‒10, 13) from the upper Visean of Yorkshire,
England, in size, outline and external ornament of the dorsal
valve. However accurate comparison is difficult for the poorly
preserved material.
Occurrence.―HK2 Unit.

42

JUN-ICHI TAZAWA

Subfamily PLICOCHONETINAE Sokolskaya, 1960
Genus PLICOCHONETES Paeckelmann, 1930
Type species.―Chonetes buchianus de Koninck, 1843.
Plicochonetes sp.
(Fig. 16A)
Material.―One specimen from locality KF201, externalmould of a ventral valve, NU-B2173.
Remarks.―The single specimen from Hikoroichi is safely
assigned to the genus Plicochonetes by the small, transverse andstrongly convex ventral valve (length 7 mm, width 10 mm), withlarge ears and external ornament consisting of coarse costellae(numbering 6‒7 in 3 mm at about midlength) and very ﬁne growthlines. This specimen most resembles the shells, described by
Litvinovich et al. (1975, p. 57, pl. 17, ﬁgs. 7, 8) as Plicochonetes
glenparkensis (Weller, 1906), from the lower Tournaisian ofcentral Kazakhstan, in having large ears on the ventral valve.
The type species, Plicochonetes buchianus (de Koninck, 1843),
reﬁgured by Muir-Wood (1962, pl. 15, ﬁgs. 1‒4), from the upper
Visean and Serpukhovian of England, differs from the present
species in the larger size and in having fewer and coarser costellaeon the ventral valve.
Occurrence.―HK1 Unit.
Subfamily SVALBARDIINAE Archbold, 1982
Genus LEIOCHONETES Roberts, 1976
Type species.―Leiochonetes salisburyensis Roberts, 1976.
Leiochonetes onimarensis sp. nov.
(Fig. 22D‒F)
Etymology.―Named after the fossil locality, Onimaru in the
Hikoroichi-cho, Ofunato City, Iwate Prefecture.
Material.―Six specimens from locality KF202: (1) external
and internal moulds of two ventral valves, NU-B2136 (holotype),2137; (2) internal mould of a ventral valve, NU-B2138; and (3)external moulds of three dorsal valves, NU-B2139‒2141.
Diagnosis.―Large-sized and moderately concavo-convex
Leiochonetes, with blunt, acute cardinal extremities.
Description.―Shell large in size for genus, transversely
semicircular in outline, with blunt, acute cardinal extremities,
widest at hinge; length 11 mm, width 20 mm in the largest dorsal
valve specimen (NU-B2139). Ventral valve moderately convex
in lateral proﬁle, most convex at about mid-length; umbo small;sulcus absent. Dorsal valve moderately concave in lateral proﬁle,with deepest at mid-length; fold absent. External surface of both
valves smooth; concentric growth lines not well preserved andobscure; hinge spines few in number near postero-lateral margins
and diverging from hinge at about 60 . Ventral interior with shortmedian septum, being one-fourth to one-ﬁfth of valve length.
Remarks.―These specimens are safely assigned to the genus
Leiochonetes by the small size, transversely semicircular outline

and moderately concavo-convex shell, with smooth externalsurface and lacking both ventral sulcus and dorsal fold. The typespecies, Leiochonetes salisburyensis Roberts (1976, p. 27, pl.
3, ﬁgs. 1‒9, text-ﬁgs. 6, 7), from the Flagstaff Sandstone (upper
Visean) of Brownmore, New South Wales, eastern Australia,differs from L. onimarensis by the smaller size and more acutecardinal extremities. Leiochonetes transversa (Carter, 1967, p.
274, pl. 14, figs. 15‒20), from the Chapel Limestone of central
Texas, differs from the present new species by the much smaller
and more transverse shell.
Occurrence.―HK2 Unit.
Suborder PRODUCTIDINA Waagen, 1883
Superfamily PRODUCTELLOIDEA Schuchert, 1929
Family PRODUCTINIDAE Muir-Wood and Cooper, 1960
Subfamily PRODUCTININAE Muir-Wood and Cooper, 1960
Genus ARGENTIPRODUCTUS Cooper and Muir-Wood, 1951
Type species.―Producta margaritacea Phillips, 1836.
Argentiproductus margaritaceus (Phillips, 1836)(Fig. 30B)
Producta margaritacea Phillips, 1836, p. 215, pl. 8, ﬁg. 8.
Productus margaritacea Phillips. Davidson, 1861, p. 159, pl. 44,
ﬁgs. 5‒8.
Productus cf. margaritaceus Phillips. Yanishevsky, 1918, p. 45,
pl. 8, ﬁg. 38.
Productus margaritaceus Phillips. Delépine, 1928, p. 22, pl. 3,
ﬁgs. 35, 36; Rotai, 1931, p. 55, pl. 3, ﬁg. 4.
Productus (Thomasina) margaritaceus Phillips. Paeckelmann,
1931, p. 184, pl. 17, ﬁgs. 10‒12.
Thomasella margaritacea (Phillips). Sarytcheva and Sokolskaya,
1952, p. 96, pl. 15, ﬁg. 106.
Argentiproductus margaritaceus (Phillips). Muir-Wood andCooper, 1960, pl. 123, figs. 11‒16 only; Litvinovich et al.,1969, p. 155, pl. 7, figs. 18, 19; Volgin and Kushnar, 1975,
p. 40, pl. 3, ﬁgs. 2‒4; Bublichenko, 1976, p. 32, pl. 4, ﬁg. 5;
Brunton et al., 1993, p. 103, figs. 13‒15; Sun and Baliński,2008, p. 506, ﬁgs. 14‒16.
Productina margaritacea (Phillips). Brunton, 1966, p. 209, pl. 8,
ﬁgs. 1‒19; pl. 15, ﬁgs. 1‒8; Roberts, 1971, p. 94, pl. 17, ﬁgs.
11‒28; Kalashnikov, 1974, p. 45, pl. 15, ﬁgs. 1‒4; Galitskaya,
1977, p. 25, pl. 5, ﬁgs. 1‒3; Donakova, 1978, p. 206, pl. 1, ﬁgs.
2, 3; Zakowa, 1985, p. 307, pl. 4, ﬁg. 3; pl. 6, ﬁg. 2.
Material.―One specimen from locality KAC1, external andinternal moulds of a dorsal valve, NU-B2235.
Remarks.―The single dorsal valve specimen from Hikoroichi
is medium in size (length 17 mm, width 28 mm), transversely
hemispherical in outline, and strongly concave but not geniculate
in lateral proﬁle. External surface of the valve is ornamented byflattened costae (numbering 5‒6 in 5 mm) on the trail, slightly
irregular concentric lamellae on the visceral disc and numerous
very fine concentric growth lines over the valve. Internal

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

43

FIGURE 17. Brachiopods of the HK1 assemblage (2). A‒F, Tolmatchoffia robusta (Tolmatchoff); A, internal mould of ventral valve, NU-B2178; B, externallatex cast (B1) and internal mould (B2) of ventral valve, NU-B2174; C, internal mould of ventral valve, NU-B2177; D, internal mould (D1, D2) and externallatex cast (D3) of dorsal valve, NU-B2181; E, external mould (E1) and internal mould (E2) of dorsal valve, NU-B2180; F, external mould (F1, F2) of dorsal
valve, NU-B2213. Scale bars represent 1 cm.

structures of the dorsal valve are obscure, except for a median
septum which is extending to half length of the valve. This
specimen is referred to Argentiproductus margaritaceus (Phillips,
1836), redescribed by Brunton et al. (1993, p. 103, ﬁgs. 13‒15)
from the upper Visean of Craven Reef Belt, Yorkshire, northern
England, in size, shape and external ornament of the dorsal valve.

Argentiproductus pectinoides (Phillips, 1836, p. 215, pl. 7, fig.
11), from the Visean of England, differs from A. margaritaceus in
the smaller size and less transverse outline.
Occurrence.―HK4 Unit.
Distribution.―Upper Tournaisian‒Serpukhovian: northeasternJapan (Hikoroichi in the South Kitakami Belt), northern Russia

44

JUN-ICHI TAZAWA(Pechora Basin), UK (England, Wales and northern Ireland),
Poland, Germany, Belgium, western Russia (Moscow Basin andDonetz Basin), central Russia (southern Urals), Kazakhstan,
Uzbekistan, Kyrgyzstan, southwestern China (Guizhou) andwestern Australia (Bonaparte Gulf Basin).
Superfamily PRODUCTOIDEA Gray, 1840
Family BUXTONIIDAE Muir-Wood and Cooper, 1960
Subfamily BUXTONIINAE Muir-Wood and Cooper, 1960
Genus TOLMATCHOFFIA Fredericks, 1933
Type species.―Productus robustus Tolmatchoff, 1924.
Tolmatchoffia robusta (Tolmatchoff, 1924)(Figs. 17A‒F, 22G‒J)
Productus robustus Tolmatchoff, 1924, p. 230, 572, pl. 13, ﬁg. 17;
pl. 14, ﬁgs. 1‒3.
Dictyoclostus robustus (Tolmatchoff). Benediktova, 1955, p. 309,
pl. 85, ﬁg. 6; Yang, 1964, p. 96, pl. 9, ﬁgs. 6, 7.
Tolmatchoffia robusta (Tolmatchoff). Muir-Wood and Cooper,
1960, pl. 95, ﬁgs. 1‒3; Sarytcheva in Sarytcheva et al., 1963,
p. 171, pl. 21, figs. 1‒3; pl. 22, figs. 1‒3; text-figs. 71, 72;
Galitskaya, 1977, p. 74, pl. 34, fig. 3; Zhang et al., 1983, p.
302, pl. 118, ﬁgs. 4, 5.
Buxtonia sp. Tazawa, 1989, p. 60, pl. 1, ﬁgs. 6‒9.
Material.―Twenty-one specimens from localities KF201 andKF202: (1) external and internal moulds of two ventral valves,
NU-B2174, 2175; (2) external mould of a ventral valve, NUB2176; (3) internal mould of four ventral valves, NU-B2043,
2177‒2179; (4) external and internal moulds of seven dorsal
valves, NU-B2044‒2047, 2180‒2182; (5) external moulds of ﬁve
dorsal valves, NU-B2048‒2051, 2183; and (6) internal moulds oftwo dorsal valves, NU-B2052, 2053.
Description.―Shell large in size for genus, slightly longer
subrectangular in outline, widest at slightly anterior to midlength;
length 58 mm, width 53 mm in the largest ventral valve specimen(NU-B2043); length 40 mm, width 66 mm in the largest dorsal
valve specimen (NU-B2180). Ventral valve strongly and unevenly
convex in lateral proﬁle, most convex at umbonal region; umbomassive, strongly incurved; ears small; sulcus moderately deep;
lateral slopes steep. Dorsal valve with large ﬂattened visceral disc,strongly geniculated and followed by short trail. External surface
of ventral valve ornamented with numerous costae, numbering
6‒7 in 5 mm at about midlength; spines dense on ears. External
ornament of dorsal valve weakly reticulate on visceral disc; spines
absent. Ventral interior poorly preserved and obscure. Dorsalinterior with large trilobate cardinal process, long, thin median
septum and elongate dendritic adductor scars.
Remarks.―These specimens can be referred to Tolmatchoffia
robusta (Tolmatchoff, 1924), redescribed by Sarytcheva (inSarytcheva et al., 1963, p. 171, pl. 21, ﬁgs. 1‒3; pl. 22, ﬁgs. 1‒3;
text-figs. 71, 72) from the lower and upper Tournaisian of theKuznetsk Basin, central Russia, in external features, particularly,

the large size, short dorsal trail and dense spine bases on ears.
Tolmatchoffia barzassica Sarytcheva (in Sarytcheva et al., 1963,
p. 174, pl. 23, ﬁgs. 1, 2, text-ﬁg. 73), from the lower and upper
Tournasian of the Kuznetsk Basin, differs from T. robusta in thesmaller size and in having no ventral sulcus.
Occurrence.―HK1 and HK2 units.
Distribution.―Lower Tournaisian‒lower Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt), central Russia(Kuznetsk Basin), Kyrgyzstan and northwestern China (Xinjiang).
Subfamily MARGINATIINAE Waterhouse, 2002
Genus MARGINATIA Muir-Wood and Cooper, 1960
Type species.―Productus fernglenensis Weller, 1909.
Marginatia burlingtonensis (Hall, 1858)(Fig. 23A‒D)
Productus flemingi var. burlingtonensis Hall, 1858, p. 598, pl. 12,
ﬁg. 3.
Productus burlingtonensis Hall. Weller, 1914, p. 104, pl. 9, ﬁgs.
1‒10; Frech, 1916, p. 239, pl. 6, fig. 1; Tolmatchoff, 1924,
p. 237, 575, pl. 14, figs. 8‒11; Girty, 1929, p. 85, pl. 9, figs.
20‒24.
Productus (Productus) burlingtonensis Hall. Nalivkin, 1937, p.
66, pl. 7, ﬁgs. 7‒11.
Productus sp. Minato, 1951, p. 366, pl. 1, ﬁg. 4.
Productus (Dictyoclostus) burlingtonensis Hall. Simorin, 1956, p.
136, pl. 9, ﬁgs. 1‒3.
Marginatia burlingtonensis (Hall). Sarytcheva in Sarytcheva
et al., 1963, p. 191, pl. 28, figs. 5‒8, text-figs. 81, 82;
Grechishnikova, 1966, p. 116, pl. 8, ﬁgs. 11‒13; Litvinovich etal., 1969, p. 213, pl. 35, ﬁgs. 2‒4; Nalivkin and Fotieva, 1973,
p. 39, pl. 8, ﬁg. 1; Bublichenko, 1976, p. 50, pl. 2, ﬁg. 12; pl. 4,
ﬁg. 6; pl. 5, ﬁgs. 4‒6; pl. 6, ﬁg. 9; Galitskaya, 1977, p. 83, pl.
22, ﬁgs. 6‒10; Nalivkin, 1979, p. 94, pl. 32, ﬁgs. 1‒10; pl. 34,
ﬁgs. 3, 4; Lee et al., 1980, p. 368, pl. 148, ﬁg. 10; Jin, 1985, p.
77, pl. 1, ﬁgs. 20‒22; Carter, 1987, p. 39, pl. 9, ﬁgs. 1‒8; Shi etal., 2005, p. 44, ﬁgs. 5D, I‒K, M; Tazawa, 2006, p. 132, ﬁgs.
6.1‒6.8; Tazawa, 2017b, p. 335, ﬁgs. 6.3‒6.5.
Dictyoclostus sp. Hase and Yokoyama, 1975, pl. 18, ﬁg. 1.
Marginatia sp. Tazawa, 1985, p. 459, ﬁgs. 2.3‒2.7; Tazawa, 1989,
p. 60, pl. 1, ﬁg. 1; Tazawa, 2002, ﬁgs. 7.1, 7.2.
Material.―Twenty-two specimens from localities KF171,
KF172, KF174, KF188, KF201 and KF202: (1) external andinternal moulds of a conjoined shell, NU-B629; (2) external andinternal moulds of a ventral valve, NU-B631; (3) external mouldof a ventral valve, NU-B632; (4) external and internal moulds ofseven dorsal valves, NU-B633‒639; and (5) external moulds oftwelve dorsal valves, NU-B640‒649, 2213, 2214.
Remarks.―These specimens can be referred to Marginatia
burlingtonensis (Hall, 1858), redescribed by Weller (1914, p.
104, pl. 9, figs. 1‒10) from the Burlington Limestone of theMississippi Valley, on the basis of the medium size, transversely

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

45

FIGURE 18. Brachiopods of the HK1 assemblage (3). A, B, Schizophoria resupinata (Martin); A, internal mould of ventral valve, NU-B2188; B, internalmould of dorsal valve, NU-B2190; C, Ovatia sp., external latex cast (C1, C2) and external mould (C3) of dorsal valve, NU-B2202; D, E, Marginatia magna
Carter; D, dorsal view (D1) and anterior view (D2) of external mould of dorsal valve, NU-B2185; E, dorsal (E1, E2), anterior (E3) and lateral (E4) views ofexternal mould of dorsal valve, NU-B2184. Scale bars represent 1 cm.

rectangular outline and strongly geniculated dorsal valve, withregular and strong reticulate ornament on the visceral disc.
Marginatia patersonensis Roberts (1965, p. 63, pl. 10, ﬁgs. 1‒5),
from the lower and upper Visean of New South Wales, eastern
Australia, differs from the present species in having ﬁner costae
and larger number of spines on the ventral valve. The typespecies, Marginatia fernglenensis Weller (1909, p. 299, pl. 12,
figs. 14‒17) from the Fern Glen Formation of Missouri, differsfrom M. burlingtonensis in having shallower ventral sulcus.

Occurrence.―HK1 and HK2 units.
Distribution.―Upper Tournaisian‒upper Visean: northeasternJapan (Hikoroichi and Yokota in the South Kitakami Belt),southwestern Japan (Hina in the Akiyoshi Belt), USA (Illinois,
Iowa and Arkansas), western Canada (Alberta), Turkey (Taurus
Mountains), central Russia (southern Urals and Kuznetsk Basin),
Kazakhstan, Kyrgyzstan and northeastern China (Liaoning).

46

JUN-ICHI TAZAWA

Marginatia magna Carter, 1968
(Fig. 18D, E)
Marginatia magna Carter, 1968, p. 1147, pl. 147, ﬁgs. 1‒9; Garanj
et al., 1975, p. 166, pl. 66, ﬁg. 8; Carter, 1999, p. 102, ﬁg. 3.
Material.―Four specimens from localities KF201 and KAC3:(1) internal mould of a conjoined shell, with external mould of thedorsal valve, NU-B2184; and (2) external moulds of three dorsal
valves, NU-B2185‒2187.
Remarks.―The Hikorichi specimens are fragmentarily
preserved, but they can be referred to Marginatia magna Carter(1968, p. 1147, pl. 147, ﬁgs. 1‒9), from the Burlington Limestoneof Missouri, in the large size (length more than 45 mm, widthabout 50 mm in the largest specimen, NU-B2184), moderately
developed ventral sulcus and dorsal fold, strongly geniculated
dorsal valve with long trail and slightly spreading lateral ﬂanks,and external ornament of dorsal valve with reticulate on visceraldisc and costate on trail and very ﬁne numerous growth lines overthe valve. The preceding species, Marginatia burlingtonensis(Hall, 1858), is readily distinguished from M. magna in thesmaller size, much more ﬁnely ribbed dorsal valve, and in having
a stronger fold‒sulcus.
Occurrence.―HK1 Unit.
Distribution.―Upper Tournaisian: northeastern Japan(Hikoroichi in the South Kitakami Belt), USA (Missouri andOklahoma) and central Russia (southern Urals).
Superfamily ECHINOCONCHOIDEA Stehli, 1954
Family ECHINOCONCHIDAE Stehli, 1954
Subfamily ECHINOCONCHINAE Stehli, 1954
Genus ECHINOCONCHUS Weller, 1914
Type species.―Productus punctatus Sowerby, 1822.
Echinoconchus punctatus (Sowerby, 1822)(Fig. 23G, H)
Productus punctatus Martin. Sowerby, 1822, p. 22, pl. 323, lower
right ﬁgure; Davidson, 1861, p. 172, pl. 44, ﬁgs. 9‒11, 16, 17.
Pustula punctata (Martin). Thomas, 1914, p. 303, pl. 17, figs.
16‒19, text-ﬁg. 11; Tolmatchoff, 1924, p. 256, 584, pl. 16, ﬁg.
9; Rotai, 1931, p. 58, pl. 4, ﬁgs. 1, 11.
Productus (Pustula) punctatus (Martin). Yanishevsky, 1918, p. 47,
pl. 3, ﬁgs. 7, 9.
Echinoconchus punctatus (Martin). Chao, 1927, p. 67, pl. 6, ﬁgs.
7, 8, 15, 16; Sarytcheva in Sarytcheva and Sokolskaya, 1952,
p. 103, pl. 18, ﬁg. 120; Dedok and Tschernjak, 1960, p. 53, pl.
1, ﬁg. 6; Pareyn, 1961, p. 197, pl. 23, ﬁgs. 1‒4; Ding in Yanget al., 1962, p. 51, pl. 19, ﬁgs. 1‒4; Yang, 1964, p. 81, pl. 4,
ﬁgs. 5, 6, 9, 10, text-ﬁg. 7; Abramov, 1965, p. 38, pl. 3, ﬁg. 2;
Litvinovich et al., 1969, p. 164, pl. 9, ﬁgs. 5, 6; pl. 10, ﬁg. 1;
Abramov, 1970, p. 117, pl. 9, ﬁg. 4; Alexandrov and Solomina,
1973, p. 93, pl. 22, ﬁgs. 1‒3; Volgin and Kushnar, 1975, p. 46,
pl. 4, ﬁg. 1; Donakova, 1978, p. 208, pl. 1, ﬁgs. 5, 6; Nalivkin,

1979, p. 78, pl. 24, figs. 8, 9; Zhang et al., 1983, p. 288, pl.
127, ﬁg. 111; pl. 128, ﬁg. 2; Jin et al., 1985, p. 192, pl. 9, ﬁgs.
11, 12; Zhan and Wu, 1987, p. 207, pl. 48, ﬁg. 38; Archbold
and Stojanović-Kuzenko, 1995, pl. 62, ﬁg. 10; Wang and Yang,
1998, p. 77, pl. 9, ﬁgs. 17, 18.
Productus (Echinoconchus) punctatus (Martin) emend. Thomas.
Paeckelmann, 1931, p. 152, pl. 15, ﬁgs. 7‒10.
Productus (Echinoconchus) punctatus (Martin). Nalivkin, 1937, p.
64, pl. 9, ﬁg. 5.
Echinoconchus punctatus (Sowerby). Muir-Wood, 1951, p. 102,
pl. 4, ﬁg. 2; Muir-Wood and Cooper, 1960, pl. 66, ﬁgs. 1, 2;
pl. 82, ﬁgs. 8‒10; pl. 83, ﬁgs. 1‒4; pl. 88, ﬁg. 11; pl. 125, ﬁg.
5; Winkler Prins, 1968, p. 89, pl. 3, ﬁgs. 12‒14; Nalivkin andFotieva, 1973, p. 35, pl. 6, ﬁg. 8; Kalashnikov, 1974, p. 48, pl.
9, ﬁgs. 1‒3; Martinez Chacon and Legrand-Blain, 1992, p. 110,
pl. 3, ﬁgs. 15‒18; Tazawa, 2017b, p. 335, ﬁgs. 6.6, 6.7.
Echinoconchus (Echinoconchus) punctatus (Sowerby). Galitskaya,
1977, p. 62, pl. 16, figs. 1‒5; pl. 18, fig. 1; text-fig. 7;
Kalashnikov, 1980, p. 34, pl. 5, ﬁg. 1.
Echinoconchus aohanensis Lee et al., 1980, p. 363, pl. 147, ﬁgs.
1, 2.
Material.―Three specimens from locality KF202: (1) internalmoulds of two ventral valves, NU-B2061, 2062; and (2) external
and internal moulds of a dorsal valve, NU-B2063.
Description.―Shell large in size for genus, transversely
subcircular in outline, hinge slightly shorter than greatest widthat midlength; length 38 mm, width about 50 mm in the better
preserved ventral valve specimen (NU-B2061); length 31 mm,
width 40 mm in the sole dorsal valve specimen (NU-B2063).
Ventral valve moderately and unevenly convex in lateral proﬁle,most convex at umbonal region; umbo rounded, massive andincurved; ears small, not clearly demarcated from visceralregion; sulcus wide and shallow; lateral slopes gently convex andspreading. Dorsal valve weakly concave, nearly ﬂat; no fold; trail
not preserved. External surface of both valves ornamented withregular, strong concentric bands; numbering 4‒5 bands in 10 mm
on ventral valve; two rows of numerous elongate spine bases oneach band of dorsal valve. Internal structures of both valves notwell preserved.
Remarks.―These specimens can be referred to Echinoconchus
punctatus (Sowerby, 1822), redescribed by Muir-Wood (1951, p.
102, pl. 4, ﬁg. 2) and reﬁgured by Muir-Wood and Cooper (1960,
pl. 66, ﬁgs. 1, 2; pl. 82, ﬁgs. 8‒10; pl. 83, ﬁgs. 1‒4; pl. 88, ﬁg. 11;
pl. 125, ﬁg. 5) from the Visean of England, in the large size andexternal ornament of both ventral and dorsal valves, consisting of
regular strong concentric bands with rows of numerous elongate
spine bases. Echinoconchus defensus (Thomas, 1914, p. 310,
pl. 17, figs. 20‒23), from the Visean of England and Scotland,differs from E. punctatus in the smaller size and in having more
numerous concentric bands on the ventral valve. Echinoconchus
alternatus (Norwood and Pratten, 1855), redescribed by
Weller (1914, p. 138, pl. 17, ﬁgs. 1‒7) from the Osagean of theMississippi Valley, USA, resembles E. punctatus in size andoutline, but the American species is distinguished from the latter

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

by the presence of broader concentric bands on the ventral valve.
Occurrence.―HK2 Unit.
Distribution.―Upper Tournaisian‒lower Permian (Asselian):
northeastern Japan (Hikoroichi and Yokota in the South KitakamiBelt), northern Russia (Verkhoyansk Range, Taimyr Peninsula andnorthern Urals), UK (Scotland, England and Wales), Germany,
Belgium, Spain, western Russia (Moscow Basin), central Russia(southern Urals), Kyrgyzstan, northwestern China (Xinjiang and
Qinghai) and northern China (Shanxi).
Genus KARAVANKINA Ramovs, 1969
Type species.―Karavankina typica Ramovs, 1969.
Karavankina jizodoensis Tazawa, 1980
(Fig. 30C, D)
Karavankina jizodoensis Tazawa, 1980, p. 362, pl. 41, ﬁgs. 8‒16;
pl. 42, ﬁg. 14; text-ﬁg. 2.
Karavankina sp. Tazawa, 1984b, p. 306, pl. 61, ﬁg. 2.
Material.―Two specimens from locality KF167: (1) internalmould of a ventral valve, NU-B2233; and (2) external and internalmoulds of a dorsal valve, IGPS97905.
Description.―Shell large in size for genus, transversely
subrectangular in outline; length 18 mm, width more than 28 mm
in the larger specimen (NU-B2233). Ventral valve gently convex
in both lateral and anterior proﬁles; no sulcus. Dorsal valve gently
concave, with nearly flat visceral disc; hinge straight, slightly
narrower than greatest width at about three-quarters length of thevalve; ears rather large, ﬂat, trigonal and not clearly demarcatedfrom visceral region; no fold. External surface of both valvesornamented with regular concentric lamellae, numbering 5‒6
in 10 mm in ventral valve, and 7‒9 in 10 mm in dorsal valve.
Moreover, numerous somewhat quincunxially arranged elongate
spine bases are preserved on the anterior portion of each lamellain the dorsal valve specimen (IGPS97905). Internally, ventral
valve having a pair of narrow, elongate and smooth adductor
scars in posterior portion of valve. Dorsal valve having a median
septum and a pair of stout buttress plates.
Remarks.―These specimens are referred to Karavankina
jizodoensis Tazawa (1980, p. 362, pl. 41, ﬁgs. 8‒16; pl. 42, ﬁg.
14; text-fig. 2), from the uppermost part of the Karaumedate
Formation (upper Visean) of the Nagasaka area, South Kitakami
Belt, on account of the large size (length 18 mm, width more than
28 mm), transversely subrectangular outline, external ornament ofboth valves consisting of numerous regular concentric lamellae,
and the presence of a pair of stout buttress plates in the dorsal
valve. Karavankina dobsinensis (Rakusz, 1932, p. 55, pl. 2, ﬁgs.
21, 22; pl. 3, ﬁg. 2) from the Moscovian of Hungary, resembles
the present species in size and shape of the ventral valve, but the
Hungarian species differs from K. jizodoensis in having more
numerous narrower concentric lamellae on the ventral valve.
Occurrence.―HK4 Unit.
Distribution.―Upper Visean: northeastern Japan (Hikoroichi

47

and Nagasaka in the South Kitakami Belt).
Family WAAGENOCONCHIDAE Muir-Wood and Cooper, 1960
Subfamily PUSTULINAE Waterhouse, 1981
Genus PUSTULA Thomas, 1914
Type species.―Producta pustulosa Phillips, 1836.
Pustula pustulosa (Phillips, 1836)(Fig. 30F)
Producta pustulosa Phillips, 1836, p. 216, pl. 7, ﬁg. 15.
Productus pustulosus Phillips. Krenkel, 1913, p. 18, 43, pl. 1, ﬁg.
7.
Pustula pustulosa (Phillips). Thomas, 1914, p. 261, pl. 17, ﬁgs.
24‒28; pl. 18, ﬁg. 1; Muir-Wood and Cooper, 1960, pl. 59, ﬁg.
4; pl. 84, figs. 1‒7; pl. 85, figs. 6‒10; Nalivkin and Fotieva,
1973, p. 36, pl. 6, ﬁgs. 11‒13; Kalashnikov, 1974, p. 51, pl. 10,ﬁgs. 6‒9; pl. 11, ﬁgs. 1, 2; pl. 13, ﬁgs. 4, 5; pl. 30, ﬁg. 6; Garanj
et al., 1975, p. 163, pl. 65, ﬁgs. 3, 4; Volgin and Kushnar, 1975,
p. 45, pl. 3, ﬁgs. 10‒12; Galitskaya, 1977, p. 67, pl. 19, ﬁgs.
1‒5; pl. 20, ﬁg. 1; Zhang et al., 1983, p. 289, pl. 109, ﬁg. 4;
Tazawa, 2017b, p. 336, ﬁgs. 7.3, 7.4.
Productus (Pustula) pustulosus Phillips emend. Thomas.
Paeckelmann, 1931, p. 138, pl. 13, ﬁgs. 2, 3.
Pustula cf. pustulosa (Phillips). Mori and Tazawa, 1980, text-ﬁg.
3.2; Tazawa, 1984b, p. 306, pl. 61, ﬁg. 8.
Material.―One specimen from locality KF152, externalmould of a dorsal valve, IGPS97904.
Remarks.―This specimen was figured by Mori and Tazawa(1980, text-fig. 3.2) and described by Tazawa (1984b, p. 306,
pl. 61, fig. 8) as Pustula cf. pustulosa (Phillips, 1836). But the
Hikoroichi species is identical with Pustula pustulosa (Phillips,
1836), redescribed by Thomas (1914, p. 261, pl. 17, ﬁgs. 24‒28;
pl. 18, ﬁg. 1) from the upper Visean of Derbyshire, England, by
the large size (length more than 35 mm, width about 76 mm),transversely subrectangular outline, well-developed dorsal fold
and external ornament of dorsal valve consisting of regularconcentric bands and numerous, quincunxially arranged elongate
spine bases. Pustula tenuipustulosa Thomas (1914, p. 288, pl. 20,
ﬁgs. 10, 11, text-ﬁg. 7), from the lower Carboniferous of Wexford,
Ireland, differs from P. pustulosa in having smaller spine bases
and less-marked bands.
Occurrence.―HK4 Unit.
Distribution.―Lower Visean‒Serpukhovian: northeasternJapan (Hikoroichi and Yokota in the South Kitakami Belt),
northern Russia (Pechora Basin), UK (England and northern
Ireland), Germany, central Russia (western and southern Urals),
Uzbekistan (Fergana), Kyrgyzstan and northwestern China(Xinjiang).
Superfamily LINOPRODUCTOIDEA Stehli, 1954
Family LINOPRODUCTIDAE Stehli, 1954
Subfamily LINOPRODUCTINAE Stehli, 1954

48

JUN-ICHI TAZAWA

Genus OVATIA Muir-Wood and Cooper, 1960
Type species.―Ovatia elongata Muir-Wood and Cooper, 1960.
Ovatia laevicosta (White, 1860)(Fig. 23E, F)
Productus laevicostus White, 1860, p. 230.
Ovatia laevicosa (White). Sarytcheva et al., 1963, p. 226, pl. 37,
ﬁgs. 3‒5, text-ﬁg. 100; Carter, 1999, p. 109, ﬁgs. 4I‒N.
Material.―Two specimens from localities KF188 and KF202:(1) external and internal moulds of a ventral valve, NU-B2064;
and (2) external mould of a ventral valve, NU-B2065.
Remarks.―These specimens are referred to Ovatia laevicosta(White, 1860), redescribed by Carter (1999, p. 109, figs. 4I‒N)
from the lower Osagean of Kenwood, northeastern Oklahoma,
USA, in the relatively large size (length about 24 mm, widthabout 39 mm in the better preserved specimen, NU-B2064) andthe number of costellae on the ventral valve (10‒11 in 5 mm at
about midlength). Ovatia elongata Muir-Wood and Cooper (1960,
p. 312, pl. 114, ﬁgs. 1‒4, 7, 11, 12), from the lower Fayetteville
Formation of Fort Gibson, Oklahoma, differs from O. laevicostain having a massive and strongly convex ventral umbo.
Occurrence.―HK2 Unit.
Distribution.―Upper Tournaisian‒lower Visean: nortrheasternJapan (Hikoroichi in the South Kitakami Belt), USA (Iowa andOklahoma) and central Russia (Kuznetsk Basin).
Ovatia cf. prolata Carter, 1987
(Fig. 30E)
cf. Ovatia prolata Carter, 1987, p. 40, pl. 5, ﬁgs. 1‒17.
Material.―One specimen from locality KAC1, external mouldof a ventral valve, NU-B2232.
Remarks.―This specimen is safely assigned to the genus
Ovatia by the small (length 24 mm, width 22 mm), subovate andstrongly convex ventral valve, ornamented with numerous finecostellae (numbering 16‒18 in 5 mm at about midlength). The
Hikoroichi species most resembles Ovatia prolata Carter (1987,
p. 40, pl. 5, figs. 1‒17), from the Banff Formation of western
Alberta, in having fine costellae on the ventral valve. Accuratecomparison is, however, difﬁcult due to lack of the dorsal valve.
Occurrence.―HK4 Unit.
Ovatia sp.
(Fig. 18C)
Material.―One specimen from locality KF201, externalmould of a dorsal valve, NU-B2202.
Remarks.―This specimen is safely assigned to the genus
Ovatia by the small size (length about 20 mm, width about 25mm), moderately concave dorsal valve, without fold, and theexternal ornament consisting of numerous regular costellae(numbering 12‒13 in 5 mm at midlength) and strong concentricrugae on ears and the anterior part of the valve. The Hikoroichi
species resembles Ovatia ovata (Hall, 1858), redescribed by
Weller (1914, pl. 16, ﬁgs. 1‒15) from the Chester Group of theMississippi Valley, in size, shape and external ornament of thedorsal valve. But accurate comparison is difﬁcult for the poorly
preserved specimen.
Occurrence.―HK1 Unit.
Genus LINOPROTONIA Ferguson, 1971
Type species.―Productus hemisphaericus Sowerby, 1822.
Linoprotonia ashfellensis Ferguson, 1971
(Fig. 24A‒C)
Linoprotonia ashfellensis Ferguson, 1971, p. 558, pl. 42, ﬁgs. 1‒7,
text-ﬁg. 2A; Pattison, 1981, p. 11, pl. 1, ﬁgs. 1, 2; pl. 7, ﬁg. 2;
pl. 9, ﬁg. 12; Tazawa and Ibaraki, 2009, p. 11, ﬁgs. 4.1‒4.3.
Gigantoproductus sp. B Tazawa, 1984a, p. 58, pl. 1, figs. 4, 5
only.
Linoprotonia sp. Tazawa, 1989, p. 61, pl. 1, ﬁg. 5 only.
Material.―Six specimens from locality KF202: (1) external
and internal moulds of a ventral valve, NU-B597; and (2) internalmoulds of ﬁve ventral valves, NU-B598‒602.
Remarks.―These specimens were previously described by
Tazawa and Ibaraki (2009, p. 11, ﬁgs. 4.1‒4.3) as Linoprotonia
ashfellensis Ferguson, 1971 on the basis of the linoproductid
external features and gigantoproductid internal structures,
particularly, in having small ears and ﬂattened visceral disc in theventral valve. Comparison with the other Linoprotonia species,
L. hemispaerica (Sowerby, 1822) and L. corrugatohemispherica(Vaughan in Dixon and Vaughan, 1911), are fully discussed by
Ferguson (1971, p. 559).
Occurrence.―HK2 Unit.
Distribution.―Lower‒upper Visean: northeastern Japan(Hikoroichi in the South Kitakami Belt) and UK (England).
Linoprotonia hikoroichiensis Tazawa and Ibaraki, 2009
(Fig. 24D‒H)
Gigantoproductus sp. B Tazawa, 1984a, p. 58, pl. 1, figs. 2, 3
only.
Linoprotonia hikoroichiensis Tazawa and Ibaraki, 2009, p. 11,ﬁgs. 5.1‒5.5.
Material.―Eighteen specimens from locality KF202: (1)external and internal moulds of three ventral valves, NU-B603(holotype), 604, 605; (2) external moulds of two ventral valves,
NU-B606, 607; (3) internal moulds of seven ventral valves, NUB608‒614; (4) external and internal moulds of a dorsal valve,
NU-B615; and (5) external moulds of five dorsal valves, NUB616‒620.
Remarks.―These specimens were described by Tazawa andMISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

49

FIGURE 19. Brachiopods of the HK1 assemblage (4). A, B, Rhynchopora sp.; A, internal mould (A1, A2) of ventral valve, NU-B2215; B, external latex cast(B1) and internal mould (B2) of dorsal valve, NU-B2216; C, Cleiothyridina harkeri Carter, internal mould (C1) and enlarged concentric lamellae (C2) of ventral
valve, NU-B2201; D‒F, Acuminothyris keokuk Carter; D, external latex cast (D1, D2) and internal mould (D3) of ventral valve, NU-B2118; E, internal mould (E1,
E2) of ventral valve, NU-B2119; F, external latex cast (F1, F2) and internal mould (F3) of dorsal valve, NU-B2120; G‒J, Unispirifer mediocris (Tolmatchoff);
G, internal mould of ventral valve, NU-B2209; H, internal mould (H1, H2) of dorsal valve, NU-B2212; I, internal mould of dorsal valve, NU-B2211; J, internalmould of ventral valve, NU-B2210; K, Tylothyris laminosa (MʼCoy), internal mould (K1, K2) of dorsal valve, NU-B2205; L, Brachythyris suborbicularis (Hall),internal mould (L1, L2) of dorsal valve, NU-B2206. Scale bars represent 1 cm, unless otherwise speciﬁed.

50

JUN-ICHI TAZAWA

FIGURE 20. Brachiopods of the HK1 assemblage (5). A, Kitakamithyris tyoanjiensis Minato, external latex cast (A1), external mould (A2) and internal mould(A3, A4) of ventral valve, NU-B2191; B, C, Kitakamithyris hikoroitiensis Minato; B, external latex cast (B1), external mould (B2) and internal mould (B3) ofventral valve, NU-B2193; C, internal mould (C1, C2) of dorsal valve, NU-B2196; D, E, Punctospirifer sp.; D, internal mould (D1, D2) of ventral valve, NUB2207; E, internal mould of dorsal valve, NU-B2208; F‒H, Syringothyris sibirica Sokolskaya; F, external latex cast (F1) and internal mould (F2) of dorsal
valve, NU-B2200; G, ventral (G1), posterior (G2) and lateral (G3) views of internal mould of ventral valve, N-B2197; H, external latex cast (H1) and internalmould (H2) of dorsal valve, NU-B2199; I, Beecheria khalfini Besnossova, ventral view of internal mould (I1, I2) of conjoined shell, NU-B2217. Scale bars
represent 1 cm.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

51

FIGURE 21. Brachiopods of the HK1 assemblage (6). Enlarged concentric rows of biramous spine bases; A, Kitakamithyris tyoanjiensis Minato, externalmould of ventral valve, NU-B2191; B, Kitakamithyris hikoroitiensis Minato, external mould of ventral valve, NU-B2193.

Ibaraki (2009, p. 11, ﬁgs. 5.1‒5.5) as Linoprotonia hikoroichiensis
Tazawa and Ibaraki, 2009. This species is medium in size forgenus, transversely semicircular to subquadrate in outline (length
36 mm, width 65 mm in the largest specimen, NU-B608), andhaving gently convex ventral valve, with large prominent ears andornamented with ﬁne costae (14‒16 in 10 mm at mid-length). L.
hikoroichiensis differs from the preceding species, L. ashfellensis
Ferguson, 1971, by the less convex ventral valve, much larger
prominent ears and ﬁner costae on the ventral valve.
Occurrence.―HK2 Unit.
Distribution.―Lower Visean: northeastern Japan (Hikoroichiin the South Kitakami Belt).
Subfamily GIGANTOPRODUCTINAE Muir-Wood and Cooper,
1960
Genus GIGANTOPRODUCTUS Prentice, 1950
Type species.―Productus giganteus Sowerby, 1822.
Gigantoproductus crassiventer (Prentice, 1949)(Fig. 25A, C)
Gigantella crassiventer Prentice, 1949, p. 257, pl. 11, ﬁg. 1; pl.
12, ﬁg. 1 only.
Gigantoproductus cf. crassiventer (Prentice). von Bitter andLegrand-Blain, 2007, p. 193, ﬁgs. 2B‒F.
Gigantoproductus cf. crassiventer (Prentice). Tazawa and Ibaraki,
2009, p. 13, ﬁgs. 6.1, 6.2.
Material.―Three specimens from locality KF202: (1) external
and internal moulds of a ventral valve, NU-B621; and (2) internalmoulds of two ventral valves, NU-B622, 623.
Remarks.―These specimens were described by Tazawa
and Ibaraki (2009, p. 13, ﬁgs. 6.1, 6.2) as Gigantoproductus cf.
crassiventer (Prentice, 1949). However, the Hikoroichi specimens
can be referred to Gigantoproductus crassiventer (Prentice,1949), from the upper Visean of England, by the ventral valve of
medium size and elongate outline (length about 62 mm, widthabout 56 mm in the best preserved specimen, NU-B622), withnumerous costae (12‒13 in 10 mm at midlength) over the valve
and weak irregular ribbing on the anterior half of the valve.
Gigantoproductus elongatus (Sarytcheva, 1928, p. 27, pl. 3, ﬁg.
4), from the upper Visean of the Moscow Basin, differs from G.
crassiventer by the more inﬂated ventral valve, having no sulcus
and more strongly developed ribbing on the ventral valve.
Occurrence.―HK2 Unit
Distribution.―Lower‒upper Visean: northeastern Japan(Hikoroichi in the South Kitakami Belt), eastern Canada (Nova
Scotia) and UK (England).
Gigantoproductus latiexpansus Sarytcheva in Sarytcheva and
Sokolskaya, 1952
(Fig. 25B)
Gigantoproductus latiexpansus Sarytcheva in Sarytcheva and
Sokolskaya, 1952, p. 130, pl. 33, ﬁg. 176.
Gigantoproductus sp. A Tazawa, 1984a, pl. 1, ﬁg. 1.
Linoprotonia sp. Tazawa, 1989, p. 61, pl. 1, ﬁg. 4 only.
Gigantoproductus cf. latiexpansus Sarytcheva. Tazawa and
Ibaraki, 2009, p. 15, ﬁg. 7.1.
Material.―Three specimens from locality KF202: (1) external
and internal moulds of a ventral valve, NU-B264; and (2) externalmoulds of two ventral valves, NU-B625, 626.
Remarks.―These specimens were described by Tazawa
and Ibaraki (2009, p. 15, fig. 7.1) as Gigantoproductus cf.
latiexpansus Sarytcheva in Sarytcheva and Sokolskaya, 1952.
But the Hikoroichi specimens are referred to Gigantoproductus
latiexpansus Sarytcheva (in Sarytcheva and Sokolskaya, 1952, p.
130, pl. 33, fig. 176), from the Venevian and Tarusian horizonsof the Moscow Basin, in the transverse outline, ill-deﬁned ears,
irregular ribbing and fine costae (numbering 16‒19 in 10 mm

52

JUN-ICHI TAZAWA

at about mid-length) on the ventral valve. Gigantoproductus
uniformis Morozov (1988, p. 32, pl. figs. 1‒5), from the upper
Visean of the Moscow Basin, differs from G. latiexpansus inhaving coarser costae on the ventral valve.
Occurrence.―HK2 Unit.
Distribution.―Lower Visean‒Serpukhovian: northeasternJapan (Hikoroichi in the South Kitakami Belt) and western Russia(Moscow Basin).
Order ORTHOTETIDA Waagen, 1884
Suborder ORTHOTETIDINA Waagen, 1884
Superfamily ORTHOTETOIDEA Waagen, 1884
Family ORTHOTETIDAE Waagen, 1884
Genus ORTHOTETES Fischer de Waldheim, 1829
Type species.―Orthotetes radiata Fischer de Waldheim, 1850.
Orthotetes keokuk (Hall, 1858)(Figs. 25D, 30H)
Orthis keokuk Hall, 1858, p. 640, pl. 19, ﬁg. 5.
Orthotetes keokuk (Hall). Weller, 1914, p. 75, pl. 7, figs. 1‒4;
Nalivkin, 1937, p. 40, pl. 3, ﬁgs. 1, 2; Minato, 1952, p. 151,
pl. 11, figs. 1, 2, 4; Simorin, 1956, p. 94, pl. 1, figs. 12, 13;
Armstrong, 1962, p. 45, pl. 6, ﬁgs. 18‒24; Litvinovich, 1962, p.
181, pl. 1, ﬁg. 6; pl. 2, ﬁg. 1; Abramov, 1965, p. 36, pl. 2, ﬁg. 6;
pl. 3, ﬁg. 1; Litvinovich et al., 1969, p. 135, pl. 3, ﬁg. 5; pl. 4,
ﬁgs. 1‒4; text-ﬁg. 40; Abramov, 1970, p. 109, pl. 2, ﬁgs. 4, 5;
pl. 3, ﬁg. 1.
Orthotetes cf. australis (Campbell). Tazawa, 1981b, p. 69, pl. 5,
ﬁg. 8.
Orthotetes sp. Tazawa, 1984b, p. 305, pl. 61, ﬁg. 11.
Material.―Four specimens from localities KF167 and KF202:(1) external cast of a ventral valve, NU-B2103; (2) external andinternal moulds of a ventral valve, NU-B2104; (3) external mouldof a ventral valve, NU-B2105; and (4) external and internalmoulds of a dorsal valve, IGPS97903.
Description.―Shell large in size for genus, transversely
semicircular in outline, hinge slightly shorter than greatest widthat about midlength; length about 45 mm, width about 78 mm in
the largest specimen (NU-B2104); length 57 mm, width about 62
mm in a relatively well preserved ventral valve specimen (NUB2105). Ventral valve nearly flat to slightly concave in lateral
proﬁle, no sulcus. External surface of ventral valve ornamented
with numerous costellae, irregular strong rugae and numerous
very fine growth lines; costellae subangular and having muchwider intercostal spaces (parvicostellate), often intercalated,
numbering 7‒9 in 5 mm near anterior margin of ventral valve.
Ventral interior with a strong median septum extending to one
third length of valve and faintly preserved large ﬂabellate muscle
scars.
Remarks.―These specimens are referred to Orthotetes keokuk(Hall, 1858), redescribed by Weller (1914, p. 75, pl. 7, ﬁgs. 1‒4)from the Keokuk Limestone of the Mississippi Valley, in size,

shape and external ornament of the ventral valve, particularly,
being widest at about midlength. The most allied species,
Orthotetes australis (Campbell, 1957, p. 45, pl. 11, figs. 1‒7),
from the upper part of the lower Burindi Group of Babbinboon,
New South Wales, eastern Australia, differs from O. keokuk inthe widest part at or slightly anterior to hinge. The type species,
Orthotetes radiata Fischer de Waldheim, 1850, redescribed by
Sokolskaya (in Sarytcheva and Sokolskaya, 1952, p. 52, pl. 7, ﬁg.
39) from the Moscovian of the Moscow Basin, western Russia, isdistinguished from the present species by the smaller size and lesstransverse outline.
Occurrence.―HK2 and HK4 units.
Distribution.―Upper Tournaisian‒upper Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt), USA (Iowa,
Missouri, New Mexico and Arizona), northern Russia(Verkhoyansk Range) and Kazakhstan.
Order ORTHIDA Schuchert and Cooper, 1932
Suborder DALMANELLIDINA Moore, 1952
Superfamily DALMANELLOIDEA Schuchert, 1913
Family RHIPIDOMELLIDAE Schuchert, 1913
Subfamily RHIPIDOMELLINAE Schuchert, 1913
Genus RHIPIDOMELLA Oehlert, 1890
Type species.―Terebratula michelini Léveillé, 1835.
Rhipidomella michelini (Léveillé, 1835)(Figs. 26A‒C, 30G)
Terebratula michelini Léveillé, 1835, p. 39, pl. 2, ﬁgs. 14‒17.
Orthis michelini (Léveillé). Davidson, 1861, p. 132, pl. 30, ﬁgs.
6‒12.
Dalmanella michelini (Léveillé). Frech, 1900, p. 201, pl. 16, ﬁg.
15.
Rhipidomella michelini (Léveillé). Rotai, 1931, p. 44, pl. 1, ﬁg. 3;
Demanet, 1934, p. 37, pl. 2, ﬁgs. 1‒9; Sarytcheva in Sarytcheva
and Sokolskaya, 1952, p. 26, pl. 1, ﬁg. 7: Litvinovich, 1962,
p. 177, pl. 1, ﬁg. 1; Zang in Yang et al., 1962, p. 19, pl. 1, ﬁgs.
1‒7; Ustritsky and Tschernjak, 1963, p. 68, pl. 1, ﬁgs. 11, 12;
Yang, 1964, p. 58, pl. 1, ﬁg. 1; Brunton, 1968, p. 17, pl. 3, ﬁgs.
1‒25, text-ﬁg. 5; Litvinovich et al., 1969, p. 127, pl. 1, ﬁgs. 9,
10; Bublichenko, 1971, p. 29, pl. 2, ﬁgs. 9‒12; Alexandrov andSolomina, 1973, p. 87, pl. 21, ﬁg. 1; Kalashnikov, 1974, p. 21,
pl. 3, ﬁgs. 7‒9; Volgin and Kushnar, 1975, p. 21, pl. 1, ﬁgs. 1, 2;
Lee and Gu, 1976, p. 231, pl. 131, ﬁgs. 1‒6; Martinez Chacon,
1979, p. 63, pl. 3, ﬁgs. 12‒15; pl. 4, ﬁgs. 1‒15; text-ﬁgs. 6, 7;
Lee et al., 1980, p. 330, pl. 145, ﬁg. 4; Ding and Qi, 1983, p.
250, pl. 88, ﬁg. 13; Zakowa, 1989, p. 115, pl. 3, ﬁg. 5; pl. 7, ﬁg.
7; Harper and Jeffrey, 1996, ﬁg. 3a; Legrand-Blain in LegrandBlain et al., 1996, p. 180, pl. 28, ﬁgs. 21, 22; Jiang, 1997, pl. 1,
ﬁg. 3; Bassett and Bryant, 2006, p. 502, pl. 1, ﬁgs. 1‒4; pl. 6,
ﬁgs. 11‒17; Sun and Baliński, 2008, p. 519, ﬁg. 26.
Rhipidomella sp. Tazawa and Katayama, 1979, p. 170, pl. 11,ﬁgs. 1‒7; Mori and Tazawa, 1980, text-ﬁgs. 3.4‒3.6; Tazawa,
1984b, p. 305, pl. 61, ﬁgs. 5‒7.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

53

FIGURE 22. Brachiopods of the HK2 assemblage (1). A, B, Leptagonia analoga (Phillips); A, external mould (A1) and external latex cast (A2, A3) of ventral
vlave, NU-B2081; B, external latex cast (B1, B2) and external mould (B3) of dorsal valve, NU-B2083; C, Rugosochonetes cf. hardrensis (Phillips), externalmould (C1, C2) and internal mould (C3) of dorsal valve, NU-B2042; D‒F, Leiochonetes onimarensis sp. nov.; D, external latex cast (D1, D2) and internal mould(D3) of ventral valve, NU-B2136 (holotype); E, external latex cast (E1, E2) and internal mould (E3) of ventral valve, NU-B2137; F, external mould (F1, F2) of
dorsal valve, NU-B2139; G‒J, Tolmatchoffia robusta (Tolmatchoff); G, internal mould of ventral valve, NU-B2043; H, internal latex cast (H1) and externalmould (H2) of dorsal valve, NU-B2047; I, internal mould of dorsal valve, NU-B2045; J, external mould (J1) and cardinal process (J2) of dorsal valve, NUB2048. Scale bars represent 1 cm.

54

JUN-ICHI TAZAWA

Material.―Eight specimens from localities KF163 andKF202: (1) external and internal moulds of a ventral valve, NUB2054; (2) internal moulds of two ventral valves, NU-B2055,
2056; (3) interior of a ventral valve, IGPS97900; (4) two abradeddorsal valves, IGPS97899, 97901; and (5) internal mould of two
dorsal valves, NU-B2057, IGPS97902.
Description.―Shell medium in size for genus, subcircular inoutline, gently dorsibiconvex, with short hinge and maximum
width at slightly anterior to midlength; length 16 mm, width 18
mm in the best preserved ventral valve specimen (NU-B2054).
Ventral valve gently convex; sulcus absent. External surface ofventral valve ornamented with numerous costellae, numbering
6‒7 in 2 mm near anterior margin. Ventral interior with large
ﬂabellate muscle scars bisected in front of adductor scars by shortmedian ridge. Dorsal interior with strong brachiophores and large
quadripartite muscle scars.
Remarks.―These specimens are referred to Rhipidomellamichelini (Léveillé, 1835), redescribed by Brunton (1968, p.
17, pl. 3, ﬁgs. 1‒25, text-ﬁg. 5) from the Visean of Fermanagh,
northern Ireland, in size and outline of the ventral valve,
particularly, in having short hinge and the widest part at slightly
anterior to midlength. Rhipidomella sp., described by Tazawa andKatayama (1979, p. 170, pl. 11, ﬁgs. 1‒7) and Tazawa (1984b, p.
305, pl. 61, ﬁgs. 5‒7) from the Odaira and Hikoroichi Formationsof the South Kitakami Belt, are conspecific with the present
species. Rhipidomella altaica Tolmatchoff (1924, p. 213, 569, pl.
13, ﬁgs. 5‒7, 9, 10) from the Tournaisian of the Kuznetsk Basin,
central Russia, differs from R. michelini in having longer hinge
and the widest part at midlength of the shell.
Occurrence.―HK2 and HK4 units.
Distribution.―Upper Tournaisian‒lower Bashkirian:northeastern Japan (Hikoroichi and Yokota in the South KitakamiBelt), northern Russia (Taimyr Peninsula and Pechora Basin), UK(England, Wales and northern Ireland), Ireland, Poland, Belgium,
France (French Pyrenees), Spain (Cantabrian Mountains), western
Russia (Moscow Basin and Donetz Basin), Iran, central Russia(southern Urals), Kazakhstan, Uzbekistan, northwestern China(Xinjiang, Qinghai, Gansu and Ningxia), northern China (Inner
Mongolia), northeastern China (Liaoning) and southwestern
China (Guizhou and Yunnan).
Superfamily ENTELETOIDEA Waagen, 1884
Family SCHIZOPHORIIDAE Schuchert and LeVene, 1929
Genus SCHIZOPHORIA King, 1850
Type species.―Conchyliolithus (Anomites) resupinatus Martin,
1809.
Schizophoria resupinata (Martin, 1809)(Figs. 18A, B, 26G, H, 31A‒E)
Conchiliolithus (Anomites) resupinatus Martin, 1809, pl. 49, ﬁgs.
13, 14.
Orthis resupinata (Martin). Davidson, 1861, p. 130, pl. 29, ﬁgs.
1‒4; pl. 30, ﬁgs. 1‒5.

Schizophoria resupinata (Martin). Yanishevsky, 1918, p. 19, pl.
1, ﬁgs. 4, 12; pl. 4, ﬁg. 2; pl. 6, ﬁg. 16; Demanet, 1934, p. 45,
pl. 3, ﬁgs. 1‒5, text-ﬁg. 9; Miloradovich, 1935, p. 6, pl. 1, ﬁgs.
11, 12; Bond, 1941，p．289，pl．21, ﬁgs. A‒C, text-ﬁgs. 33,
34; Minato, 1952, p. 150, pl. 5, ﬁg. 3; pl. 6, ﬁg. 4; Sarytcheva
in Sarytcheva and Sokolskaya, 1952, p. 29, pl. 2, fig. 12;
Parkinson, 1954, p. 368, text-figs. 1, 2; Litvinovich, 1962,
p. 178, pl. 1, fig. 2; Besnossova in Sarytcheva et al., 1963,
p. 77, pl. 3, ﬁgs. 5‒8, text-ﬁg. 24; Ustritskiy and Tschernjak,
1963, p. 69, pl. 1, figs. 13‒16; Yang, 1964, p. 59, pl. 1, figs.
2, 3; Abramov, 1965, p. 35, pl. 2, ﬁg. 3; Brunton, 1968, pl. 2,
ﬁgs. 1‒6; Pocock, 1968, p. 80, pl. 18, ﬁg. 7, text-ﬁgs. 13‒15;
Besnossova et al. in Sarytcheva, 1968, p. 53, pl. 1, ﬁgs. 11‒13;
Lazarev, 1969, pl. 10, ﬁgs. 1‒5, text-ﬁgs. 1, 2; Litvinovich etal., 1969, p. 129, pl. 2, ﬁg. 1; Abramov, 1970, p. 107, pl. 1, ﬁgs.
5‒7; Aisenverg and Poletaev, 1971, pl. 60, ﬁg. 1; Nalivkin andFotieva, 1973, p. 20, pl. 1, ﬁgs. 6‒8; Yanagida, 1973, p. 101, pl.
16, ﬁgs. 3‒9; Kalashnikov, 1974, p. 22, pl. 3, ﬁgs. 1‒3; Garanj
et al., 1975, p. 155, pl. 62, ﬁg. 2; Volgin and Kushnar, 1975, p.
23, pl. 1, ﬁgs. 3‒5; Litvinovich et al., 1975, p. 52, pl. 16, ﬁg. 7;
Lazarev, 1976, pl. 2, ﬁgs. 3, 4; pl. 3, ﬁgs. 1‒5; text-ﬁg. 58; table
11; Lee and Gu, 1976, p. 229, pl. 131, ﬁgs. 7, 9‒11; Martinez
Chacon, 1979, p. 54, pl. 2, ﬁgs. 1‒15; pl. 3, ﬁgs. 1‒10; textﬁgs. 3‒5; Minato et al., 1979a, pl. 22, ﬁgs. 1, 2; Tazawa andKatayama, 1979, p. 169, pl. 11, ﬁgs. 8‒14; Kalashnikov, 1980,
p. 24, pl. 2, figs. 2, 3; Mori and Tazawa, 1980, text-fig. 3.3;
Tazawa, 1981b, p. 67, pl. 5, ﬁgs. 3‒5; Tazawa et al., 1981, pl. 1,
ﬁgs. 4‒6; Ding and Qi, 1983, p. 245, pl. 88, ﬁg. 7; Zhang et al.,1983, p. 265, pl. 107, ﬁgs. 1‒3; Tazawa, 1984b, p. 304, pl. 61,
ﬁg. 9; Abramov and Grigorjeva, 1986, p. 74, pl. 1, ﬁgs. 15‒18;
Yanai et al., 1988, pl. 1, ﬁgs. 9, 10; Zakowa, 1989, p. 103, pl.
1, figs. 1‒5; pl. 2, figs. 1‒5; pl. 3, figs. 1‒4; text-figs. 2‒10;table 2; Jiang, 1997, pl. 1, ﬁgs. 1, 2; Bassett and Bryant, 2006,
p. 504, pl. 6, ﬁgs. 1‒10; pl. 7, ﬁgs. 1‒16; text-ﬁgs. 5‒7; Butts,
2007, p. 55, ﬁgs. 5.3‒5.10; Ibaraki et al., 2014, p. 73, ﬁgs. 4.1,
4.2.
Schizophoria aff. resupinata (Martin). Yanagida, 1962, p. 122, pl.
21, ﬁgs. 4‒13, text-ﬁg. 22; Hase and Yokoyama, 1975, pl. 16,
ﬁgs. 6, 7.
Schizophoria (Schizophoria) resupinata (Martin). Sun andBaliński, 2008, p. 521, ﬁgs. 27F‒L.
Material.―Sixteen specimens from localities KAC1, KF167,
KF188, KF200 and KF201: (1) external and internal moulds offive ventral valves, NU-B2188, 2189, 2218‒2220; (2) internalmould of ﬁve ventral valves, IGPS97898, NU-B2058, 2221‒2223;(3) external and internal moulds of a dorsal valve, NU-B2224;
and (4) internal moulds of ﬁve dorsal valves, NU-B2059, 2060,
2190, 2225, 2226.
Remarks.―These specimens are referred to Schizophoria
resupinata (Martin, 1809), redescribed by Pocock (1968, p. 80,
pl. 18, ﬁg. 7, text-ﬁgs. 13‒15) from the upper Tournaisian‒upper
Visean of Belgium and the UK (England, Wales and Isle of Man),in the medium size (length about 42 mm, width about 55 mm in
the largest specimen, NU-B2218), wider subrectangular outline,

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

55

FIGURE 23. Brachiopods of the HK2 assemblage (2). A‒D, Marginatia burlingtonensis (Hall); A, external latex cast of dorsal valve, NU-B629; B, internallatex cast (B1) and internal mould (B2) of dorsal valve, NU-B633; C, external mould of dorsal valve, NU-B634; D, external latex cast (D1) and external mould (D2)of dorsal valve, NU-B643; E, F, Ovatia laevicosta (White); E, external latex cast of ventral valve, NU-B2065; F, external latex cast (F1, F2) and ventral view(F3), anterior view (F4), posterior view (F5) and lateral view (F6) of external mould of ventral valve, NU-B2064; G, H, Echinoconchus punctatus (Sowerby); G,internal mould of ventral valve, NU-B2061; H, external mould (H1) and internal mould (H2) of dorsal valve, NU-B2063; I, Tolmatchoffia robusta (Tolmatchoff),
external latex cast (I1), internal latex cast (I2), external mould (I3) and cardinal process (I4) of dorsal valve, NU-B2044. Scale bars represent 1 cm.

56

JUN-ICHI TAZAWA

moderately convex dorsal valve and the ventral muscle field
moderately incised, ﬂabellate. Schizophoria connivens (Phillips,
1836), redescribed by Pocock (1968, p. 64, pl. 18, ﬁgs. 1, 2, textﬁgs. 1‒4), from the Tournaisian‒Visean of Belgium and the UK(Scotland, England, Wales and Isle of Man), differs from thepresent species in the smaller size and in having more strongly
incised, elliptical muscle ﬁeld in the ventral valve.
Occurrence.―HK1, HK2 and HK4 units.
Distribution.―Upper Tournaisian‒upper Visean: northeasternJapan (Hikoroichi and Yokota in the South Kitakami Belt), central
Japan (Kanto Mountains), southwestern Japan (Akiyoshi Belt),
USA (Idaho), northern Russia (Verkhoyansk Range, Taimyr
Peninsula, Novaya Zemlya and Pechora Basin), UK (England,
Wales, and Isle of Man), Poland, Belgium, Spain, western Russia(Moscow Basin and Donetz Basin), central Russia (southern
Urals and Kuznetsk Basin), Kazakhstan, Uzbekistan (Fergana),
northwestern China (Xinjiang and Shaanxi), northern China (Inner
Mongolia) and southwestern China (Guizhou and Yunnan).
Order RHYNCHONELLIDA Kuhn, 1949
Superfamily RHYNCHOPOROIDEA Muir-Wood, 1955
Family RHYNCHOPORIDAE Muir-Wood, 1955
SubfamilyRHYNCHOPORINAE Muir-Wood, 1955
Genus RHYNCHOPORA King, 1865
Type species.―Terebratula genitziana de Verneuil, 1845.
Rhynchopora sp.
(Fig. 19A, B)
Material.―Two specimens from locality KF201: (1) internalmould of a ventral valve, NU-B2215; and (2) external and internalmoulds of a dorsal valve, NU-B2216.
Description.―Shell large in size for genus, transversely
subpentagonal in outline, with greatest width at about midlength;
length 16 mm, width about 22 mm in the larger specimen (NUB2216); length 15 mm, width 18 mm in the smaller specimen(NU-B2215). Ventral valve gently convex in both lateral andanterior proﬁles; umbo small, pointed; sulcus wide and shallow;
external ornament consisting of numerous simple rounded costae,
numbering seven on sulcus and ﬁve on each lateral slope. Internal
structure of ventral valve not well preserved. Dorsal valve much
more convex than ventral valve; fold wide and low; external
ornament consisting of 17 or more simple, broad and rounded
costae. Interior of dorsal valve with a short and thin median
septum.
Remarks.―These specimens are safely assigned to the genus
Rhynchopora by the transversely subpentagonal outline. The
Hikoroichi species most resembles Rhynchopora sansabensis
Carter (1967, p. 314, pl. 26, figs. 1‒8, text-figs. 8, 9), from the
Chappel Limestone of Texas, in size, shape and external ornamentof both valves, although the number of costae are slightly fewer
than the Texan species. Rhynchopora persinuata (Winchell,1865), redescribed by Carter (1999, p. 115, ﬁgs. 8E‒bb, 9), fromthe lower Osagean of Oklahoma, differs from the present species

in the smaller size and in having more numerous ﬁne costae on
the ventral sulcus.
Occurrence.―HK1 Unit.
Superfamily RHYNCHOTETRADOIDEA Licharew in
Rzhonsnitskaya, 1956
Family TETRACAMERIDAE Licharew in Rzhonsnitskaya, 1956
Genus ROTAIA Rzhonsnitskaya, 1959
Type species.―Rhynchonella subtrigona Meek and Worthen,
1860.
Rotaia hikoroichiensis Tazawa, 2006
(Figs. 26E, F, 30I)
Rotaia sp. Tazawa, 1996, p. 8, ﬁg. 4; Tazawa, 2002, ﬁgs. 7.3‒7.5.
Rotaia hikoroichiensis Tazawa, 2006, p. 135, ﬁgs. 8.1‒8.3.
Material.―Three specimens from localities KF156 andKF202, external and internal moulds of three ventral valves, NUB49, 2173, 2227.
Remarks.―Of the two specimens, the larger one (NU-B49)
was described by Tazawa (2006, p. 135, ﬁgs. 8.1‒8.3) as Rotaia
hikoroichiensis Tazawa, 2006. The smaller one also is referred
to R. hikoroichiensis by the large ventral valve with ill-definedsulcus and coarse costae. Comparison with the other species ofRotaia has been discussed by Tazawa (2006, p. 136).
Occurrence.―HK2 and HK4 units.
Distribution.―Lower‒upper Visean: northeastern Japan(Hikoroichi in the South Kitakami Belt).
Order ATHYRIDIDA Boucot, Johnson and Staton, 1964
Suborder ATHYRIDINA Boucot, Johnson and Staton, 1964
Superfamily ATHYRIDOIDEA Davidson, 1881
Family ATHYRIDIDAE Davidson, 1881
Subfamily ATHYRIDINAE Davidson, 1881
Genus ACTINOCONCHUS MʼCoy, 1844
Type species.―Actinoconchus paradoxus MʼCoy, 1844.
Actinoconchus expansus (Phillips, 1836)(Fig. 26I, J)
Spirifera expansa Phillips, 1836, p. 220, pl. 10, ﬁg. 8.
Athyris expansa (Phillips). Davidson, 1859, p. 82, pl. 16, ﬁgs. 14,
16; pl. 17, ﬁg. 3 only.
Athyris (Actinoconchus) expansa (Phillips). Licharew, 1934, p.
45, 115, pl. 8, ﬁgs. 4, 5.
Actinoconchus expansus (Phillips). Nalivkin and Fotieva, 1973, p.
75, pl. 24, ﬁg. 7.
Actinoconchus? expansus (Phillips). Poletaev, 1975, p. 89, pl. 21,
ﬁgs. 1‒6, text-ﬁg. 19.
Cleiothyridina expansa (Phillips). Jin et al., 1979, p. 124, pl. 36,
ﬁgs. 35, 36.
Actinoconchus expansus expansus (Phillips). Brunton, 1980, p.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

57

FIGURE 24. Brachiopods of the HK2 assemblage (3). A‒C, Linoprotonia ashfellensis Ferguson; A, external latex cast (A1), ventral (A2), anterior (A3),posterior (A4) and lateral (A5) views of internal moud of ventral valve, NU-B597; D‒H, Linoprotonia hikoroichiensis Tazawa and Ibaraki; D, external latex cast(D1), ventral (D2), anterior (D3), posterior (D4) and lateral (D5) views of ventral valve, NU-B603; E, external latex cast (E1) and internal mould (E2) of ventral
valve, NU-B604; F, external mould of dorsal valve, NU-B616; G, internal mould of ventral valve, NU-B608; H, internal mould (H1) and external mould (H2) of
dorsal valve, NU-B615. Scale bars represent 1 cm.

58

JUN-ICHI TAZAWA

FIGURE 25. Brachiopods of the HK2 assemblage (4). A, C, Gigantoproductus crassiventer (Prentice); A, ventral (A1), anterior (A2), posterior (A3) and lateral(A4) views of internal mould of ventral valve, NU-B622; C, internal mould (C1) and external latex cast (C2) of ventral valve, NU-B621; B, Gigantoproductus
latiexpansus Sarytcheva, external latex cast (B1), and ventral (B2), anterior (B3), posterior (B4) and lateral (B5) views of internal mould of ventral valve, NUB624; D, Orthotetes keokuk (Hall), external mould of ventral valve, NU-B2105. Scale bars represent 1 cm.

222, ﬁg. 10.
Material.―Three specimens from locality KF202: (1) external
and internal moulds of a ventral valve, NU-B2110; and (2)external and internal moulds of two dorsal valves, NU-B2111,
2112.
Description.―Shell large in size for genus, transversely oval
to elliptical in outline, with greatest width at about midlength;
length about 25 mm, width about 29 mm in the single ventral

valve specimen (NU-B2110); length about 23 mm, width about
32 mm in the larger dorsal valve specimen (NU-B2111). Ventral
valve gently convex in both lateral and anterior profiles; umbo
small; sulcus absent. Dorsal valve slightly convex in both proﬁles;
fold absent. External surface of both valves ornamented withnumerous concentric lamellae with short ﬂanges, numbering 5‒6lamellae in 3 mm near anterior margin of ventral valve. Internalsurface of ventral valve sculptured with faint radial vascularmarkings. Dorsal interior with a pair of stout diverging socket

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION59

FIGURE 26. Brachiopods of the HK2 assemblage (5). A‒C, Rhipidomella michelini (Léveillé); A, internal mould (A1, A2) of ventral valve, NU-B2054; B,internal mould of ventral valve, NU-B2056; C, internal mould of dorsal valve, NU-B2057; D, Hustedia sp., internal mould (D1, D2) of ventral valve, NUB2109; E, F, Rotaia hikoroichiensis Tazawa; E, external latex cast (E1) and internal mould (E2) of ventral valve, NU-B49; F, external latex cast (F1) and internalmould (F2) of ventral valve, NU-B2173; G, H, Schizophoria resupinata (Martin); G, internal mould of ventral valve, NU-B2058; H, internal mould of dorsal
valve, NU-B2059; I, J, Actinoconchus expansus (Phillips), I, external latex cast (I1) and internal mould (I2) of ventral valve, NU-B2110; J, external latex cast(J1), enlarged concentric lamellae (J2) and internal mould (J3) of dorsal valve, NU-B2111; K, L, Actinoconchus paradoxus MʼCoy; K, external latex cast (K1),
enlarged concentric lamellae (K2) and internal mould (K4) of dorsal valve, and internal mould (K3) of ventral valve, NU-B2113; L, external latex cast of ventral
valve, NU-B2114; M, N, Lamellosathyris qaidamensis Chen, Shi and Zhan; M, internal mould of ventral valve, NU-B2036; N, internal mould of dorsal valve,
NU-B2039. Scale bars represent 1 cm, unless otherwise speciﬁed.

60

JUN-ICHI TAZAWA

ridges.
Remarks.―These specimens are referred to Actinoconchus
expansus (Phillips, 1836), redescribed by Brunton (1980, p. 222,
fig. 10) as Actinoconchus expansus expansus (Phillips, 1836)
from the lower Visean of Bolland, Yorkshire, in the large size andtransversely elliptical outline. Actinoconchus patulus Brunton(1980, p. 222, ﬁgs. 7‒9), from the Visean of England, is readily
distinguished from A. expansus by the lower and wider outline.
Occurrence.―HK2 Unit.
Distribution.―Lower Visean‒Serpukhovian: northeasternJapan (Hikoroichi in the South Kitakami Belt), northern Russia(Kolyma), UK (England), western Russia (Donetz Basin), central
Russia (western Urals) and northwestern China (Qinghai).
Actinoconchus paradoxus MʼCoy, 1844
(Fig. 26K, L)
Actinoconchus paradoxus MʼCoy, 1844, p. 150, pl. 21, fig. 6;
Brunton, 1980, p. 221, ﬁgs. 4‒6.
Athyris planosulcata (Phillips). Davidson, 1859, p. 80, pl. 16, ﬁgs.
9‒11 only.
Material.―Two specimens from locality KF202: (1) internalmould of a conjoined shell, with external mould of the dorsal
valve, NU-B2113; and (2) external mould of a ventral valve, NUB2114.
Remarks.―These specimens can be referred to Actinoconchus
paradoxus MʼCoy, 1844, redescribed by Brunton (1980, p. 221,ﬁgs. 4‒6) on the type specimens from the lower Visean of County
Kildare, Ireland, by the small (length 18 mm, width about 23
mm in the better preserved specimen, NU-B2113), subcircularand moderately convex ventral valve without sulcus and theexternal ornament consisting of numerous concentric lamellae(numbering 5‒6 in 3 mm near anterior valve margin) with faintradiating striations. Actinoconchus planosulcatus (Phillips, 1836),
redescribed by Brunton (1980, p. 224, ﬁgs. 13, 14) from the lower
Visean of Bolland, Yorkshire, differs from A. paradoxus in having
a very shallow sulcus on the ventral valve.
Occurrence.―HK2 Unit.
Distribution.―Lower Visean: northeastern Japan (Hikoroichiin the South Kitakami Belt) and Ireland (Kildare).
Genus LAMELLOSATHYRIS Jin and Fang, 1983
Type species.―Spirifer lamellosus Léveillé, 1835.
Lamellosathyris qaidamensis Chen, Shi and Zhan, 2003
(Figs. 26M, N, 27A)
Athyris lamellosa (Léveillé). Hall and Clarke, 1895, p. 46, ﬁgs.
16‒20; Weller, 1914, p. 465, pl. 78, ﬁgs. 1‒5, 15‒20; Zhang etal., 1983, p. 375, pl. 124, ﬁgs. 1, 2.
Actinoconchus cf. lamellosa (Léveillé). Minato, 1951, p. 380, pl.
1, ﬁg. 6.
Actinoconchus lamellosa (Léveillé). Minato, 1952, p. 173, pl. 11,

ﬁg. 6.
Cleiothyridina lamellosa (Léveillé). Yang, 1964, p. 144, pl. 22,
ﬁgs. 1, 2.
Athyris lamellosus (Léveillé). Litvinovich in Litvinovich et al.,1969, p. 281, pl. 69, ﬁg. 6.
Actinoconchus lamellosus (Léveillé). Minato et al., 1979a, pl. 19,
ﬁg. 1.
Lamellosathyris lamellosa (Léveillé). Tazawa, 1989, p. 61, pl. 1,
ﬁg. 3; Carter, 1999, p. 121, ﬁgs. 13, 14.
Lamellosathyris qaidamensis Chen, Shi and Zhan, 2003, p. 849,
ﬁgs. 4, 5.1‒5.6, 5.8, 5.9, 5.11, 5.12, 5.16, 5.17, 6.
Material.―Twelve specimens from locality KF202: (1)external and internal moulds of a conjoined shell, NU-B2029;(2) internal moulds of two conjoined shells, NU-B2030, 2031;(3) external moulds of four ventral valves, NU-B2032‒2035; (4)internal moulds of three ventral valves, NU-B2036‒2038; and (5)internal moulds of two dorsal valves, NU-B2039, 2040.
Description.―Shell large in size for genus, transversely
subelliptical in outline; dorsibiconvex in lateral proﬁle; cardinal
extremities rounded; hinge much shorter than greatest width atmidlength; length 37 mm, width 55 mm in the largest specmen(NU-B2029). Ventral valve moderately and unevenly convex inlateral proﬁle, strongly convex in posterior half, and gently convex
to nearly ﬂat in anterior half; umbo small, slightly incurved andhaving large rounded foramen; interarea very narrow, convex;
sulcus originating near beak, shallow and narrow, with roundedbottom. Dorsal valve a little more convex than ventral valve, withgreatest convexity posterior to midlength; umbo incurved beneath
opposite valve; fold low and narrow on anterior portion of valve;
External surface of both valves ornamented with regular, strong
concentric lamellae, numbering 2‒3 in 10 mm length on ventral
valve. Ventral interior with thick short dental plates, large broadteeth, poorly deﬁned ﬂabellate diductor scars and narrow adductor
scars. Dorsal interior with small, triangular cardinal plates;
spiralia directed laterally.
Remarks.―These specimens were described by Tazawa(1989, p. 61, pl. 1, ﬁg. 3) as Lamellosathyris lamellosa (Leveillé,1835). But the Hikoroichi specimens can be referred toLamellosathyris qaidamensis Chen, Shi and Zhan, 2003, from theupper Tournaisian‒lower Visean of the Qaidam Basin, Qinghai,
northwestern China, on account of the large size, dorsibiconvex
proﬁle and strong broad concentric lamellae on the both ventral
and dorsal valves. The type species, Lamellosathyris lamellosa(Léveillé, 1835), redescribed by Brunton (1980, p. 225, figs.
15‒17) from the upper Tournaisian of Tournai, Belgium, differsfrom L. qaidamensis in the smaller size and in having more
numerous and narrower concentric lamellae on the both valves.
Shells of Lamellosathyris species from the lower Carboniferousof the South Kitakami Belt, Actinoconchus cf. lamellosa (Léveillé,1835) described by Minato (1951, p. 380, pl. 1, fig. 6) fromthe Jumonji Stage (=middle part of the Arisu Formation) of the
Shimoarisu area and Actinoconchus lamellosa (Léveillé, 1835)
described by Minato (1952, p. 173, pl. 11, ﬁg. 6) from the Jumonji
Stage of the Shimoarisu and Yokota areas, are conspeciﬁc with L.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

61

FIGURE 27. Brachiopods of the HK2 assemblage (6). A, Lamellosathyris qaidamensis Chen, Shi and Zhan, ventral (A1) and dorsal (A2) views of externallatex cast, and ventral (A3) and dorsal (A4, A5) views of internal mould of conjoined shell, NU-B2029; B, C, Spirifer karagai (Litvinovich); B, external latex
cast (B1) and internal mould (B2) of ventral valve, NU-B2132; C, external latex cast (C1) and internal mould (C2) of dorsal valve, NU-B2134; D‒G, Unispirifer
striatoconvolutus (Dun and Benson); D, external latex cast (D1) and internal mould (D2) of ventral valve, NU-B2096; E, internal mould of ventral valve, NUB2099; F, internal mould of dorsal valve, NU-B2101; G, external latex cast (G1) and internal mould (G2) of dorsal valve, NU-B2100. Scale bars represent 1 cm.

62

JUN-ICHI TAZAWA

FIGURE 28. Brachiopods of the HK2 assemblage (7). A‒E, Unispirifer kozuboensis (Minato); A, external latex cast of ventral valve, NU-B2086; B, internalmould of ventral valve, NU-B2088; C, internal mould of ventral valve, NU-B2089; D, external latex cast of ventral valve, NU-B2087; E, external latex cast (E1)and internal mould (E2) of dorsal valve, NU-B2093; F‒K, Tegulocrea incerta (Hall), F, external latex cast (F1, F2) and internal mould (F3) of ventral valve, NUB2066; G, external latex cast of ventral valve, NU-B2071; H, external latex cast of dorsal valve, NU-B2075; I, external latex cast (I1, I2) and internal mould (I3)of dorsal valve, NU-B2068; J, internal mould of ventral valve, NU-B2080; K, external latex cast (K1, K2) and internal mould (K3) of ventral valve, NU-B2067;
L, Acuminothyris triangularis Roberts, internal mould of dorsal valve, NU-B2121; M, N, Pseudocyrinx cf. missouriensis Weller; M, external mould of ventral
interarea, NU-B2041; N, internal mould of dorsal valve, NU-B2108. Scale bars represent 1 cm.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

qaidamensis.
Occurrence.―HK2 Unit.
Distribution.―Upper Tournaisian‒lower Visean: northeasternJapan (Hikoroichi, Shimoarisu, and Yokota in the South KitakamiBelt), USA (Ohio, Indiana, Missouri and Oklahoma), Kazakhstan
and northwestern China (Xinjiang and Qinghai).
Subfamily CLEIOTHYRIDININAE Alvarez, Rong and Boucot,
1998
Genus CLEIOTHYRIDINA Buckman, 1906
Type species.―Atrypa pectinifera Sowerby, 1840.
Cleiothyridina harkeri Carter, 1987
(Fig. 19C)
Cleiothyridina obmaxima (McChesney)?. Nelson, 1961, pl. 4,
ﬁgs. 5, 6; pl. 7, ﬁg. 14.
Cleiothyridina harkeri Carter, 1987, p. 60, pl. 19, ﬁgs. 24‒27.
Material.―One specimen from locality KF200, internal mouldof a ventral valve, with partly preserved shell substance, NUB2201.
Remarks.―This specimen is referred to Cleiothyridina
harkeri Carter (1987, p. 60, pl. 19, ﬁgs. 24‒27), from the upper
part of the Banff Formation of western Alberta, Canada, by thelarge size (length 28 mm, width about 50 mm), transversely
subelliptical outline and gently convex ventral valve, with weak
shallow sulcus. The external surface of the ventral valve iscovered by very dense, narrow growth lamellae which are fringed
with numerous flattened spines. Carter (1987, p. 60) noted that
Cleiothyridina obmaxima (McChesney)?, figured by Nelson(1961, pl. 4, figs. 5, 6; pl. 7, fig. 14) from the Banff Formationof Alberta, is conspeciﬁc with C. harkeri. Cleiothyridina royssii(Léveillé, 1835), redescribed by Brunton (1980, p. 228, ﬁgs. 23,24), from the upper Tournaisian of Tournai, Belgium, differs fromC. harkeri in the smaller size and in having deeper ventral sulcus.
Occurrence.―HK1 Unit.
Distribution.―Lower‒upper Tournaisian: northeastern Japan(Hikoroichi in the South Kitakami Belt) and northern Canada(Alberta).
Cleiothyridina sp.
(Fig, 33C)
Cleiothyridina sp. Tazawa, 1984b, p. 307, pl. 61, ﬁg. 5.
Material.―One specimen from locality KF167, externalmould of a valve fragment, IGPS97907.
Remarks.―The fragmentarily preserved specimen from
Hikoroichi is safely assigned to the genus Cleiothyridina by thecharacteristic external ornament of regular concentric lamellae,edges of which are fringed with numerous ﬂattened spines. But
the Hikoroichi specimen is too poorly preserved for specificdetermination.
Occurrence.―HK4 Unit.

63

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 sp.
(Fig. 26D)
Material.―One specimen from locality KF202, internal mouldof a ventral valve, NU-B2109.
Remarks.―This specimen is safely assigned to the genus
Hustedia by the small, elongate-oval ventral valve (length about
12 mm, width about 7 mm) with shallow sulcus and the external
ornament consisting of relatively few strong costae (numbering
6 on one side of lateral flanks). The Hikoroichi species mostresembles Hustedia aff. radialis (Phillips, 1836), described by
Nalivkin (1937, p. 120, pl. 34, figs. 5, 6) from the Kassin and
Rusakov beds of northeastern Kazakhstan, in size, shape andexternal ornament of the ventral valve. Hustedia radialis (Phillips,
1836), redescribed by Brunton (1984, p. 44, ﬁgs. 30‒38) from the
Visean of Yorkshire, England, differs from the present species in
the wider outline and more numerous costae on the ventral valve.
Occurrence.―HK2 Unit.
Order SPIRIFERIDA Waagen, 1883
Suborder SPIRIFERIDINA Waagen, 1883
Superfamily SPIRIFEROIDEA King, 1846
Famiy SPIRIFERIDAE King, 1846
Subfamily SPIRIFERINAE King, 1846
Genus SPIRIFER Sowerby, 1816
Type species.―Conchyliolithus (Anomia) striatus Martin,
1793.
Spirifer liangchowensis Chao, 1929
(Fig. 31G)
Spirifer liangchowensis Chao, 1929, p. 6, pl. 1, figs. 1‒7, textﬁg. 1; Yang in Yang et al., 1962, p. 99, pl. 39, ﬁg. 1; Tazawa,
2017b, p. 340, ﬁgs. 8.1‒8.5.
Neospirifer liangchowensis (Chao, 1929): Wang et al., 1964, p.
508, pl. 89, ﬁgs. 14‒16; Ding and Qi, 1983, p. 401, pl. 135, ﬁg. 5.
Spirifer sp. Tazawa, 1984b, p. 307, pl. 61, ﬁg. 1.
Material.―One specimen from locality KF151, an imperfectconjoined shell, IGPS97908.
Remarks.―The single specimen from Hikoroichi wasdescribed by Tazawa (1984b, p. 307, pl. 61, ﬁg. 1) as Spirifer sp.
But the specimen can be referred to Spirifer liangchowensis Chao
(1929, p. 6, pl. 1, figs. 1‒7, text-fig. 1), from the Chouniugou
Formation of Gansu, northwestern China, on account of the

64

JUN-ICHI TAZAWA

average-sized, less transverse shell (length more than 39 mm,
width about 50 mm), with narrow and deep but not clearly
demarcated ventral sulcus. The Hikoroichi specimen resembles
well the shells, described by Tazawa (2017b, p. 340, ﬁgs. 8.1‒8.5)
as S. liangchowensis from the uppermost part of the Odaira
Formation (upper Visean) in the Yokota area, South Kitakami
Belt. Spirifer karagai (Litvinovich, 1962, p. 273, pl. 33, fig.
2), from the upper Visean of central Kazakhstan, resembles S.
liangchowensis in size and outline of the shell, but differs fromthe latter in having shallower ventral sulcus and lower dorsal fold.
Occurrence.―HK4 Unit.
Distribution.―Lower‒upper Visean: northeastern Japan(Hikoroichi and Yokota in the South Kitakami Belt) andnorthwestern China (Gansu).
Spirifer karagai (Litvinovich, 1962)(Fig. 27B, C)
Neospirifer karagai Litvinovich, 1962, p. 273, pl. 33, ﬁg. 2.
Spirifer karagai (Litvinovich). Litvinovich et al., 1969, p. 259, pl.
62, ﬁgs. 1‒4.
Material.―Four specimens from locality KF202: (1) external
and internal moulds of a ventral valve, NU-B2132; (2) internalmould of a ventral valve, NU-B2133; (3) external and internalmoulds of a dorsal valve, NU-B2134; and (4) internal mould of adorsal valve, NU-B2135.
Description.―Shell medium in size for genus, slightly
transverse, subrectangular in outline, widest at hinge; cardinal
extremities blunt, angular, not mucronate; length 38 mm, widthabout 58 mm in the largest specimen (NU-B2132). Ventral valve
strongly and unevenly convex in lateral proﬁle, most convex inumbonal region; umbo small, strongly incurved; sulcus narrowand shallow, not clearly demarcated from ﬂanks. Dorsal valve lessinﬂated than ventral valve, moderately convex in umbonal region
and flanks, flattened near cardinal extremities; fold narrow andlow, deﬁned by fold-bounding grooves. External surface of both
valves ornamented with numerous, ﬁne, rounded and bifurcating
costae, irregular concentric rugae and very ﬁne numerous growthlines; costae numbering 4‒5 in 5 mm at midlength of ventral
ﬂanks. Internal structures of both valves not well preserved.
Remarks.―These specimens are referred to Spirifer karagai(Litvinovich, 1962, p. 273, pl. 33, ﬁg. 2), from the upper Viseanof central Kazakhstan, on account of the middle-sized, lesstransverse shell, with shallow ventral sulcus and low dorsal fold.
The preceding species, Spirifer liangchowensis Chao (1929), isdistinguished from S. karagai in possessing deeper ventral sulcus
and higher dorsal fold. Spirifer gregeri Weller (1914, p. 359,
pl. 55, figs. 1‒8), from the lower Burlington Limestone of theMississippi Valley, differs from S. karagai in having longer shellwith shorter hinge.
Occurrence.―HK2 Unit.
Distribution.―Lower‒upper Visean: northeastern Japan(Hikoroichi in the South Kitakami Belt) and Kazakhstan.

Subfamily PROSPIRINAE Carter, 1974
Genus UNISPIRIFER Campbell. 1957
Type species.―Spirifer striatoconvolutus Dun and Benson,
1920.
Unispirifer striatoconvolutus (Dun and Benson, 1920)(Fig. 27D‒G)
Spirifera striato-convoluta Dun and Benson, 1920, p. 350, pl. 20,
ﬁgs. 7, 8.
Unispirifer striatoconvolutus (Dun and Benson). Campbell, 1957,
p. 68, pl. 14, ﬁgs. 1‒9, text-ﬁgs. 10‒12; Jiang, 1993, p. 216, pl.
1, ﬁgs. 1‒14, text-ﬁgs. 1, 2; Jiang, 1997, pl. 3, ﬁgs. 12‒14; pl. 4,
ﬁg. 13.
Unispirifer (Unispirifer) striatoconvoluts (Dun and Benson). Shi
et al., 2005, p. 55, ﬁgs. 12G‒K.
Material.―Eight specimens from locality KF202: (1) external
and internal moulds of two ventral valves, NU-B2095, 2096;(2) external moulds of two ventral valves, NU-B2097, 2098; (3)internal mould of a ventral valve, NU-B2099; (4) external andinternal moulds of two dorsal valves, NU-B2100, 2101; and (5)internal mould of a dorsal valve, NU-B2102.
Description.―Shell large in size for genus, transversely
semicircular in outline, with greatest width at hinge, cardinal
extremities mucronate; length 32 mm, width about 70 mm in
the largest ventral valve specimen (NU-B2097). Ventral valve
moderately convex in lateral profile, most convex at umbonal
region; umbo small, incurved; sulcus narrow and shallow inposterior half, becoming broader and not clearly demarcatedfrom flanks anteriorly. Dorsal valve less convex than opposite
valve; fold very shallow, indistinct. External surface of both
valves ornamented with numerous moderately strong costae,
few irregular concentric rugae and very fine closely spaced
growth lines; costae flatly rounded, simple on flanks and often
bifurcated on both sulcus and fold, numbering 7‒8 costae in 10mm at anterior margin of ventral ﬂanks. Ventral interior with stout
divergent dental plates and heart-shaped muscle field. Dorsalinterior not well preserved.
Remarks.―These specimens are referred to Unispirifer
striatoconvolutus (Dun and Benson, 1920), redescribed byCampbell (1957, p. 68, pl. 14, ﬁgs. 1‒9, text-ﬁgs. 10‒12) from theMerlewood Formation of Babbinboon, New South Wales, eastern
Australia, on account of the large, transverse and mucronate shells
with very shallow ventral sulcus and low indistinct dorsal fold.
Unispirifer fluctuosus (Glenister, 1955, p. 68, pl. 7, figs. 1‒14;
pl. 8, fig. 1‒8), from the Moogooree Limestone of the NorthWest Basin, western Australia, resembles U. striatoconvolutus ingeneral shape, but differs in the smaller size and in having coarser
costae on the both valves.
Occurrence.―HK2 Unit.
Distribution.―Upper Tournaisian‒upper Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt), southwestern
China (Yunnan) and eastern Australia (New South Wales).

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

65

FIGURE 29. Brachiopods of the HK2 assemblage (8). A, B, Kitakamithyris hikoroitiensis Minato; A, external latex cast (A1, A2), external moud (A3), enlarged
concentric rows of biramous spine bases (A4) and internal mould (A5) of ventral valve, and external latex cast (A6) and internal mould (A7) of dorsal valve
in a conjoined shell, NU-B2084; B, internal mould of ventral valve, showing numerous spines on the antero-lateral margins of the valve, NU-B2085; C,
Punctospirifer gigantiformis Lee and Gu, external latex cast of ventral valve, NU-B2107; D, Punctospirifer plicatosulcatus Glenister, external latex cast ofventral valve, NU-B2106. Scale bars represent 1 cm, unless otherwise speciﬁed.

Unispirifer mediocris (Tolmatchoff, 1924)(Fig. 19G‒J)
Spirifer mediocris Tolmatchoff, 1924, p. 181, 561, pl. 11, ﬁgs. 13,
14.
Spirifer (Spirifer) marionensis Shumard. Nalivkin, 1937, p. 100,
pl. 28, ﬁg. 1.
Fusella mediocris (Tolmatchoff). Besnossova, 1959, p. 71, pl.

3, fig. 10; pl. 11, fig. 10; text-figs. 24, 25; Besnossova in
Sarytcheva et al., 1963, p. 289, pl. 52, ﬁg. 8; Abramov, 1965,
p. 48, pl. 9, ﬁg. 1; Abramov, 1970, p. 137, pl. 25, ﬁgs. 12‒17;
Kalashnikov, 1974, p. 121, pl. 40, figs. 5, 6, text-fig. 21;
Litvinovich et al., 1975, p. 89, pl. 28, ﬁgs. 12, 13.
Unispirifer? corpulentus Carter, 1968, p. 1149, pl. 147, figs.
10‒22.

66

JUN-ICHI TAZAWA

Material.―Four specimens from locality KF201: (1) internalmoulds of two ventral valves, NU-B2209, 2210; and (2) internalmoulds of two ventral valves, NU-B2211, 2212.
Description.―Shell small in size for genus, slightly transverse
subelliptical in outline, with greatest width at hinge; length 26mm, width 34 mm in the better preserved dorsal valve specimen(NU-B2212). Ventral valve moderately convex in lateralslopes; umbo small, incurved; ears small, mucronate; interarea
subtrigonal, concave; sulcus shallow, bearing four or ﬁve sulcul
costae; lateral ﬂanks ornamented with numerous simple rounded
costae. Dorsal valve gently convex in lateral proﬁle; fold low but
clearly demarcated from lateral slopes; lateral slopes ornamented
with numerous rounded costae on whole valve and a few strong
irregular concentric rugae near anterior margin. Interior of ventral
valve with stout diverging dental plates, enclosing posteriror
portion of muscle ﬁeld. Internal structure of dorsal valve not wellpreserved.
Remarks.―These specimens are referred to Unispirifer
mediocris (Tolmatchoff, 1924, p. 181, 561, pl. 11, figs. 13,14), from the Tournaisian of the Kuznetsk Basin, on accountof size, shape and external ornament of both ventral and dorsal
valves, in particular, the shallow ventral sulcus, low dorsal fold
and mucronate cardinal extremities. Unispirifer? corpulentus
Carter (1968, p. 1149, pl. 147, ﬁgs. 10‒22), from the Burlington
Limestone of Missouri, is deemed a junior synonym of U.
mediocris. Unispirifer kozuboensis (Minato, 1952, p. 155, pl. 5,
ﬁg. 7), from the Jumonji Stage of the Yokota area, South Kitakami
Belt, differs from U. mediocris in the larger size and moretransverse outline.
Occurrence.―HK1 Unit.
Distribution.―Lower Tournaisian‒lower Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt), USA (Missouri),
northern Russia (Verkhoyansk Range and northern Urals), central
Russia (Kuznetsk Basin) and Kazakhstan.

produced, triangular and nearly ﬂat. Dorsal valve gently convex
in both lateral and anterior proﬁles; fold narrow and low, clearly
demarcated from lateral flanks by a pair of distinct grooves
in juveniles, but indistinct in adults. External surface of both
valves ornamented with numerous moderately strong costae,
irregular concentric rugae and very fine closely spaced growthlines; costae becoming broad and flat anteriorly, mostly simple
but rarely bifurcated; numbering 7‒8 in 10 mm on lateral ﬂanks
near anterior margin of ventral valve. Ventral interior with stout
divergent dental plates. Dorsal interior not well preserved andobscure.
Remarks.―These specimens are referred to Unispirifer
kozuboensis (Minato, 1952), originally described by Minato(1952, p. 155, pl. 5, ﬁg. 7) as Spirifer kozuboensis from the Arisu
Formation of Kozubo (Otsubo), Yokota area, South Kitakami Belt,in size, shape and external ornament of both valves, particularly,
in having mucronate cardinal extremities. Unispirifer tornacensis(de Koninck, 1883), redescribed by Sartenaer and Plodowski(1996, p. 60, pl. 1, ﬁgs. 1‒10; pl. 2, ﬁgs. 11‒19; pl. 3, ﬁgs. 27‒41;
pl. 4, ﬁgs. 42‒58; pl. 5, ﬁgs. 60‒73) as Atylephorus tornacensis(de Koninck, 1883) from the upper Tournaisian of Belgium,
somewhat resembles U. kozuboensis, but the Belgian species
differs from the Kitakami species in the larger dimensions and inpossessing less numerous, stronger costae on the both valves.
Occurrence.―HK2 Unit.
Distribution.―Upper Tournaisian‒lower Visean: northeasternJapan (Hikoroichi, Yokota and Okuhinotsuchi in the South
Kitakami Belt).

Unispirifer kozuboensis (Minato, 1952)(Fig. 28A‒E)

Angiospirifer bisulcatus (Sowerby, 1825)(Fig, 31F)

Spirifer kozuboensis Minato, 1952, p. 155, pl. 5, ﬁg. 7; Minato etal., 1979a, pl. 18, ﬁg. 5.
Unispirifer kozuboensis (Minato). Tazawa and Kurita, 1986, p.
167, ﬁg. 2.1.

Spirifer bisulcatus Sowerby, 1825, p. 152, pl. 494, figs. 1, 2;
Scupin, 1900, p. 315, pl. 10, fig. 6; Gröber, 1908, p. 222,
239, pl. 26, figs. 3, 8‒10, text-figs. 3, 4; Yanishevsky, 1918,
p. 82, pl. 6, ﬁgs. 1, 15 only; Frech, 1916, p. 252, pl. 8, ﬁg. 7;
Grabau, 1924, p. 238, text-ﬁg. 152; Rotai, 1931, p. 78, pl. 6,
ﬁgs. 3‒5; Grabau, 1931, p. 26, ﬁg. 6; Demanet, 1938, p. 88, pl.
8, ﬁgs. 17‒19; Dedok and Tschernjak, 1960, p. 65, pl. 8, ﬁg.
3; Litvinovich, 1962, p. 279, pl. 35, ﬁg. 2; Litvinovich et al.,1969, p. 262, pl. 72, ﬁgs. 2‒4; Volgin, 1965, p. 53, pl. 3, ﬁgs. 7,
8.
Spirifera bisulcata (Sowerby). Davidson, 1858, p. 31, pl. 5, ﬁg.
1; pl. 6, ﬁgs. 6‒16 only; Dun and Benson, 1920, p. 347, pl. 21,
ﬁg. 1.
Spirifer trigonalis var. bisulcatus Sowerby. Gröber, 1909, p. 366,
pl. 3, ﬁgs. 2, 4.
Choristites bisulcatus (Sowerby). Grabau, 1932, p. 96, pl. 2, ﬁg.
13; Mori and Tazawa, 1980, text-fig. 3.1; Tazawa, 1984b, p.

Material.―Nine specimens from locality KF202: (1) external
and internal moulds of a ventral valve, NU-B2086; (2) externalmould of a ventral valve, NU-B2087; (3) internal moulds of ﬁveventral valves, NU-B2088‒2092; and (4) external and internalmoulds of two dorsal valves, NU-B2093, 2094.
Description.―Shell medium in size for genus, transversely
subtrigonal in outline, with greatest width at hinge; cardinal
extremities mucronate; length about 35 mm, width about 60 mm
in the largest specimen (NU-B2088). Ventral valve moderately
and unevenly convex in lateral proﬁle, most convex at umbonal
region; umbo small, incurved; sulcus narrow and shallow, notclearly demarcated from lateral flanks; ears moderately large,

Family CHORISTITIDAE Waterhouse, 1968
Subfamily ANGIOSPIRIFERINAE Legrand-Blain, 1985
Genus ANGIOSPIRIFER Legrand-Blain, 1985
Type species.―Spirifer trigonalis Martin, 1809.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

67

FIGURE 30. Brachiopods of the HK4 assemblage (1). A, Leptagonia analoga (Phillips), external mould of dorsal valve, NU-B2239; B, Argentiproductus
margaritaceus (Phillips), external latex cast (B1, B2), internal mould (B3) and internal latex cast (B4) of dorsal valve, NU-B2235; C, D, Karavankina
jizodoensis Tazawa; C, internal mould of ventral valve, NU-B2233; D, external latex cast (D1, D2), internal latex cast (D3) and internal mould (D4) of dorsal
valve, IGPS97905; E, Ovatia cf. prolata Carter, external latex cast of ventral valve, NU-B2232; F, Pustula pustulosa (Phillips), external mould of dorsal
valve, IGPS97904; G, Rhipidomella michelini (Léveillé), internal mould of dorsal valve, IGPS97902; H, Orthotetes keokuk (Hall), external latex cast (H1)and internal mould (H2) of dorsal valve, IGPS97903; I, Rotaia hikoroichiensis Tazawa, external latex cast of fragmented ventral valve, NU-B2227. Scale bars
represent 1 cm.

308, pl. 61, ﬁg. 4.
Spirifer ex gr. bisulcatus Sowerby. Korejwo, 1986, pl. 9, ﬁg. 6;
Abramov, 1970, p. 142, pl. 22, ﬁg. 8.
Angiospirifer bisulcatus (Sowerby). Angiolini et al., 2011, p. 74,
ﬁgs. 1, 2, 6.

Material.―One specimen from locality KF153, external andinternal moulds of a ventral valve, IGPS97909.
Description.―Shell medium in size for genus, wider
subtrigonal in outline, with greatest width at hinge; length 28mm, width 34 mm in the sole specimen (IGPS97909). Ventral

68

JUN-ICHI TAZAWA

valve strongly and unevenly convex in lateral profile, mostconvex at umbonal region; umbo narrow, incurved; cardinal
extremities blunt, produced; sulcus narrow, moderately deep andhaving a pair of simple primary costae and 1 or 2 simple costae
on outside the primary costae. External surface of ventral lateralslopes ornamented with numerous rounded costae with narrowintercostal grooves; costae mostly simple but a few of thembifurcated near anterior valve margin, numbering 9 costae on eachlateral slope. Interior of ventral valve not well preserved. Shellsubstance thick at umbonal region.
Remarks.―This specimen was described by Tazawa (1984b,
p. 308, pl. 61, ﬁg. 4) as Choristites bisulcatus (Sowerby, 1825).
But the genus of the species is now assigned to Angiospirifer
Legrand-Blain, 1985. Angiospirifer bisulcatus most resembles
Angiospirifer pseudobisulcatus Rotai (1951, p. 48, pl. 7, fig.
9; pl. 8, figs. 1, 2), from the lower Bashkirian of the Donetz
Basin, western Russia, but the Russian species differs from thepresent species in having less produced cardinal extremities. The
type species, Angiospirifer trigonalis (Martin, 1809), is readily
distinguished from A. bisulcatus in having anteriorly elevateddorsal fold.
Occurrence.―HK4 Unit.
Distribution.―Upper Visean‒Bashkirian: northeasternJapan (Hikoroichi in the South Kitakami Belt), northern Russia(Verkhoyansk Range and Taimyr Peninsula), UK (Scotland,
England, Isle of Man and northern Ireland), Belgium, Germany,
Poland, western Russia (Donetz Basin), Kazakhstan, Uzbekistan,
Kyrgyzstan, northwestern China (Xinjiang) and eastern Australia(New South Wales).

strongly convex in lateral proﬁle; umbo small, incurved; sulcus
narrow and moderately deep; lateral slopes gently convex. Dorsal
valve slightly less convex than opposite valve; fold narrow andmoderately high; lateral slopes gently convex to nearly ﬂat. Entire
surface of both valves ornamented with numerous ﬁne imbricate
costae; costae often bifurcate, numbering 7‒9 costae in 5 mm
at about midlength of ventral lateral slope; imbricate growthlamellae regularly developed. Ventral interior with short, strong
dental adminicula. Other internal structures of both valves notwell preserved.
Remarks.―These specimens are referred to Tegulocrea
incerta (Hall, 1858), redescribed by Carter (1992, p. 331, ﬁgs. 3,4) from the St. Joe Formation (upper Tournaisian) of Kenwood,
Mayes County, Oklahoma, in the small to medium size, wider
semicircular to subquadrate outline and in possessing numerous
fine imbricate costae on both valves. Shells described by
Minato and Kato (1977, p. 614, pl. 1, ﬁgs. 2‒11) as Imbrexia cf.
incertus (Hall) from the upper part of the Hikoroichi Formation(= Karoyama Formation of Tazawa et al., 1981) at north of
Karoyama, Nisawa area, South Kitakami Belt, differs fromthe present species by the much larger dimensions. Tegulocrea
tenuicostata (Hall, 1858), redescribed by Weller (1914, p. 328, pl.
42, ﬁgs. 4‒16) from the Keokuk, Warsaw and Salem Formationsof the Mississippi Valley, is distinguished from T. incerta by thelarger size and more transverse outline.
Occurrence.―HK2 Unit.
Distribution.―Upper Tournaisian‒lower Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt) and USA (Iowa
and Oklahoma).

Family IMBREXIIDAE Carter, 1992
Genus TEGULOCREA Carter, 1992

Superfamily PAECHELMANELLOIDEA Ivanova, 1972
Family STROPHOPLEURIDAE Carter, 1974
Subfamily STROPHOPLEURINAE Carter, 1974
Genus ACUMINOTHYRIS Roberts, 1963

Type species.―Spirifer incertus Hall, 1858.
Tegulocrea incerta (Hall, 1858)(Fig. 28F‒K)
Spirifer incertus Hall, 1858, p. 602, pl. 13, ﬁg. 3.
Tegulocrea incerta (Hall). Carter, 1992, p. 331, ﬁgs. 3, 4; Carter,
1999, p. 125, ﬁgs. 18A‒D.
Material.―Fifteen specimens from locality KF202: (1)external and internal moulds of two ventral valves, NU-B2066,
2067; (2) external and internal moulds of a dorsal valve, NUB2068; (3) external moulds of five ventral valves, NU-B2069,
2071‒2074; (4) external moulds of three dorsal valves, NUB2070, 2075, 2076; and (5) internal moulds of four ventral valves,
NU-B2077‒2080.
Description.―Shell small to medium in size for genus,transversely semicircular to subquadrate in outline, with greatest
width at slightly anterior to hinge; cardinal extremities obtuse to
blunt, angular; length 19 mm, width 31 mm in the largest dorsal
valve specimen (NU-B2075); length 18 mm, width 22 mm in the
average-sized ventral valve specimen (NU-B2071). Ventral valve

Type species.―Acuminothyris triangularis Robets, 1963.
Acuminothyris triangularis Roberts, 1963
(Fig. 28L)
Acuminothyris triangularis Roberts, 1963, p. 15, pl. 5, ﬁgs. 1‒8,
text-ﬁgs. 7, 8.
Spirifer mundulus Rowley. Minato and Kato, 1977, p. 615, pl. 1,
ﬁg. 1; Minato et al., 1979a, pl. 14, ﬁg. 2.
Material.―One specimen from locality KF202, internal mouldof a dorsal valve, NU-B2121.
Remarks.―The Hikoroichi specimen can be referred to
Acuminothyris triangularis Roberts (1963, p. 15, pl. 5, figs.
1‒8, text-figs. 7, 8), from the Bingleburra Formation (upper
Tournaisian) of Lewinsbrook, New South Wales, eastern Australia,
by the large size (length 8 mm, width 37 mm), very transverse
outline and external ornament of the dorsal valve, consisting of
numerous costae and ﬁne numerous concentric growth lamellae.
The single ventral valve specimen, described by Minato and KatoMISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

69

FIGURE 31. Brachiopods of the HK4 assemblage (2). A‒E, Schizophoria resupinata (Martin); A, external latex cast (A1) and internal mould (A2) of ventral
valve, NU-B2218; B, internal mould of ventral valve, NU-B2219; C, internal latex cast (C1) and internal mould (C2) of ventral valve, NU-B2222; D, internal
nould of dorsal valve, NU-B2225; E, internal latex cast (E1) and internal mould (E2) of dorsal valve, NU-B2226; F, Angiospirifer bisulcatus (Sowerby),
external latex cast (F1) and internal mould (F2) of ventral valve, IGPS97909; G, Spirifer liangchowensis Chao, ventral (G1), anterior (G2), posterior (G3), lateral(G4) and interior (G5, G6) views of conjoined shell, IGPS97908. Scale bars represent 1 cm.

(1977, p. 615, pl. 1, ﬁg. 1) as Spirifer mundulus Rowley (1893)
from the lower part of the Hikoroichi Formation at Omori in the
Hikoroichi area, is identical with the present species in size, shapeand external ornament of the ventral valve.

Occurrence.―HK2 Unit.
Distribution.―Upper Tournaisian‒lower Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt) and eastern
Australia (New South Wales).

70

JUN-ICHI TAZAWA

FIGURE 32. Brachiopods of the HK4 assemblage (3). A, B, Torynifer asiatica Besnossova, external latex cast of ventral valve (A1, A2), external latex castof dorsal valve (A3), internal mould of ventral valve (A4), internal mould of dorsal valve (A5) and enlarged concentric rows of biromous spine bases on theexternal mould of ventral valve of conjoined shell, NU-B2228; B, internal mould of ventral valve, NU-B2229. Scale bars represent 1 cm, unless otherwise
speciﬁed.

Acuminothyris keokuk Carter, 1990
(Fig. 19D‒F)
Acuminothyris keokuk Carter, 1990, p. 236, figs. 8.1‒8.9; Chen
and Archbold, 2000, p. 193, ﬁgs. 5.1‒5.4.
Material.―Six specimens from locality KF201: (1) an abradedconjoined shell, NU-B2115; (2) external mould of a conjoined
shell, NU-B2116; (3) internal mould of a conjoined shell, NUB2117; (4) external and internal moulds of a ventral valve, NUB2118; (5) internal mould of a ventral valve, NU-B2119; and (6)external and internal moulds of a dorsal valve, NU-B2120.

Description.―Shell medium in size for genus, transversely
fusiform in outline, widest at hinge; cardinal extremities
subangular, alate; length 11 mm, width about 28 mm in the largest
specimen (NU-B2120). Ventral valve moderately convex in lateralprofile; umbo small, incurved; interarea low, concave; lateralﬂanks ﬂattened; sulcus narrow and deep, with prominent sulcusbounding costae, Dorsal valve moderately convex as opposite
valve; fold narrow and low, but well differentiated. Externalsurface of ventral valve ornamented with simple rounded costae
and fine concentric lamellae; numbering 11‒12 costae on eachﬂank; 8‒9 lamellae in 3 mm. External ornament of dorsal valve
same as ventral valve. Internal structures of both valves not well

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

preserved.
Remarks.―These specimens are referred to Acuminothyris
keokuk Carter (1990, p. 236, figs. 8.1‒8.9), from the Keokuk
Limestone of Missouri, in size, shape and external ornamentof the both ventral and dorsal valves. The type species,
Acuminothyris triangularis Roberts (1963, p. 15, pl. 5, ﬁgs. 1‒8,
text-ﬁgs. 7, 8) from the upper Tournaisian of New South Wales,
eastern Australia, differs from A. keokuk in the larger and moretransverse shell.
Occurrence.―HK1 Unit.
Distribution.―Upper Tournaisian‒lower Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt), USA (Missouri)
and northwestern China (Xinjiang).
Superfamily BRACHYTHYRIDOIDEA Fredericks, 1924
Family BRACHYTHYRIDIDAE Fredericks, 1924
Genus BRACHYTHYRIS MʼCoy, 1844
Type species.―Spirifera ovalis Phillips, 1836.
Brachythyris suborbicularis (Hall, 1858)(Fig. 19L)
Spirifer suborbicularis Hall, 1858, p. 644.
Brachythyris suborbicularis (Hall). Weller, 1914, p. 374, pl. 61,
ﬁgs. 1‒8; pl. 62, ﬁgs. 1‒12; Nalivkin, 1937, p. 111, pl. 27, ﬁgs.
5, 11, 12; Lapina, 1962, p. 136, pl. 3, ﬁg. 1; Abramov, 1965,
p. 53, pl. 9, fig. 6; Abramov, 1970, p. 150, pl. 22, figs. 4‒7;
Nalivkin and Fotieva, 1973, p. 70, pl. 23, ﬁgs. 1‒3; Garanj etal., 1975, p. 198, pl. 91, ﬁg. 3; Nalivkin, 1979, p. 141, pl. 52,
ﬁgs. 3, 4, 8; pl. 56, ﬁg. 2.
Material.―One specimen from locality KF201, internal mouldof a dorsal valve, NU-B2206.
Remarks.―This specimen can be referred to Brachythyris
suborbicularis (Hall, 1858), redescribed by Weller (1914, p.
374, pl. 61, ﬁgs. 1‒8; pl. 62, ﬁgs. 1‒12) from the Burlington and
Keokuk limestones of the Mississippi Valley, by the medium tolarge size (length more than 30 mm, width about 40 mm), slightly
wider orbicular outline and gently convex dorsal valve, with low
and wide fold and 5‒6 simple, broad and rounded costae on eachside of lateral slope. Brachythyris chouteanensis Weller (1909, p.
305, pl. 13, ﬁg. 11), from the Chouteau Limestone of Missouri,
differs from B. suborbicularis in the smaller size and in having
lower ventral interarea.
Occurrence.―HK1 Unit.
Distribution.―Upper Tournaisian‒upper Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt), USA (Indiana,
Iowa and Missouri), northern Russia (Verkhoyansk Range),
central Russia (western Urals and southern Urals) andKazakhstan.
Suborder DELTHYRIDINA Ivanova, 1972
Superfamily DELTHYRIDOIDEA Phillips, 1841
Family MUCROSPIRIFERIDAE Boucot, 195971

Subfamily TYLOTHYRIDINAE Carter, 1972
Genus TYLOTHYRIS North, 1920
Type species.―Cyrtia laminosa MʼCoy, 1844.
Tylothyris laminosa (MʼCoy, 1844)(Fig. 19K)
Cyrtia laminosa MʼCoy, 1844, p. 137, pl. 21, ﬁg. 4.
Spirifera laminosa (MʼCoy). Davidson, 1858, p. 36, pl. 7, figs.
17‒19 only; Garwood, 1916, pl. 15, ﬁgs. 2, 3.
Syringothyris laminosa forma 2 Vaughan, 1915, p. 43, pl. 6, ﬁgs.
1‒3.
Tylothyris laminosa (MʼCoy). North, 1920, p. 197, pl. 13, ﬁgs. 1,
2, 12, 13, 16, text-ﬁgs. 1m, n, 4a, b; Dehée, 1929, p. 23, pl. 3,
ﬁgs. 12‒14; Plodowski, 1970, p. 21, pl. 1, ﬁgs. 3, 4, text-ﬁg.
9; Nalivkin, 1979, p. 131, pl. 49, ﬁg. 15; Brunton, 1984, p. 77,
ﬁgs. 108‒119; Tazawa et al., 1984, p. 355, pl. 67, ﬁgs. 8‒11;
Sun and Baliński, 2011, p. 833, ﬁgs. 7C, 31.
Spirifer (Tylothyris) laminosa (MʼCoy). Nalivkin, 1937, p. 109, pl.
33, ﬁgs. 3, 4.
Delthyris aff. clarksvillensis (Winchell). Minato, 1952, p. 162, pl.
7, ﬁgs. 2, 5.
Spirifer (Tylothyris) laminosa (MʼCoy). Simorin, 1956, p. 189, pl.
18, ﬁgs. 12, 13.
D e l t h y r i s c l a r k s v i l l e n s i s ( Wi n c h e l l ) . M i n a t o e t a l . ,1979a, pl. 17, ﬁg. 4.
Tylothyris laminosa laminosa (MʼCoy). Bassett and Bryant, 2006,
p. 518, pl. 12, ﬁgs. 1‒6; pl. 14, ﬁgs. 1‒18: pl. 11. ﬁgs. 1‒19: pl.
16, ﬁgs. 1, 2.
Material.―One specimen from locality KAC3, internal mouldof a dorsal valve, NU-B2205.
Remarks.―The material available is lacking the ventral
valve, but it resembles well the specimens of Tylothyris laminosa(MʼCoy, 1844), redescribed by Brunton (1984, p. 77, figs.
108‒119) from the upper Visean of County Fermanagh, northern
Ireland. The single specimen from Hikoroichi is characterized
by the medium size (length 16 mm, width 29 mm), transverse
outline, moderately convex dorsal valve with seven or morecostae on either side of prominent fold and partly preserved
ornament consisting of regular lamellose growth lines. Delthyris
aff. clarksvillensis (Winchell), described by Minato (1952, p.
162, pl. 7, figs. 2, 5) from the Hikoroichi Series (= Karoyama
Formation of Tazawa et al., 1981) at 808 m hill, Okuhinotsuchiin the South Kitakami Belt, is considered to be a synonym of thepresent species. Tylothyris transversa Roberts (1971, p. 224, pl.
50, ﬁgs. 1‒19), from the upper Tournaisian of the Bonaparte Gulf
Basin, northwestern Australia, differs from T. laminosa in being
shorter and more transverse outline.
Occurrence.―HK1 Unit.
Distribution.―Upper Tournaisian‒upper Visean: northeasternJapan (Hikoroichi, Shimoarisu, Okuhinotsuchi and Soma in
the South Kitakami Belt), UK (England and northern Ireland),Belgium, Afghanistan, central Russia (western Urals), Kazakhstan

72

JUN-ICHI TAZAWA

and southwestern China (Guizhou).
Superfamily RETICULARIOIDEA Waagen, 1883
Family ELYTHIDAE Fredericks, 1924
Subfamily ELYTHINAE Fredericks, 1924
Genus KITAKAMITHYRIS Minato, 1951
Type species.―Torynifer (Kitakamithyris) tyoanjiensis Minato,
1951.
Kitakamithyris tyoanjiensis Minao, 1951
(Figs. 20A, 21A)
Torynifer (Kitakamithyris) tyoanjiensis Minato, 1951, p. 374, pl. 1,
ﬁg. 3; pl. 4, ﬁg. 7.
Kitakamithyris tyoanjiensis Minato, 1952, p. 170, pl. 6, ﬁg. 5; pl.
9, ﬁg. 4; Minato, 1953, text-ﬁg. 2; Minato et al., 1979a, pl. 17,
ﬁg. 1.
Material.―Four specimens from localities KF200 and KF201:(1) external and internal moulds of a ventral valve, NU-B2191;
and (2) external moulds of three valves (ventral valve or dorsal
valve unclear), NU-B2192, 2203, 2204.
Description.―Shell medium to large in size for genus,
subelliptical in outline, with greatest width slightly anterior
to midlength; length about 28 mm, width about 27 mm in thebest preserved specimen (NU-B2191). Ventral valve gently andunevenly convex in lateral profile, most convex in umbonal
region; sulcus narrow and shallow. External surface of ventral
valve ornamented with regular concentric lamellae, with a row
of fine biramouse spine bases on anterior margin; numbering
4 lamellae in 5 mm, and 15‒16 spine bases in 5 mm at about
midlength of valve. Ventral interior with a median septum
extending to one-third valve length and a pair of slightly divergent
dental adminicula and covered with numerous ﬁne radial grooves.
Remarks.―These specimens are referred to Kitakamithyris
tyoanjiensis Minato (1951, p. 374, pl. 1, ﬁg. 3; pl. 4, ﬁg. 7), fromthe upper part of the Choanji Formation (upper Famennian)of Choanji, South Kitakami Belt, in having small and closely
arranged spine bases on the ventral valve. Kitakamithyris
hikoroitiensis Minato (1951, p. 375, pl. 1, ﬁg. 1), from the upper
part of the Choanji Formation of Choanji, is readily distinguishedfrom K. thoanjiensis in having wider-spaced and larger spine
bases on the ventral valve.
Occurrence.―HK1 Unit.
Distribution.―Upper Devonian (upper Famennian)‒upper
Tournaisian: northeastern Japan (Choanji and Hikoroichi in theSouth Kitakami Belt).
Kitakamithyris hikoroitiensis Minato, 1951
(Figs. 20B, C, 21B, 29A, B)
Torynifer (Kitakamithyris) hikoroitiensis Minato, 1951, p. 375, pl.
1, ﬁg. 1.
Kitakamithyris hikoroitiensis Minato, 1952, p. 171, pl. 7, ﬁg. 3;

pl. 8, ﬁg. 6; Minato et al., 1979a, pl. 16, ﬁg. 1.
Kitakamithyris semicircularis Minato, 1952, p. 171, pl. 7, ﬁg. 6;
pl. 8, ﬁg. 5; pl. 10, ﬁg. 3; Minato et al., 1979a, pl. 15, ﬁg. 1.
Material.―Five specimens from localities KF188 and KF201:(1) external and internal moulds of a conjoined shell, NU-B2084;(2) external and internal moulds of three ventral valves, NUB2193‒2195; and (3) internal mould of a ventral valve, NUB2085.
Description.―Shell medium in size for genus, subelliptical in
outline, with maximum width at midlength; length 31 mm, widthabout 29 mm in the largest specimen (NU-B2193). Ventral valve
gently convex in lateral proﬁle, most convex in umbonal region,
nearly flat in anterior portion; sulcus narrow and very shallow.
Dorsal valve gently convex in both lateral and anterior proﬁles,without fold. External surface of both valves ornamented withregular concentric lamellae bearing a row of biramous spine
bases; numbering 2 lamellae in 5 mm and 4‒5 spine bases in 5mm at about midlength of ventral valve. Ventral interior with along median septum and a pair of divergent dental adminicula.
Dorsal interior not well preserved. Inner surfaces of both valves
sculptured with ﬁne radiating grooves.
Remarks.―These specimens are referred to Kitakamithyris
hikoroitiensis Minato (1951, p. 375, pl. 1, ﬁg. 1), from the upper
part of the Choanji Formation at Choanji, in size, shape andexternal ornament of the shell, particularly in having rather large
and sporadically arranged spine bases on the both ventral anddorsal valves. Kitakamithyris semicircularis Minato (1952, p.
171, pl. 7, ﬁg. 6; pl. 8, ﬁg. 5; pl. 10, ﬁg. 3), from the Choanji and
Arisu Formations of the South Kitakami Belt, is deemed a juniorsynonym of K. hikoroitiensis. The type species, Kitakamithyris
tyoanjiensis Minato, 1951, is readily distinguished from thepresent species by the smaller and more closely arranged spine
bases on the both valves.
Occurrence.―HK1 and HK2 units.
Distribution.―Upper Devonian (upper Famennian)‒lower
Visean: northeastern Japan (Choanji, Hikoroichi and Yokota in theSouth Kitakami Belt).
Subfamily TORYNIFERINAE Carter in Carter et al., 1994
Genus TORYNIFER Hall and Clarke, 1894
Type species.―Torynifer criticus Hall and Clarke, 1894.
Torynifer asiatica Besnossova in Besnossova et al., 1962
(Fig. 32A, B)
Reticularia lineata Martin. Tolmatchoff, 1924, p. 191, 563, pl. 12,ﬁgs. 1‒3, 5.
Reticularia sinuata Tolmatchoff, 1924, p. 194, 564, pl. 12, ﬁgs. 4,
7.
Reticularia pseudolineata Hall. Nalivkin, 1937, p. 115, pl. 2,
ﬁgs. 5, 6; pl. 33, ﬁg. 6; Rotai, 1941, p. 114, pl. 28, ﬁgs. 2‒4;
Simorin, 1956, p. 198, pl. 17, ﬁgs. 10, 11.
Torynifer pseudolineatus asiaticus Besnossova in Besnossova etMISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION73

FIGURE 33. Brachiopods of the HK4 assemblage (4). A, B, Dimegelasma sp., A, internal latex cast (A1) and internal mould (A2) of ventral valve, IGPS97911;
B, internal latex cast (B1) and internal mould (B2) of ventral valve, IGPS97910; C, Cleiothyridina sp., external latex cast (C1, C2) of valve (ventral or dorsal
uncertain), IGPS97907; D, Punctospirifer iwatensis sp. nov., external latex cast (D1, D2) and internal mould (D3) of ventral valve, NU-B2231 (holotype). Scalebars represent 1 cm.

al., 1962, p. 176, pl. C-23, ﬁgs. 4‒5; Besnossova in Sarytcheva
et al., 1963, p. 303, pl. 58, figs. 1‒6, text-figs. 133‒135;
Kotlyar and Popeko, 1967, p. 173, pl. 48, ﬁgs. 3‒6; Plodowski,
1970, p. 106, pl. 8, ﬁg. 2; Yang and Fan, 1983, p. 279, pl. 4, ﬁg.
17; Zhang et al., 1983, p. 368, pl. 124, ﬁg. 7.
Torynifer pseudolineata (Hall). Litvinovich et al., 1969, p. 270,
pl. 66, ﬁgs. 7, 8.
Torynifer asiatica Besnossova. Pavlova, 1969, p. 63, pl. 4, ﬁgs.
1‒6; pl. 11, ﬁgs. 6, 7; pl. 13, ﬁg. 3; text-ﬁgs. 37‒39.
Material.―Three specimens from locality KAC1: (1) external
and internal moulds of a conjoined shell, NU-B2228; and (2)external and internal moulds of two ventral valves, NU-B2229,
2230.
Description.―Shell large in size for genus, transversely
subelliptical in outline, with greatest width at about two-thirds
length; cardinal extremities rounded; anterior commissure slightly
uniplicate; length 34 mm, width 50 mm in the largest specimen(NU-B2228). Ventral valve gently and unevenly convex inlateral profile, most convex in umbonal region; sulcus shallow
but distinct in anterior half of valve. Dorsal valve gently convex
in lateral profile; fold very low near anterior margin of valve.

External surface of ventral valve ornamented with slightly
irregular concentric lamellae bearing a row of ﬁne biramous spine
bases on anterior margin; numbering 2 lamellae in 5 mm at justanterior to umbo, 4 lamellae in 5 mm at midlength, and 5 lamellaein 5 mm near anterior valve margin; spine bases regularly
arranged on front of each lamella with a density of 7 in 5 mm at
about midlength. External ornament of dorsal valve similar to
opposite valve. Ventral interior with a median septum extending
to one third valve length and a pair of strong dental adminicula.
Dorsal vave interior with a long thin median septum extending tohalf length of valve; other internal structures not well preserved.
Internal surfaces of both valves sculptured with numerous fineradial grooves.
Remarks.―These specimens are referred to Torynifer asiatica
Besnossova in Besnossova et al. (1962, p. 176, pl. C-23, figs.
4‒5), from the lower Tersinsk Horizon (upper Tournaisian) of
the Kuznetsk Basin, on account of the large, gently biconvex
shell, with shallow ventral sulcus and low dorsal fold. Torynifer
pseudolineata (Hall, 1858), redescribed by Weller (1914, p. 429,
pl. 74, ﬁgs. 1‒11; pl. 75, ﬁg. 20) from the Keokuk Limestone of
the Mississippi Valley, differs from T. asiatica in having more
prominent ventral sulcus and dorsal fold.

74

JUN-ICHI TAZAWA

Occurrence.―HK4 Unit.
Distribution.―Lower Tournaisian‒Serpukhovian: northeasternJapan (Hikoroichi in the South Kitakami Belt), Afghanistan,central Russia (Kuznetsk Basin and Zabaykal), Kazakhstan andnorthwestern China (Xinjiang and Tibet).
Order SPIRIFERINIDA Ivanova, 1972
Suborder SPIRIFERINIDINA Ivanova, 1972
Superfamily SYRINGOTHYRIDOIDEA Fredericks, 1926
Family SYRINGOTHYRIDIDAE Fredericks, 1926
Subfamily SYRINGOTHYRIDINAE Fredericks, 1926
Genus SYRINGOTHYRIS Winchell, 1863
Type species.―Syringothyris typa Winchell, 1863.
Syringothyris sibirica Sokolskaya in Sarytcheva et al., 1963
(Fig. 20F‒H)
Syringothyris sibirica Sokolskaya in Sarytcheva et al., 1963, p.
277, pl. 49, ﬁgs. 1‒10.
Material.―Four specimens from localities KAC3 and KF201:(1) internal mould of two ventral valves, NU-B2197, 2198; and(2) external and internal moulds of two dorsal valves, NU-B2199,
2200.
Description.―Shell medium in size for genus, transversely
subsemicircular in outline, with greatest width at hinge; length
about 35 mm, width more than 70 mm in the largest dorsal valve
specimen (NU-B2199). Ventral valve having a high, slightly
concave interarea (height about 24 mm in the larger ventral valve,
NU-B2198). Dorsal valve gently convex in lateral profile; foldwide and low. External surface of dorsal valve ornamented withsimple rounded costae on ﬂanks, but smooth on fold; numbering
5‒6 costae in 10 mm at about midlength. Internally, ventral valve
having a syrinx and a pair of strong divergent dental adminicura.
Remarks.―These specimens are fragmentarily preserved,
but can be referred to Syringothyris sibirica Sokolskaya (inSarytcheva et al., 1963, p. 277, pl. 49, ﬁgs. 1‒10) from the upper
Tournaisian‒lower Visean of the Kuznetsk Basin, central Russia,
by the rather low ventral interarea and wide, low dorsal fold.
Syringothyris transversa Minato (1951, p. 377, pl. 5, ﬁg. 1), fromthe Arisu Formation of the Shimoarisu area, South Kitakami Belt,
differs from the present species in having a highly elevated fold in
the dorsal valve.
Occurrence.―HK1 Unit.
Distribution.―Upper Tournarisian‒lower Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt) and central Russia(Kuznetsk Basin).
Subfamily PERMASYRINXINAE Waterhouse, 1986
Genus PSEUDOSYRINX Weller, 1914
Type species.―Pseudosyrinx missouriensis Weller, 1914.
Pseudosyrinx cf. missouriensis Weller, 1914
(Fig. 28M, N)

cf. Pseudosyrinx missouriensis Weller, 1914, p. 406, pl. 65, ﬁgs.
5‒9; pl. 66, ﬁgs. 11‒13.
Material.―Two specimens from locality KF202: (1) external
and internal moulds of a ventral interarea, NU-B2041; and (2)internal mould of a dorsal valve, NU-B2108.
Remarks.―The specimens from Hikoroichi are fragmentarily
preserved, but they can be assigned to the genus Pseudosyrinx
in possessing a high, triangular and flattened ventral interarea
and in lacking syrinx. The Hikoroichi species most resembles
Pseudosyrinx missouriensis Weller (1914, p. 406, pl. 65, figs.
5‒9; pl. 66, figs. 11‒13), from the Burlington Limestone of theMississippi Valley, in the large size (length 40 mm, width about
72 mm in the dorsal valve, NU-B2108) and external ornament of
numerous simple rounded costae on the ventral valve. However,accurate comparison is difficult for the poorly preserved
specimens.
Occurrence.―HK2 Unit.
Family DIMEGELASMIDAE Carter in Carter et al., 1994
Genus DIMEGELASMA Cooper, 1942
Type species.―Spirifer neglectus Hall, 1858.
Dimegelasma sp.
(Fig. 33A, B)
Dimegelasma sp. Tazawa and Katayama, 1979, p. 171, pl. 11,ﬁgs. 15, 16.
Tomiopsis sp. Tazawa, 1984b, p. 308, pl. 61, ﬁg. 10.
Material.―Two specimens from locality KF163, internalmoulds of two ventral valves, IGPS97910, 97911.
Remarks.―These specimens were described by Tazawa(1984b, p. 308, pl. 61, ﬁg. 10) as Tomiopsis sp. But the Hikoroichi
species can be assigned to the genus Dimegelasma by the medium
to large size (length about 46 mm, width about 39 mm in thelarger specimen, IGPS97910), gently convex ventral valve, withrounded cardinal extremities and by the presence of very long thindental plates in the ventral valve. Dimegelasma sp. Tazawa andKatayama (1979, p. 171, pl. 11, ﬁgs. 15‒16), from the middle partof the Odaira Formation in the Yokota area, South Kitakami Belt,
is deemed conspecific with the present species. The Hikoroichi
species superficially resembles Dimegelasma grandicosta(Abramov and Grigorjeva, 1986, p. 162, pl. 29, ﬁgs. 4‒6; pl. 30,
ﬁgs. 1, 2, 10), from the middle‒upper Visean of the Verkhoyansk
Range, northern Russia. Speciﬁc identiﬁcation, however, remains
difﬁcult due to the poor preservation of the material.
Occurrence.―HK4 Unit.
Superfamily PENNOSPIRIFERINOIDEA Dagys, 1972
Family PUNCTOSPIRIFERIDAE Waterhouse, 1975
Genus PUNCTOSPIRIFER North, 1920
Type species.―Punctospirifer scabricosta North, 1920.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

Punctospirifer plicatosulcatus Glenister, 1955
(Fig. 29D)
Punctospirifer plicatosulcatus Glenister, 1955, p. 69, pl. 6, ﬁgs.
26‒29; pl. 8, figs. 10‒12; Roberts, 1971, p. 268, pl. 58, figs.
1‒16; Thomas, 1971, p. 168, pl. 25, ﬁgs. 3‒7, text-ﬁgs. 63, 64.
Material.―One specimen from locality KF202, externalmould of a ventral valve, NU-B2106.
Remarks.―The single ventral valve specimen from Hikoroichi
can be referred to Punctospirifer plicatosulcatus Glenister (1955,
p. 69, pl. 6, ﬁgs. 26‒29; pl. 8, ﬁgs. 10‒12), from the Moogooree
Limestone of the Carnarvon Basin, western Australia, in thesmall, transverse shell (length about 10 mm, width about 18 mm),a few strong costae with numerous closely-spaced concentriclamellae and in having a distinct median sulcal costa in the ventral
valve. The most allied species, Punctospirifer solidirostris (White,
1860), redescribed by Carter (1972, p. 490, pl. 4, figs. 24‒29)
from the Gilmore City Limestone of north-central Iowa, differsfrom P. plicatosulcatus in the less transverse outline.
Occurrence.―HK2 Unit.
Distribution.―Lower Tournaisian‒lower Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt), northwestern
Australia (Bonaparte Gulf Basin and Canning Basin) and western
Australia (Carnarvon Basin).
Punctospirifer gigantiformis Lee and Gu in Lee et al., 1980
(Fig. 29C)
Punctospirifer gigantiformis Lee and Gu in Lee et al., 1980, p.
420, pl. 157, ﬁgs. 4, 13, 15, 17; Gu, 1992, p. 252, pl. 66, ﬁgs.
10, 12; pl. 72, ﬁg. 13; pl. 73, ﬁgs. 4‒6, 8, 10, 20.
Material.―One specimen from locality KF202, externalmould of a ventral valve, NU-B2107.
Remarks.―The single ventral valve specimen from Hikoroichi
is referred to Punctospirifer gigantiformis Lee and Gu in Lee etal. (1980, p. 420, pl. 157, ﬁgs. 4, 13, 15, 17), from the Lujuantun
Formation of Jilin, northeastern China, by the large size (length
about 25 mm, width more than 30 mm) and in having deep sulcus
with V-shaped bottom and subangular costae on lateral flanks.
This species is distinguished from most species of the genus by
the large size.
Occurrence.―HK2 Unit.
Distribution.―Lower‒upper Visean: northeastern Japan(Hikoroichi in the South Kitakami Belt) and northeastern China(Jilin).
Punctospirifer iwatensis sp. nov.
(Fig. 33D)
Etymology.―Named after the fossil locality, Iwate Prefecture.
Material.―One specimen from locality KF148, external andinternal moulds of a ventral valve, NU-B2231 (holotype).
Diagnosis.―Large, transverse Punctospirifer, with ventral

75

sulcus bearing a median costa on anterior two-thirds and 9 simple
subangular costae on each lateral slope of ventral valve.
Description.―Shell large in size for genus, transversely
subelliptical in outline; cardinal extremities acute; length 13mm, width 30 mm in the holotype (NU-B2231). Ventral valve
moderately and unevenly convex in lateral profile, strongly
convex at umbonal region, becoming gently convex anteriorly;interarea moderately high and slightly concave; delthyrium open,
broad, measuring 5 mm wide at base; sulcus wide and deep, about
10 mm in width at anterior margin, and bearing a median costa
on anterior two-thirds; lateral slopes gently convex and nearly ﬂatin anterior proﬁle. External ornament of ventral valve consistingof simple subangular costae and closely-spaced imbricate growthlaminae; numbering 9 costae on each lateral slope, 5‒6 laminae in
3 mm at about midlength. Internally, ventral valve having a pairof short, slightly divergent dental adminicula and a stout, shortmedian septum.
Remarks.―Punctospirifer iwatensis sp. nov. resembles
Punctospirifer magnus Poletaev (2000, p. 36, pl. 5, ﬁgs. 15‒17),from the upper Tournaisian of the Donets Basin, western Russia,
in size and outline, but differs from the Russian species in thesmaller size and in having wider sulcus and fewer costae on theventral valve. Punctospirifer transversus (McChesney, 1860),
redescribed by Butts (2007, p. 60, ﬁgs. 5.54‒5.58) from the Arco
Hills Formation (Chesterian) of Idaho, differs from P. iwatensis
in the smaller size and in having alate cardinal extremities.
Punctospirifer plicatosulcatus Glenister (1955, p. 69, pl. 6, ﬁgs.
26‒29; pl. 6, figs. 10‒12), from the Moogooree Limestone of
the Carnarvon Basin, western Australia, differs from the present
species in the smaller size and in having fewer costae on theventral lateral slopes.
Occurrence.―HK4 Unit.
Punctospirifer sp.
(Fig. 20D, E)
Material.―Two specimens from localities KF200 and KF201:(1) internal mould of a ventral valve, NU-B2207; and (2) internalmould of a dorsal valve, NU-B2208.
Remarks.―These specimens are safely assigned to the genus
Punctospirifer by the small, transverse (length about 14 mm,
width more than 22 mm in the larger specimen, NU-B2207)
and punctate shell, ornamented with few strong costae andnumerous closely-spaced concentric lamellae. However, speciﬁc
identiﬁcation is difﬁcult for the poorly preserved material.
Occurrence.―HK1 Unit.
Order TEREBRATULIDA Waagen, 1883
Suborder TEREBRATULIDINA Waagen, 1883
Superfamily DIELASMATOIDEA Schuchert, 1913
Family BEECHERIIDAE Smirnova, 2004
Genus BEECHERIA Hall and Clarke, 1893
Type species.―Beecheria davidsoni Hall and Clarke, 1893.

76

JUN-ICHI TAZAWA

Beecheria khalfini Besnossova in Sarytcheva et al., 1963
(Fig. 20I)
Beecheria khalfini Besnossova in Sarytcheva et al., 1963, p. 338,
pl. 64, ﬁgs. 4‒6; Zhang et al., 1983, p. 383, pl. 124, ﬁg. 12.
Material.―One specimen from locality KF200, internal mouldof a conjoined shell, NU-B2217.
Description.―Shell medium in size for genus, elongately
suboval in outline, with rectimarginate anterior commissure androunded front margin, widest at midlength; length 32 mm, widthabout 18 mm. Ventral valve gently convex in lateral profile,
except for moderately convex umbonal region; umbo narrow,with small permesothyrid foramen. Dorsal valve gently convex;
interior of valve with a pair of diverging hinge plates and a low
median ridge.
Remarks.―This specimen can be referred to Beecheria
khalfini Besnossova (in Sarytcheva et al., 1963, p. 338, pl.
64, figs. 4‒6), from the lower Tournaisian‒lower Visean of
the Kuznetsk Basin, on account of size and shape of the shell,particularly in having rather strongly convex umbo in the ventral
valve. Beecheria chouteauensis (Weller, 1914, p. 257, pl. 32, ﬁgs.
1‒17), from the Chouteau and Pierson limestones of Missouri,
differs from the present species in having more ﬂattened shell andless convex ventral umbo.
Occurrence.―HK1 Unit.
Distribution.―Lower Tournaisian‒lower Visean: northeasternJapan (Hikoroichi in the South Kitakami Belt) and central Russia(Kuznetsk Basin).
ACKNOWLEDGEMENTS
I sincerely thank Kenji Konishi (Professor Emeritus of
Kanazawa University, Kanazawa) and Soichiro Kawabe(Fukui Prefectural Dinosaur Museum, Katsuyama) for editing
the memoir including this study; Shu-zhong Shen (Nanjing
Institute of Geology and Palaeontology, Chinese Academy of
Sciences, Nanjing) and an anonymous reviewer for their valuablecomments and suggestions on the manuscript; Mitsugi Harashinai(Mizusawa-ku, Oshu City, Iwate Prefecture), Toshio Kawamura(Miyagi University of Education, Sendai) and Atsushi Kaneko(Higashinada-ku, Kobe City, Hyogo Prefecture) for providing partof the brachiopod specimens; and Yousuke Ibaraki (Fossa Magna
Museum, Itoigawa) and Naotomo Kaneko (AIST, Geological
Survey of Japan, Tsukuba) for their help in drawing the ﬁgures.
REFERENCES
Abramov, B. S. 1965. Lower Carboniferous brachiopods ofSette-Daban; pp. 31‒59 in V. F. Vozin (ed.), Palaeontology
and Biostratigraphy of Palaeozoic and Triassic Deposits in
Yakutsk. Nauka, Moskva.***Abramov, B. S. 1970. Biostratigraphy of the Carboniferous
Deposits of Sette-Daban (Southern Verkhoyansk). Nauka,
Moskva, 176 pp.***Abramov, B. S., and A. D. Grigorjeva. 1986. Biostratigraphy
and Brachiopods of the Lower Carboniferous of Verkhoyansk.
Nauka, Moskva, 192 pp.***Aisenverg, D. E., and V. I. Poletaev. 1971. Description of new
species; Brachiopoda; pp. 66‒91 in D. E. Aisenverg (ed.),Atlas of the Tournaisian Fauna in the Donetz Basin (With the
Description of the New Species). Naukova Dumka, Kiev.***Alexandrov, V. A., and R. V. Solomina. 1973. Description oforganic remains: Brachiopoda; pp. 86‒131 in O. L. Einor(ed.), Stratigraphy and Fauna of the Carboniferous Depositsof the Shartym River Region (Southern Urlas). Kievskiy
Ordena Lenina Gosudarstvenny Universitet, Lvov.***
Alvarez, F., and J.-Y. Rong. 2002a. Athyridida; pp. 1475‒1583
in R. L. Kaesler (ed.), Treatise on Invertebrate Paleontology,
Part H Brachiopoda Revised, Volume 4: Rhynchonelliformea(Part). Geological Society of America, Boulder andUniversity of Kansas, Lawrence.
Alvarez, F., and J.-Y. Rong. 2002b. Retziidina; pp. 1584‒1601
in R. L. Kaesler (ed.), Treatise on Invertebrate Paleontology,
Part H Brachiopoda Revised, Volume 4: Rhynchonelliformea(Part). Geological Society of America, Boulder andUniversity of Kansas, Lawrence.
Alvarez, F., J.-Y. Rong. and A. J. Boucot. 1998. The classiﬁcationof athyridid brachiopods. Journal of Paleontology 72: 827‒855.
Angiolini, L., S. Long and I. Davies. 2011. Revision of Sowerbyʼs
species Spirifer bisulcatus, Spirifer pinguis and Spirifer
rotundatus from the Tournaisian‒Visean of Great Britain.
Memoirs of the Association of Australasian Palaeontologists
41: 71-85.
Archbold, N. W. 1982. Classification and evolution of thebrachiopod family Rugosochonetidae Muir-Wood, 1962.
Proceedings of the Royal Society of Victoria 94: 1‒10.
Archbold, N. W., and S. Stojanović-Kuzenko. 1995. Biostratigraphy;
Brachiopoda; pp. 59‒64 in I. Filipović (ed.), The Carboniferousof Northwestern Serbia. Gemini, Beograd.
Armstrong, A. 1962. Stratigraphy and paleontology of theMississippian System in southwestern New Mexico andadjacent southern Arizona. State Bureau of Mines andMineral Resources New Mexico Institute of Mining andTechnology, Memoir 8: 1‒99.
Asama, K., T. Asano, E. Sato and Y. Yamada. 1985. Early
Carboniferous plants from the Hikoroichi Formation,
Southern Kitakami Massif, Northeast Japan (Preliminary
report). Journal of the Geological Society of Japan 91:
425‒426.*
Asama, K., Y. Yamada and E. Sato. 1989. Pteridophytes; pp.
109‒121 in Ofunato City Museum (ed.), Fosssils of Onimaru.
Kawashima Insatsu, Ichinoseki.*
Bassett, M. G., and C. Bryant. 2006. A Tournaisian brachiopod
fauna from South-east Wales. Palaeontology 49: 485‒535.
Benediktova, R. N. 1955. Brachiopoda; pp. 307‒312 in L. L.
Khalfina (ed.), Atlas of Leading Species of Fossil Fauna
and Flora in the Western Siberia, Vol. 1. Gosgeoltekhizdat,
Moskva.***Besnossova, G. A. 1959. Lower Carboniferous brachiopods

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATIONof the Kuznetsk Basin (Families Cyrtospiriferidae andSpiriferidae). Trudy Paleontologicheskogo Instituta,
Akademiya Nauk SSSR 75: 1‒134.***
Besnossova, G. A., R. N. Benediktova, T. G. Sarytcheva andA. N. Sokolskaya. 1962. Brachiopoda; pp. 143‒183 in L. L.
Khalﬁna (ed.), Biostratigraphy of the Paleozoic of the Sayan‒
Altai Mountains, Volume 3. Upper Paleozoic. Sniiggims,
Novosibirsk.***von Bitter, P. H., and M. Legrand-Blain. 2007. Gigantoproductid
brachiopods from the Musquodoboit Limestone of Nova
Scotia (upper Windsor Group, Mississippian): Late Asbian‒
Brigantian paleobiogeographic connections between Eastern
Canada and Central England; pp. 191‒203 in T. E. Wong(ed.), Proceedings of the 15th International Congress on
Carboniferous and Permian Stratigraphy, Utrecht, theNetherlands, 10‒16 August 2003. Royal Netherlands
Academy of Arts and Sciences, Amsterdam.
Bond, G. 1941. Species and variation in British and Belgium
Carboniferous Schizophoriidae. Proceedings of the Geologists
Association 52: 285‒303.
Boucot, A. J. 1959. Brachiopods of the Lower Devonian rocks
at Highland Mills, New York. Journal of Paleontology 33:
727‒769.
Boucot, A. J., J. G. Johnson and R. D. Staton. 1964. On someatrypoid, retzioid, and athyridoid Brachiopoda. Journal of
Paleontology 38: 805‒822.
Brand, P. J. 1972. Some British Carboniferous species ofthe brachiopod genus Leptagonia McCoy. Bulletin of theGeological Survey of Great Britain 39: 57‒79.
Branson, E. B. 1938. Stratigraphy and paleontology of theLower Mississippian of Missouri. University Missouri
Studies 13: 1‒208.
Bronn, H. G. 1862. Die Klassen und Ordnungen der Weichthiere(Malacozoa), Vol. 3. C. F. Winterʼsche Verlagshandlung,Leipzig and Heidelberg, 518 pp.
Brunton, C. H. C. 1966. Siliciﬁed productoids from the Viseanof County Fermanagh. Bulletin of the British Museum(Natural History), Geology 12: 175‒243.
Brunton, C. H. C. 1968. Siliciﬁed brachiopods from the Viseanof County Fermanagh (2). Bulletin of the British Museum
Natural History (Geology) 16: 1‒70.
Brunton, C. H. C. 1980. Type species of some Upper Palaeozoic
athyridide brachiopods. Bulletin of the British Museum
Natural History (Geology) 34: 219‒234.
Brunton, C. H. C. 1984. Siliciﬁed brachiopods from the Viseanof County Fermanagh, Ireland (3), Rhynchonellids, spiriferids
and terebratulids. Bulletin of the British Museum Natural
History (Geology) 38: 27‒130.
Brunton, C. H. C., S. S. Lazarev, R. E. Grant and Y.-G. Jin.
2000. Productidina; pp. 424‒609 in R. L. Kaesler (ed.),Treatise on Invertebrate Paleontology, Part H Brachiopoda
Revised, Volume 3: Linguliformea, Craniiformea, andRhynchonelliformea (Part). Geological Society of America,
Boulder and University of Kansas, Lawrence.
Brunton, C. H. C., D. J. C. Mundy and S. S. Lazarev. 1993.

77

Productellid and plicatiferid (productoid) brachiopods fromthe Lower Carboniferous of the Craven Reef Belt, northYorkshire. Bulletin of the British Museum Natural History(Geology) 49: 99‒119.
Bublichenko, N. L. 1971. Lower Carboniferous Brachiopods of
the Rudny Altai (Tarkham Formation). Nauka, Alma-Ata, 189
pp.***
Bublichenko, N. L. 1976. Lower Carboniferous Brachiopods of
the Rudny Altai (Vukhtarminsk, Ulbinsk and Pravoloktevsk).
Nauka, Alma-Ata, 211 pp.***Buckman, S. S. 1906. Brachiopod nomenclature: Epithyris,
Hypothyris, Cleiothyris Phillips, 1841. Annals and Magazine
of Natural History, Series 7, 18: 321‒327.
Butts, S. 2007. Silicified Carboniferous (Chesterian)Brachiopoda of the Arco Hills Formation, Ohio. Journal of
Paleontology 81: 48‒63.
Campbell, K. S. W. 1957. A Lower Carboniferous brachiopodcoral fauna from New South Wales. Journal of Paleontology
31: 34‒98.
Carter, J. L. 1967. Mississippian brachiopods from the
Chappel Limestone of central Texas. Bulletin of American
Paleontology 53: 253‒488.
C a r t e r, J . L . 1 9 6 8 . N e w g e n e r a a n d s p e c i e s o f E a r l yMississippian brachiopods from the Burlington Limestone.
Journal of Paleontology 42: 1140‒1152.
Carter, J. L. 1972. Two new genera of lamellose spiriferaceanbrachiopods. Journal of Paleontology 46: 729‒734.
Carter, J. L. 1974. New genera of spiriferid and brachythyridid
brachiopods. Journal of Paleontology 48: 674‒696.
Carter, J. L. 1987. Lower Carboniferous brachiopods from the
Banff Formation of Western Alberta. Geological Survey ofCanada, Bulletin 378: 1‒183.
Carter, J. L. 1990. New brachiopods (Brachiopoda: Articulata)
from the Late Osagean of the Upper Mississippi Valley.
Annals of Carnegie Museum 59: 219‒247.
Carter, J. L. 1992. New genera of Lower Carboniferous
spiriferid brachiopods (Brachiopoda, Spiriferida). Annals of
Carnegie Museum 61: 327‒338.
Carter, J. L. 1999. Tournaisian (early Osagean) brachiopods
from a bioherm in the St. Joe Formation near Kenwood,
Oklahoma. Annals of Carnegie Museum 68: 91‒149.
Carter, J. L. 2006a. Spiriferinidina; pp. 1897‒1909 in R. L.
Kaesler (ed.), Treatice on Invertebrate Paleontology, Part H
Brachiopoda Revised, Volume 5: Rhynchonelliformea (Part).
Geological Society of America, Boulder and University of
Kansas, Lawrence.
Carter, J. L. 2006b. Pennospiriferinoidea; pp. 1910‒1929 in R. L.
Kaesler (ed.), Treatise on Invertebrate Paleontology, Part H
Brachiopoda Revised, Volume 5: Rhynchonelliformea (Part).
Geological Society of America, Boulder and University of
Kansas, Lawrence.
Carter, J. L., J. G. Johnson, R. Gourvennec and H.-F. Hou. 1994.
A revised classiﬁcation of the spiriferid brachiopods. Annalsof Carnegie Museum 63: 327‒374.
Chao, Y. T. 1927. Productidae of China, Part 1: Producti.

78

JUN-ICHI TAZAWA

Palaeontologia Sinica, Series B, 5: 1‒244.
Chao, Y. T. 1929. Carboniferous and Permian spiriferids ofChina. Palaeontologia Sinica, Series B, 11: 1‒133.
Chen, Z.-Q., and N. W. Archbold. 2000. Tournaisian‒Viseanbrachiopods from the Gancaohu area of southern Tienshan
Mountains, Xinjiang, NW China. Geobios 33: 183‒199.
C h e n , Z . - Q . , G . R . S h i a n d L . - P. Z h a n . 2 0 0 3 . E a r l y
Carboniferous athyridid brachiopods from Qaidam Basin,
northwest China. Journal of Paleontology 77: 844‒862.
C o o p e r, G . A . 1 9 4 2 . N e w g e n e r a o f N o r t h A m e r i c a nbrachiopods. Journal of the Washington Academy of Sciences
32: 228‒235.
Cooper, G. A., and H. M. Muir-Wood. 1951. Brachiopod
homonyms. Journal of the Washington Academy of Sciences
41: 195‒196.
Cvancara, A. M. 1958. Invertebrate fossils from the Lower
Carboniferous of New South Wales. Journal of Paleontology
32: 846‒888.
Dagys, A. S. 1972. Morphology and systematics of Mesozoic
retzioid brachiopods. Trudy Sibirskoe Otdelenie, Institut
Geologii i Geoﬁziki 112: 94‒105.***Davidson, T. 1858‒1863. British Fossil Brachiopoda, Vol. 2.
Permian and Carboniferous Species. PalaeontographicalSociety, London, 280 pp.
Davidson, T. 1881. On genera and species of spiral-bearing
Brachiopoda, from specimens developed by the Rev. Norman
Glass: with notes on the results obtained by Mr. George Mawfrom extensive washing of the Wenlock and Ludlow shales of
Shropshire. Geological Magazine, New Series, 8: 1‒13.
Dedok, T. A., and G. E. Tschernjak. 1960. Lower Carboniferous
brachiopods of the Taymir Peninsula. Trudy NIIGA 111:
52‒72.***
Dehée, R. 1929. Description de la Faune dʼEtroengt: Faune dePassage du Dévonien au Carbonifère. Mémoires de la Société
Géologique de France (Nouvelle Série) 11: 1‒62.
Delépine, G. 1928. Les brachiopodes du Marbre Noir de Dinant(Viséan inférieur). Memoires du Musée Royal dʼHistoire
Naturelle de Belgique 37: 1‒37.
Demanet, F. 1934. Les Brachiopodes du Dinantien de laBelgique, Premier Volume: Atremata, Neotremata, Protremata(Pars). Mémoires du Musée Royal dʼHistoire Naturelle deBelgique 61: 1‒116.
Demanet, F. 1938. La Faune des Couches de passage duDinantien au Namurien dans le synclinorium de Dinant.
Mémoires du Musée Royal dʼHistoire Naturelle de Belgique
84: 1‒201.
Ding, P., and W. Qi. 1983. Phylum Brachiopoda (Carboniferous‒
Permian); pp. 44‒425 in Xian Institute of Geology andMineral Resources (ed.), Palaeontological Atlas of Northwest
China; Shaanxi, Gansu and Ningxia Volume, Part 2. Upper
Palaeozoic. Geological Publishing House, Beijing.**
Dixon, E. E. L., and A. Vaughan. 1911. The Carboniferous
succession in Gower (Glamogranshire), with notes on its
fauna and conditions of deposition. Quaterly Journal of theGeological Society of London 67: 477‒571.

Donakova, L. M. 1978. Visean brachiopods of the eastern
slope of the Southern Urals (Magnitogorsk Synclinorium).
Ezhegudanik, VPO 21: 205‒227.***Dun, W. S., and B. A. Benson. 1920. The geology and petrologyof the Great Serpentine Belt of New South Wales, Part 9. The
geology, palaeontology and petrography of the Currabubula
district, with notes on adjacent regions; descriptive portion.
Proceedings of the Linnean Society of New South Wales 45:
337‒374.
Etheridge, R. 1872. Description of the Palaeozoic and Mesozoic
fossils of Queensland. Quarterly Journal of the GeologicalSociety of London 28: 317‒360.
Ferguson, J. 1971. Linoprotonia, a new genus of Lower
Carboniferous productoid. Proceedings of the Yorkshire
Geological Society 38: 549‒564.
Fischer de Waldheim, G. 1825. Notice sur la Choristite.
Programme dʼinvitation à la Sociéte Impériale des Naturalistes
de Moscou. Sociéte Impériale des Naturalistes de Moscou,
Moscow, 12 pp.
Fischer de Waldheim, G. 1829. Quelques fossiles du gouverment
de Moscou. Société Impériale des Naturalistes de Moscou
Bulletin 1: 375‒376.
Fischer de Waldheim, G. 1850. Orthotetes, genre de la famille
des brachiopodes. Société Impériale des Naturalistes deMoscou Bulletin 23: 491‒494.
Frech, F. 1900. Zur Kenntnis der mittleren Palàozoicum in
Hocharmenien und Persien; pp. 183‒207 in F. Frech and
G. Arthaber (ed.), Über das Paläozoicum in Hocharmenien
und Persien, mit einem Anhang über die Kreide von Sirab in
Persien, Beiträge zur Paläontologie und Geologie ÖsterreichUngarns und des Orients, Band 12. Wilheim Braumüller,
Wien und Leipzig.
Frech, F. 1916. Geologie Klainasiens im Bereich derBagdadbahn: Ergebnisse eigener Reisen und paläontologische
Untersuchungen. Zeitschrift der Deutschen Geologischen
Gesellschaft 68: 1‒325.
Fredericks, G. 1924. Paleontological studies, 2: On Upper
Carboniferous spiriferids from the Urals. Izvestyia
Geologicheskogo Komiteta 38: 295‒324.***
Fredericks, G. 1926. Table for classiﬁcation of the genera of thefamily Spiriferidae King. Izvestiya Akademia Nauk SSSR,
Series 6 20: 393‒423.***
Fredericks, G. 1933. Paleontological notes, 4. On some Upper
Paleozoic brachiopods of Eurasia. Material TSNIGRI,
Paleontologiya i Stratigraﬁya 2: 24‒33.***Gaetani, M. 1968. Lower Carboniferous brachiopods from
central Elburz, Iran. Revista Italiana di Paleontologia e
Stratigraﬁa 70: 665‒744.
Galitskaya, A. Ya. 1977: Early and Middle Carboniferous
Productids of Northern Kirgiz. Ilim, Frunze, 297 pp.***
Garanj, I. M., S. N. Guseva, V. V. Devingtal, L. M. Donakova,
N. V. Enokyan, N. V. Kalashnikov, N. N. Lapina, E. N.
Mikhaylova, D. V. Nalivkin, S. V. Semichatova, D. L.
Stepanov, G. A. Stepanova, M. F. Shestakova and O. L. Einor.
1975: Brachiopoda; pp. 154‒203 in D. L. Stepanov, A. K.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

Krylova, L. P. Grozdnilova, V. M. Pozner and A. A. Syltanaev
1975. Palaeontological Atlas of the Carboniferous Deposits of
the Urals. Nedra, Leningrad.***Garwood, E. J. 1916. The faunal succession in the Lower
Carboniferous of Westmorland and North Lancashire.
Proceedings of the Geologists Association 27: 1‒43.
Girty, G. H. 1920. Carboniferous and Triassic faunas. UnitedStates Geological Survey Professional Paper 111: 641‒648.
Girty, G. H. 1927. Descriptions of new species of Carboniferous
and Triassic fossils. United States Geological Survey
Professional Paper 152: 411‒446.
Girty, G. H. 1929. The fauna of the Middle Boone near
Batesville, Arkansas. United States Geological Survey
Professional Paper 154: 73‒103.
Glenister, B. F. 1955. Devonian and Carboniferous spiriferids
from the North-West Basin, western Australia. Journal of theRoyal Society of Western Australia 29: 46‒71.
Grabau, A. W. 1923‒1924. Stratigraphy of China, Part 1.
Palaeozoic and Older. Geological Survey, Ministry ofAgriculture and Commerce, Peking, 528 pp.
G r a b a u , A . W. 1 9 3 1 . S t u d i e s f o r s t u d e n t s , S e r i e s 1 .
Palaeontology: The Brachiopoda, Part 2. Science Quaterly of
the National University of Peking 2: 397‒422.
Grabau, A. W. 1932. The significance of the sinal formulain Devonian and post-Devonian spirifers. Bulletin of theGeological Society of China 11: 93‒95.
Gray, J. E. 1840. Synopsis of the Contents of the Britich
Museum, 42nd Edition. British Museum, London, 370 pp.
Grechishnikova, I. A. 1966. Stratigraphy and brachiopodsof the Lower Carboniferous of the Rudny Altai. Trudy
Moskovskogo Obschestva Ispytateley Prirody 20: 1‒184.***
Gröber, P. 1908. Ueber die Faunen des untercarbonischen
Transgressionsmeeres des zentralen Tian-schan, die in der
Umgebung des Sart-dschol-Passes gefunden worden sind.
Neues Jahrbuch für Mineralogie, Geologie und Palaeontologie
26: 213‒248.
Gröber, P. 1909. Carbon und Carbonfossilien des nordlichen
und zentralen Tian-schan. Abhandlungen der königlich
Bayerisschen Akademie der Wissenschaften 2 Klass 24:341‒384.
Grunt, T. A., 1986. Classification of brachiopods of the order
Athyridida. Trudy Paleontologicheskogo Instituta, Akademiya
Nauk SSSR 215: 1‒200.***Gu, F. 1992. Carboniferous and Permian Brachiopoda; pp.
210‒255 in Jilin Institute of Geology and Mineral Resources(ed.), Paleontological Atlas of Jilin. Jilin Science andTechnology Publishing House, Changchun.**Hall, J. 1858. Report on the geological survey of the State of
Iowa, embracing the results of investigations made during
portions of the year 1855‒1857; pp. 473‒724 in J. Hall and J.
D. Whitney (eds.), Palaeontology, Vol. 1, Part 2. Published by
Authority of the Legistrature of Iowa, Des Moines.
Hall, J., and J. M. Clarke. 1893‒1895. An Introduction to the
Study of the Genera of Palaeozoic Brachiopoda. Natural
History of New York, Palaeontology, Vol. 8, Pt. 2, Charles

79

van Benthuysen and Sons, Albany, 394 pp.
Harper, D. A. T., and A. L. Jeffrey. 1996. Mid-Dinantianbrachiopod biofacies from western Ireland; pp. 427‒436 in
P. Strogen, L. D. Somerville and G. L. Jones (eds.), Recent
Advances in Lower Carboniferous Geology. GeologicalSociety Special Publication, No. 107, Geological Society of
London, London.
Hase, A., and M. Yokoyama. 1975. Geological age and structureof the Hina Limestone, Okayama Prefecture, Southwest
Japan. Journal of Science of the Hiroshima University, Series
C, 7: 167‒182.
Ibaraki, Y., Y. Miyake and J. Tazawa. 2014. Early Carboniferous(late Visean) brachiopods from the Koyama Limestone of
Kamiotake in the Oga area, Okayama Prefecture, southwest
Japan. Earth Science (Chikyu Kagaku) 68: 69‒79.
Ivanova, E. A. 1972. Main features of spiriferid evolution(Brachiopoda). Paleontologicheskiy Zhurnal, 1922 3:
28‒42.***
Jiang, J.-J. 1993. A primary study on ontogeny of Unispirifer
striatoconvolutus (Benson et Dun). Stratigraphy andPaleontology of China 2: 215‒220.
Jiang, J.-J. 1997. Early Carboniferous biostratigraphy of West
Yunnan. Tethyan Geology 21: 182‒197.**
Jin, S. 1985. Early Carboniferous (Tournaisian) fossil
brachiopods from Qingshuigou of Baoshan County, Yunnan;
pp. 75‒85 in Chengdu Institute of Geology and Mineral
Resources (ed.), Contribution to the Geology of the Qinghai‒
Xizang (Tibet) Plateau, No. 16. Geological Publishing House,
Beijing.**
Jin, Y.-G., and R.-S. Fang. 1983. Early Carboniferous
brachiopods from Shidian, Yunnan. Acta Palaeontologica
Sinica 22: 139‒152.**
Jin, Y.-G., S.-L. Ye, H.-K. Xu and D.-L. Sun. 1979. Phylum
Brachiopoda; pp. 60‒217 in Nanjing Institute of Geology
and Palaeontology, Academia Sinica and Qinghai Geological
Science Institute (eds.), Palaeontological Atlas of Northwest
China, Qinghai Section, Part 1. Geological Publishing House,
Beijing.**
Jin, Y.-G., Y. Wang, D.-L. Sun and Q. Shi. 1985. Late Palaeozoic
and Triassic brachiopods from the east of the Qinghai‒
Xiznang Plateau; pp. 182‒237 in Regional Geological
Surveying Team of Sichuan Province and Nanjing Instituteof Geology and Palaeontolgy, Academia Sinica (eds.),
Stratigraphy and Palaeontology in W. Sichuan and E. Xizang,
China, Part 3. Sichuan Science and Technology Publishing
House, Chengdu.**Kaesler, R. L. 2000a. Treatise on Invertebrate Paleontology,
Part H Brachiopoda Revised, Volume 2: Linguliformea,
Craniiformea, and Rhynchonelliformea (Part). GeologicalSociety of America, Boulder and University of Kansas,
Lawrence. 423 pp. (1‒423).
Kaesler, R. L. 2000b. Treatise on Invertebrate Paleontology,
Part H Brachiopoda Revised, Volume 3: Linguliformea,
Craniiformea, and Rhynchonelliformea (Part). GeologicalSociety of America, Boulder and University of Kansas,

80

JUN-ICHI TAZAWA

Lawrence. 496 pp. (424‒919).
Kaesler, R. L. 2002. Treatise on Invertebrate Paleontology, Part
H Brachiopoda Revised, Volume 4: Rhynchonelliformea(Part). Geological Society of America, Boulder andUniversity of Kansas, Lawrence. 768 pp. (921‒1688).
Kaesler, R. L. 2006. Treatise on Invertebrate Paleontology, Part
H Brachiopoda Revised, Volume 5: Rhynchonelliformea(Part). Geological Society of America, Boulder andUniversity of Kansas, Lawrence. 632 pp. (1689‒2320).
Kalashnikov, N. V. 1974. Early Carboniferous Brachiopods fromthe Petchora Urals. Nauka, Leningrad, 166 pp.**
Kalashnikov, N. V. 1980. Brachiopods of the Upper Palaeozoic
of European Siberia, USSR. Nauka, Leningrad, 132 pp.***
Kaneko, A. 1983. A new species of Conophillipsia (Proetinae
trilobite) from the lower Carboniferous Hikorocihi Formationin Japan. Earth Science (Chikyu Kagaku) 37: 61‒68.
Kaneko, A. 1986. Trilobites from the upper part (H4 Member)of the lower Carboniferous Hikorocihi Formation, Kitakami
Mountains, Northeast Japan. Journal of the GeologicalSociety of Japan 92: 69‒71.*
Kaneko, A. 1989. Trilobites; pp. 87‒107 in Ofunato City
Museum (ed.), Fossils of Onimaru. Kawashima Insatsu,
Ichinoseki.*Kase, T. 1988. Early Carboniferous (Viséan) gastropods from the
Hikoroichi Formation of the Kitakami Mountains, northeast
Japan. Transactions and Proceedings of the PalaeontologicalSociety of Japan, New Series, 149: 361‒370.
Kase, T. 1989. Gastropoda; pp. 67‒77 in Ofunato City Museum(ed.), Fossils of Onimaru. Kawashima Insatsu, Ichinoseki.*Kato, M., and T. Nakamura. 1979. Chapter 2i4. Upper
Carboniferous; Correlation; pp. 140‒141 in M. Minato,
M. Hunahashi, J. Watanabe and M. Kato (eds.), Variscan
Geohistory of Northern Japan: The Abean Orogeny. Tokai
University Press, Tokyo.
Kato, M., I. Niikawa, T. Kawamura, Y. Ezaki and E. Sato. 1989.
Corals; pp. 13‒35 in Ofunato City Museum (ed.), Fossils of
Onimaru. Kawashima Insatsu, Ichinoseki.*
Kawamura, T. 1983. The Lower Carboniferous formations in the
Hikoroichi region, southern Kitakami Mountains, northeast
Japan (Part 1): Stratigraphy of the Hikoroichi Formation.
Journal of the Geological Society of Japan 89: 707‒722.*
Kawamura, T., T. Uchino, M. Kawamura, K. Yoshida, M.
Nakagawa and H. Nagata. 2013. Geology of Hayachine San
District; Quadrangle Series, 1:50,000. Geological Survey ofJapan, AIST, Maeda Insatsu, Tsukuba, 101 pp.*King, W. 1846. Remarks on certain genera belonging to the class
Palliobranchiata. Annals and Magazine of Natural History,
London 18: 26‒42, 83‒94.
King, W. 1850. A Monograph of the Permian Fossils of England.
Palaeontographical Society Monograph, Volume 3, London,
258 pp.
King, W. 1865. Remarks on the history of two specimensof Rhynchopora geinitziana de Verneuil, from near theRiver Oukhla, Province of Archangel, and belonging to thecollection of the Corps des Mines of St. Petersburg. Annals

and Magazine of Natural History, Series 3, 16: 124‒128.
Kobayashi, T., and T. Hamada. 1980. Carboniferous trilobitesof Japan in comparison with Asian, Paciﬁc and other faunas.
Palaeontological Society of Japan, Special Papers 23: 1‒132.
de Koninck, L. G. 1842‒1844. Description des Animaux
Fossiles qui se Trouvent dans le Terrain Carbonifère deBelgique. H. Dessain, Liége, 650 pp.
de Koninck, L. G. 1883. Sur le Spirifer mosquensis et sur ses
affinitiés avec quelques autres espèces du mème genre.
Bulletin du Musée Royal dʼHistoire Naturelle de Belgique 2:
371‒395.
Korejwo, K. 1986. Biostratigraphy of the Carboniferous depositsof the Świdnik blocks (Lublin Coal Basin). Acta Geologica
Polonica 36: 339‒346.
Kotlyar, G. V., and L. I. Popeko. 1967. Biostratigraphy, Bryozoa
and Brachiopoda of the Upper Paleozoic of Zabaykal.
Zabayckal Branch, Ministry of Geography, SSSR, Chita, 323
pp.***
Krenkel, E. 1913. Faunen aus dem Unterkarbon des südlichen
und östlichen Tian-Schan. Abhandlungen der Königlich
Bayerischen Akademie der Wissenschaften Mathematisch‒Physikalische Klasse 26 1‒ 44.
Kuhn, O. 1949. Lehrbuch der Palaozoologie. Schweizerbartʼsche
Verlagsbuchhandlung, Stuttgart. 326 pp.
Lapina, N. N. 1962. Brachiopods from the Krestyakhosk
Conglomerate of mouth of the Lena River. Trudy VNIGRI
196: 125‒142.***
Lazarev, S. S. 1969. Canal system of brachiopod Schizophoria
and Orthotichia. Paleontologicheskii Zhurnal, 1969 2:
66‒72.***
Lazarev, S. S. 1976. Morphology and Evolution of Brachiopods(of the Superfamily Enteletacea). Nauka, Moskva, 167 pp.***
Lee, L., and F. Gu. 1976. Carboniferous and Permian
Brachiopoda; pp. 228‒306 in Geological Bureau of Nei
Mongol and Geological Institute of Northeast China (eds.),
Palaeontological Atlas of Northeast China; Nei Mongol,
Part 1. Palaeozoic Volume. Geological Publishing House,
Beijing.**
Lee, L., F. Gu and Y.-Z. Su. 1980. Carboniferous and Permian
Brachiopoda; pp. 327‒428 in Shenyang Institute of Geology
and Mineral Resources (ed.), Paleontological Atlas ofNortheast Asia (1) Paleozoic Volume. Geological Publishing
House, Beijing.**
Legrand-Blain, M. 1985. A new genus of Carboniferous
spiriferid brachiopod from Scotland. Palaeontology 28:567‒576.
Legrand-Blain, M., J.-J. Delvolvé and M. Hansotte. 1996.
Carboniferous brachiopods from the Arize Massif (Ariègo),French Pyrenées. Geobios 29: 177‒207.
Léveillé, C. 1835. Aperçue géologique de quelques localités très
riches en coquilles sur les frontières de France et de Belgique.
Mémoires de la Société Géologique de France 2: 29‒40.
Licharew, B. K. 1934. Die Fauna der Permischen Ablagerungendes Kolyma-Gebietes. Akademii Nauk SSSR, Leningrad, 148pp.***MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

Litvinovich, N. V. 1962. Carboniferous and Permian Depositsof the Western Part of Central Kazakhstan. Moskovskogo
Universiteta, Moscow, 389 pp.***Litvinovich, N. V., G. G. Aksenova and M. V. Martynova.
1975. Descriptions of Fauna: Brachiopoda; pp. 50‒96 in T.
A. Gorokhova (ed.), Fauna of the Devonian‒Carboniferous
Boundary Beds in Central Kazakhstan. Nedra, Moskva.***Litvinovich, N. V., G. G. Aksenova and T. P. Razina. 1969.
Stratigraphy and Lithology of the Lower Carboniferous
Deposits in the West-Central Kazakhstan. Nedra, Moskva,
447 pp. ***Marcou, J. 1858. Geology of North America, with Two Reportson the Prairies of Arkansas and Texas, the Rockey Mountainsof New Mexico and the Sierra Nevada of California. Printed
for the author by Züricher and Furrer, Zürich, 144 pp.
Martin, W. 1793. Figures and Descriptions of Petrifications
Collected in Derbyshire. Wigan, London, 29 pp.
Martin, W. 1809. Petrificata Derbiensia; or Figures and
Descriptions of Petrefactions Collected in Derbyshire. Wigan,
London, 28 pp.
Martinez Chacon, M. L. 1979. Braquiopodos Carboniferosde la Cordillera Cantabrica (Orthida, Strophomenida y
Rhynchonellida). Memoria del Instituto Geologico y Minerode España, Servico de Publicationes, Madrid, 283 pp.
Martinez Chacon, M. L., and M. Legrand-Blain. 1992.
Braquiópodos. Coloquios de Paleontologia 44: 91‒144.
MʼCo y, F. 1 844. A Synopsis of the Characters of the
Carboniferous Limestone of Ireland. Williams and Norgate,
London, 207 pp.
McChesney, J. H. 1860. Descriptions of new species of fossils
from the Palaeozoic rocks of the western states. Transactionsof the Chicago Academy of Sciences 1: 1‒87.
Meek, F. B., and A. H. Worthen. 1860. Descriptions of new
Carboniferous fossils from Illinois and other westernstates. Proceedings of the Academy of Natural Sciences ofPhiladelphia, Series 2, 12: 447‒472.
Miloradovich, B. V. 1935. Materials to the study of the Upper
Paleozoic Brachiopoda from the Northern Island of Novaya
Zemlya. Trudy Arkticheskogo Instituta 19: 1‒166.***
Minato, M. 1950a. Zur Orogenese und zum Vulkanismus im
jüngeren Palaeozoikum des Kitakami-Gebirges, Honshu,
Japan. Journal of the Faculty of Science, Hokkaido
University, Series 4, 7: 277‒302.
Minato, M. 1950b. Stratigraphische Stellung der Hikoroiti-Serie.
Journal of the Geological Society of Japan 56: 143‒148.*
Minato, M. 1951. On the Lower Carboniferous fossils of theKitakami Massif, Northeast Honsyu, Japan. Journal of
the Faculty of Science, Hokkaido University, Series 4 7:355‒382.
Minato, M. 1952. A further note on the Lower Carboniferous
fossils of the Kitakami Mountainland, northeast Japan.
Journal of the Faculty of Science, Hokkaido University,
Series 4, 8: 136‒174.
Minato, M. 1953. On some reticulate Spiriferidae. Transactions
and Proceedings of the Palaeontological Society of Japan,

81

New Series, 11: 65‒73.
Minato, M. 1955. Zur stratigraphischen Lücke der Prä-OnimaruSerie (Ober Visé). Journal of the Faculty of Science,
Hokkaido University, Series 4, 9: 31‒41.
Minato, M. 1966. Stratigraphie des Palaeozoikums desKitakami-Gebirges und Abe-Orogenese in Japan; pp. 143‒159
in Editorial Committee of Prof. Susumu Matushita Memorial
Volume (ed.), Professor Susumu Matsushita Memorial
Volume. Nippon Insatsu, Osaka.*
Minato, M., S. Hashimoto, K. Suyama, H. Takeda, Y. Suzuki, S.
Kimura, K. Yamada, T. Kakimi, T. Ichikawa and H. Suetomi.
1953. Biostratigraphie des Karbons im Kitakami-Gebirge,
Nordöstliches Honshu, Japan. Journal of the GeologicalSociety of Japan 59: 385‒399.*
Minato, M., M. Hunahashi, J. Watanabe and M. Kato. 1979a.
Variscan Geohistory of Northern Japan: Abean Orogeny.
Tokai University Press, Tokyo, 427 pp.
Minato, M., and M. Kato. 1977. Two spiriferids from the
Hikoroichi Formation (Tournaisian) in the Kitakami
Mountains, Japan. Journal of the Faculty of Science,
Hokkaido University, Series 4, 17: 613‒617.
Minato, M., and M. Kato. 1979. Chapter 2h4. Upper Viséan;Correlation; pp. 136 in M. Minato, M. Hunahashi, J.
Watanabe and M. Kato (eds.), Variscan Geohistory ofNorthern Japan: The Abean Orogeny. Tokai University Press,
Tokyo.
Minato, M., M.Kato, I. Niikawa and T. Nakamura. 1979b.
Chapter 2h2. Upper Viséan; Stratigraphy; pp. 133‒134 in
M. Minato, M., Hunahashi, J. Watanabe and M. Kato (eds.),
Variscan Geohistory of Northern Japan: The Abean Orogeny.
Tokai University Press, Tokyo.
Minato, M., H. Takeda, M. Kato, H. Suetomi, Y. Hirata, M.
Kawamura and S. Haga. 1979c. Chapter 2f2. Tournaisian;Stratigraphy; pp. 74‒79 in M. Minato, M., Hunahashi,
J. Watanebe and M. Kato (eds.), Variscan Geohistory ofNorthern Japan: The Abean Orogeny. Tokai University Press,
Tokyo.
Moore, R. C. 1952. Brachiopoda; pp. 197‒267 in R. C. Moore, C.
G. Lalicker and A. G. Fischer, Invertebrate Fossils. McGrawHill, New York.
Mori, K., and J. Tazawa. 1980. Discovery and significance ofViséan rugose corals and brachiopods from the type localityof the Lower Carboniferous Hikoroichi Formation. Journal of
the Geological Society of Japan 86: 143‒146.*
Morozov, P. E. 1988. On the systematic composition of the
Moscow area brachiopods of the family Gigantoproductidae.
Paleontologicheskiy Zhurnal, 1988, 1: 22‒33.***Muir-Wood, H. M. 1951. The Brachiopoda of Martinʼs
Petrificata Derbiensia . Annals and Magazine of Natural
History, Series 12, 4: 97‒118.
Muir-Wood, H. M. 1955. A History of the Classiﬁcation of the
Phylum Brachiopoda. British Museum (Natural History),
London, 124 pp.
Muir-Wood, H. M. 1962. On the Morphology and Classiﬁcationof the Brachiopod Suborder Chonetoidea. British Museum

82

JUN-ICHI TAZAWA(Natural History), London, 132 pp.
Muir-Wood, H. M., and G. A. Cooper. 1960. Morphology,
classification and life habits of the Productoidea(Brachiopoda). Geological Society of America, Memoir 81:
1‒447.
Nalivkin, D. V. 1937. Brachiopoda of the Upper and Middle
Devonian and Lower Caroniferous of north-eastern
Kazakhstan.Trudy TSNIGRI 99: 1‒200.***
Nalivkin, D. V. 1979. Tournaisian Brachiopods of the Urals.
Nauka, Leningrad, 248 pp.***Nalivkin, D. V., and N. N. Fotieva. 1973. Brachiopods fromthe Boundary Beds of Tournaisian and Visean in the Western
Slope of the Urals. Nauka, Moskva, 118 pp.***Nelson, S. J. 1961. Mississippian faunas of Western Canada.
Geological Association of Canada, Special Paper 2: 1‒39.
Niko, S. 1989. Cephalopoda; pp. 79‒85 in Ofunato City
Museum (ed.), Fossils of Onimaru. Kawashima Insatsu,
Ichinoseki.*North, F. J. 1920. On Syringothyris Winchell, and certain
Carboniferous Brachiopoda referred to Spiriferina dʼOrbigny.
Quaterly Journal of the Geological Society of London 76:
162‒227.
Norwood, J. G., and H. Pratten. 1855. Notice of Producti
found in the western states and territories, with descriptionsof twelve new species. Philadelphia Academy of Natural
Sciences Journal, New Series, 3: 1‒21.
Oehlert, D. P. 1890. Note sur différents groupes établis dans legenre Orthis et en particulier sur Rhipidomella Oehlert (=Rhipidomys Oehlert, olim). Journal de Conchyliologie 30:366‒374.
Okubo, M. 1951. Zur Hikoroichi-Serie und der bedeutenden
Diskordanz vor der Hikoroichi-Epoche. Journal of theGeological Society of Japan 57: 95‒209.*O n u k i , Y. 1 9 3 7 . O n t h e P a l a e o z o i c f o r m a t i o n n e a r
Sakamotozawa, Kitakami Mountainland. Journal of theGeological Society of Japan 44: 168‒186.*
Onuki, Y. 1956. Geology of the Kitakami Massif; pp. 1‒189 in
Iwate Prefecture (ed.), Explanation Text of Geology of Iweate
Prefecture. Sasaki Shuppan, Sendai.*
Onuki, Y. 1969. Geology of the Kitakami Massif, northeast
Japan. Contributions from the Institute of Geology andPaleontology, Tohoku University 69: 1‒239.*
Öpik, A. A. 1934. Über Klitamboniten. Universitatis Tartuensis(Dorpatensis) Acta et Commentationes, Series A, 26: 1‒239.
Paeckelmann, W. 1930. Die Brachiopoden des deutschen
Unterkarbons, 1 Teil: Die Orthiden, Strophomeniden
und Choneten des mitteleren und oberen Unterkarbons.
Abhandlungen der Preussischen Geologischen Landesanstalt,
Neue Folge, 122: 143‒326.
Paeckelmann, W. 1931. Die Fauna des deutschen Unterkarbons,
2 Teil: Die Productinae und Productus-ähnlichen Chonetinae.
Abhandlungen der Preussischen Geologischen Landesanstalt,
Neue Folge, 136: 1‒441.
Pareyn, C. 1961. Les Massifs Carbonifères du Sahara SudOranais, Tome 2, Paléontologie Stratigraphique. CNRS, Paris,

224 pp.
Parkinson, D. 1954. Quantitative studies of brachiopods fromthe Lower Carboniferous reef limestones of England, 1.
Schizophoria resupinata (Martin). Journal of Paleontology
28: 367‒381.
Pattison, J. 1981. The stratigraphical distribution ofgigantoproductoid brachiopods in the Viséan and Namurian
rocks of some areas in northern England. Report of theInstitute of Geological Sciences 81/9: 1‒30.
Pavlova, E. E. 1969. The development of brachiopods of thefamily Reticulariidae. Trudy Paleontologicheskogo Instituta,
Akademiya Nauk SSSR 120: 1‒130.***Phillips, J. 1836. Illustrations of the Geology of Yorkshire; or a
Description of the Strata and Organic Remains: Accompanied
by a Geological Map, Sections and Diagrams and Figures of
the Fossils, Part 2. The Mountain Limestone District. John
Murray, London, 253 pp.
Phillips, J. 1841. Figures and Descriptions of the Palaeozoic
Fossils of Cornwall, Devon, and West Somerset. Geological
Survey of Great Britain, Memoir 1, Longman and Co.,
London, 231 pp.
Plodowski, G. 1970. Stratigraphie und Spiriferen (Brachiopoda)
des Paläozoikums der Dascht-e-Nawar/SW (Afganistan).
Palaeontographica, Abteilung A, 134: 1‒132.
Pocock, Y. P. 1968. Carboniferous schizophoriid brachiopods
from western Europe Palaeontology 11: 64‒93.
Poletaev, V. I. 1975. Early Carboniferous and Bashkirian smoothspiriferids and athyrides of the Donetz Basin. Naukova
Dumka, Kiev, 138 pp. ***Poletaev, V. I. 2000. New species of Early Carboniferous
and Bashkirian spiriferids of the Donetz Basin.
Paleontologicheskiy Zhurnal, 2000 1: 31‒37.***
Prentice, J. E. 1949. The hemisphaericus-like gigantellids of thesouthern Pennines. Proceedings of the Yorkshire GeologicalSociety 27: 247‒369.
Prentice, J. E. 1950. The genus Gigantella Sarycheva.
Geological Magazine 87: 436‒438.
Qiao, L., and S.-Z. Shen. 2014. Global paleobiogeography ofbrachiopods during the Mississippian: Response to the global
tectonic reconfiguration, ocean circulation, and climatechanges. Gondwana Research 26: 1173‒1185.
Racheboeuf, P. R. 2000. Chonetidina; pp. 362‒423 in R.
L. Kaesler (ed.), Treatise on Invertebrate Paleontology,
Part H Brachiopoda Revised, Volume 2: Linguliformea,
Craniiformea, and Rhynchonelliformea (Part). GeologicalSociety of America, Boulder and University of Kansas,
Lawrence.
Rakusz, G. 1932. Die oberkarbonische Fossilien von Dobsina(Dobšina) und Nasyvisnyȯ. Geologica Hungarica, Serie
Palaeontologie, 8: 1‒223.
Ramovš, A. 1969. Karavankininae, eine neue Unterfamilie der
Productiden (Brachiopoda) aus den alpinen Oberkarbonund Permischichten. Jeseniski Zbornik Jekle in Ljudje 2:
251‒268.
Roberts, J. 1963. A Lower Carboniferous fauna from

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

Lewinsbrook, New South Wales. Journal and Proceedings,
Royal Society of New South Wales 97: 1‒29.
Roberts, J. 1965. A Lower Carboniferous fauna from Trevallyn,
New South Wales. Palaeontology 8: 54‒81.
Roberts, J. 1971. Devonian and Carboniferous brachiopods fromthe Bonaparte Gulf Basin, northwestern Australia. Bulletinof the Bureau of Mineral Resources, Australia, Geology andGeophysics 122: 1‒319.
Roberts, J. 1976. Carboniferous chonetacean and productaceanbrachiopods from eastern Australia. Palaeontology 19: 17‒77.
Rotai, A. P. 1931. Brachiopods and stratigraphy of Lower
Carboniferous of the Donetz Basin. Trudy Izvestiya Glavnogo
Geologo-Razvedochnogo Upravleniya 73: 35‒144.***
Rotai, A. P. 1941. Phylum Brachiopoda; pp. 85‒117 in L. S.
Librovicha (ed.), Atlas of Index Fossils of USSR, Volume 4.
Lower Carboniferous System. VSEGEI, Leningrad.***Rotai, A. P. 1951. Brachiopods of the Middle Carboniferous of
the Donetz Basin. Gosgeolizdad, Moskva, 179 pp.***Rowley, R. R. 1893. Description of some new species ofcrinoids, blastoids and brachiopods from the Devonian and
Sub-Carboniferous rocks of Missouri. American Geologists
12: 303‒309.
Rzhonsnitskaya, M. A. 1956. Systematization of
Rhynchonellida; pp. 125‒126 in E. Guzman and others(eds.), Resumenes de Los Trabajos Presentodos. International
Geological Congress, Mexico, Report 20.
Rzhonsnitskaya, M. A. 1959. On the systematics of therhynchonellids. Paleontologichesii Zhurnal 1: 25‒36.***
Sakagami, S. 1962. Lower Carboniferous Bryozoa from the
Hikoroichi Series. Transactions and Proceedings of the
Palaeontological Society of Japan, New Series, 46: 227‒242.
Sakagami, S. 1989. Bryozoa; pp. 37‒49 in Ofunato City
Museum (ed.), Fossils of Onimaru. Kawashima Insatsu,
Ichinoseki.*
Sartenaer, P., and G. Plodowski. 1996. Restatement of the late
Tournaisian Spirifer tornacensis de Koninck, 1883 on thebase of the original collection. Bulletin de lʼInstitut Royal desSciences Naturelles de Belgique 66: 53‒71.
Sarytcheva, T. G. 1928. The Podmoskovian productid group
Productus giganteus Martin (Gigantella gen. nov.). Trudy
MGU 1: 1‒71.***
Sarytcheva, T. G. 1968. Brachiopods of the Upper Palaeozoicof the eastern Kazakhstan. Trudy Paleontologicheskogo
Instituta, Akademiya Nauk SSSR 121: 1‒212.***
Sarytcheva, T. G., and A. N. Sokolskaya. 1952. Description of
the Palaeozoic Brachiopoda of the Moscow Basin. Trudy
Paleontologicheskogo Instituta, Akadamiya Nauk SSSR 38:
1‒307.***
Sarytcheva, T. G., and A. N. Sokolskaya. 1959. On theclassification of pseudopunctate brachiopods. Doklady,
Akademii Nauk SSSR 125: 181‒184.***
Sarytcheva, T. G., A. N. Sokolskaya, G. A. Besnossova
and S. V. Maksimova. 1963. Carboniferous brachiopods
and palaeogeography of the Kuznetsk Basin. Trudy
Paleontologicheskogo Instituta, Akadamiya Nauk SSSR 95:

83

1‒547.***
Savage, N. M. 2002. Rhynchoporoidea; pp. 1232‒1235 in R. L.
Kaesler (ed.), Treatise on Invertebrate Paleontology, Part H
Brachiopoda Revised, Volume 4: Rhynchonelliformea (Part).
Geological Society of America, Boulder and University of
Kansas, Lawrence.
Schuchert, C. 1893. Classiﬁcation of the Brachiopoda. American
Geologist 11: 141‒167.
Schuchert, C. 1913. Class Brachiopoda; pp. 290‒449 in C.
Schuchert, C. K. Swarts, T. P. Maynard and R. B. Rowe, The
Lower Devonian Deposits of Maryland. Maryland Geological
Survey, Baltimore.
Schuchert, C. 1929. Classiﬁcation of brachiopod genera, fossil
and recent; pp. 10‒25 in J. F. Pompeckj (ed.), Fossilium
Catalogus, Vol. 1. W. Junk, Berlin.
Schuchert, C., and G. A. Cooper. 1932. Brachiopod genera of
the suborders Orthoidea and Pentameroidea. Memoirs of the
Peabody Museum of Natural History 4: 1‒270.
Schuchert, C., and C. M. Le Vene. 1929. Brachiopoda (generum
et genotyporum index et bibliographia); pp. 1‒140 in J. F.
Pompekj (ed.), Fossilim Catalogus, Vol. 1, Animalia, Part 42.
W. Junk, Berlin.
Scupin, H. 1900. Die Spiriferen Deutschland. Palaeontologische
Abhandlungen, Neue Folge 4: 207‒344.
Selden, P. A. 2007. Treatise on Invertebrate Paleontology, Part
H Brachiopoda Revised, Volume 6: Supplement. GeologicalSociety of America, Boulder and University of Kansas,
Lawrence, 906 pp. (2321‒3226).
S h i , G . R . , Z . - Q . C h e n a n d L . - P. Z h a n . 2 0 0 5 . E a r l y
Carboniferous brachiopod faunas from the Baoshan block,west Yunnan, southwest China. Alcheringa 29: 31‒85.
Simorin, A. M. 1956. Stratigraphy and Brachiopods of the
Karaganda Basin. Akademii Nauk Kazakhskoy SSR, AlmaAta, 296 pp.***Smirnova, T. N. 2004. The lower Kazanian (Upper Permian)
terebratulid assemblage of the family Beecheriidae fam.
nov. (Brachiopoda) from the east of the Russian Platform.
Palaeontologicheskiy Zhurnal, 2004, 2: 50‒54.***
Sokolskaya, A. N. 1950. Chonetidae of the Russian Platform.
Trudy Paleontologicheskogo Instituta, Akadamiya NaukSSSR 27: 1‒107.***
Sokolskaya, A. N. 1960. Order Strophomenida; pp. 206‒220 in
T. G. Sarytcheva (ed.), Bryozoa and Brachiopoda. Osnovy
Paleontologii, Vol. 7, Akademii Nauk SSSR, Moskva.***Sommer, K. 1909. Die Fauna des Culms von Königsberg bei
Giessen. Neues Jahrbuch für Mineralogie, Geologie undPaläontologie 28: 611‒660.
Sowerby, J. 1815‒1818. The Mineral Conchology of GreatBritain, Vol. 2. Published by the author, London, 235 pp.
Sowerby, J. 1821‒1822. The Mineral Conchology of GreatBritain, Vol. 4. Published by the author, London, 184 pp.
Sowerby, J. 1823‒1825. The Mineral Conchology of GreatBritain, Vol. 6. Richar Taylor, London, 230 pp.
Sowerby, J. 1840‒1846. The Mineral Conchology of GreatBritain, Vol. 7. Published by the author, London, 80 pp.

84

JUN-ICHI TAZAWA

Stehli, F. G. 1954. Lower Leonardian Brachiopoda of the Sierra
Diablo. Bulletin of the American Museum of Natural History
105: 257‒358.
Sun, Y.-L., and A. Baliński. 2008. Silicified Mississippianbrachiopods from Muhua, southern China: Lingulids,
craniids, strophomenids, productids, orthotetids, and orthids.
Acta Palaeontogica Polonica 53: 485‒524.
Sun, Y.-L., and A. Baliński. 2011. Silicified Mississippianbrachiopods from Muhua, southern China: Rhynchonellides,
athyridides, spiriferides, spiriferinides, and terebratulides.
Acta Palaeontogica Polonica 56: 793‒842.
Tazawa, J. 1980. Viséan brachiopods from the Karaumedate
Formation, southern Kitakami Mountains. Transactions andProceedings of the Palaeontological Society of Japan, New
Series, 119: 359‒370.
Tazawa, J. 1981a. Notes on some Carboniferous brachiopods
from the Kitakami Mountains, northeast Japan. Saito Ho-on
Kai Museum, Natural History, Research Bulletin 49: 53‒61.
Tazawa, J. 1981b. An Early Carboniferous brachiopod faunafrom the Karoyama Formation in the Kitakami Mountains,
northeast Japan. Saito Ho-on Kai Museum, Natural History,
Research Bulletin 49: 63‒78.
Ta z a w a , J . 1 9 8 4 a . O c c u r r e n c e o f G i g a n t o p ro d u c t u s(Carboniferous Brachiopoda) from the lower Hikoroichi
Formation, Kitakami Mountains and its significance. Earth
Science (Chikyu Kagaku) 38: 132‒134.*
Tazawa, J. 1984b. Early Carboniferous (Visean) brachiopods
from the Hikoroichi Formation of the Kitakami Mountains,
northeast Japan. Transactions and Proceedings of the
Palaeontological Society of Japan, New Series, 133: 300‒312.
Tazawa, J. 1985. Carboniferous brachiopods Marginatia andUnispirifer from the Hikoroichi and Arisu Formations,
Kitakami Mountains, Northeast Japan. Earth Science (Chikyu
Kagaku) 39: 459‒462.*
Tazawa, J. 1989. Brachiopoda; pp. 51‒65 in Ofunato City
Museum (ed.), Fossils of Onimaru. Kawashima Insatsu,
Ichinoseki.*Tazawa, J. 1996. Rotaia (Rhynchonellida, Brachiopoda)
from the Lower Carboniferous of northeast Japan and its
palaeobiogeographical significance. Science Reports ofNiigata University, Series E, 11: 1‒11.
Tazawa, J. 2002. Late Paleozoic brachiopod faunas of the South
Kitakami Belt, northeast Japan, and their paleobiogeographic
and tectonic implications. Island Arc 11: 287‒301.
Tazawa, J. 2004. The strike-slip model: A synthesis on the origin
and tectonic evolution of the Japanese Islands. Journal of theGeological Society of Japan 110: 503‒517.*
Tazawa, J. 2006. The Marginatia‒Syringothyris‒Rotaia
brachiopod assemblage from the Lower Carboniferousof the South Kitakami Belt, northeast Japan, and its
palaeobiogeographical implications. Paleontological Research
10: 127‒139.
Tazawa, J. 2017a. Discovery of Cyrtospirifer (Late Devonian
Brachiopoda) from Choanji in the South Kitakami Belt,
northeastern Japan. Journal of the Geological Society ofJapan 123: 101‒105.
Tazawa, J. 2017b. An early Carboniferous (late Visean)brachiopod fauna from Tairagai in the Yokota area, South
Kitakami Belt, Japan. Paleontological Research 21: 329‒346.
Tazawa, J., Y. Gunji and K. Mori. 1984. A Visean brachiopod
fauna from the Mano Formation, Soma district, Abukuma
Mountains, Northeast Japan. Transactions and Proceedingsof the Palaeontological Society of Japan, New Series, 134:
347‒360.
Tazawa, J., and Y. Ibaraki. 2009. Linoprotonia and Gigantoproductus(Linoproductoidea, Brachiopoda) from the Lower Carboniferous
in the Onimaru quarry, Hikoroichi, southern Kitakami
Mountains, NE Japan. Science Reports of Niigata University(Geology) 24: 7‒19.
Tazawa, J., Y. Ibaraki and I. Nishikawa. 2016. Early Carboniferous(Visean) brachiopods from the Hina Limestone, Okayama
Prefecture, SW Japan, and their palaeobiogeographicalimplications. Science Reports of Niigata Univerisyty(Geology) 31: 45‒68.
Tazawa, J., F. Itabashi and K. Mori. 1981. Lower Carboniferous
System in the Nisawa district, southern Kitakami Mountains,
Japan. Contributions from the Institute of Geology andPaleontology, Tohoku University 83: 21‒37.*
Tazawa, J., and T. Katayama. 1979. Lower Carboniferous
brachiopods from the Odaira Formation in the southern
Kitakami Mountains. Science Reports of the Tohoku
University, Second Series, 49: 165‒173.
Tazawa, J., and H. Kurita. 1986. Brachiopods and correlation of
the Lower Carboniferous Shittakazawa Formation, southern
Kitakami Mountains, Northeast Japan. Journal of theGeological Society of Japan 92: 167‒170.*
Tazawa, J., and I. Niikawa. 2018. Desquamatia (Seratrypa)
from the Upper Devonian Choanji Formation in the South
Kitakami Belt, Japan, and its stratigraphical significance.
Journal of the Geological Society of Japan 124: 111‒116.*
Thomas, G. A. 1971. Carbonifeous and Early Permianbrachiopods from western and northern Australia. Bulletin of
the Bureau of Mineral Resources, Geology and Geophysics
56: 1‒277.
Thomas, I. 1914. The British Carboniferous Producti, 1. Genera
Pustula and Overtonia. Memoirs of the Geological Survey ofGreat Britain, Palaeontology 1: 197‒366.
Tolmatchoff, I. P. 1924. Lower Carboniferous fauna of theKuznetsk Basin. Materialy po Obschey i Prikladnoy Geologii
25: 1‒663.***Ustritsky, V. I., and G. E. Tschernjak. 1963. Biostratigraphy andbrachiopods of the Upper Palaeozoic of Taimyr. Trudy NIIGA
134: 1‒139.***Vaughan, A. 1915. Correlation of Dinantian and Avonian.
Quarterly Journal of the Geological Society of London 71:
1‒52.
de Verneuil, E. 1845. Paléontologie, Mollusques, Brachiopodes;
pp. 17‒395 in R. I. Murchison, E. de Verneuil and A.
Keyserling (eds.), Géologie de la Russie dʼEurope et desMontagnes de lʼOural, Vol. 2. John Murray, London.

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATION

Volgin, V. I. 1965. Brachiopods of the Gazsk Formation, South
Fergana. Leningrad University, Leningrad, 95 pp. ***Volgin, V. I., and L. V. Kushnar. 1975. Late Visean Brachiopods
and Bivalves of Southern Fergana. Leningradskogo
Universiteta, Leningrad, 112 pp. ***Waagen, W. 1883‒1884. Salt Range fossils, 1. ProductusLimestone fossils: Brachiopoda. Palaeontologia Indica, Series
13, 1: 391‒546 (1883), 547‒728 (1884).
Wang, C.-W., and S.-P. Yang. 1998. Late Carboniferous‒
Early Permian Brachiopods of Central Xinjiang and Their
Biostratigraphical Studies. Geological Publishing House,
Beijing, 156 pp.**Wang, Y., Y.-G. Jin and D.-W. Fang. 1964. Brachiopod Fossilsof China, Part 2. Science Press, Beijing. pp. 355‒777.**
Waterhouse, J. B. 1968. The classification and descriptionsof Permian Spiriferida (Brachiopoda) from New Zealand.
Palaeontographica, Abteilung A 129: 1‒94.
Waterhouse, J. B. 1975. New Permian and Triassic brachiopod
taxa. Papers of the Department of Geology, University of
Queensland 7: 1‒23.
Waterhouse, J. B. 1981. Early Permian brachiopods from Ko
Yao Noi and near Krabi, southern Thailand; pp. 45‒124 in J.
B. Waterhouse, K. Pitakpaivan and N. Mantajit, The Permian
Stratigraphy and Palaeontology of Southern Thailand.
Geological Survey Memoir 4, Geological Survey Division,
Department of Mineral Resources, Bangkok.
Waterhouse, J. B. 1986. New Late Palaeozoic invertebrate taxa.
Bulletin of the Indian Geologistsʼ Association 19: 1‒8.
Waterhouse, J. B. 2002. Classiﬁcation within Productidina andStrophalosiidina (Brachiopoda). Earthwise 5: 1‒60.
Weller, S. 1906. Kinderhook faunal studies, 4. The fauna of
the Glen Park Limestone. Transactions of the Academy of
Science of St. Louis 16: 435‒471.
Weller, S. 1909. Kinderhook faunal studies, 5. The fauna of theFern Glen Formation. Bulletin of the Geological Society ofAmerica 20: 265‒332.
Weller, S. 1914. The Mississippian Brachiopoda of theMississippi Valley Basin. Illinois State Geological Survey
Monograph 1, Illinois State Geological Survey, Urbana, 508
pp.
White, C. A. 1860. Observations upon the geology andpaleontology of Burlington, Iowa, and its vicinity. Boston
Society of Natural History Journal 7: 209‒235.
Winchell, A. 1863. Descriptions of fossils from the Yellow
Sandstone lying beneath the Burlington Limestone at
Burlington, Iowa. Proceedings of the Academy of Natural
Sciences of Philadelphia, Series 2, 15: 2‒25.
Winchell, A. 1865. Descriptions of new species of fossils, from
the Marshall Group of Michigan, and its supposed equivalent,in other states; with notes on some fossils of the same age
previously described. Proceedings of the Academy of Natural
Sciences of Philadelphia, Series 2, 17: 109‒133.
Winkler Prins, C. F. 1968. Carboniferous Productidina andChonetidina of the Cantabrian Mountains (NW Spain):
Systematics, stratigraphy and palaeontology. Leidse

85

Geologische Mededelingen 43: 41‒155.
Xu, H.-K., and Z.-G. Yao. 1988. Brachiopoda; pp. 263‒326
in C.-M. Yu (ed.), Devonian‒Carboniferous Boundary in
Nanbiancun, Guilin, China: Aspects and Records. Science
Press, Beijing.
Yabe, H., and T. Sugiyama. 1937. Preliminary report on thefossiliferous Gotlandian and Devonian deposits newly
discovered in the Kitakami Mountainland. Proceedings of theImperial Academy of Tokyo 13: 417‒420.
Yanagida, J. 1962. Carboniferous brachiopods from Akiyoshi,
southwest Japan, Part 1. Memoirs of the Faculty of Science,
Kyushu University, Series D, 12: 87‒127.
Yanagida, J. 1973. Early Carboniferous Viséan faunas discoveredfrom Mitsuzawa, Part 2. Brachiopods. Transactions andProceedings of the Palaeontological Society of Japan, New
Series, 90: 99‒112.
Yanai, S., O. Koseki, S. Niko and T. Hamada. 1988. Tateishi
nappe and Upper Devonian to Lower Carboniferous innorthern area of the Abukuma Mountains, NE Japan. Earth
Science (Chikyu Kagaku) 42: 179‒186.*
Yang, D.-L. 1984. Systematic description of Palaeontology:
Brachiopoda; pp. 203‒239, 330‒333 and 387‒396 in
Yichan Institute of Geology and Mineral Resources(ed.), Biostratigraphy of the Yangtze Gorge Area (3) Late
Palaeozoic Era. Geologocal Publishing House, Beijing.**Yang, D.-L., S.-Z.Ni, M.-L. Chang and R.-X. Zhao. 1977.
Phylum Brachiopoda; pp. 303‒470 in Geological Institute of
Hubei et al. (eds.), Palaeontological Atlas of South-Central
China, Part 2. Late Palaeozoic Volume. Geological Publishing
House, Beijing.**Yang, F. 1994. Chapter 7 Carboniferous; pp. 131‒162 in H.F. Yin (ed.), The Palaeobiogeography of China. Oxford
University Press, New York.
Yang, S.-P. 1964. Lower and Middle Carboniferous Brachiopods
from the Northern Slope of Mt. Borochoro, Xinjiang, China,
and their Stratigraphical Signiﬁcane. Science Press, Beijing,
179 pp. **Yang, S.-P. 1983. Palaeozoogeographic provinces of theLower Carboniferous brachiopods of China; pp. 64‒73 in
Editorial Committee on Basic Theory of Palaeontology (ed.),
Palaeobiogeographic Provinces of China. Science Press,
Beijing.**Yang, S.-P., and Y.-N. Fan. 1983. Carboniferous brachiopods
from Xizang (Tibet) and their faunal provinces; pp. 265‒289
in CGQXP Editional Committee, Ministry of Geology
and Mineral Resources, PRC (eds.), Contribution to the
Geology of the Qinghai‒Xizang (Tibet) Plateau, Volume 11.
Geological Publishing House, Beijing.**Yang, Z.-Y., P.-Z. Ting (Ding), H.-F. Yin, S.-X. Zhang and J.-S.
Fang. 1962. Carboniferous, Permian and Triassic brachiopod
faunas from the Chilianshan region; pp. 1‒129 in Institute ofGeology and Paleontology, Geological Institute, Academia
Sinica and Beijing University of Geology (eds.), Monographon Geology of the Chilianshan Mountains, Vol. 4, Pt. 4.
Science Press, Beijing.**

86

JUN-ICHI TAZAWA

Yanishevsky, M. 1918. Materials for the study of the Lower
Carboniferous fauna of Fergana. Trudy Geologicheskago
Komiteta, Novaya Seriya 162: 1‒145.***
Zakowa, H. 1985. Some Productidina (Brachiopoda) from theUpper Visean of Galęzice. Kwartalnik Geologiczny, 29:301‒328.
Zakowa, H. 1989. Orthid brachiopods from the Upper Visean(Carboniferous) of the Świetokrzyskie Mts., Poland. Acta
Palaeontologica Polonica 34: 91‒124.
Zhan, L.-P., and S.-Z. Wu. 1987. Brachiopoda; pp. 201‒232
in Institute of Geology, Xinjiang Geological Bureau andInstitute of Geology, China Academy of Geological Sciences(eds.), The Carboniferous and Permian Stratigraphy and Biota
in Kalpin Region, Xinjiang. Ocean Press, Beijing.**＜
Arisu Formation ……………… 有住層
Choanji ……………………… 長安寺
Higuchizawa ………………… 樋口沢Hikami Granite ………… 氷上花崗岩
Hikoroichi Formation ……… 日頃市層
Hina …………………………… 日南
Kanenokura Stage ………… 金ノ倉階Karaumedate Formation …… 唐梅館層Karoyama Formation ……… 加労山層

Zhang, C., F.-M. Zhang, Z.-X. Zhang and Z. Wang. 1983.
Phylum Brachiopoda; pp. 262‒386 in Regional Geological
Surveying Team of Xinjiang, Institute of Geoscience ofXinjiang, and Geological Surveying Group of Petroleum
Bureau of Xinjiang (eds.), Palaeontological Atlas ofNorthwest China; Xinjiang Autonomous Region, Part 2. Late
Palaeozoic Volume. Geological Publishing House, Beijing.**
*
**
***

地名・地層名: in Japanese: in Chinese: in Russian＞

Kawauchi Formation ………… 川内層
Mt. Goyosan ………………… 五葉山
Nagaiwa Formation ………… 長岩層
Nagasaka ……………………… 長坂
Nakazato Formation ………… 中里層
Nisawa ………………………… 荷沢
Odaira Formation …………… 大平層
Ofunato ……………………… 大船渡
Okuhinotsuchi …………… 奥火の土

Omori …………………………… 大森
Onimaru Formation ………… 鬼丸層
Otsubo ………………………… 小坪
Sakamotozawa ……………… 坂本沢Shimoarisu …………………… 下有住
Shizu …………………………… 清水
Soma …………………………… 相馬
Yokota ………………………… 横田

MISSISSIPPIAN BRACHIOPODS FROM HIKOROICHI FORMATIONAPPENDIX
Topographic and stratigraphic locations, lithologies andcontaining brachiopod species of the 16 fossil localities (KAC1,
KAC3, KF148, KF151, KF152, KF153, KF156, KF163, KF167,
KF171, KF172, KF174, KF188, KF200, KF201 and KF202) inthe Hikoroichi area.
KAC1: Upper stream of small valley at SE of Onimaru (39 06
57 N, 141 39 07 E), dark grey ﬁne-grained sandstoneof the upper part of the HK4 Unit, with Leptagonia
analoga, Argentiproductus margaritaceus, Ovatia cf.
prolata, Schizophoria resupinata and Torynifer asiatica.
KAC3: NE of Omori (39 06 37 N, 141 39 38 E), dark grey
fine-grained sandstone of the lower part of the HK1
Unit, with Marginatia magna, Tylothyris laminosa and
Syringothyris sibirica.
KF148: Near (60 m SW) the summit of Mt. Senokidayama (3906 49 N, 141 39 19 E), medium-grained tuffaceous
sandstone of the upper part of the HK4 Unit, with
Punctospirifer iwatensis.
KF151: Northern ridge of Mt. Senokidayama, at 590 m N of thesummit (39 07 10 N, 141 39 17 E), grey, ﬁne-grained,
calcareous sandstone of the top of the HK4 Unit, withSpirifer liangchowensis.
KF152: Northern ridge of Mt. Senokidayama, at 15 m SW of
KF151 (39 07 09 N, 141 39 16 E), grey ﬁne-grainedsandstone of near the top of the HK4 Unit, with Pustula
pustulosa.
KF153: Northern ridge of Mt. Senokidayama, at 20 m SW of
KF151 (39 07 08 N, 141 39 17 E), grey, ﬁne-grained,
calcareous sandstone of the top of the HK4 Unit, with
Angiospirifer bisulcatus.
KF156: Road cut at 150 m SW of junction of the Sakamotozawa
and Onimarusawa valleys (39 07 43 N, 141 39 26 E),
greenish-grey, ﬁne-grained tuffaceous sandstone of near
the base of the HK4 Unit, with Rotaia hikoroichiensis.
KF163: Upper Higuchizawa Valley (39 07 09 N, 141 39 22E), grey, arenaceous limestone of the upper part of the
HK4 Unit, with Rhipidomella michelini and Dimegelasmasp.
KF167: Eastern slope facing the Onimarusawa Valley, 280 m
SW of junction of the Sakamotozawa and Onimarusawa
valleys (39 07 43 N, 141 39 19 E), grey, finegrained sandstone of the lower part of the HK4 Unit,
with Karavankina jizodoensis, Orthotetes keokuk,

87

Schizophoria resupinata and Cleiothyridina sp.
KF171: Hill along the eastern side of the Onimarusawa Valley (3907 26 N, 141 39 22 E), black shale of near the base of
the HK2 Unit, with Marginatia burlingtonensis.
KF172: Hill along the eastern side of the Onimarusawa Valley(39 07 27 N, 141 39 18 E), black shale of the lower
part of the HK2 Unit, with Leptagonia analoga andMarginatia burlingtonensis.
KF174: Hill along the eastern side of the Onimarusawa Valley (3907 34 N, 141 39 29 ), black shale of the upper part of
the HK2 Unit, with Marginatia burlingtonensis.
KF188: Hill along the eastern side of the Onimarusawa Valley (3907 35 N, 141 39 31 E), black shale of the upper partof the HK2 Unit, with Marginatia burlingtonensis, Ovatia
laevicosta, Schizophoria resupinata and Kitakamithyris
hikoroichiensis.
KF200: Southern upper stream of the Higuchizawa Valley (3906 43 N, 141 39 35 E), grey fine-grained sandstoneof the lower part of the HK1 Unit, with Schizophoria
resupinata, Cleiothyridina harkeri, Kitakamithyris
tyoanjiensis, Punctospirifer sp. and Beecheria khalfini.
KF201: Eastern slope at 120 m NE of KF200 (39 6 47 N, 141
39 39 E), grey ﬁne-grained sandstone of the lower partof the HK1 Unit, with Leptagonia analoga, Plicochonetessp., Tolmatchoffia robusta, Marginatia burlingtonensis,
M. magna, Ovatia sp., Schizophoris resupinata,
Rhynchopora sp., Unispirifer mediocris, Acuminothyris
keokuk, Brachythyris suborbicularis, Kitakamithyris
tyoanjiensis, K. hikoroitiensis, Syringothyris sibirica andPunctospirifer sp.
KF202: Onimaru quarry, NE of Onimaru (39 07 28 N, 141 39
20 E), dark grey, fine-grained calcareous sandstoneof the upper part of the HK2 Unit, with Leptagonia
analoga, Rugosochonetes cf. hardrensis, Leiochonetes
onimarensis, Tolmatchoffia robusta, Marginatia
burlingtonensis, Echinoconchus punctatus, Ovatia
laevicosta, Linoprotonia ashfellensis, L. hikoroichiensis,
Gigantoproductus crassiventer, G. latiexpansus,
Orthotetes keokuk, Rhipidomella michelini, Rotaia
hikoroichiensis, Actinoconchus expansus, A. paradoxus,
Lamellosathyris qaidamensis, Hustedia sp., Spirifer
karagai, Unispirifer striatoconvolutus, U. kozuboensis,
Tegulocrea incerta, Acuminothyris triangularis,
Pseudosyrinx cf. missouriensis, Punctospirifer
plicatosulcatus, and P. gigantiformis.