Two Late Pennsylvanian conodont species, Gondolella sublanceolata Gunnell and Idiognathodus sulciferus Gunnell, were extracted from limestone boulders in the Mizuboradani Valley, Fukuji district, central Japan. These provide the first evidence of Missourian (Kasimovian) cosmopolitan conodonts in the Akiyoshi and Hida Gaien belts, Inner Zone of Japan. The limestone boulders might be derived from the Ichinotani Formation and/or from limestone clasts in conglomerates of the Permian Sorayama Formation that crop out in the Mizuboradani Valley.
Introduction
The Fukuji district, Takayama City, Gifu Prefecture, central Japan, is well known as a locality of Paleozoic sedimentary rocks that form a component of the Hida Gaien belt (Figures 1.1, 1.2). The Devonian Fukuji, Carboniferous Ichinotani, and Permian Mizuyagadani formations comprise mainly fossiliferous limestone beds that have been investigated for over 60 years for their paleontological and lithological signature (e.g. Igo, 1956; Niikawa, 1978, 1980; Harayama, 1990). Both macro- and microfossils, including fusulinids, corals, brachiopods, cephalopods, trilobites, ostracods and conodonts have been reported from the three formations (Igo, 1956; Niikawa, 1978, 1980; Kuwano, 1987; Niko and Hamada, 1987; Tazawa et al., 2010; Tanaka et al., 2013; Stocker et al., 2018). In the Mizuboradani Valley (Figure 1.3), located on the northwest side of Okuhidaonsen Village, Carboniferous macro- and microfossils are reported from gray, greenish gray and dark gray limestone boulders (Goto and Okura, 2004; Tazawa et al., 2010; Isaji and Okura, 2014; Stocker et al., 2018). Goto and Okura (2004) reported 14 isolated teeth of Carboniferous and Permian cartilaginous fish. Isaji and Okura (2014) described well preserved molluscan larval shells, including gastropods and bivalves, from the Ichinotani Formation. Stocker et al. (2018) reported an ostracod assemblage from the Carboniferous Ichinotani Formation consisting of silicified carapaces of Amphissites, Kirkbya, Bairdia, Aechmina and Healdia species. According to Stocker et al. (2018), the ostracods indicate a shallow-marine ‘Eifelian type’ ecological assemblage of Becker (1971), which is characteristic of mid-Palaeozoic fore-reef ecosystems.
In this paper, we report newly discovered Carboniferous conodonts from float limestone boulders in the Mizuboradani Valley. Provenance of the boulders and the biostratigraphic distribution of the species/fauna are also discussed herein.
Geological setting
The three southern islands of Japan (Honshu, Shikoku and Kyushu) are comprised predominantly of Paleozoic, Mesozoic and Cenozoic accretionary complexes (Isozaki and Maruyama, 1991; Isozaki et al., 2010). The Jurassic to Cenozoic accretionary complexes form elongate tectonic belts bounded by major structural lineaments. The Median Tectonic Line, one of the major tectonic lineaments of Japan, subdivides rocks of the ‘Outer Zone’, composed of the Sambagawa metamorphic belt, Chichibu belt, Kurosegawa belt, and Shimanto belt, from those of the ‘Inner Zone’ consisting mainly of the Ryoke metamorphic belt, and Mino, Akiyoshi, Hida and Hida Gaien tectonic belts on Honshu Island (Taira, 1990; Isozaki et al., 2010).
Figure 1.
1, map showing southwest and central Japan with distribution of limestone blocks in the Akiyoshi and Hida Gaien belts, modified from Ishida et al. (2013). Abbreviations: MTL, Median Tectonic Line; ISTL, Itoigawa-Shizuoka Tectonic Line. 2, distribution of Hida Gaien belt, modified from Tsukada et al. (2004). 3, map showing the study area with the locality of conodont-bearing limestone boulders (marked by the star), Gifu Prefecture, central Japan modified from 1:25,000 scale topographic map ‘Yakedake’ of the Geospatial Information Authority of Japan (GSI). 4, geological map of the Fukuji area modified from Harayama (1990).

The Fukuji district is situated in the eastern part of the Fukuji-Hongo-Furukawa area, which is the stratotype area of the Hida Gaien belt (Kamei and Igo, 1957; Tsukada et al., 2004). According to Tsukada et al. (2004), in the Fukuji-Hitoegane district, Ordovician to Permian sedimentary rocks are divided into six formations in ascending order: the Ordovician to Silurian Hitoegane Formation, the Silurian Yoshiki Formation, the Devonian Fukuji Formation, the Carboniferous Ichinotani Formation, the Permian Mizuyagadani Formation, and the Permian Sorayama Formation (Figure 1.4). The physical stratigraphical relationships between these formations are summarized by Niikawa (1978, 1980), Harayama (1990), Tsukada et al. (1999, 2004) and Williams et al. (2014). The Ichinotani Formation, mainly composed of fossiliferous marine carbonate rocks, is subdivided into six fusulinid zones, namely, Eostaffella, Profusulinella, Lower Fusulinella-Fusulina, Beedeina, Upper Fusulinella-Fusulina, and Triticites zones in ascending order. The Mizuyagadani Formation consists of carbonates and siliciclastic rocks, in which the Pseudoschwagerina and Pseudofusulina fusulinid zones are recognised (Figure 1.4; Niikawa, 1980; Harayama, 1990). The Sorayama Formation conformably overlies the Mizuyagadani Formation, and is dominated by massive pyroclastic rocks and intercalating conglomerates, sandstones and mudstones (Tsukada et al., 1999).
In the Mizuboradani Valley, carbonate and siliciclastic rocks of the Lower Fusulinella-Fusulina Zone to Triticites Zone of the Ichinotani Formation, carbonate rocks of the Pseudoschwagerina Zone of the Mizuyagadani Formation, and tuffaceous rocks of the Sorayama Formation crop out (Figure 1.4; Harayama, 1990). The Ichinotani and Mizuyagadani formations dominate the southern part of the valley, with the Sorayama Formation in the northern part. In the downstream of the valley, Carboniferous and Permian float limestone boulders that contain abundant macro- and microfossils have been reported (e.g. Goto and Okura, 2004). In this study, we collected float limestone boulders from the upstream of the valley to extract conodont microfossils.
Figure 2.
1, polished slab of conodont-bearing greenish gray float limestone boulder (T-01). Abbreviations: C, crinoid stems; B, brachiopod shell. 2–5, photomicrographs of thin section of float limestone boulders. 2, bioclastic limestone (right side) covered by tuffaceous mudstone (left side) including fusulinid foraminifera (white arrow) (T-01). Dashed line shows the boundary between limestone and mudstone. 3, bioclastic limestone (T-01). 4, bioclastic limestone covered by tuffaceous mudstone (I-02). Dashed line shows the boundary between limestone and mudstone. 5, bioclastic limestone (K-01).

Material and methods
We analyzed four greenish-gray limestone boulders (numbered I-01, I-02, K-01, T-01) collected as float in the downstream of the Mizuboradani Valley (Figures 1–3). These limestone boulders, 10–40 cm in diameter, associated with greenish-gray tuff and tuffaceous mudstone (Figures 2.2, 2.4). Goto and Okura (2004), and Isaji and Okura (2014) yielded well preserved microfossils from I-01, and Stocker et al. (2018) reported well preserved ostracods from three boulders (I-01, I-02, K-01: Figure 3). In the laboratory, these limestones were crushed into pellets 2–3 cm in diameter and immersed in 5–6% acetic acid to extract the phosphatic microfossils. Conodont elements were obtained from I-01, K-01 and T-01 (Figure 3). This process also recovered abundant silicified fusulinid foraminifers, gastropods, bivalves, brachiopods and ostracods from the limestones.
We used the Olympus SZX7 light microscope (Olympus Co., Ltd., Tokyo) for picking microfossils. Conodonts were coated with platinum using the magnetron sputter MSP-1S (Vacuum Device Co., Ltd., Ibaraki) for capturing images with an Ace-700-S Scanning Electron Microscope (Sanyu Electron Co., Ltd., Tokyo).
Systematic paleontology
Species were identified based on the P1 element. The specimens studied in this paper are stored in the Micropaleontology Collection (MPC) of the National Museum of Nature and Science, Tsukuba, Japan. Orientation, terminology and measurement of P1 elements are shown in Figure 4, following Clark and Mosher (1966), Sweet (1988), Purnell et al. (2000), and Rosscoe and Barrick (2009). All elements are well preserved and of a dark-gray color.
Order Prioniodinida Sweet, 1988
Superfamily Gondolelloidea (Lindström, 1970)
Family Gondolellidae Lindström, 1970
Genus Gondolella Stauffer and Plummer, 1932
Type species.—Gondolella elegantula Staufler and Plummer, 1932.
Remarks.—Genus Gondolella, a distinctive platform type, was described on the basis of the gondola-shaped P1 element (Stauffer and Plummer, 1932; Clark and Mosher, 1966). According to Henderson and Orchard (1991), the genus evolved from Mesogondolella clarki (Koike, 1967), which has a wide segminiplanate element and lacks ornamentation on the platform, and is of early Moscovian, Early Pennsylvanian age. The oldest known species of Gondolella (Gondolella laevis Kossenko and Kozitskaya in Kossenko, 1975), which was described from early Moscovian strata, lacks transverse ridges on the platform, but late Moscovian to Gzhelian species (e.g. Gondolella bella Stauffer and Plummer, 1932) have conspicuous transverse ridges on the platform (Clark and Mosher, 1966; von Bitter and Merrill, 1980). The egminate types (e.g. Gondolella gymna Merrill and King, 1971) range from the early Moscovian to Gzhelian, and subsequently evolved into the segminiplanate type and genus Gondolelloides Henderson and Orchard, 1991 in the early Permian (Henderson and Orchard, 1991).
Multielement apparatuses of some species of Gondolella (e.g. Gondolella sublanceolata) are reconstructed by von Bitter (1976), von Bitter and Merrill (1980), and Merrill and von Bitter (2007).
Figure 3.
Conodonts and other fossil from the four float limestone boulders (I-01, I-02, K-01 and T-01) in the downstream of the Mizuboradani Valley (this and previous studies), and from a dark gray limestone boulder reported by Tazawa et al. (2010). Open circles show our original data. Solid circles show the fossils figured in previous studies. Double circles show the fossils observed by Stocker et al. (2018).

Figure 4.
Orientation, terminology and measurement for P1 elements of Gondolella and Idiognathodus. Abbreviations: L, length; H, height; W, width; b, blade; bc, basal cavity; bg, basal groove; c, cusp; ca, carina; cr, crest; fb, free blade; iar, inner adcarinal ridge; il, inner lobe; k, keel; me, medial carina; oar, outer adcarinal ridge; ol, outer lobe; pf, platform; pt, posterior tip; tr, transverse ridge.

Gondolella sublanceolata
Gunnell, 1933
Figure 5
Gondolella bella Stauffer and Plummer, 1932. Clark and Mosher, 1966, p. 383, pl. 45, figs. 5–9.
Gondolella sublanceolata Gunnell, 1933, p. 287, pl. 32, figs. 53–55; Ellison, 1941, p. 122, pl. 21, figs. 18, 19, 22, 23, 34, 35; Lindström, 1964, fig. 55C; Clark and Mosher, 1966, p. 387, pl. 45, figs. 20–22, 24–30; von Bitter, 1976, p. 5, 7, 10, fig. 3G–K.
Gondolella symmetrica Ellison, 1941, p. 121, pl. 21, figs. 3, 4, 17, 21, 26, 30; Lindström, 1964, p. 61, figs. 21K, 40C.
Gondolella dubia Ellison, 1941, p. 124, pl. 21, figs. 8, 12, 16.
Material examined.—Three specimens, MPC32956-32958, from I-01, and one specimen, MPC32959, from K-01.
Description.—Four slender gondola-like segminiplanate elements; platform shows isosceles triangle-like outline with sharply pointed anterior end and rounded or square posterior end. 1.02–1.22 mm, average 1.11 mm in length; 0.24–0.32 mm, average 0.28 mm in height; 0.17– 0.27 mm, average 0.23 mm in width; giving a length to width ratio of 4.2–4.8 in four specimens. In upper view, carina bears 13–17 short pointed and erect denticles which contain a posterior node-like cusp; surface of denticulation covered by striations; platform ornamented with amorphous transverse ridges which makes denticulation on the platform margin; weak lateral furrow; posterior end of the platform fused with cusp. In lateral view, highest point situated in anterior one-fourth of length; height of denticulation gradually decreases to cusp; straight basal margin downturned just anterior to flange; cusp declined posteriorly; platform margin undulated. In lower view, wide keel (one-third of element width) runs from oval or rectangular flange to anterior end; groove splits the keel, starts from basal pit; width of flange achieves that of element. Largest specimen (MPC32958) has a well developed free blade and transverse ridges on the platform; these ridges form crests along the lateral side of the carina.
Remarks.—Gondolella sublanceolata is characterized by a slender lanceolate segminiplanate P1 element with developed free blade (Clark and Mosher, 1966). These features distinguish this species from the other species of Gondolella (e.g. Gondolella bella). In this study, both small and large specimens of G. sublanceolata were collected, but the free blade is only developed in the largest specimen. However, the other features: outline of the platform, transverse platform ridges, denticulation of carina, downturned flange, straight and wide keel with furrow, are shared by all specimens.
Gondolella cf. sublanceolata reported by Kusunoki et al. (2004) is characterized by distinct denticulation of the carina on a broad platform and is distinguished from G. sublanceolata by lack of transverse platform ridges and free blade. The form of the former species is similar to that of the species of genus Mesogondolella.
Occurrence.—Missourian of North America (Stauffer and Plummer, 1932; Gunnell, 1933; Ellison, 1941; Clark and Mosher, 1966; von Bitter, 1976) and Carboniferous limestone boulders from the Mizuboradani Valley, Gifu Prefecture, central Japan (this study).
Figure 5.
SEM images of P1 elements of Gondolella sublanceolata Gunnell, 1933. 1, MPC32956; 2, MPC32957; 3, MPC32958; 4, MPC32959. 1–3 from I-01; 4 from K-01. For 1–4; a, d, lateral views; b, upper view; c, lower view.

Order Ozarkodinida Dzik, 1976
Family Idiognathodontidae Harris and Hollingsworth, 1933
Genus
Idiognathodus
Gunnell, 1931
Type species.—Idiognathodus claviformis Gunnell, 1931.
Remarks.—Idiognathodus was established by Gunnell (1931) based on segminiscaphate P1 elements, showing complex ornamentation on the platform surface. The type material is from the Fort Scott Limestone, Missouri, USA. According to Sweet (1988), larger P1 elements are characterized by a broader platform, transverse ridges on the posterior tip, and lobate areas occupied by nodes developed on the anterolateral shoulders of the platform. These features distinguish Idiognathodus from Streptognathodus Stauffer and Plummer, 1932. The biostratigraphic range of species in Idiognathodus is from the Pennsylvanian to the Permian (Sweet, 1988).
Idiognathodus sulciferus
Gunnell, 1933
Figure 6
Idiognathodus sulciferus Gunnell, 1933, p. 271, pl. 31, fig. 16; Barrick and Boardman, 1989, p. 185, pl. 1, figs. 9, 23, 24; Barrick and Walsh, 1999, p. 154, fig. 6.1; Ritter et al., 2002, p. 508, figs. 8.21, 8.25; Barrick et al., 2004, p. 241, pl. 4, fig. 13; Rosscoe and Barrick, 2009, p. 127, 128; Barrick et al., 2013, pl. 3, fig. 12.
Idiognathodus sulciferus sulciferus Gunnell, 1933. Rosscoe and Barrick, 2009, p. 127, pl. 3, figs. 12–17, pl. 6, fig. 2a.
Idiognathodus chiriformis Gunnell, 1933, p. 272, pl. 31, fig. 23. Idiognathodus cuneiformis Gunnell, 1933, p. 270, pl. 31, fig. 8; Barrick and Walsh, 1999, p. 153, fig. 5.2.
Idiognathodus swadei Rosscoe and Barrick, 2009. Qi et al., 2013, fig. 9i, j.
Idiognathodus n. sp. 1. Goreva et al., 2009, fig. 6B–F.
Material examined.—Six specimens, MPC32960, 32961, 32963–32965, 32967, from I-01, and six specimens, MPC32962, 32966, 32968–32971, from K-01.
Description.—Medium- to large-sized elements (MPC32962-32971): ten segminiscaphate elements; 1.21–1.49 mm in length; 0.42–0.48 mm in height; 0.35– 0.4.0 mm in width; length to width ratio 3.5–4.0 (measurements based on three complete specimens). Largest specimen (MPC32971) probably achieves 2.0 mm in length. Laterally compressed free blade bears 10–12 triangular-shaped denticles which continue to medial carina, and length of blade achieves over one-third of element length; highest point situated in anterior. Both adcarinal ridges extend from central part of platform to blade; bearing short pointed or node-like denticles in anterior; constricted to central side at both the inner and outer lobes; gradually disrupted on platform. Width of inner adcarinal ridge is twice that of outer one. Platform surface covered by microreticulation. Outer lobe moderately restricted; bears one or two rows of nodes, which align with medial carina. Laterally subrounded or pointed inner lobe bears one to two rows of nodes or ring of discrete nodes. Relatively narrow pointed posterior tip; covered by 5–7 widely spaced transverse ridges; shows normal surface or weak eccentric groove. Basal margin straight under free blade; arched under platform; strongly downturned to platform end. Deep basal cavity continues to anterior basal groove. Attachment surface shows facet of concentric layer.
Small-sized elements (MPC32960, 32961): segminiscaphate elements; 0.70 and 0.77 mm in length; 0.21 and 0.24 mm in height; 0.13 and 0.16 mm in width; length to width ratio 4.0 and 4.9 in two specimens. Laterally compressed free blade bears 8 triangular-shaped pointed denticles, which continue to medial carina. The carina continues to half to two-thirds of platform length. Both adcarinal ridges bear pointed denticles and continue to over half the length of the platform. Inner ridge slightly flanged. Inner and outer lobes are incompletely developed. Slender subrounded posterior tip; covered by 2–4 transverse ridges; shows normal surface. Basal margin straight and gradually upturned anteriorly under free blade; arched under platform. Basal cavity continues to anterior basal groove.
Remarks.—According to Rosscoe and Barrick (2009), Idiognathodus sulciferus is characterized by a moderately restricted outer lobe, better developed inner lobe which protrudes in a triangular shape, pointed posterior tip covered by coarse transverse ridges, and moderately outward flared inner adcarinal ridge forming a high collar around free blade. Idiognathodus expansus Stauffer and Plummer, 1932 is identified by its restricted outer lobe, and subrounded posterior tip covered by a closely spaced transverse ridge. Idiognathodus swadei Rosscoe and Barrick, 2009, bears an expanded and highly developed outer lobe covered by discrete hemispherical nodes.
Idiognathodus sulciferus is distinguished from I. expansus and I. swadei by its moderately restricted outer lobe, outward flared adcarinal ridge and widely spaced transverse ridges on the platform. Medium-sized specimens of I swadei (Rosscoe and Barrick, 2009, pl. 2, figs. 5, 6, 12) are discriminated by their larger number of spherical nodes on both the inner and outer lobes.
Idiognathodus n. sp. 1 from Russia (Goreva et al., 2009, fig. 6B–F), and I. swadei from South China (Qi et al., 2013, fig. 9i, j), are both characterized by their moderately restricted lobes and widely spaced transverse ridges on the posterior tip. These are considered conspecific with I. sulciferus.
Occurrence.—The first occurrence datum of this species is in the Lower Checkerboard Limestone of the Helper Formation, South Mound Checkerboard Cyclothem, USA (uppermost Desmoinesian); its last occurrence datum is in the Swope Cyclothem, USA (lower Missourian) (Rosscoe and Barrick, 2009; Barrick et al., 2013). This range encompasses the Idiognathodus eccentricus, I. turbatus, and I. cancellosus conodont zones and corresponds to the lower Kasimovian, Upper Pennsylvanian. Idiognathodus sulciferus is also known from the lower Kasimovian in South China (Qi et al., 2013) and Russia (Goreva et al., 2009).
Figure 6.
SEM images of P1 elements of Idiognathodus sulciferus Gunnell, 1933. 1, MPC32960; 2, MPC32961; 3, MPC32962; 4, MPC32963; 5, MPC32964; 6, MPC32965; 7, MPC32966; 8, MPC62967; 9, MPC32968; 10, MPC32969; 11, MPC32970; 12, MPC32971. 1, 2, 4–6, 8 from I-01; 3, 7, 9–12 from K-01. For 1–12; a, lateral view; b, upper view; c, lower view; d, postero-upper view.

Biostratigraphy of the conodonts
The conodont species Gondolella sublanceolata and Idiognathodus sulciferus from limestone boulders I-01, K-01 and T-01, derived from the upstream of the Mizuboradani Valley (Figure 1.3), indicate an early Missourian (Kasimovian, Late Pennsylvanian) age. Previously, Kuwano (1987) reported Devonian and a few Carboniferous conodonts from the Fukuji and Ichinotani formations, in the Ichinotani Valley located about 500 m south of the Mizuboradani Valley (Figure 1.4): the Carboniferous conodonts he reported are Idiognathodus sinuosus Ellison and Graves, 1941 and Streptognathodus lateralis Higgins and Bouckaert, 1968, from the Ichinotani Formation. These conodonts indicate an Early Pennsylvanian, Bashkirian age (Barrick et al., 2013). Niikawa (1978, 1980) established fusulinid range zones in the Ichinotani and Mizuyagadani formations in the Fukuji district, and estimated the age to be from Visean (Middle Mississippian) to Asserian (earliest Permian). However, Kasimovian strata have not been reported, because the Moscovian (Middle Pennsylvanian) and Gzhelian (uppermost Pennsylvanian) parts of the lithostratigraphy are structurally juxtaposed (Niikawa, 1980). Thus, this is the first report of Kasimovian conodonts in the Hida Gaien belt.
In Japan, Carboniferous conodonts have mainly been reported from Carboniferous to Permian massive limestone olistoliths in the Akiyoshi belt by several authors: Mississippian to Middle Pennsylvanian conodont assemblages were reported from the Omi Limestone, Niigata (Igo and Koike, 1964), the Akiyoshi Limestone, Yamaguchi (Igo, 1973), the Atetsu Limestone, Okayama (Koike, 1967), the Hina Limestone, Okayama (Mizuno, 1997), and the Ko-yama Limestone, Okayama (Ishida et al., 2012, 2013). An Early Pennsylvanian conodont assemblage was also reported from limestone blocks in the Chichibu belt (Ishida et al., 2005), and Gzhelian (latest Pennsylvanian) conodonts were reported from the Tamba belt (Kusunoki et al., 2004). However, prior to this study, Kasimovian conodonts have not been reported, and their discovery is important for biostratigraphic correlation with other global areas.
Possible source of the conodont-bearing limestone boulders
In the Mizuboradani Valley, Carboniferous marine fossils have been documented from limestone boulders (Goto and Okura, 2004; Tazawa et al., 2010; Isaji and Okura, 2014; Stocker et al., 2018). The dark gray limestone blocks reported by Tazawa et al. (2010) contain fusulinids (Eostaffella? sp., Millerella aff. marblensis Thompson, 1942, Ozawainella sp.) that indicate the upper Bashkirian (Lower Pennsylvanian), probably within the Profusulinella Zone that exists in the Ichinotani Formation. Isaji and Okura (2014) reported larval shells of gastropods and bivalves from greenish-gray limestone boulders (I-01) that contain fusulinids, brachiopods, ostracods, trilobites, crinoids, conodonts and isolated teeth of cartilaginous fish described by Goto and Okura (2004). Isaji and Okura (2014) suggested that the limestone was possibly derived from the interval of the Fusulinella-Fusulina Zone within the Ichinotani Formation. Sample I-02, reported by Stocker et al., (2018) contains the fusulinid Ozawainella mosquensis Rauser-Chemoussova in Rauser-Chemoussova et al. (1951), which was only reported from the Beedeina Zone, just below the Upper Fusulinella-Fusulina Zone of Niikawa (1980) in the upstream of the Mizuboradani Valley.
We have been unable to find exposures of greenish-gray tuff and tuffaceous mudstone in the Beedeina Zone of the Ichinotani Formation. Thus, the Missourian conodont-bearing limestone boulders might be derived from uninvestigated exposures and sections in the valley.
In this study, we focused on the Sorayama Formation as one of the possible sources of the conodont-bearing float limestone boulders. According to Igo (1956), Tsukada and Takahashi (1998), and Tsukada et al. (1999), the Sorayama Formation is widely distributed in the Fukuji district and consists mainly of mafic to intermediate pyroclastic rocks with intercalated conglomerates, sandstones and mudstones. The conglomerates are clast-supported and contain pebble- to boulder-sized limestone clasts. The limestones contain Devonian corals Favosites and Heliolites, and Carboniferous fusulinids such as Ozawainella sp., Beedeina ichinotaniensis (Igo, 1957), Quasifusulina longissima (Möller, 1878), and Triticites sp. (Igo, 1956; Tsukada et al., 1999). In contrast, the matrix (tuffaceous mudstone and/or calcareous sandstone) of the conglomerates contains middle Permian fusulinids such as Nankinella sp., Parafusulina aff. gigantea (Deprat, 1913) and Russiella pulchra Miklukho-Maclay, 1958. Tsukada et al. (1999) suggested that the limestone clasts of the Sorayama Formation were derived from both the Devonian Fukuji and Carboniferous Ichinotani formations. The limestone boulders documented for our paper are lithologically similar to the above-mentioned limestone clasts of Carboniferous age. According to Tsukada et al. (1999), tuffaceous clastic materials which are components of the matrix of the conglomerate of the Sorayama Formation fill the chambers of some Permian fusulinids. This feature is observed in our sample T-01 (Figure 2.2). These observations enable us to consider the conglomerates of the Sorayama Formation as one of the likely sources of our Carboniferous-age limestone boulders (Figure 7).
Figure 7.
1, stylised image of a conglomerate bed of the Permian Sorayama Formation. These beds are clast-supported and contain abundant pebble- to boulder-sized subangular and subrounded clasts of sandstone, mudstone, various igneous rocks, and Devonian and Carboniferous fossiliferous limestones in a tuffaceous mudstone and/or calcareous sandstone matrix. The limestones were probably derived from the Devonian Fukuji and Carboniferous Ichinotani formations. 2, float limestone boulder as it would appear if reworked from the conglomerates of the Sorayama Formation.

Biogeography
The Missourian (Kasimovian, Late Pennsylvanian) conodont species Gondolella sublanceolata and Idiognathodus sulciferus, are commonly reported in the Pennsylvanian of the USA (Gunnell, 1933; Clark and Mosher, 1966; Barrick et al., 2004; Rosscoe and Barrick, 2009). Idiognathodus sulciferus is also known from the Moscow Basin of Russia (Goreva et al., 2009), and from Guizhou, South China (Qi et al., 2013). The similarity of the Kasimovian conodonts of Japan with those of southern China and the USA continues a trend that is also apparent from Early and Middle Pennsylvanian faunas. Ishida et al. (2012, 2013) reported Bashkirian to early Moscovian (Early to Middle Pennsylvanian) conodonts from the Ko-yama Limestone Group, Akiyoshi Belt, Okayama, Japan. The Moscovian conodont assemblage consists of Mesogondolella clarki (Koike, 1967), Idiognathoides attenuatus (Harris and Hollingsworth, 1933). Id. convexus (Ellison and Graves, 1941), Id. macer (Wirth, 1967), Idiognathodus delicatus Gunnell, 1931, Streptognathodus suberectus Dunn, 1966, Neognathodus bothrops Merrill, 1972 and N. symmetricus (Lane, 1967). Seven species (M. clarki, Id. attenuatus, Id. convexus, I. delicatus, S. suberectus, N. bothrops and N. symmetricus) from the Ko-yama Limestone are reported from the Midcontinent area of the western USA (Ellison and Graves, 1941; Dunn, 1966, 1970; Merrill, 1972; von Bitter and Merrill, 1977; Barrick et al., 2013). Mesogondolella clarki, Id. convexus, I. delicatus, S. suberectus, N. bothrops and N. symmetricus are also known from Guizhou, South China (Qi et al., 2013). Mesogondolella clarki, N. bothrops and N. symmetricus appear to be cosmopolitan species in the Moscovian (Middle Pennsylvanian).
Concluding remarks
In this study, we report the Missourian (Carboniferous) conodonts Gondolella sublanceolata Gunnell, 1933 and Idiognathodus sulciferus Gunnell, 1933 from float limestone boulders collected in the downstream of the Mizuboradani Valley, Fukuji district, central Japan. According to the conodont age, the float limestone boulders might be from the Triticites Zone of the Ichinotani Formation, or derived from boulder-sized limestone clasts in the conglomerates of the Permian Sorayama Formation.
Gondolella sublanceolata has previously been reported only from the Carboniferous of North America, and this is the first report of the species from a western Panthalassic setting. This suggests that Gondolella was a widespread pan-Panthalassic Ocean species during the Middle to Late Pennsylvanian, and thus of biostratigraphical value for international correlation.
Author contributions
T. Maekawa described conodonts collected from limestone boulders and discussed their biostratigraphic age. T. Komatsu, G. Tanaka, M. Okura, and A. Umayahara investigated the Ichinotani Formation in the Mizuboradani Valley and collected limestone boulders. T. Maekawa, C. P. Stocker and M. Okura collected microfossils from prepared residues of the boulders. T. Maekawa, T. Komatsu, M. Williams and C. P. Stocker identified the biogeographical significance of the Pennsylvanian conodonts. All authors contributed to writing the paper.
Acknowledgements
We are grateful to Tatsuya Hayashi (Kyushu University) and Naoki Ikegami (Mifune Dinosaur Museum) for taking SEM images of conodonts. We thank Keisuke Ishida (Tokushima University), an anonymous reviewer and the associate editor Yasunari Shigeta (National Museum of Nature and Science) for valuable comments on the draft. Part of this study was supported by the JSPS-VAST Joint Research Program, Grants-in-Aid from the Japan Society for the Promotion of Science (16K05593 to Komatsu). Mark Williams and Christopher P. Stocker thank the Leverhulme Trust (International Network Grant IN-2014-025 ‘Assembling the Early Palaeozoic terranes of Japan’).