名古屋大学博物館報告
Bull. Nagoya Univ. Museum
No. 21, 57–125, 2005

Tabulate corals from the Devonian Fukuji Formation,
Hida Gaien belt, central Japan— Part 1 —Kazuhiro Tsukada
The Nagoya University Museum, Chikusa-ku, Nagoya 464-8601, Japan

Abstract
The Fukuji area, in the Hida Gaien belt (HGB), is one of the most significant areas for Paleozoic
stratigraphy in Japan. The Devonian Fukuji Formation of this area yield abundant, well-preserved coralfossils, however, the biostratigraphy and paleobiogeographical significance of this fauna still remain
unclear and precise details of the Devonian situation of the HGB have yet to be revealed. In order tounderstand the Paleozoic history and geological framework of the HGB better, the stratigraphy andfaunal assemblages of this formation have been reexamined.
Limestone and muddy limestone samples amounting to some 420 kg from seven horizons (Locs. 1 to7) along the Ichinotani Valley, Fukuji area, Gifu Prefecture, central Japan have been studied here. The
tabulate coral fauna from the Loc. 1 (N36˚13.17’ E137˚31.37’) comprises the following 17 species:
Favosites goldfussi, F. flexuosus, F. sp. A, Squameopora hidensis, S. zhanwaensis fukujiensis subsp.
nov., S. cf. zhanwaensis, Sapporipora kamitakaraensis sp. nov., Sa. karatanioum sp. nov., Heliolites
wenxianicus, H. cf. gemina, H. ichinotaniensis sp. nov., H. ? sp., Helioplasma takayamaensis sp. nov.,
Pseudoplasmopora okuhidaensis sp. nov., P. cf. arguta, Striatopora sp., Gertholites ? sp.
This paper describes these tabulate corals from the Loc. 1.
Introduction
The Fukuji area, in the Hida Gaien belt (HGB), is one of the most significant areas for Paleozoic
stratigraphy in Japan. It was first defined as a complex zone between the Hida and the Mino belts by
Kamei (1955a) and was redefined as a belt composed of fault-bound blocks of Paleozoic-Mesozoic shelf
facies rocks (Tsukada et al., 2004). In recent work, the Paleozoic shelf facies rocks of the HGB havebeen divided into the “Fukuji type” and the “Moribu type” based on their lithostratigraphy (Tsukada,
1999; Tsukada, 2003; Tsukada et al., 2004).
Devonian strata of the “Fukuji type” yield abundant, well-preserved fossils which have been studied
by many geologists and paleontologists (e.g. Fujimoto et al., 1962; Hamada, 1959a, b; Kamei, 1952,
1955b; Kamei and Igo, 1955; Kobayashi and Igo, 1956; Koizumi and Kakegawa, 1970; Kozu, 1911;
Kuwano, 1986, 1987; Niikawa, 1980; Niko, 2001; Ohno, 1977; Okazaki, 1974; Research Group for the
Paleozoic of Fukuji, 1973). Nevertheless, the paleobiogeographical significance of this fauna still remains unclear and precise details of the Paleozoic tectonic history of the HGB have yet to be revealed.
In order to understand the Paleozoic history and geological framework of East Asia better, the stratigraphy and faunal assemblages of the Devonian Fukuji Formation have been reexamined in the Fukuji
area of Gifu Prefecture, central Japan. This paper describes the tabulate corals from the Upper Memberof that formation.
Geological framework
Paleozoic rocks in the Fukuji area are divided into the following five formations (Text-Fig. 1.): (1)Lower to Middle (?) Devonian Yoshiki Formation, (2) Lower to Middle (?) Devonian Fukuji Formation,— 57 —

b

a

137

E

138

E

Sea of
Japan
37

N

66

ToyamaGamatagawa River

70
Takaharagawa
River

Takayama

70

80

60

Hida Gaien
belt
36

N

75

70

56
80

Mt. Sorayama

75

Ichinotani Valley
80

70

FukujiFig. 2.
Osobudani
Valley65

45

Mizuyagadani
ValleyLimestone

1 km

Undividedstrata80

Tuffaceous clastic rocks
and clastic rocks
Mafic volcanic rocks
Serpentinite

Quaternary
volcaniclastic rocks
Lower Cretaceous
Tedori Group
Funatsu Granite
Rocks ofthe Mino belt
Kashiate Formation(age-unknown )Middle PermianSorayama Formation
Lower to Middle PermianMizuyagadani Formation
Carboniferous
Ichinotani Formation
Devonian
Fukuji Formation
Devonian
Yoshiki Formation

Mafic intrusive rocks
Ordovician to Silurian
Hitoegane Formation
Ordovician ? Iwatsubodani
Formation"Imami greenstones"(age-unknown )Gamata Crystalline Schists(age-unknown)

Strike and dip of tilted strata(tops unknown)
Strike of vertical strata (tops unknown)
Strike and dip of normal strata
Strike of vertical strata
Strike and dip of overturned strata
Fault

Text-Fig. 1. Index (a) and geological (b) maps of the Fukuji area (after Tsukada and Takahashi, 2000).

(3) upper Lower to Upper Carboniferous Ichinotani Formation, (4) Lower Permian Mizuyagadani Formation, and (5) Lower Permian Sorayama Formation. The Yoshiki Formation is overlain by the Fukuji
Formation (Igo, 1990). Although the Fukuji, Ichinotani and Mizuyagadani Formations are in fault contact with each other, they are likely to form a conformable or unconformable succession primarily because the Mizuyagadani and Sorayama Formations include many limestone clasts that might be derived
from the Fukuji and Ichinotani Formations. The Sorayama Formation conformably overlies the
Mizuyagadani Formation (Tsukada et al., 1999; Tsukada and Takahashi, 2000).
The Fukuji Formation, which consists largely of massive limestone with felsic tuff intercalations in
its middle horizon, is well exposed along the Ichinotani Valley in the Fukuji area (Text-Fig. 1). This
formation is divided into the Lower, Middle and Upper Members in ascending order (Kamei, 1952;
1955b). The Lower and Upper Members, composed mostly of white, black and light to dark grey limestone, is particularly rich in well-preserved fossils such as corals, stromatoporids, brachiopods, trilobites, bivalves, ostracods, conodonts and others (e.g. Hamada, 1959a, b; Kamei, 1952, 1955b; Kamei
and Igo, 1955; Kobayashi and Igo, 1956; Koizumi and Kakegawa, 1970; Kuwano, 1986, 1987; Niikawa,
1980; Ohno, 1977; Okazaki, 1974; Research Group for the Paleozoic of Fukuji, 1973). The limestone is— 58 —

partly well-bedded with muddy limestone or mudstone. Much of the grey limestone is Folk’s (1959)
micrite and biomicrite. This grey limestone is rich in corals and stromatoporids which form the main
allochemical constituent. The Middle member is composed of alternating beds of black mudstone, darkgrey muddy limestone and pale green felsic tuff.
Five coral zones are proposed within this formation on the basis of the biostratigraphy of the genus
Favosites (Kamei, 1955b). The formation trends east and dips steeply north along the Ichinotani Valley.
Limestone and muddy limestone samples amounting to some 420 kg from seven horizons (Locs. 1 to 7)
along the Ichinotani Valley have been studied here (Text-Fig. 2). This paper describes the tabulate corals
from Loc. 1 (N36°13.17′ E137°31.37′, Text-Figs. 2 and 3).

Loc. 7

Loc. 6

Loc. 5N

Loc. 4
Loc. 3Loc. 2

Loc. 1

10 m

Fukuji
Formation
YoshikiFormation

N60 E 60 N

Text-Fig. 2. Route map along the Ichinotani Valley. See Text-Fig. 1 for the locality. ss: sandstone, ms: mudstone, ls:
limestone, alt: alternation.
— 59 —Text-Fig. 3. Photographs showing the field occurrence of Loc. 1. Most samples at this locality werefrom float stones derived from the immediately neighbouring cliff. (a) Float stones at
Loc. 1. (b) The cliff beside Loc. 1. The upper part of the cliff had collapsed to provide
the source of the float stones.

The tabulate coral fauna from the Loc. 1, which can be assigned to upper part of the bed 9 (F2b
subzone) of Kamei (1955b), comprises the following 17 species:
Favositidae:
Favosites goldfussi d’Orbigny 1850
Favosites flexuosus Kamei, 1955b
Favosites sp. A
Squameopora hidensis (Kamei, 1955b)
Squameopora zhanwaensis fukujiensis subsp. nov.
Squameopora cf. zhanwaensis Lin and Huang, 1987
Sapporipora kamitakaraensis sp. nov.
Sapporipora karatanioum sp. nov.
Heliolitidae:
Heliolites wenxianicus Zhang, 1981
Heliolites cf. gemina (Tchi, 1976)
Heliolites ichinotaniensis sp. nov.
Heliolites ? sp.
Helioplasma takayamaensis sp. nov.
Pseudoplasmopora okuhidaensis sp. nov.
Pseudoplasmopora cf. arguta Bondarenko, 1963
Pachyporidae:
Striatopora sp.
Gertholites ? sp.

— 60 —

SYSTEMATIC DESCRIPTION
Order FAVOSITIDA Wedekind, 1937
Suborder FAVOSITINA Wedekind, 1937
Superfamily FAVOSITOIDE Dana, 1846
Family FAVOSITIDAE Dana, 1846
Definition. Corallum cerioid, hemispherical, nodular, rarely branching or tollinoid, with prismaticcorallites; calices perpendicular to distal surface; walls thin, with pores arranged in regular longitudinalrows; septa short, equal, with spines in longitudinal rows or squamulae; tabulae usually complete, perpendicular to inner wall surface, rarely incomplete, concave or convex.
Remarks. This family comprises four subfamilies, Favositinae, Paleofavositinae, Pachyfavositinae
and Emmonsinae (Hill, 1981). Young and Elias (1995, p. 77) considered the position and form of muralpores in the Favositidae to be very variable and therefore not a character to be used to distinguish
Favositinae from Paleofavositinae.
Subfamily FAVOSITINAE Dana, 1846
Definition. Favositidae with septa represented by septal spines; longitudinally arranged rows of muralpores on wall face.
Genus FAVOSITES Lamarck, 1816
Type species. Favosites gothlandicus Milne-Edwards and Haime, 1850, p. 256.
Diagnosis (after Hill, 1981, p. F541). Corallum cerioid, tabular, massive; corallites prismatic, thinwalled; septa represented by longitudinal rows of spines or absent; tabulae complete, subhorizontal;
mural pores on corallite faces, in one or several longitudinal rows.
Favosites goldfussi d’Orbigny, 1850 Plate 1, Figs. 1-6
v.1936 Favosites goldfussi; Jones, p. 19, Pl. 2, Figs 8-10.

Reg. No.
Samples: NUM-Fz-010811-21 and NUM-Fz-010811-34. Two samples.
Specimens: NUM-Fz-010811-21-1 and NUM-Fz-010811-34-9. Two specimens.
Thin sections: Seven (010811-21-1-1, 010811-21-1-2, 010811-21-1-3, 010811-21-1-4, 010811-211-5, 010811-21-1-6 and 010811-21-1-7) and one (010811-34-9-1) thin sections cut from NUM-Fz010811-21-1 and NUM-Fz-010811-34-9 respectively.
Material, horizons and localities
All specimens were from the Upper Member of the Fukuji Formation along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Description
Specimens mostly embedded in limestone matrix. Corallum cerioid, tabular, hemispherical, pyriform, ca. 22mm wide and 23-32mm high; corallites prismatic and polygonal (3- to 8-sided), fanning out
from basal portion to lateral surface of corallum at obtuse angles and to top of corallum mostly at rightangles. Calice deep, maximum depth 5.2mm. Corallites bimodal in size are randomly distributed incross section. Immature (3- and 4-sided) width 0.5-1.24mm (0.785mm in average) and mature (5- to 8sided) width 1.0-3.0mm (1.95mm in average). Walls straight, moderately thick (0.1-0.2mm). Each corallite
in contact with thin (generally less than 0.02mm), distinct median suture. Several longitudinal rows of
slender or thick septal spines up to 0.67mm long developed especially in the mud-filled part wheretabulae distantly spaced. Spines horizontal, pointing upwards or downwards, 4-7 per 3mm. Squamulaeabsent. Mural pores circular or ellipsoidal, large (0.17-0.38mm in diameter) with very thin pore plate— 61 —

(less than 0.01mm thick). Pores arranged longitudinally in 1-2 rows on the face, 0.62-1.6mm apart.
Tabulae complete, generally planar, rarely concave or wavy, perpendicular to inner surface of corallite
wall, thin (less than 0.02mm), 3-8 per 5mm.
Remarks
The specimens here, characterized by large corallites (1.0 to 3.0mm in diameter), distantly spaced,thin tabulae (3-8 per 5mm) and numerous thick or slender septal spines (4-6 per 3mm), are identical to
Favosites goldfussi (including F. eifelensis Nicolson) from Auburg, Gerolstein, Eifel, Germany, British
Museum (Natural History) collection Reg. Nos. R26565 and R26568, described by Jones (1936). F.
eifelensis is considered to be a young form of F. goldfussi (Jones, 1936). F. goldfussi in the BM (NH)collection has large corallites (the average diameter of ten 5- to 8-sided corallites is 2.0mm), thin tabulae
spaced at 6-7 per 5mm, and thick or slender septal spines spaced longitudinally at 4-7 per 3mm. In
addition, other characters of these BM (NH) specimens such as moderately thick walls (0.08-0.17mm)
with a thin distinct median suture, circular or ellipsoidal mural pores 0.17-0.26mm in diameter arranged
in one or two rows on the face, 0.75-1.2mm apart longitudinally, are also identical to those of the present
specimens.
Favosites flexuosus Kamei, 1955b

Plate 2, Figs. 1-3

1934 Favosites cf. gotlandicus Lamarck; Ozaki in Shimizu et al., p. 69, Pl. XII, Figs. 1-6.
1955b Favosites flexuosus Kamei; Kamei, p. 47, Pl. III, Fig. 2.

Reg. No.
Samples: NUM-Fz-010809-c, NUM-Fz-010809-e and NUM-Fz-010811-15. Three samples.
Specimens: NUM-Fz-010809-c-1, NUM-Fz-010809-e-1 and NUM-Fz-010811-15-1. Three specimens.
Thin sections: Four (010809-c-1-1, 010809-c-1-2, 010809-c-1-3 and 010809-c-1-4), four (010809e-1-1, 010809-e-1-2, 010809-e-1-3 and 010809-e-1-4), and two (010811-15-1-1 and 010811-15-1-2)
thin sections cut from NUM-Fz-010809-c-1, NUM-Fz-010809-e-1 and NUM-Fz-010811-15-1, respectively.
Material, horizons and localities
NUM-Fz-010811-15 was from the Upper Member of the Fukuji Formation, along the Ichinotani
Valley, Takayama City, Gifu Prefecture, central Japan (Loc.1). Two specimens, NUM-Fz-010809-c and
NUM-Fz-010809-e, were from sub-rounded pebbles at Loc. 1.
Diagnosis (after Kamei, 1955b)
Species of Favosites with large corallites, strongly wrinkled wall, distantly spaced thin tabulae, septal spine absent.
Description
Specimens mostly embedded in limestone matrix. Corallum cerioid; massive, hemispherical, morethan 23mm wide and 55mm high; corallites prismatic and polygonal (3- to 8-sided), fanning out frombasal portion to surface of corallum at obtuse or right angles. Calice shallow or deep (0.05-3.5mm indepth). Corallites bimodal in size are randomly distributed in cross section. Mature (5-8-sided) width
generally 2.0-3.0mm (1.5mm in min., 3.5mm in max.) and immature (3- and 4-sided) width 0.5-1.2mm(0.7mm in average). Walls thin or moderately thick (0.06-0.1mm), strongly wrinkled. Each corallite incontact with thin (0.02-0.06mm) median suture. Septal spines generally absent or rarely a row of shortspines (up to 0.14mm long). Mural pores large (0.2-0.32mm in diameter), circular with very thin poreplate (less than 0.02mm thick). Pores distantly arranged in one or two longitudinal rows. Tabulae complete, horizontal, mostly planar or slightly concave, very thin (less than 0.02mm), 5-10 per 5mm. Growth
wrinkles on corallum surface.
— 62 —

Remarks
The specimens assigned here to F. flexuosus are characterized by large corallites, a strongly wrinkled
wall, distantly spaced thin tabulae, and the absence of septal spines. The description above coincides
completely with that given for F. flexuosus by Kamei (1955b). Kamei’s description is of a specimen
from a slightly higher horizon to the one studied in this paper.
Favosites sp. A
Plate 2, Figs. 4-5; Plate 3, Figs. 1-3
Reg. No.
Sample: NUM-Fz-010809-a. One sample.
Specimen: NUM-Fz-010809-a-1. One specimen.
Thin sections: 010809-a-1-1 and 010809-a-1-2.
Material, horizons and localities
This specimen was from a sub-rounded pebble at Loc. 1, the Ichinotani Valley, Takayama City, Gifu
Prefecture, central Japan.
Description
Specimens embedded in limestone matrix. Corallum massive, cerioid, pyriform, 32mm wide andmore than 37mm high. Corallites, prismatic, polygonal (4-, 5- or 6-sided), fanning out from basal portion to lateral surface of corallum at obtuse angle and to top of corallum mostly at right angles. Calice
shallow, 0.9-1.5mm in depth. Corallites bimodal in size are randomly distributed in cross section. Immature (4-sided) width from 0.3 to 0.8mm, mature (5- or 6-sided) width 1.0-1.2 mm. Epitheca walls ca.
0.02mm, dark wall 0.03-0.1mm, straight and moderately thin (less than 0.3mm). Thin (less than 0.02mm)
median suture present. Peripheral wall thickening absent. Several longitudinal rows of very short (lessthan 0.1mm) thick, blunt septal spines developed in peripheral part. Spines horizontal or pointing upwards, 6-7 per 3mm. Squamulae absent. Mural pores circular, 0.2-0.25mm in diameter, with very thin(less than 0.01mm) pore plate. Pores arranged longitudinally in 1-3 rows alternating at mid-faces or nearedges of corallites, 0.4-0.9 mm apart. Tabulae, 6-10 per 5mm, thin (less than 0.02mm), complete, generally planar but rarely concave, perpendicular to inner surface of corallite wall.
Remarks
This specimen is characterized by 1-3 longitudinal rows of large mural pores, moderately thick wallswith a distinct median suture, distantly spaced thin tabulae (6-10 per 5mm) and numerous thick, blunt
septal spines.
White and Shengwu (2004, p. 45) summarize the problems in defining species of tabulate corals.
Their simple structure and variable morphology has been pointed out by many authors (e. g. Scrutton,
1989, p. 40), and has led to a proliferation of synonymous taxa. Scrutton (1989) advocated that no new
species should be described based on unsatisfactory materials. Although the specimen Reg. No. 010809a-1 might well be a new species of the genus Favosites, an open nomenclature is used in this paper,
following Scrutton’s advice.
Genus SQUAMEOPORA Preobrazhenskiy, 1967
Type species. Favosites hidensis Kamei, 1955b
Other species. S. hidensiformis (Mironova, 1961); S. sichanensis Lin and Huang, 1987; S. zhanwaensis
Lin and Huang, 1987; S. ertangensis Tchi and Wang, 1989.
Diagnosis (after Preobrazhenskiy, 1967, p. 4). Corallum cylindrical with narrow base; corallites
prismatic and polygonal, radiating from central part and opening out at surface of corallum at right oracute angle. Walls moderately thick, peripheral wall-thickening present, surrounding visceral cavities ofcorallites. Mural pores circular, large, displaced towards costae, without pore ridge. Tabulae of conven— 63 —tional type. Septa represented by squamulae or spines in the peripheral part.
Squameopora hidensis (Kamei, 1955b)

Plate 3, Figs. 4-5; Plate 4, Figs. 1-5; Plate 5, Figs. 1-2

v.1955b Favosites hidensis Kamei; Kamei, p. 53, Pl. III, Figs. 4a-c; Pl. IV, Fig 7.
1959b Favosites hidensis Kamei; Hamada, p. 208, Pl. XVI, Figs. 1-12.
1967 Squameopora hidensis (Kamei); Preobrazhenskiy, p. 8.
1974 Striatoporella hidensis (Kamei); Mironova, p. 49.
1981 ?Squameopora hidensis (Kamei); Hill, p. F546, Figs. 359 1a, b.
1987 Squameopora sichanensis Lin and Huang; Lin and Huang, p. 212, Pl. 42, Fig. 2.

Reg. No.
Samples: NUM-Fz-010811-3, NUM-Fz-010811-18, NUM-Fz-010811-25 and NUM-Fz-010811-31.
Four samples.
Specimens: NUM-Fz-010811-3-2b, NUM-Fz-010811-18-2, NUM-Fz-010811-25-1, NUM-Fz010811-25-2, NUM-Fz-010811-25-4 and NUM-Fz-010811-31-1. Six specimens.
Thin sections: One (010811-3-2-1b), three (010811-18-2-1, 010811-18-2-2 and 010811-18-2-3), two(010811-25-1-1 and 010811-25-1-2), one (010811-25-2-1b), three (010811-25-4-1, 010811-25-4-2 and
010811-25-4-3), and four (010811-31-1-1, 010811-31-1-2, 010811-31-1-3 and 010811-31-1-4) cut from
NUM-Fz-010811-3-2, NUM-Fz-010811-18-2, NUM-Fz-010811-25-1, NUM-Fz-010811-25-2, NUMFz-010811-25-4 and NUM-Fz-010811-31-1, respectively.
Material, horizons and localities
All six specimens were from the Upper Member of the Fukuji Formation, along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Diagnosis (based on Kamei, 1955b and Hamada, 1959b)
Small corallum, thin walled with peripheral wall thickening, closely spaced, thin tabulae, 1-2 longitudinal rows of mural pores on wall face. Kamei (1955b) reported that septal spines are absent in thetype specimen, but Hamada (1959b) described longitudinal rows of septal spines at the calice.
Description
Specimens entirely embedded in limestone matrix. Corallum cerioid, small, 15-35mm wide, morethan 10mm high, pyriform or conical, circular or ellipsoidal in cross-section. Corallites prismatic andpolygonal (3- to 8-sided), rounded in the peripheral part, fanning out from central or basal portion tocorallum surface at obtuse or right angles. Calice generally shallow, 0.5-1.6mm deep. Corallites bimodal in size are randomly distributed in cross section. Immature (3-4-sided) width ranges from 0.28 to
0.75mm (0.5mm in average), mature (5-8-sided) from 0.51 to 1.6mm (1.0mm in average). Walls moderately thick (ca. 0.1mm or more), straight. Peripheral wall thickened with fine fibrous sclerenchymal
deposit of calcite (0.08-0.27mm thick) perpendicular to wall, clearly seen in cross section. Each corallite
is in contact with thin (generally less than 0.02mm) median suture. Intracalicular offsets increasing
peripherally. Several longitudinal rows of slender or thick septal spines (up to 0.4mm in length) developed at peripheral mud-filled part of corallites. Spines pointing upwards, downwards or wavy. Squamulae
generally absent but squamula-like very thick wedge-shaped spines may be present. Mural pores circular, medium or large (0.08-0.22mm in diameter) with very thin pore plate (less than 0.02mm thick).
Pores arranged in 1 or 2 longitudinal rows on face, ca. 0.35-0.77mm apart. Tabulae complete or rarely
incomplete, perpendicular or oblique to inner surface of corallite wall, planar or concave, closely spaced,
8-18 per 5mm, thin or thick (0.04-0.22mm thick).
Remarks
This species is characterized by a small corallum, walls thickening towards the periphery, closely
spaced tabulae and septal spines in the peripheral part. The specimens compare very well with Sqameopora— 64 —

hidensis in wall structure and size of corallum. Hamada (1959b) pointed out the “peripheral wall-thickening” as an important character of this species. Although Kamei (1955b) mentioned that the corallumis small in this species (about 1cm in cross section and 3cm in longitudinal section) and generallyclavate or pyriform in shape, the holotype (G.I.S.U.L 30119 in Shinshu University, Japan) is more than
7cm in cross section and globular in shape. The present specimens are similar to the paratype (G.I.S.U.L
30120 in Shinshu University, Japan) in corallum size and shape.
Squameopora zhanwaensis fukujiensis subsp. nov.
Plate 5, Figs. 3-7
Name
Derived from the hamlet of Fukuji, Takayama City, central Japan.
Reg. No.
Samples: NUM-Fz-010811-1 and NUM-Fz-010811-4. Two samples.
Specimens:
Holotype. NUM-Fz-010811-4-1 (thin sections: 010811-4-1-1 and 010811-4-1-2).
Paratype. NUM-Fz-010811-1-1, NUM-Fz-010811-1-2 and NUM-Fz-010811-1-3.
Thin sections: Two (010811-1-1-1 and 010811-1-1-2), three (010811-1-2-1, 010811-1-2-2 and 0108111-2-3) and three (010811-1-3-1, 010811-1-3-2 and 010811-1-3-3) thin sections cut from NUM-Fz-0108111-1, NUM-Fz-010811-1-2 and NUM-Fz-010811-1-3 respectively.
Material, horizons and localities
All four specimens, NUM-Fz-010811-1-1, NUM-Fz-010811-1-2, NUM-Fz-010811-1-3 and NUMFz-010811-4-1 were from the Upper Member of the Fukuji Formation along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Diagnosis
Species of Squameopora with thin-walled corallites fanning out from basal portion, tabulae distantly
spaced, septal spines in rows.
Description
Specimens entirely embedded in limestone matrix. Corallum cerioid, massive; more than 50 mm
wide and 33mm high; corallites prismatic, polygonal (3- to 8-sided), fanning out from basal portion tosurface of corallum at obtuse or right angles. Calice generally shallow, 0.1-1.3mm in depth. Corallites
bimodal in size are randomly distributed in cross section. Immature (3- and 4-sided) width 0.4 to 1.1mm(0.67mm in average), mature (5- to 8-sided) width 0.8 to 2.0mm (1.41mm in average). Walls thin (lessthan 0.1mm), straight or rarely wrinkled in peripheral part. Each corallite in contact with thin (generally
less than 0.02mm) median suture. Sclerenchymal fine fibrous calcite deposit perpendicular to wall issometimes observed in cross section. Several longitudinal rows of slender or thick septal spines (up to
0.35mm long) pointing upwards are particularly developed in peripheral, mud-filled part of corallites.
Wedge-shaped spines are occasionally seen in cross section. Mural pores circular, medium or large (0.10.26mm in diameter) with very thin pore plate (less than 0.02mm thick). Pores arranged in 1 or 2 longitudinal rows on face or rarely at wall junction, spaced ca. 0.4-1.25mm apart. Tabulae normally complete, planar and perpendicular to inner surface of corallite wall, rarely incomplete, concave and obliqueat peripheral part, normally thin (less than 0.07mm), closely spaced (8-14 per 5mm).
Remarks
The specimens here are clearly different from S. hidensis (Kamei, 1955b; Hamada, 1959b), S.
hidensiformis (Mironova, 1961), S. sichuanensis Lin and Huang, 1987 and S. ertangensis Tchi andWang, 1989 in the size and shape of the corallum. In addition, the peripheral wall-thickening which
Hamada (1959b) noted as an important character of S. hidensis is uncommon in these specimens. The
present specimens compare very well with S. zhanwaensis described and figured by Lin and Huang— 65 —

(1987) from the Lower Member of the Yanglugou Formation, Yanglugou, west Qinling Mountains, China,
in having a large corallum and corallites which fan out from the basal portion. They are, however,distinguished from that species in having more distantly spaced tabulae. Lin and Huang (1987) described that S. zhanwaensis has tabulae of 14-16 in 5mm or sometimes 24-26 in. 5mm.
Squameopora cf. zhanwaensis Lin and Huang, 1987
Plate 6, Figs. 1-5; Plate 7, Figs. 1-7; Plate 8, Figs. 1-2
cf.1987 Squameopora zhanwaensis Lin and Huang; Lin and Huang, p. 212, Pl. 43, Figs. 2-4.

Reg. No.
Samples: NUM-Fz-010811-10, NUM-Fz-010811-15, NUM-Fz-010811-16, NUM-Fz-010811-17,
NUM-Fz-010811-24, NUM-Fz-010811-26, NUM-Fz-010811-27, NUM-Fz-010811-33 and NUM-Fz010811-34. Nine samples.
Specimens: NUM-Fz-010811-10-1, NUM-Fz-010811-15-2, NUM-Fz-010811-16-1, NUM-Fz010811-17-1, NUM-Fz-010811-17-2, NUM-Fz-010811-17-3, NUM-Fz-010811-17-4, NUM-Fz-01081124-4, NUM-Fz-010811-26-2, NUM-Fz-010811-27-2, NUM-Fz-010811-33-1, NUM-Fz-010811-34-8
and NUM-Fz-010811-34-13. Thirteen specimens.
Thin sections: Three (010811-10-1-1, 010811-10-1-2 and 010811-10-1-3), one (010811-15-2-1), four
(010811-16-1-1, 010811-16-1-2, 010811-16-1-3 and 010811-16-1-4), one (010811-17-1-5b), four
(010811-17-2-1, 010811-17-2-2, 010811-17-2-3 and 010811-17-2-4), one (010811-17-3-1), two (01081117-4-1 and 010811-17-4-2), two (010811-24-4-1 and 010811-24-4-2), three (010811-26-2-1, 01081126-2-2 and 010811-26-2-3), three (010811-27-2-1, 010811-27-2-2 and 010811-27-2-3), three (01081133-1-1, 010811-33-1-2 and 010811-33-1-3), two (010811-34-8-1 and 010811-34-8-2), and two (01081134-13-1 and 010811-34-13-2) thin sections cut from NUM-Fz-010811-10-1, NUM-Fz-010811-15-2,
NUM-Fz-010811-16-1, NUM-Fz-010811-17-1, NUM-Fz-010811-17-2, NUM-Fz-010811-17-3, NUMFz-010811-17-4, NUM-Fz-010811-24-4, NUM-Fz-010811-26-2, NUM-Fz-010811-27-2, NUM-Fz010811-33-1, NUM-Fz-010811-34-8 and NUM-Fz-010811-34-13 respectively.
Material, horizons and localities
All specimens were from the Upper Member of the Fukuji Formation along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Diagnosis
Corallum comparatively large; corallites fanning out from basal portion; wall usually thin; tabulae
planar, closely spaced; septal spines numerous.
Description
Specimens entirely embedded in limestone matrix. Corallum cerioid, massive; conical, pyriform, 1873mm wide, 20-50mm high; corallites prismatic and polygonal (3- to 8-sided), radiate or fanning out
from basal or central portion to lateral surface of corallum at obtuse to acute angle and to top of corallum
mostly at right angles. Calice generally shallow, maximum depth 6.5mm. Corallites bimodal in size are
randomly distributed in cross section. Immature (3- and 4-sided) width 0.2-1.18mm (0.6mm in average)
and mature (5- to 8-sided) width 0.8-2.22mm (1.27mm in average). Walls normally thin (less than 0.1mm),
rarely thickened (0.22-0.3mm) by fine fibrous sclerenchymal calcite deposit, straight or rarely wrinkled
in peripheral part. Each corallite in contact with thin (normally less than 0.02mm) median suture. Several longitudinal rows of slender or thick septal spines (up to 1.2mm long) developed particularly in
mud-filled part where tabulae are distantly spaced. Spines horizontal, upward or downward pointing orwavy, 7-11 per 3mm. Squamulae absent though very thick squamula-like wedge spines occasionallyseen in cross section. Mural pores circular or ellipsoidal, medium or large (0.08-0.3mm in diameter)with very thin pore plate (less than 0.01mm thick). Pores arranged longitudinally in 1 or 2 rows alternat— 66 —

ing on face or rarely at wall junction, spaced ca. 0.11-1.0mm (usually 0.3-0.5mm) apart. Tabulae complete, generally planar, rarely concave or sagging, perpendicular or oblique to inner surface of corallite
wall, thin to thick (0.02-0.2mm), usually 8-16 (rarely up to 20) per 5mm.
Remarks
The specimens assigned here to S. cf. zhanwaensis compare well with figures of the holotypes in Lin
and Huang (1987) from the Lower Member of the Yanglugou Formation, Yanglugou, west Qinling Mountains, China, except that the Japanese specimens have large, distantly spaced mural pores.
Genus SAPPORIPORA Ozaki, 1934
Type species. Sapporipora favositoides Ozaki, 1934 (in Shimizu et al., 1934, p. 74)
Other species. Favosites (S.) favositoides var. biserialis Zhizhina and Smirnova, 1957; Favosites (S.)
deversor Yanet; S. tarbagataica Barskaja in Bogdanov ed. (1963); S. baoxingensis Kim in Xinan geological institute ed. (1978); S. jingmenensis Chou in Xian and Chow (1978); S. qinlingnensis Zhang,
1981; S. liuhuiensis, Zhou, 1986; S. gannanensis Tchi, 1987; S. shaanxiensis Lin in Xian Institute ofGeological and Mineral Resources, Geological and Mineral Resources Section ed. (1982); S. xinjiangensis
Wang in Wang et al. (2001); S. kamitakaraensis sp. nov.; S. karataniorum sp. nov.
Diagnosis (after Shimizu et al., 1934, p. F603). Corallum massive with small corallites. Corallites
polygonal, prismatic, united, their walls perforated by single row of very large pores. Walls thick. Tabulae complete, horizontal, spaced at regular intervals. Number of corallites often increased at junction offour walls.
Remarks. Lang et al. (1940) suggested that this genus is probably synonymous with the genus
Favosites. Sapporipora can, however, be distinguished from Favosites by its small slender corallites andmuch larger mural pores.
Sapporipora kamitakaraensis sp. nov.
Plate 8, Figs. 3-5; Plate 9, Figs. 1-5
Name
After Kamitakara Village, now part of Takayama City, central Japan.
Reg. No.
Samples: NUM-Fz-010811-5, NUM-Fz-010811-12, NUM-Fz-010811-17, NUM-Fz-010811-24,
NUM-Fz-010811-25, NUM-Fz-010811-26 and NUM-Fz-010811-31. Seven samples.
Specimens:
Holotype. NUM-Fz-010811-17-1 (thin sections: 010811-17-1-1, 010811-17-1-2, 010811-17-1-3,
010811-17-1-4 and 010811-17-1-5).
Paratype. NUM-Fz-010811-5-1, NUM-Fz-010811-12-1, NUM-Fz-010811-17-2b, NUM-Fz-01081124-7, NUM-Fz-010811-25-2, NUM-Fz-010811-26-3, NUM-Fz-010811-31-2 and NUM-Fz-010811-314.
Thin sections: Two (010811-5-1-1 and 010811-5-1-2), one (010811-12-1-1), one (010811-17-2-4b),
three (010811-25-2-1, 010811-25-2-2 and 010811-25-2-3), two (010811-24-7-1 and 010811-24-7-2),
two (010811-26-3-1 and 010811-26-3-2), one (010811-31-2-1) and three (010811-31-4-1, 010811-314-2 and 010811-31-4-3) thin sections cut from NUM-Fz-010811-5-1, NUM-Fz-010811-12-1, NUMFz-010811-17-2b, NUM-Fz-010811-25-2, NUM-Fz-010811-24-7, NUM-Fz-010811-26-3, NUM-Fz010811-31-2 and NUM-Fz-010811-31-4 respectively.
Material, horizons and localities
All specimens were from the Upper Member of the Fukuji Formation, along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Diagnosis— 67 —

A species of Sapporipora with small, slender corallites, distinctive median suture, distantly spaced
planar tabulae and row of large mural pores.
Description
Specimens entirely embedded in limestone matrix. Corallum massive, cerioid; spherical or hemispherical, circular or ellipsoidal in cross section; small, 17-50mm wide by 14mm to more than 40mm
high. Corallites very slender, prismatic and polygonal (3- to 8-sided), sub-matched in size in cross section, radiate from basal or central portion to surface of corallum and opening out at obtuse or rightangles. Calice deep, (up to 3.1mm). Immature (3- and 4-sided) width 0.3-0.82mm (0.6mm in average),mature (5- to 8-sided) 0.6-1.33mm (0.9mm in average). Wall straight, thin (ca. 0.1mm), rarely thickened
and rounded by fine, sclerenchymal calcite deposit, peripheral wall thickening absent. Each corallite incontact with distinctive median suture (0.01-0.06mm thick). Septal spine absent or rarely present inperipheral part, maximum length 0.26mm. Spines feeble, slender or thick. Squamulae absent. Mural
pores circular, 0.15-0.78mm in diameter (usually 0.15-0.45mm) with very thin pore plate (less than
0.01mm). Each wall face with one longitudinal row of large pores 0.5-1.33mm apart. Tabulae complete,
generally planar, rarely concave or sagging, perpendicular or oblique to inner surface of corallite wall,very thin (less than 0.02mm), distantly spaced (4-10 per 5mm).
Remarks
The specimens assigned here to S. kamitakaraensis are characterized by their small, slender corallites,
Table 1. Main morphological features of genus Sapporipora (after Shimizu et al., 1934; Zhizhina and Smirnova, 1958;
Bogdanov ed., 1963; Xinan Geological Institute ed., 1978; Xian and Chow, 1978; Zhang, 1981; Zhou, 1986; Tchi,
1987; Xian Institute of Geological and Mineral Resources, Geological and Mineral Resources Section ed., 1982;
Wang et al., 2001)
S. kamitakarensis

S. karataniorum

S. favositoides
Ozaki

S. deversor Yanet

S. tarbagataica
Barskaja

S. baoxingensis
Kimcorallites

size in diam.

0.6-1.33mm

0.75-1.55mm

small

0.5-1.0mm

0.36-0.47mm

0.4-0.8mm

wallthicknessless than 0.2mm

less than 0.2mm

thick

0.05-0.1mm

0.04-0.11mm

0.02-0.03mm

size in diam.

0.15-0.45mm

0.15-0.42mm

very large

mural porespacetabulae

0.15×0.20.2×0.3mm

0.5-1.33mm

0.44-1.15mm

ca. 0.3mm

0.3-0.4mm

arrangement

1 row, mid faces

1 row, mid faces

1 row, mid faces

1 row, mid faces

shapeplanar or curved

planar or curved

planar or curved

spacing

4-10 in 5mm

7-10 in 5mm

more 9 in 3mmrare or absent

well-developed

absentS. jingmenensis
Chow

S. qinlingnensis
Zhang

S. liuhuiensis
Zhou

S. gannanensis
Tchi

S. shaanxiensis
Lin

S. xinjiangensis
Wang

septal spine

slightly concave
3.33-10 in 5mm

8-10 in 3mm

6-12 in 5mm
absentcorallites

size in diam.

0.8-1.0mm

generally
0.5-0.6mm

0.4-1.0mm

generally
0.5-0.6mm

generally
0.7-1.05mm

generally
0.7-1.0mm

wall

thickness

0.1-0.25mm

0.03-0.04mm(rarely 0.1-0.15mm)

0.04-0.2mm

0.05-0.1mm

0.03-0.1mm

0.1-0.3mm

mural poresize in diam.

0.2x0.30.23×0.35mm

0.3×0.15-0.4×0.3

0.2-0.3mm or
0.2×0.3-0.3×0.4mm

0.2-0.3×
0.15-0.2mm

0.15mm

0.2mm

space

0.6-1.6mm

0.2-0.5mm(commonly
0.3-0.4mm)

0.7-0.8mm

0.2-0.7mm

0.3mm

arrangement

1 row, mid faces

generally 1 row,
mid or edge of faces

1 row, midfaces or angles

2 row, midfaces or angles

1 row, mid faces

shapeplanar or curved

planarplanar or curved

planar or curved

saggedspacing

5-8 in 5mm(rarely10-12.5
in 5mm)5 in 5mm

8-10 in 5mm

7.1-25 in 6mm

12-13 in 5mm

13-15 in 5mm

absentabsent or rareabsentrare

absentwell-developed

tabulaeseptal spine— 68 —

1 row

the distinctive median suture, the distantly spaced tabulae and the very large mural pores. Ten species of
the genus Sapporipora were compared with the specimens examined in this paper: S. favositoides Ozaki(in Shimizu et al., 1934, p. 74), S. deversor Yanet (in Khodalevich et al., 1959, p. 101), S. tarbagataica
Barskaja (in Bogdanov ed., 1963, p. 132), S. baoxingensis Kim (in Xinan Geological Institute ed., 1978,
p. 66), S. jingmenensis Chow (in Xian and Chow, 1978, p. 182), S. qinlingnensis Zhang (in Zhang, 1981,
p. 91), S. liuhuiensis Zhou (in Zhou, 1986, p. 83), S. gannanensis Tchi (in Tchi, 1987, p. 250), S.
shaanxiensis Lin (in Xian Institute of Geological and Mineral Resources, Geological and Mineral Resources Section ed, 1982, p. 64), S. xinjiangensis Wang (in Wang et al., 2001, p. 73) (Table 1).
S. kamitakaraensis is clearly distinguished from S. favositoides and S. tarbagataica by its distantly
spaced tabulae, from S. baoxingensis by its thicker walls, and from S. xinjiangensis in having planar
tabulae. In some ways, S. kamitakaraensis is similar to S. shaanxiensis. It differs, however in having
much larger mural pores and distantly spaced tabulae. S. deversor has much thicker tabulae than S.
kamitakaraensis. S. liuhuiensis and S. qinlingnensis are distinguishable from S. kamitakaraensis by theirregular shape of its corallites. S. jingmenensis is very similar to S. kamitakaraensis but the latter differsin having ellipsoidal mural pores.
Sapporipora karataniorum sp. nov. Plate 9, Fig. 6; Plate 10, Figs. 1-6
Name
The name honours Mr. and Mrs. Karatani who have strongly supported this study.
Reg. No.
Samples: NUM-Fz-010811-25, NUM-Fz-010811-33 and NUM-Fz-010811-34-5. Three samples.
Specimens:
Holotype. NUM-Fz-010811-33-2 (thin sections: 010811-33-2-1 and 010811-33-2-2).
Paratype. NUM-Fz-010811-25-3, NUM-Fz-010811-34-1 and NUM-Fz-010811-34-5a.
Thin sections: One (010811-25-3-1), one (010811-34-1-1), and two (010811-34-5-1a and 01081134-5-2a) thin sections cut from NUM-Fz-010811-25-3, NUM-Fz-010811-34-1-1 and NUM-Fz-01081134-5a, respectively.
Material, horizons and localities
All specimens were from the Upper Member of the Fukuji Formation, along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Diagnosis
Corallites small and slender; wall generally thin, with distinctive median suture, generally straight
but wrinkled in the peripheral part; tabulae planar, distantly spaced; mural pores large, in one row onface of corallites; septal spines numerous at calice.
Description
The specimens are mostly embedded in a limestone matrix. The corallum is massive, cerioid, spherical, hemispherical, pyriform or conical, circular or ellipsoidal in cross-section, small (17-25mm wide
and 12-25mm high). Corallites slender, prismatic and polygonal (3- to 8-sided), sub-matched in size incross section, radiate from basal or central portion to surface of corallum and opening out at obtuse or
right angles. Calice deep (up to 8mm in depth). Immature corallites (3- and 4-sided) width 0.4 - 0.66mm,
mature (5- to 8-sided) width 0.75 - 1.55mm. Walls straight but slightly wavy in peripheral part, generallythin (less than 0.1mm), rarely thick (0.1-0.15mm). Walls rarely thickened and rounded by finesclerenchymal calcite deposit. Each corallite in contact with distinctive median suture (0.01-0.04mm).
Septal spines common in peripheral part (up to 0.42mm long). Spines slender or thick, in longitudinalrows, 0.25-0.53mm apart. Squamulae absent. Mural pores very large (0.15-0.42mm in diameter) withvery thin pore plate (less than 0.02mm thick). Each wall face with one longitudinal row of pores 0.44— 69 —

1.15mm apart. Tabulae complete, generally planar but rarely concave or sagging, perpendicular or oblique to inner surface of corallite wall, thin (less than 0.1mm thick), distantly spaced 7-10 per 5mm.
Remarks
The small and slender corallites with distantly spaced tabulae and very large mural pores suggest that
this species belongs to the genus Sapporipora. The specimens are similar to S. kamitakaraensis sp. nov.,
but they differ in having wrinkled walls and numerous septal spines in the peripheral part of the corallum(Table 1). S. xinjiangensis, the only previously known species, has long septal spines (Wang et al., 2001)
and is clearly distinguished from S. karataniorum by having deeply sagging tabulae (Table 1).
Order HELIOLITIDA Frech, 1897
Suborder HELIOLITINA Frech, 1897
Superfamily HELIOLITICAE Lindström, 1876
Family HELIOLITIDAE Lindström, 1876
Definition (after Hill, 1981). Corallum massive with cylindrical tabularia surrounded by coenenchymeof prismatic tubules with complete or incomplete diaphragms; tabularia with 12 septal laminae commonly with spinose axial edges, or with 12 longitudinal radial rows of discrete spines, or aseptate;tabulae complete, commonly flat and horizontal, but strongly convex or conical in Saaremolites.
Remarks. This family comprises nine genera: Heliolites, Ningqiangolites, Saaremolites, Dnestrites,
Helioplasmolites, Helioplasma, ?Paeckelmannopora, Okopites and Pachycanalicula (Hill, 1981).
Heliolites and the last four genera being reported from the Lower to Middle Devonian (Hill, 1981).
Flügel (1956, p. 75) considered the type species of Pachycanalicula to be a subspecies of Heliolites(Heliolites) porosus (Goldfuss).
Genus HELIOLITES Dana, 1846
Type species. Astraea porosa Goldfuss, 1826, p. 64.
Diagnosis (after Hill, 1981, p. F603). Corallum variable in form, massive; tabularia cylindrical, rounded
to polygonal or substellate in transverse section; septa developed with 12 axially spinose laminae orspines; tabulae horizontal, coenenchyme of prismatic tubules with transverse diaphragms.
Heliolites wenxianicus Zhang, 1981

Plate 11, Figs. 1-5

1981 Heliolites wenxianicus Zhang; Zhang, p. 175-176, Pl. 70, Fig. 2.
1987 Heliolites wenxianicus Zhang; Tchi, p. 276, Pl. 80, Fig. 6.

Reg. No.
Samples: NUM-Fz-010809-b, NUM-Fz-010811-3, NUM-Fz-010811-4, NUM-Fz-010811-13, NUMFz-010811-25, NUM-Fz-010811-28 and NUM-Fz-010811-30. Seven samples.
Specimens: NUM-Fz-010809-b-1, NUM-Fz-010811-3-1, NUM-Fz-010811-4-1b, NUM-Fz-01081113-3, NUM-Fz-010811-13-4, NUM-Fz-010811-25-4b, NUM-Fz-010811-28-7, NUM-Fz-010811-2810 and NUM-Fz-010811-30-5. Nine specimens.
Thin sections: Eight (010809-b-1-1, 010809-b-1-2, 010809-b-1-3, 010809-b-1-4, 010809-b-1-5,010809-b-1-6, 010809-b-1-7 and 010809-b-1-8), one (010811-3-1-1), one (010811-4-1-1b), one (01081113-3-1), one (010811-13-4-1), two (010811-25-4-1b and 010811-25-4-2b), one (010811-28-7-2b), three(010811-28-10-1, 010811-28-10-2 and 010811-28-10-3), and two (010811-30-5-1 and 010811-30-5-2)
thin sections cut from NUM-Fz-010809-b-1, NUM-Fz-010811-3-1, NUM-Fz-010811-4-1, NUM-Fz010811-13-3, NUM-Fz-010811-13-4, NUM-Fz-010811-25-4, NUM-Fz-010811-28-7, NUM-Fz-01081128-10, and NUM-Fz-010811-30-5, respectively.
Material, horizons and localities— 70 —

All specimens except for NUM-Fz-010809-b-1 were from the Upper Member of the Fukuji Formation, along the Ichinotani Valley, Takayama City, Gifu Prefecture, central Japan (Loc.1). NUM-Fz-010809b-1 was from a sub-rounded pebble at Loc.1.
Description
Corallum of variable form, heliolitoid. Tabularia circular or subpolygonal in transverse section, cylindrical, straight, diameter 0.6-1.25mm; walls 0.025-0.12 thick; septal spines commonly well-developed in 12 longitudinal rows, slender, horizontal or curved upwards; tabulae complete, horizontal or
oblique, flat, convex or concave, generally very thin (less than 0.04mm), 10-15 per 5mm. Tabularia
separated by 0.2-1.42mm of coenenchymal tubules. Coenenchymal tubules 4- to 6-sided in transverse
section, 0.1-0.47mm in diameter; coenenchymal walls 0.025-0.12mm thick (usually 0.025-0.075mm
thick); diaphragms generally flat, 20-32 per 5mm.
Remarks
These specimens are identified as belonging to the genus Heliolites. 61 species of the genus Heliolites
from East Asia were compared with the specimens examined in this paper: H. anfuiensis Yu and Deng;
H. arboreus Sugiyama; H. assuetus Bondarenko; H. baichengensis Wang; H. (Pachycanalicura) cf.
barrandei (Hoernes); H. bohemicus Wentzel; H. (Paraheliolites) bulongensis (Lin et Huang); H.
compactiformis Tchi; H. daguanensis Kim; H. decipiens (McCoy); H. densitabulatus Wang; H. decipiens
var. minimus Rukhin; H. fenggangensis Yang; H. fovetabulatus Tchi; H. funingensis King; H. (Stelliporella)gemina (Tchi); H. guiyangensis Chow; H. guizhouensis Yang; H. huanghuachongensis Chow; H. insolens
Tchernychev; H. insolens dunbeiensis Dubatolov; H. interstinctus (Linne); H. interstinctus-intermedis
Wentzel; H. (Paraheliolites) jilinensis Tchi; H. lindstromi Kovalevsky; H. luorepingensis Wu; H.
luxarboreus Yang; H. magnus Yu; H. mileensis King; H. minutissimus Chow; H. multitabulatus
Bondarenko; H. obliquus Yu; H. onukii Sugiyama; H. orientalis Yu; H. (Paraheliolites) obvellatus Deng
and Cheng; H. orientalis spinosus Yang; H. paradoxicus Lin and Wang; H. paravulgaris Kim; H. porosus(Goldfuss); H. pseudobohemicus Bondarenko; H. regularis var. kuznetskiensis Tchernychev; H. repkinae
Kovalevsky; H. (Paraheliolites) salairicus Tchernychev; H. sinkiangensis Yu; H. stella King; H.
tashanensis Lin and Chow; H. tchernychevi Bondarenko; H. tomensis Tchernychev; H. uksunayensis
Mironova; H. vesicotabulatus Tchi; H. volutus Yu and Deng; H. vulgaris irregularis Tchernychev; H.
waicunensis Lin and Chow; H. wenxianicus Zhang; H. wudangensis Chow; H. yanbianensis Kim; H.
yanglugouensis Lin and Fuan; H. yassensis obliquespinus Chu; H. zhanwabeigouensis Lin and Fuan; H.
zhanwaensis Lin and Fuan; H. zoigeensis Lin and Fuan (Xian Institute of Geological and Mineral Resources, Geological and Mineral Resources Section ed, 1982; Xinjiang Geological Department, Xinjiang
Geological Institute and Xinjiang Petroleum Department eds., 1981; Nanjing Institute of Geology andMineralogy, Geology and Mineralogy Section ed., 1983; Tchi, 1980; Xiong and Gu, 1978; Wang et al.,
2001; Tchi, 1987; Xinan Geological Institute ed., 1978; Lin and Huang, 1987; Lin and Wang, 1987;
Geological Department of Inner Mongolia Autonomous Region (GDIMAR) and Dongbei Institute ofGeology (DIG) eds., 1976; Fubei Institute of Geology, Geological Department of Hubei, Geological
Department of Guandong, Geological Department of Henan, Geological Department of Hunan andGeological Department of Guangxi Autonomous Region eds., 1977; Geological Department of Yunnaned., 1974; Regional Geological Surveying Team of Hubei ed., 1994; Guizhou Geological and Paleontological Team ed, 1978; Deng, 1999; Yeh and Chow, 1995; Jin, 1984; Deng and Zheng, 2000; Sugiyama,
1940; Yu, 1962; Lin and Chow, 1977; Lin and Huang, 1986; Kamei, 1955b; Zhang, 1981).
Those assigned here to H. wenxianicus are characterized by closely spaced diaphragms. They are
similar to H. orientalis spinosus (in Guizhou Geological and Paleontological Team ed, 1978; p. 237), H.
guiyangensis (in Guizhou Geological and Paleontological Team ed, 1978; p. 237), H. repkinae (inGDIMAR and DIG eds., 1976; p. 123), H. regularis var. kuznetskiensis (in GDIMAR and DIG eds.,— 71 —

(mm)
1.0

0.9

0.8

diameterof tubule

0.7

0.6

1: H. (Stelliporella) gemina (in Deng, 1999)
2: H. orientalis spinosus (in Guizhou geological and paleontological team ed, 1978)
3: H. guiyangensis (in Guizhou geological and paleontological team ed, 1978)
4: H. wudangensis (in Guizhou geological and paleontological team ed, 1978)
5: H. minutissimus (in Guizhou geological and paleontological team ed, 1978)
6: H. luxarboreus (in Guizhou geological and paleontological team ed, 1978)
7: H. tomensis (in Guizhou geological and paleontological team ed, 1978)
8: H. porosus (in Guizhou geological and paleontological team ed, 1978)
9: H. insolens (in Geological department of Yunnan ed., 1974)
10: H. stella (in Geological department of Yunnan ed., 1974)
11: H. mileensis (in Geological department of Yunnan ed., 1974)
12: H. funingensis (in Geological department of Yunnan ed., 1974)
13: H. repkinae (in Geological Department of Inner Mongolia Autonomous
Region (GDIMAR) and Dongbei institute of geology (DIG) eds., 1976)
14: H. regularis var. regularis (in GDIMAR and DIG eds., 1976)
15: H. compactiformis (in GDIMAR and DIG eds., 1976)
16: H. vesicotabulatus (in GDIMAR and DIG eds., 1976)
17: H. fovetabulatus (in GDIMAR and DIG eds., 1976)
18: H. uksunayensis (in GDIMAR and DIG eds., 1976)
19: H. yassensis obliquespinus (in GDIMAR and DIG eds., 1976)
20: H. (S.) gemina (in GDIMAR and DIG eds., 1976)

15

10

12
12

19
11

H. cf. gemina

0.5

17

3

7

0.4

1
16

9

2

8

6

0.3

4
14

20

0.2

18

13

5

H. wenxianicus

0.1

0.5

1.0

1.5

2.0

2.5
(mm)

diameter of tabularia

(mm)
0.9

0.8

0.7

diameterof tubule

0.6

0.5

21: H. interstinctus (in Lin and Huang, 1987)
22: H. cf. pseudobohemicus (in Lin and Huang, 1987)
23: H. zhanwabeigouensis (in Lin and Huang, 1987)
24: H. pseudobohemicus (in Lin and Huang, 1987)
25: H. zhanwaensis (in Lin and Huang, 1987)
26: H. tchernychevi (in Lin and Huang, 1987)
27: H. multitabulatus (in Lin and Huang, 1987)
28: H. yanglugouensis (in Lin and Huang, 1987)
29: H. wenxianicus (in Tchi, 1987)
30: H. insolens var. dunbeiensis (in Tchi, 1987)
31: H. salairicus (in Xiong and Gu, 1978)
32: H. jilinensis (in Tchi, 1980)
33: H. lindstromi (in Tchi, 1980)
34: H. insolens (in Tchi, 1980)
35: H. insolens dunbeiensis (in Tchi, 1980)
36: H. waicunensis (in Li and Lin, 1982)
37: H. insolens dunbeiensis (in Yeh and Chow, 1995)
38: H. daguanensis (in Jin, 1984)
39: H. (Paraheliolites) obvellatu (in Deng and Zheng, 2000)
40: H. cf. interstinctus (in Sugiyama, 1940)
41: H. cf. barrandei (in Sugiyama, 1940)
42: H. onuki (in Sugiyama, 1940)

22

31

30
25
37

H. cf. gemina

26
23

24
39

31

27

0.4

(no description
on diameter of tubulae)

38
35

34
36

0.3

40
33

29
21
41

32

0.2

28

H. wenxianicus

0.1

42

0.5

1.0

1.5

2.0

2.5
(mm)

diameter of tabularia
Text-Fig. 4. Diameters of tabularia (x-axis) and tubules (y-axis) of Heliolites from Asia (having septal structure only).
GDIMAR: Geological Department of Inner Mongolia Autonomous Region, DIG: Dongbei Institute ofGeology, XIGMRS; Xian Institute of Geological and Mineral Resources Section. (continue to next page)— 72 —

(mm)
1.0

0.9

0.8

diameterof tubule

0.7

0.6

1: H. (Stelliporella) gemina (in Deng, 1999)
2: H. orientalis spinosus (in Guizhou geological and paleontological team ed, 1978)
3: H. guiyangensis (in Guizhou geological and paleontological team ed, 1978)
4: H. wudangensis (in Guizhou geological and paleontological team ed, 1978)
5: H. minutissimus (in Guizhou geological and paleontological team ed, 1978)
6: H. luxarboreus (in Guizhou geological and paleontological team ed, 1978)
7: H. tomensis (in Guizhou geological and paleontological team ed, 1978)
8: H. porosus (in Guizhou geological and paleontological team ed, 1978)
9: H. insolens (in Geological department of Yunnan ed., 1974)
10: H. stella (in Geological department of Yunnan ed., 1974)
11: H. mileensis (in Geological department of Yunnan ed., 1974)
12: H. funingensis (in Geological department of Yunnan ed., 1974)
13: H. repkinae (in Geological Department of Inner Mongolia Autonomous
Region (GDIMAR) and Dongbei institute of geology (DIG) eds., 1976)
14: H. regularis var. regularis (in GDIMAR and DIG eds., 1976)
15: H. compactiformis (in GDIMAR and DIG eds., 1976)
16: H. vesicotabulatus (in GDIMAR and DIG eds., 1976)
17: H. fovetabulatus (in GDIMAR and DIG eds., 1976)
18: H. uksunayensis (in GDIMAR and DIG eds., 1976)
19: H. yassensis obliquespinus (in GDIMAR and DIG eds., 1976)
20: H. (S.) gemina (in GDIMAR and DIG eds., 1976)

15

H. ichinotaniensis

10

12
12

19
11

0.5

17

3

0.4

7
1

2

16

9

8

6

0.3
14

20

0.2

18

4

13

5

H. ? sp.

0.1

0.5

1.0

1.5

2.0

2.5
(mm)

diameter of tabularia
(mm)
0.9

0.8

0.7

diameterof tubule

0.6

0.5

21: H. interstinctus (in Lin and Huang, 1987)
22: H. cf. pseudobohemicus (in Lin and Huang, 1987)
23: H. zhanwabeigouensis (in Lin and Huang, 1987)
24: H. pseudobohemicus (in Lin and Huang, 1987)
25: H. zhanwaensis (in Lin and Huang, 1987)
26: H. tchernychevi (in Lin and Huang, 1987)
27: H. multitabulatus (in Lin and Huang, 1987)
28: H. yanglugouensis (in Lin and Huang, 1987)
29: H. wenxianicus (in Tchi, 1987)
30: H. insolens var. dunbeiensis (in Tchi, 1987)
31: H. salairicus (in Xiong and Gu, 1978)
32: H. jilinensis (in Tchi, 1980)
33: H. lindstromi (in Tchi, 1980)
34: H. insolens (in Tchi, 1980)
35: H. insolens dunbeiensis (in Tchi, 1980)
36: H. waicunensis (in Li and Lin, 1982)
37: H. insolens dunbeiensis (in Yeh and Chow, 1995)
38: H. daguanensis (in Jin, 1984)
39: H. (Paraheliolites) obvellatu (in Deng and Zheng, 2000)
40: H. cf. interstinctus (in Sugiyama, 1940)
41: H. cf. barrandei (in Sugiyama, 1940)
42: H. onuki (in Sugiyama, 1940)

H. ichinotaniensis
22

31

30
25
37
26
23

24

(no description
on diameter of tubulae)

39
31

27

0.4

38
29
34
36

0.3

35
40

33
21

28
41
32

0.2

H. ? sp.

0.1

0.5

42

1.0

1.5

diameter of tabularia— 73 —

2.0

2.5
(mm)

frequency (number in 5mm)
50

5

40

diaphragm(no description
on diaphragm frequency)

30

17

H. wenxianicus
89

H. ? sp.
4

1: H. (Stelliporella) gemina (in Deng, 1999)
2: H. orientalis spinosus (in Guizhou geological and paleontological team ed, 1978)
3: H. guiyangensis (in Guizhou geological and paleontological team ed, 1978)
4: H. wudangensis (in Guizhou geological and paleontological team ed, 1978)
5: H. minutissimus (in Guizhou geological and paleontological team ed, 1978)
6: H. luxarboreus (in Guizhou geological and paleontological team ed, 1978)
7: H. tomensis (in Guizhou geological and paleontological team ed, 1978)
8: H. porosus (in Guizhou geological and paleontological team ed, 1978)
9: H. insolens (in Geological department of Yunnan ed., 1974)
10: H. stella (in Geological department of Yunnan ed., 1974)
11: H. mileensis (in Geological department of Yunnan ed., 1974)
12: H. funingensis (in Geological department of Yunnan ed., 1974)
13: H. repkinae (in GDIMAR and DIG eds., 1976)
14: H. regularis var. regularis (in GDIMAR and DIG eds., 1976)
15: H. compactiformis (in GDIMAR and DIG eds., 1976)
16: H. vesicotabulatus (in GDIMAR and DIG eds., 1976)
17: H. fovetabulatus (in GDIMAR and DIG eds., 1976)
18: H. uksunayensis (in GDIMAR and DIG eds., 1976)
19: H. yassensis obliquespinus (in GDIMAR and DIG eds., 1976)
20: H. (S.) gemina (in GDIMAR and DIG eds., 1976)

6

14
20

20

3

2
21

19

13

1

10

12

11

H. ichinotaniensis
15

10

7
16

H. cf. gemina
37

18

5

10

15

20

25

30

35

40

50

frequency (number in 5mm)

tabulafrequency (number in 5mm)50

21: H. interstinctus (in Lin and Huang, 1987)
22: H. cf. pseudobohemicus (in Lin and Huang, 1987)
23: H. zhanwabeigouensis (in Lin and Huang, 1987)
24: H. pseudobohemicus (in Lin and Huang, 1987)
25: H. zhanwaensis (in Lin and Huang, 1987)
26: H. tchernychevi (in Lin and Huang, 1987)
27: H. multitabulatus (in Lin and Huang, 1987)
28: H. yanglugouensis (in Lin and Huang, 1987)
29: H. wenxianicus (in Tchi, 1987)
30: H. insolens var. dunbeiensis (in Tchi, 1987)
31: H. salairicus (in Xiong and Gu, 1978)
32: H. jilinensis (in Tchi, 1980)
33: H. lindstromi (in Tchi, 1980)
34: H. insolens (in Tchi, 1980)
35: H. insolens dunbeiensis (in Tchi, 1980)
36: H. waicunensis (in XIGMRS, 1982)
37: H. insolens dunbeiensis (in Yeh and Chow, 1995)
38: H. daguanensis (in Jin, 1984)
39: H. (Paraheliolites) obvellatu (in Deng and Zheng, 2000)
40: H. cf. interstinctus (in Sugiyama, 1940)
42: H. onuki (in Sugiyama, 1940)(no description
on diaphragm frequency)
40
32diaphragm

33

30

38

H. wenxianicus
H. ? sp.

35
30

29

20

20

25

28

25

23

21
39

26
27
31

24

36

H. ichinotaniensis

22

34

10
H. cf. gemina
37

40

42

5

10

15

20

tabula25

30

35

frequency (number in 5mm)

Text-Fig. 5. Tabular (x-axis) and diaphragm (y-axis) frequencies of Heliolites from Asia (having septal structure only).

— 74 —

1976; p. 123), H. wenxianicus (in Tchi, 1987; p. 276), H. lindstromi (in Tchi, 1980; p. 185) and H.
insolens (in Tchi, 1980; p. 185) in their septal structure, the size of the tabularia, the size of thecoenenchymal tubules, the tabular frequency and the diaphragm frequency (Text-Figs. 4 and 5). However, the coenenchymal wall of the specimens examined here is generally thicker than that of H. repkinae,
H. regularis var. kuznetskiensis or H. lindstromi. H. orientalis var. spinosus differs in having substellate
tabularia. Although GDIMAR and DIG eds. (1976) described spines pointing upwards from thecoenenchymal wall in H. guiyangensis, such spines were not observed in the specimens examined here
which compare very well with H. wenxianicus as described and figured by Zhang (1981) from the upper
part of the Middle Devonian Xiaputonggou Formation, Putonggou, south Qinling Mountains, China,
and by Tchi (1987) from the upper part of the Lower Devonian Xiaputonggou Formation, Putonggou,
west Qinling Mountains, China.
Heliolites cf. gemina (Tchi, 1976)

Plate 12, Figs. 1-3

1976 Stelliporella gemina Tchi; GDIMAR and DIG eds., p. 125, Pl. 50, Fig. 3.
1999 Heliolites gemina (Tchi); Deng, p. 229, Pl. 40, Fig. 1.

Reg. No.
Samples: NUM-Fz-010809-f, NUM-Fz-010811-15, NUM-Fz-010811-30 and NUM-Fz-010811-34.
Four samples.
Specimens: NUM-Fz-010809-f-1a, NUM-Fz-010811-15-3, NUM-Fz-010811-30-4 and NUM-Fz010811-34-2a. Four specimens.
Thin sections: Three (010809-f-1-1, 010809-f-1-2 and 010809-f-1-3), two (010811-15-3-1 and 01081115-3-2), one (010811-30-4-1), and one (010811-34-2-1a) thin sections cut from NUM-Fz-010809-f-1,
NUM-Fz-010811-15-3, NUM-Fz-010811-30-4, and NUM-Fz-010811-34-2a, respectively.
Material, horizons and localities
All specimens except for NUM-Fz-010809-f-1 were from the Upper Member of the Fukuji Formation along the Ichinotani Valley, Takayama City, Gifu Prefecture, central Japan (Loc.1). NUM-Fz-010809f-1 was from a sub-rounded pebble at Loc.1.
Description
Corallum of variable form, heliolitoid. Tabularia circular, oval or subpolygonal in transverse section,cylindrical, straight, diameter 0.75-1.25mm; walls 0.04-0.2mm thick; septa commonly represented by
12 slender spines with thick bases, rarely by septal laminae with axially spinose edges, horizontal or
curved upwards; tabulae complete, horizontal or oblique, flat, convex or concave, generally very thin(less than 0.05mm), 7-10 per 5mm. Tabularia separated by 0.3-1.67mm of coenenchymal tubules.
Coenenchymal tubules 3- to 8-sided in transverse section, 0.23-0.51mm in diameter; coenenchymal
walls 0.025-0.12mm thick; diaphragms generally flat, 8-18 per 5mm.
Remarks
These specimens are identified as belonging to the genus Heliolites by their circular tabularia andcoenenchymal tubules with transverse diaphragms. 61 species of the genus Heliolites from East Asia
were compared with the specimens examined in this paper: H. anfuiensis Yu and Deng; H. arboreus
Sugiyama; H. assuetus Bondarenko; H. baichengensis Wang; H. (Pachycanalicura) cf. barrandei(Hoernes); H. bohemicus Wentzel; H. (Paraheliolites) bulongensis (Lin et Huang); H. compactiformis
Tchi; H. daguanensis Kim; H. decipiens (McCoy); H. densitabulatus Wang; H. decipiens var. minimus
Rukhin; H. fenggangensis Yang; H. fovetabulatus Tchi; H. funingensis King; H. (Stelliporella) gemina(Tchi); H. guiyangensis Chow; H. guizhouensis Yang; H. huanghuachongensis Chow; H. insolens
Tchernychev; H. insolens dunbeiensis Dubatolov; H. interstinctus (Linne); H. interstinctus-intermedis
Wentzel; H. (Paraheliolites) jilinensis Tchi; H. lindstromi Kovalevsky; H. luorepingensis Wu; H.
— 75 —

luxarboreus Yang; H. magnus Yu; H. mileensis King; H. minutissimus Chow; H. multitabulatus
Bondarenko; H. obliquus Yu; H. onukii Sugiyama; H. orientalis Yu; H. (Paraheliolites) obvellatus Deng
and Cheng; H. orientalis spinosus Yang; H. paradoxicus Lin and Wang; H. paravulgaris Kim; H. porosus(Goldfuss); H. pseudobohemicus Bondarenko; H. regularis var. kuznetskiensis Tchernychev; H. repkinae
Kovalevsky; H. (Paraheliolites) salairicus Tchernychev; H. sinkiangensis Yu; H. stella King; H.
tashanensis Lin and Chow; H. tchernychevi Bondarenko; H. tomensis Tchernychev; H. uksunayensis
Mironova; H. vesicotabulatus Tchi; H. volutus Yu and Deng; H. vulgaris irregularis Tchernychev; H.
waicunensis Lin and Chow; H. wenxianicus Zhang; H. wudangensis Chow; H. yanbianensis Kim; H.
yanglugouensis Lin and Fuan; H. yassensis obliquespinus Chu; H. zhanwabeigouensis Lin and Fuan; H.
zhanwaensis Lin and Fuan; H. zoigeensis Lin and Fuan (Xian Institute of Geological and Mineral Resources, Geological and Mineral Resources Section ed, 1982; Xinjiang Geological Department, Xinjiang
Geological Institute and Xinjiang Petroleum Department eds., 1981; Nanjing Institute of Geology andMineralogy, Geology and Mineralogy Section ed., 1983; Tchi, 1980; Xiong and Gu, 1978; Wang et al.,
2001; Tchi, 1987; Xinan Geological Institute ed., 1978; Lin and Huang, 1987; Lin and Wang, 1987;
Geological Department of Inner Mongolia Autonomous Region (GDIMAR) and Dongbei Institute ofGeology (DIG) eds., 1976; Fubei Institute of Geology, Geological Department of Hubei, Geological
Department of Guandong, Geological Department of Henan, Geological Department of Hunan andGeological Department of Guangxi Autonomous Region eds., 1977; Geological Department of Yunnaned., 1974; Regional Geological Surveying Team of Hubei ed., 1994; Guizhou Geological and Paleontological Team ed, 1978; Deng, 1999; Yeh and Chow, 1995; Jin, 1984; Deng and Zheng, 2000; Sugiyama,
1940; Yu, 1962; Lin and Chow, 1977; Lin and Huang, 1986; Kamei, 1955b; Zhang, 1981).
In respect of the size of their tabularia, the size of the coenenchymal tubules, tabular frequency anddiaphragm frequency, the specimens examined here are similar to H. (Stelliporella) gemina (in GDIMAR
and DIG eds, 1976; p. 125 and Deng, 1999; p. 229), H. orientalis spinosus (in Guizhou Geological andPaleontological Team ed, 1978; p. 237), H. stella (in Geological department of Yunnan ed., 1974; p.
247), H. repkinae (in GDIMAR and DIG eds., 1976; p. 123), H. regularis var. kuznetskiensis (in GDIMAR
and DIG eds., 1976; p. 123) and H. insolens (in Tchi, 1980; p. 185) (Text-Figs. 4 and 5).
H. (S.) gemina described and figured in GDIMAR and DIG eds (1976) from the Upper Silurian Xibiehe
Formation, Inner Mongolia, China, is similar to the specimens examined here in having axially spinose
septal laminae and septal spines. However, the Japanese specimens have more closely spaced tabulae anddiaphragms than the Chinese specimens. In H. gemina, described by Deng (1999) from the Middle Silurian Shaebuer Formation, Inner Mongolia, China, the septa are only represented by septal laminae.
Heliolites ichinotaniensis sp. nov. Plate 12, Figs. 4-7
Name
The name is derived from the Ichinotani Valley, Fukuji, central Japan.
Reg. No.
Samples: NUM-Fz-010811-14, NUM-Fz-010811-18, NUM-Fz-010811-20, NUM-Fz-010811-21,
NUM-Fz-010811-29 and NUM-Fz-010811-32. Six samples.
Specimens:
Holotype. NUM-Fz-010811-18-1 (thin sections: 010811-18-1-1, 010811-18-1-2 and 010811-18-1-3).
Paratype. NUM-Fz-010811-14-1, NUM-Fz-010811-20-1, NUM-Fz-010811-21-2, NUM-Fz-01081129-1 and NUM-Fz-010811-32-1.
Thin sections: One (010811-14-1-1), two (010811-20-1-1 and 010811-20-1-2), six (010811-21-2-1,
010811-21-2-2, 010811-21-2-3, 010811-21-2-4, 010811-21-2-5 and 010811-21-2-6), two (010811-291-1 and 010811-29-1-2), and nine (010811-32-1-4, 010811-32-1-5, 010811-32-1-7, 010811-32-1-8,— 76 —

010811-32-1-9a, 010811-32-1-10, 010811-32-1-11a, 010811-32-1-12a and 010811-32-1-13a) thin sections cut from NUM-Fz-010811-14-1, NUM-Fz-010811-20-1, NUM-Fz-010811-21-2, NUM-Fz-01081129-1 and NUM-Fz-010811-32-1 respectively.
Material, horizons and localities
All specimens were from the Upper Member of the Fukuji Formation along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Description
Corallum of variable form, heliolitoid. Tabularia generally polygonal or substellate rarely circular intransverse section, cylindrical, straight, diameter 0.9-1.5mm; walls 0.025-0.1 thick; septal spines welldeveloped in 12 longitudinal rows, slender, long, horizontal to sharply curved upwards; tabulae complete, horizontal or oblique, flat, convex or concave, rarely sagging, generally very thin (less than 0.05mm),
usually 8-11 per 5mm, rarely 12-16 per 5mm. Tabularia separated by 0.5-2.5mm of coenenchymal tubules. Coenenchymal tubules 4- to 8-sided in transverse section, 0.15-0.75mm in diameter (mostly 0.40.5mm); coenenchymal walls 0.02-0.66mm thick with distinct median suture; diaphragms generally
flat, but rarely convex or concave, 11-19 per 5mm (usually 13-15 per 5mm),
Remarks
This species is characterized by thin-walled polygonal or substellate tabularia, distantly spaced diaphragms and long slender septal spines. In the size of the tabularia, the size of the coenenchymal tubules, tabula frequency and diaphragm frequency, the specimens studied here are similar to H. (S.)
gemina (in GDIMAR and DIG eds., 1976; p. 125 and Deng, 1999; p. 229), H. regularis var. kuznetskiensis(in GDIMAR and DIG eds., 1976; p. 123), H. fovetabulatus (in GDIMAR and DIG eds., 1976; p. 124),
H. tchernychevi (in Lin and Huang, 1987; p. 229), H. insolens dunbeiensis (in Tchi, 1980; p. 185 andTchi, 1987; p. 276) and H. insolens (in Tchi, 1980; p. 185) (Text-Figs. 4 and 5). Although H. regularisvar. kuznetskiensis and H. fovetabulatus are similar to this new species in having long slender spines,
they differ in having circular tabularia. H. tchernychevi has stellate to substellate tabularia, and in H.
ichinotaniensis the diaphragms are much more closely spaced.
Heliolites ? sp.
Plate 13, Figs. 1-4
Reg. No.
Samples: NUM-Fz-010811-7, NUM-Fz-010811-13, NUM-Fz-010811-25, NUM-Fz-010811-28,
NUM-Fz-010811-33 and NUM-Fz-010811-34. Six samples.
Specimens: NUM-Fz-010811-7-1, NUM-Fz-010811-13-1, NUM-Fz-010811-25-4, NUM-Fz-01081128-1, NUM-Fz-010811-33-2b, NUM-Fz-010811-34-2b and NUM-Fz-010811-34-12. Seven specimens.
Thin sections: Two (010811-7-1-1 and 010811-7-1-2), two (010811-13-1-1 and 010811-13-1-2),
two (010811-25-4-1b and 010811-25-4-2b), two (010811-28-1-1 and 010811-28-1-2), one (010811-332-2b), one (010811-34-2-1b), and five (010811-34-12-1, 010811-34-12-2, 010811-34-12-3, 01081134-12-4 and 010811-34-12-5) thin sections cut from NUM-Fz-010811-7-1, NUM-Fz-010811-13-1, NUMFz-010811-25-4, NUM-Fz-010811-28-1, NUM-Fz-010811-33-2, NUM-Fz-010811-34-2 and NUM-Fz010811-34-12 respectively.
Material, horizons and localities
All specimens were from the Upper Member of the Fukuji Formation along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Description
Corallum of variable form, heliolitoid. Tabularia circular in transverse section, cylindrical, straight,
diameter 0.75-1.12mm; walls 0.025-0.2 thick; septal spines well developed in 12 longitudinal rows,
slender or rarely very thick, horizontal or curved upwards; tabulae complete, horizontal or oblique, flat,— 77 —

convex or concave, generally very thin (less than 0.02mm), 10-11 per 5mm. Each tabularium is separated by 0.25-1.35mm of coenenchymal tubules. Coenenchymal tubules 4- to 6-sided in transverse section, 0.125-0.46mm in diameter; coenenchymal walls 0.02-0.15mm thick, often with lenticular thickening; diaphragms generally flat, but rarely slightly convex or concave, 20-25 per 5mm where closely
spaced and 16-19 per 5mm where distantly spaced.
Remarks
The specimens assigned here to Heliolites ? sp. A are similar to those referred to H. wenxianicus inthis paper in having closely spaced tabulae and diaphragms (Text-Figs. 4 and 5), but they can be distinguished from H. wenxianicus by the lenticular thickening of the coenenchymal walls and likely to be
assigned to genus Helioplasma as they occasionally have incomplete and overlapping convex diaphragms.
Genus HELIOPLASMA Kettnerová, 1933
Type species. Helioplasma kolihai Kettnerová, 1933, p. 181.
Other species. H. domestica Bondarenko, 1966; H. (?) indotata Bondarenko, 1966; H. propinguus(Bondarenko, 1966); H. aliena Galle, 1973; H. dnestriensis (Bondarenko, 1981); H. urjupica Bondarenko;
H. (?) squameoformis Bondarenko; H. delicatum Yanet and Lobanov, 1992 (in Lobanov, 1992).
Bondarenko (1966) was uncertain whether H. domestica was a member of the genus Helioplasma,
while Galle (1969) included it in that genus. In addition, Galle (1969) included Bogimbailites propinguus
Bondarenko, 1966 in the genus Helioplasma.
Diagnosis (after Hill, 1981, p. F603). Corallum heliolitoid; tabularial walls slightly thickened; septa
represented by long septal spines; tabulae complete, not evenly horizontal; coenenchymal tubules normally prismatic with horizontal diaphragms, or occasionally elongate in transverse section with incomplete overlapping convex diaphragms as well as horizontal, complete diaphragms.
Helioplasma takayamaensis sp. nov.
Plate 13, Figs. 5-8
Name
The name is derived from Takayama City, Gifu Prefecture, central Japan.
Reg. No.
Samples: NUM-Fz-010811-26-1 and NUM-Fz-010811-28-2. Two specimens.
Specimens:
Holotype. NUM-Fz-010811-26-1 (thin sections: 010811-26-1-1 and 010811-26-1-2).
Paratype. NUM-Fz-010811-28-2.
Thin sections: Two (010811-28-2-1 and 010811-28-2-2) thin sections cut from NUM-Fz-01081128-2.
Material, horizons and localities
All specimens were from the Upper Member of the Fukuji Formation, along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Description
Corallum massive, heliolitoid. Tabularia circular or polygonal in transverse section, cylindrical, straight,
diameter 1.1-1.5mm; walls 0.025-0.125 thick; septal spines well developed in 12 longitudinal rows,
slender, long, curved upwards; tabulae complete, concave or sagging, rarely flat or convex, generally
very thin (less than 0.04mm), 6-10 per 5mm. Tabularia separated by 0.25-1.35mm of coenenchymal
tubules. Coenenchymal tubules 4- to 8-sided in transverse section, 0.2-0.67mm in diameter; coenenchymal
walls 0.03-0.1mm thick; diaphragms sometimes deeply sagging sometimes conical, incomplete andconvex, also some horizontal complete diaphragms, 11-16 per 5mm.
Remarks— 78 —

(mm)
0.7

0.6

H. takayamaensis

diameterof tubule

0.5

6
7
5

0.4

4
9
0.3
8
10

1: H. kolihai (in Galle, 1969, 1973)
2: H. (?) domestica (in Bondarenko, 1966)
3: H. domestica (in Galle, 1969)
4: H. (?) indotata (in Bondarenko, 1966)
5: H. (?) indotata (in Galle, 1969)
6: B. propinguus (in Bondarenko, 1966)
7: H. propinguus (in Galle, 1969)
8: H. aliena (in Galle, 1973)
9: H. dnestriensis (in Bondarenko, 1981)
10: H. delicatum (in Lobanov, 1992)

3
2

0.2
1
0.1
0.5

1.0

1.5

2.0

2.5

3.0

diameter of tabularium

3.5
(mm)

Text-Fig. 6. Diameters of tabularia (x-axis) and tubules (y-axis) of Helioplasma.

frequency (number in 5mm)
70

diaphragm

60

50

1: H. kolihai (in Galle, 1969, 1973)
2: H. (?) domestica (in Bondarenko, 1966)
3: H. domestica (in Galle, 1969)
4: H. (?) indotata (in Bondarenko, 1966)
5: H. (?) indotata (in Galle, 1969)
6: B. propinguus (in Bondarenko, 1966)
7: H. propinguus (in Galle, 1969)
8: H. aliena (in Galle, 1973)
9: H. dnestriensis (in Bondarenko, 1981)
10: H. delicatum (in Lobanov, 1992)

10

9

40

1
30
3

4

2

20
8
10

6
4

H. takayamaensis

5

5

10

15

20

tabulae25

30

Text-Fig. 7. Tabular (x-axis) and diaphragm (y-axis) frequencies of Helioplasma.

— 79 —

35

frequency (number in 5mm)

These specimens are characterized by thick-walled tabularia, and deeply sagging or overlapping
convex diaphragms. These features place them clearly within the genus Helioplasma. The specimens
studied here are clearly distinguished from H. kolihai (in Galle, 1969; p. 170 and 1971; p. 34), H.
domestica (in Bondarenko, 1966; p. 194), H. (?) indotata (in Bondarenko, 1966; p. 193), H. aliena (inGalle, 1973; p. 35), and H. delicatum (in Lobanov, 1992; p. 9) in having widely spaced tabulae anddiaphragms (Text-Figs. 6 and 7). H. propinguus (in Bondarenko, 1966; p. 192) has much larger tabularia
than the present specimens (Text-Fig. 6). The tabula of H. dnestriensis (in Bondarenko, 1981; p. 16) are
not concave or sagging, and are thus different from those of the present specimens.
Family PSEUDOPLASMOPORIDAE Bondarenko, 1963
Definition (Hill, 1981). Corallum variable in form; walls of tabularia smooth (with no trace of facets)or longitudinally folded; septa either laminae or longitudinal rows of spines, or absent; tabulae complete, horizontal; an aureola of 12 tubules surrounding each tabularium, variable in radius; remaining
coenenchyme also of tubules with complete, horizontal diaphragms or rarely with oblique and incomplete diaphragms.
Remarks. This family is composed of four genera, Pseudoplasmopora, Amphilites, Pachyhelioplasma
and Visbylites (Hill, 1981). The former three genera were reported from the Lower to ?Middle Devonian(Hill, 1981).
Genus PSEUDOPLASMOPORA Bondarenko, 1963
Type species. Pseudoplasmopora conspecta Bondarenko, 1963, p. 47.
Other species. P. follis (Milne-Edwards and Haime, 1851); P. gippslandica (Chapman, 1914); P.
aseptata (Regnell, 1941); P. cargoensis (Hill, 1957); P. arguta arguta Bondarenko, 1963; P. plasmopolides
Lin, 1963; P. septosa Bondarenko, 1966; P. subdecipiens (Kovalevsky, 1956); P. bella (Kovalevsky,1956); P. arguta festivus Bondarenko, 1966; P. isenica Bondarenko, 1966; P. regularis (Dun, 1927); P.
lojopingensis Xiong in Xiong and Gu (1978); P. sanxiaensis Xiong in Xiong and Gu (1978); P.
yichangensis Xiong in Xiong and Gu (1978); P. hubeiensis Xiong in Xiong and Gu (1978); P. microsa
Wang in Xinjiang Geological Department et al. eds. (1981); P. longisepta Klaamann, 1984; P. yaokengensis
Deng and Zheng, 2000.
Although Bondarenko (1963) included Plasmopora follis Milne-Edwards and Haime, 1851 in his
new genus Pseudoplasmopora, whether or not this decision was correct is debatable. Bondarenko (1966)
considered P. follis described by Shimizu et al. (1934) from the Ken-niho conglomerate, North Korea tobe a synonym of P. septosa.
Diagnosis (after Hill, 1981, p. F603). Pseudoplasmoporidae with septa consisting of septal spines orabsent, walls of tabularia and tubules unthickened, diaphragms of tubules rarely oblique and incomplete.
Pseudoplasmopora okuhidaensis sp. nov. Plate 14, Figs. 1-7
Name
The name is derived from Okuhida-onsen-go (Okuhida Spa), Takayama City, Gifu Prefecture, central Japan.
Reg. No.
Samples: NUM-Fz-010811-28-4, NUM-Fz-010811-33-4 and NUM-Fz-010811-34. Three samples.
Specimens:
Holotype. NUM-Fz-010811-28-4 (thin sections: 010811-28-4-1, 010811-28-4-2, 010811-28-4-3 and
010811-28-4-4).
Paratype. NUM-Fz-010811-33-4 and NUM-Fz-010811-34-5b.
— 80 —

Thin sections: One (010811-33-4-1), and two (010811-34-5-1b and 010811-34-5-2b) thin sections
cut from NUM-Fz-010811-33-4, and NUM-Fz-010811-34-5b respectively.
Material, horizons and localities
All specimens were from the Upper Member of the Fukuji Formation, along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Description
Corallum massive, heliolitoid. Tabularia circular with an aureola of 12 elongated coenenchymal tubules in transverse section, cylindrical, straight, diameter 1.125-2.0mm; walls 0.025-0.15mm thick (commonly 0.075-0.15mm); septa well developed, represented by 12 laminae with axially spinose edges or
longitudinal rows of slender, long septal spines. Spinose edges and spines generally curved upwards.
Tabulae complete, mostly concave or sagging, generally very thin (less than 0.05mm), 6-8 per 5mm.
Tabularia separated by 0.425-1.5mm of coenenchymal tubules. Coenenchymal tubules 4- to 6-sided intransverse section, 0.1-1.15mm in diameter; coenenchymal walls 0.025-0.125mm thick; diaphragms
generally flat, complete or rarely incomplete, 7-13 per 5mm.
Remarks
These specimens are referable to the genus Pseudoplasmopora by having an aureola of 12 tubules
around each tabularium and tubules normally with flat diaphragms. Pseudoplasmopora okuhidaensis ischaracterized by large corallites, distantly spaced tabulae and the presence of diaphragms (Text-Figs. 8
and 9). Tabulae are generally concave or rarely deeply sagging.
The specimens are clearly distinguished from any other species of the genus by having distantly
spaced tabulae and diaphragms (Text-Fig. 9). They are similar to P. subdecipiens (in Bondarenko, 1966;
p. 160) and P. cargoensis (in Bondarenko and Minjin, 1981; p. 116) in tabular frequency, but are quite
different from those species in having much smaller tabularia (Text-Figs. 8 and 9).
Pseudoplasmopora cf. arguta Bondarenko, 1963

Plate 15, Figs. 1-6

cf.1963 Pseudoplasmopora arguta Bondarenko: Bondarenko, p. 48, Pl. IV, Fig. 2.
cf.1966 Pseudoplasmopora arguta Bondarenko: Bondarenko, p. 162, Pl. XX, Fig. 4.

Reg. No.
Samples: NUM-Fz-010811-33 and NUM-Fz-010811-34. Two samples.
Specimens: NUM-Fz-010811-33-3, NUM-Fz-010811-34-2b and NUM-Fz-010811-34-3. Three specimens.
Thin sections: Three (010811-33-3-2, 010811-33-3-1 and 010811-33-3-3), one (010811-34-2-1b),
and five (010811-34-3-3, 010811-34-3-4, 010811-34-3-5a, 010811-34-3-6a and 010811-34-3-7a) thinsections cut from NUM-Fz-010811-33-3, NUM-Fz-010811-34-2b and NUM-Fz-010811-34-3 respectively.
Material, horizons and localities
All specimens were from the Upper Member of the Fukuji Formation, along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Description
Corallum massive, heliolitoid. Tabularia circular with an aureola of 12 elongated coenenchymal tubules in transverse section, cylindrical, straight, diameter 0.75-1.25mm; walls 0.025-0.125mm thick;septa well-developed, represented by 12 longitudinal rows of long, slender septal spines, curved upwards. Tabulae complete, horizontal, flat or concave or rarely wavy, moderately thick (less than 0.1mm),
7-20 per 5mm. Tabularia separated by 0.65-1.175mm of coenenchymal tubules. Coenenchymal tubules
4- to 5-sided in transverse section, 0.175-0.925mm in diameter; coenenchymal walls 0.025-0.075mm
thick; diaphragms flat or rarely concave or convex, 12-20 per 5mm.
— 81 —

(mm)1.4

diameterof tubule

1.2

1: P. conspecta (in Bondarenko, 1963, 1966)
2: P. septosa (in Bondarenko, 1966)
3: P. subdecipiens (in Bondarenko, 1966)
4: P. bella (in Bondarenko, 1966)
5: P. arguta arguta (in Bondarenko, 1963, 1966)
6: P. arguta festivus (in Bondarenko, 1966)
7: P. isenica (in Bondarenko, 1966)
8: P. regularis (in Bondarenko, 1966)
9: P. cf. isenica (in Wang et al., 2001)
10: P. lojopingensis (in Xiong and Gu, 1978)
11: P. sanxiaensis (in Xiong and Gu, 1978)
12: P. yichangensis (in Xiong and Gu, 1978)
13: P. hubeiensis (in Xiong and Gu, 1978)

13

P. okuhidaensis

1.0
P. cf. arguta
12

0.8
9

11

10

10
0.6

7
3
2

0.4

8
6

1
5

4

0.2

0.2

0.4

0.6

0.8

1.0

1.2

1.4

1.6

1.8

diameter of tabularium
(mm)
1.4

diameterof tubule

1.2

14: P. microsa (in Xinjiang geological department et al. eds., 1981)
15: P. longisepta (in Klaamann, 1984)
16: P. yaokengensis (in Deng and Zheng, 2000)
17: P. cargoensis (in Hill, 1957)
18: P. cargoensis (in Bondarenko and Minjin, 1981)
19: H. interstinctica gippslandica (in Chapman, 1914)
20: P. gippslandica (in Hill and Jell, 1969)
21: P. follis (in Shimizu et al., 1934)
22: P. follis (in Noble and Young, 1984)
23: P. aseptata (in Xinjiang geological department et al. eds., 1981)
24: P. aff. aseptata (in Deng and Zheng, 2000)(no description
on diameter of tubulae)(no description
on diameter of tubulae)(no description
on diameter of tubulae)(no description
on diameter of tubulae)

1.0

2.0
(mm)

P. okuhidaensis

P. cf. arguta
0.8

0.6
24
23

22

0.4
18
14
15
0.2
20
16
17
0.2

0.4

0.6

21
0.8

1.0

19
1.2

1.4

1.6

1.8

diameter of tabularium
Text-Fig. 8. Diameters of tabularia (x-axis) and tubules (y-axis) of Pseudoplasmopora.
— 82 —

2.0(mm)

frequency (number in 5mm)
35

30

diaphragm

25

11

20

10

6
4

12

15

1

6

1

13

10

8

5

3

7
2
P. cf. arguta
4910

1: P. conspecta (in Bondarenko, 1963, 1966)
2: P. septosa (in Bondarenko, 1966)
3: P. subdecipiens (in Bondarenko, 1966)
4: P. bella (in Bondarenko, 1966)
5: P. arguta arguta (in Bondarenko, 1963, 1966)
6: P. arguta festivus (in Bondarenko, 1966)
7: P. isenica (in Bondarenko, 1966)
8: P. regularis (in Bondarenko, 1966)
9: P. cf. isenica (in Wang et al., 2001)
10: P. lojopingensis (in Xiong and Gu, 1978)
11: P. sanxiaensis (in Xiong and Gu, 1978)
12: P. yichangensis (in Xiong and Gu, 1978)
13: P. hubeiensis (in Xiong and Gu, 1978)

P. okuhidaensis
5

5

10

15

20

25

30

35

40

45

50

frequency (number in 5mm)

tabulafrequency (number in 5mm)
35
(no description
on diaphragm frequency)

30(no description
on diaphragm frequency)

19
21

diaphragm

25

14
P. cf. arguta

20

15(no description
on diaphragm frequency)

23
16

P. okuhidaensis

24
10
14: P. microsa (in Xinjiang geological department et al. eds., 1981)
15: P. longisepta (in Klaamann, 1984)
16: P. yaokengensis (in Deng and Zheng, 2000)
18: P. cargoensis (in Bondarenko and Minjin, 1981)
19: H. interstinctica gippslandica (in Chapman, 1914)
21: P. follis (in Shimizu et al., 1934)
22: P. follis (in Noble and Young, 1984)
23: P. aseptata (in Xinjiang geological department et al. eds., 1981)
24: P. aff. aseptata (in Deng and Zheng, 2000)

5
15
18

22
5

10

15

20

25

tabula

30

35

40

45

Text-Fig. 9. Tabular (x-axis) and diaphragm (y-axis) frequencies of Pseudoplasmopora.
— 83 —

50

frequency (number in 5mm)

Remarks
These specimens are referable to the genus Pseudoplasmopora by having an aureola of 12 tubules
around each tabularium and tubules usually with flat diaphragms. The specimens here are clearly different from Pseudoplasmopora okuhidaensis in having much smaller tabularia and coenenchymal tubules,
closely spaced diaphragms and lenticular coenenchymal wall-thickening.
The present specimens are clearly distinguished from P. follis (in Shimizu, 1934; p. 66 and Noble andYoung, 1984; p. 874), P. gippslandica (in Chapman, 1914; p. 311 and Hill and Jell, 1969; p. 23), P.
aseptata (in Xinjiang Geological Department et al. eds., 1981; p. 66 and Deng and Zheng, 2000; p. 221),
P. lojopingensis (in Xiong and Gu, 1978; p. 257), P. sanxiaensis (in Xiong and Gu, 1978; p. 258), P.
yichangensis (in Xiong and Gu, 1978; p. 258), P. hubeiensis (in Xiong and Gu, 1978; p. 258), P. microsa(in Xinjiang Geological Department et al. eds., 1981; p. 66) and P. yaokengensis (in Deng and Zheng,
2000; p. 221) in having numerous long septal spines. In addition, they can be distinguished from P.
subdecipiens (in Bondarenko, 1966; p. 160), P. yichangensis, P. hubeiensis, P. aseptata, P. yaokengensis
and P. cargoensis (in Hill, 1957; p. 104, and Bondarenko and Minjin, 1981; p. 116) by the size of thetabularium and the size of the coenenchymal tubules (Text-Fig. 8). The specimens are quite different
from P. conspecta (in Bondarenko, 1963; p. 47, and 1966; p. 161) in the spacing of the tabulae anddiaphragms (Text-Fig. 9.). P. isenica (in Bondarenko, 1966; p. 164, and Wang et al., 2001; p. 82) differsin having a larger distance between each tabularium. P. regularis (figured in Bondarenko, 1966; p. 165)
usually seems to have oblique, incomplete diaphragms. Pseudoplasmopora cf. arguta here is similar to
P. arguta described by Bondarenko (1963; p. 48, and 1966; p. 162), but his description of the typespecimen did not mention the lenticular coenenchymal wall-thickening.
Superfamily PACHYPORICAE Gerth, 1921
Family PACHYPORIDAE Gerth, 1921
Definition (after Hill, 1981, p. F575). Corallum cerioid and branching; corallites polygonal or roundedpolygonal in cross section and diverging from axis of branch to open usually normal to surface of branch,
but toward apex of branch at an acute angle; corallite walls thin and favositoid in axial zones, more orless markedly developed in peripheral zone of branch; original microstructure, when retained, withstereozone consisting of radial fibers deposited in concentric growth laminae; septal spines or rarely
squamulae may project from stereozone into tabularium; mural pores and tabulae present.
Genus STRIATOPORA Hall, 1851
Type species. Striatpora flexuosa Hall, 1851, p. 97.
Diagnosis (after Hill, 1981, p. F582). Corallum, ramose with cylindrical or slightly compressedbranches; corallites curving gently away from axial longitudinal direction and opening obliquely tosurface on small branches; calices may show septal ridges. Walls thin in axial zone, thickening to become wide stereozones distally, with distinct growth lamellation; corallites polygonal in section but
tabularium cylindrical distally because of wall thickening; mural pores common; septal spines may
project into lumen; tabulae complete; new corallites originate in one of two position, either near axis ofbranch or near boundary between inner thin-walled and outer thick-walled zones; the latter do not produce new corallites.
Striatopora sp.
Plate 16, Figs. 1-11; Plate 17, Figs. 1-8
Reg. No.
Samples: NUM-Fz-010809-e, NUM-Fz0o10809-h, NUM-Fz-010811-24, NUM-Fz-010811-28, NUMFz-010811-29, NUM-Fz-010811-30, NUM-Fz-010811-31, NUM-Fz-010811-32, NUM-Fz-010811-33— 84 —

and NUM-Fz-010811-34. Ten samples.
Specimens: NUM-Fz-010809-e-1, NUM-Fz-010809-e-2, NUM-Fz-010809-h-1, NUM-Fz-01081124-1, NUM-Fz-010811-24-2, NUM-Fz-010811-24-3, NUM-Fz-010811-24-6, NUM-Fz-010811-28-3,
NUM-Fz-010811-28-5, NUM-Fz-010811-28-6, NUM-Fz-010811-28-7, NUM-Fz-010811-29-1, NUMFz-010811-30-1, NUM-Fz-010811-31-5, NUM-Fz-010811-31-7, NUM-Fz-010811-32-1, NUM-Fz010811-32-2, NUM-Fz-010811-33-5, NUM-Fz-010811-34-2, NUM-Fz-010811-34-6, NUM-Fz-01081134-7, NUM-Fz-010811-34-8, NUM-Fz-010811-34-11 and NUM-Fz-010811-34-16. Twenty-four specimens.
Thin sections: Three (010809-e-1-1, 010809-e-1-2 and 010809-e-1-3), three (010809-e-2-1, 010809e-2-2 and 010809-e-2-3), three (010809-h-1-1, 010809-h-1-2 and 010809-h-1-3), one (010811-24-1-1),
one (010811-24-2-1), one (010811-24-1-3-1), two (010811-24-1-6-1 and 010811-24-1-6-3), one (01081128-3-1), two (010811-28-5-1 and 010811-28-5-2), two (010811-28-6-1 and 010811-28-6-2), one (01081128-7-1), one (010811-29-1-1b), one 010811-30-1-1), one (010811-31-5-1), one (010811-31-7-1), six(010811-32-1-1, 010811-32-1-2, 01081-32-1-6, 010811-32-1-11b, 010811-32-1-12b, 010811-32-113band), two (010811-32-2-1 and 010811-32-2-2), two (010811-33-5-1 and 010811-33-5-2), one(010811-34-2-1), one (010811-34-6-1), two (010811-34-7-1 and 010811-34-7-2), one (010811-34-81b), one (010811-34-11-1) and two (010811-34-16-1 and 010811-34-16-2) thin sections cut from NUMFz-010809-e-1, NUM-Fz-010809-e-2, NUM-Fz-010809-h-1, NUM-Fz-010811-24-1, NUM-Fz-01081124-2, NUM-Fz-010811-24-3, NUM-Fz-010811-24-6, NUM-Fz-010811-28-3, NUM-Fz-010811-28-5,
NUM-Fz-010811-28-6, NUM-Fz-010811-28-7, NUM-Fz-010811-29-1, NUM-Fz-010811-30-1, NUMFz-010811-31-5, NUM-Fz-010811-31-7, NUM-Fz-010811-32-1, NUM-Fz-010811-32-2, NUM-Fz010811-33-5, NUM-Fz-010811-34-2, NUM-Fz-010811-34-6, NUM-Fz-010811-34-7, NUM-Fz-01081134-8, NUM-Fz-010811-34-11 and NUM-Fz-010811-34-16 respectively.
Material, horizons and localities
All specimens except for NUM-Fz-010809-e and NUM-Fz-010809-h were from the Upper Memberof the Fukuji Formation, along the Ichinotani Valley, Takayama City, Gifu Prefecture, central Japan(Loc.1). NUM-Fz-010809-e and NUM-Fz-010809-h were from sub-rounded pebbles at Loc.1.
Description
Specimens mostly embedded in limestone matrix. Corallum cerioid, branching, cylindrical, ramose;branches 3-8mm in diameter; corallites rounded or rounded-polygonal in cross section. Corallites diverging from axis to open generally at acute angles to surface but at obtuse angles to surface in a specimen NUM-Fz-010809-e-1, width 0.2-0.9mm. Walls thin and favositoid in axial part (0.025-0.1mm thick)but slightly thickened to form stereozones in peripheral part (0.05-0.2mm thick). Concentric growthlaminae in stereozones. Each corallite in contact with thin distinct median suture. Thick, short septal
spines present especially at calice. Spines pointing slightly upwards. Squamulae absent. Mural poresnumerous, circular or ellipsoidal, large (0.125-0.25mm in diameter). Pores arranged longitudinally on
face, ca. 0.6-1.2mm apart. Tabulae absent or very rare.
Genus GERTHOLITES Sokolov, 1955
Type species. Pachypora curvata Waagen and Wentzel, 1886 of Gerth, 1921, p. 107.
Other species. G. xizangensis Lin, 1984; G.? tebaga Frugel, 1997.
Diagnosis (after Hill, 1981, p. F582). Corallum branching; branches of radially diverging corallites
increasing notably in diameter during growth and opening normal to surface of branch: edge of tabularium
raised in calice, variable in size and distant; septal spines visible in calices; skeletal thickening great;mural tunnels vermiform, anastomosing; tabulae rare to absent.

— 85 —

Gertholites ? sp. Plate 17, Figs. 9-10
Reg. No.
Sample: NUM-Fz-010811-34. One sample.
Specimen: NUM-Fz-010811-34-15.
Thin section: One NUM-Fz-010811-34-15-1.
Material, horizons and localities
The specimen was from the Upper Member of the Fukuji Formation, along the Ichinotani Valley,
Takayama City, Gifu Prefecture, central Japan (Loc.1).
Description
Corallum branching, cylindrical; branch 2.5mm in diameter, 11.6mm in height; corallites diverging
from axis to open at acute angle to surface, width 0.175-0.3mm. Walls in axial part 0.05-0.075mm thick,with peripheral stereozones 0.05-0.275mm thick. Each corallite in contact with thin, distinct mediansuture. Septal spines, squamulae and tabulae absent. Mural tunnels anastomosing; skeletal thickening
heavy.
Acknowledgements
I would like to thank Prof. M. Adachi and Associate Prof. H. Yoshida of Nagoya University for theirhelpful advice. I am indebted to Dr. N. MacLeod and Dr. B. Rosen of the British Museum (Natural
History) for their help and advice during the course of the study. Special thanks go to Ms. J. Darrell, Dr.
D. White and Dr. G. Clarke of the British Museum (Natural History), and Prof. C. Minjin of the Mongolian University of Science and Technology for their critical reading of the manuscript. I am grateful to
Mr. and Mrs. Karatani of Takayama City, Japan for their support during the field survey. Part of the work
was supported by a Grant-in-Aid from the Daiwa Anglo-Japanese Foundation.
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飛騨外縁帯デボン系福地層の床板サンゴ化石について
束田― その 1 ―和弘（TSUKADA Kazuhiro）名古屋大学博物館

飛騨外縁帯福地地域のデボン系福地層は保存良好なサンゴ化石を多産し，日本の古生界研究の上で重要な地層の一つである．筆者は現在，本層の床板サンゴ化石群集について検討を進めているが，その一部が明らかになったのでここに報告する．
本研究では，岐阜県高山市福地の一ノ谷沿いで福地層の７地点（（Locs. 1 to 7）より，合計 420kg の含サンゴ化石試料を採取した．そのうち１地点（Loc. 1）の約 63kg の試料について検討した結果，以下の床板サンゴ化石が同定された．Favosites goldfussi, F. flexuosus, F. sp. A, Squameopora hidensis, S.

zhanwaensis fukujiensis subsp. nov., S. cf. zhanwaensis, Sapporipora kamitakaraensis sp. nov., Sa.
karatanioum sp. nov., Heliolites wenxianicus, H. cf. gemina, H. ichinotaniensis sp. nov., H. ? sp.,
Helioplasma takayamaensis sp. nov., Pseudoplasmopora okuhidaensis sp. nov., P. cf. arguta, Striatopora
sp., Gertholites ? sp.

(Received 25 October 2005, accepted 15 November 2005)— 90 —

PLATE— 91 —

Plate 1
Favosites goldfussi d’Orbigny, 1850
Figs. 1 and 2 Transverse and longitudinal thin section. Fig. 3 Longitudinal thin section. Fig. 4 Transverse thin section.
Fig. 5 Longitudinal thin section. Fig. 6 Transverse thin section.

— 92 —

Plate 1— 93 —

Plate 2
Favosites flexuosus Kamei, 1955
Fig. 1 Longitudinal thin section. Figs. 2-3 Transverse thin section.
Favosites sp. A
Fig. 4 Longitudinal thin section. Fig. 5 Transverse thin section.

— 94 —

Plate 2— 95 —

Plate 3
Favosites sp. A
Fig. 1 Longitudinal thin section. Fig. 2 Close up view of Fig. 1. Fig. 3 Transverse thin section.
Squameopora hidensis (Kamei, 1955)
Fig. 4 Close up view of Fig. 5. Fig. 5 Transverse and longitudinal thin section.

— 96 —

Plate 3— 97 —

Plate 4
Squameopora hidensis (Kamei, 1955)
Figs. 1-4 Longitudinal thin section. Fig. 5 Transverse thin section.

— 98 —

Plate 4— 99 —

Plate 5
Squameopora hidensis (Kamei, 1955)
Fig. 1 Transverse thin section. Fig. 2 Transverse and longitudinal thin section.
Squameopora zhanwaensis fukujiensis subsp. nov.
Figs. 3-4 Longitudinal thin section. Figs. 5-7 Transverse thin section.

— 100 —

Plate 5— 101 —

Plate 6
Squameopora cf. zhanwaensis Lin and Huang, 1987
Figs. 1, 3 and 5 Longitudinal thin section. Figs. 2 and 4 Transverse thin section. The arrow in Fig. 2 shows squamula.

— 102 —

Plate 6— 103 —

Plate 7
Squameopora cf. zhanwaensis Lin and Huang, 1987
Figs. 1-5 Longitudinal thin section. Figs. 6-7 Transverse thin section.

— 104 —

Plate 7— 105 —

Plate 8
Squameopora cf. zhanwaensis Lin and Huang, 1987
Figs. 1-2 Longitudinal thin section.
Sapporipora kamitakaraensis sp. nov.
Figs. 3-4 Longitudinal thin section. Fig. 5 Transverse thin section.

— 106 —

Plate 8— 107 —

Plate 9
Sapporipora kamitakaraensis sp. nov.
Figs. 1-2 and 5 Longitudinal thin section. Fig. 3 Transverse thin section. Fig. 4 Close up view of Fig. 3.
Sapporipora karatanioum sp. nov.
Fig. 6 Transverse and longitudinal thin section.

— 108 —

Plate 9— 109 —

Plate 10
Sapporipora karataniorum sp. nov.
Fig. 1 Transverse thin section. Figs. 2-3 Close up view of Fig. 1. Fig. 4 Transverse and longitudinal thin section. Fig. 5
Close up view of Fig. 4. Fig. 6 Longitudinal thin section.

— 110 —

Plate 10— 111 —

Plate 11
Heliolites wenxianicus Zhang, 1981
Figs. 1 and 5 Longitudinal thin section. Figs. 2-4 Transverse thin section.

— 112 —

Plate 11— 113 —

Plate 12
Heliolites cf. gemina (Tchi, 1976)
Fig. 1 Transverse thin section. Figs. 2-3 Longitudinal thin section.
Heliolites ichinotaniensis sp. nov.
Fig. 4 Transverse thin section. Fig. 5 Close up view of Fig. 4. Figs. 6-7 Longitudinal thin section.

— 114 —

Plate 12— 115 —

Plate 13
Heliolites ? sp.
Fig. 1 Transverse thin section. Figs. 2-4 Longitudinal thin section.
Helioplasma sp. A
Fig. 5 Transverse thin section. Figs. 6-8 Longitudinal thin section.

— 116 —

Plate 13— 117 —

Plate 14
Pseudoplasmopora okuhidaensis sp. nov.
Fig. 1-2 Transverse and longitudinal thin section. Figs. 3 and 7 Transverse thin section. Figs. 4-6 Longitudinal thinsection.

— 118 —

Plate 14010811-34-5-2b010811-28-4-1

010811-33-4-1

010811-33-4-1— 119 —

Plate 15
Pseudoplasmopora cf. arguta Bondarenko, 1963
Figs. 1-2 and 5 Longitudinal thin section. Figs. 3-4 Transverse and longitudinal thin section. Fig. 6 Transverse thinsection.

— 120 —

Plate 15— 121 —

Plate 16
Striatopora sp.
Figs. 1, 2, 6, 8 and 9 Longitudinal thin section. Fig. 4 Close up view of Fig. 6. Figs. 3 and 5 Transverse thin section. Figs.
7, 10 and 11 Tangential section.

— 122 —

Plate 16— 123 —

Plate 17
Striatopora sp.
Figs. 1 and 6 Transverse thin section. Fig. 2 Close up view of Fig. 1. Fig. 3 Close up view of Fig. 5. Figs. 4-5 and Figs.
7-8 Longitudinal thin section.
Gertholites ? sp.
Fig. 9 Longitudinal thin section. Fig. 10 Close up view of Fig. 9.

— 124 —

Plate 17— 125 —