A monospecific assemblage of terebratulide brachiopods
in the Upper Cretaceous seep deposits of Omagari,
Hokkaido, Japan
ANDRZEJ KAIM, MARIA ALEKSANDRA BITNER, ROBERT G. JENKINS, and YOSHINORI HIKIDA
Kaim, A., Bitner, M.A., Jenkins, R.G., and Hikida, Y. 2010. A monospecific assemblage of terebratulide brachiopods in
the Upper Cretaceous seep deposits of Omagari, Hokkaido, Japan. Acta Palaeontologica Polonica 55 (1): 73–84.
The Campanian (Upper Cretaceous) seep carbonate at Omagari (Hokkaido, Japan) yields a monospecific association of
the terebratulide brachiopod Eucalathis methanophila Bitner sp. nov. The association is the only occurrence of brachio−
pods known from the post−Early Cretaceous history of chemosynthesis−based communities. Unlike many earlier
rhynchonellide−dominated hydrocarbon seep associations—which disappeared in Aptian times—this association is com−posed of chlidonophorid terebratulides. It is hypothesised here that large rhynchonellide brachiopods have beenoutcompeted from chemosynthesis−based associations by large chemosymbiotic bivalves (especially lucinids) and that
this seep association containing numerous terebratulide brachiopods originated as a result of immigration from the back−ground fauna settling in a seep that lacked numerous large bivalves but offered some hard substrates for brachiopod at−tachment. Some living chlidonophorids are known to settle around seep/vent localities or more generally in deep−waterhard−substrate settings. We review occurrences of brachiopods in chemosynthesis−based associations and show thatbrachiopods immigrated repeatedly to seep/vent environments. Eucalathis methanophila Bitner sp. nov. represents theoldest and single Mesozoic record of the genus. The new species is similar in ornamentation to three living species,
Indo−Pacific E. murrayi, eastern Atlantic E. tuberata, and Caribbean E. cubensis but differs in having a higher beak andwider loop. Additionally the studied species is nearly twice as large as E. tuberata.
Key wo r d s: Brachiopoda, Chlidonophoridae, Eucalathis, hydrocarbon seep, chemosynthesis−based community, Cam−panian, Cretaceous, Mesozoic, Japan.
Andrzej Kaim [kaim@twarda.pan.pl] and Maria Aleksandra Bitner [bitner@twarda.pan.pl], Instytut Paleobiologii
PAN, ul. Twarda 51/55, PL−00−818 Warszawa, Poland;
Robert G. Jenkins [robertgj@ori.u−tokyo.ac.jp], HADEEP, Ocean Research Institute, University of Tokyo, 1−15−1
Minamidai, Nakano−ku, Tokyo 164−8639, Japan;
Yoshinori Hikida [nmhikida@coral.ocn.ne.jp], Nakagawa Museum of Natural History, Hokkaido 068−0835, Japan.
Received 9 June 2009, accepted 23 October 2009, available online 27 October 2009.

Introduction
Although present in the oldest−known Silurian hydrothermalvent deposits (Little et al. 1999b) brachiopods seem to be ofdecreasing importance in chemosynthesis−based associations
through time. After an Early Cretaceous acme of brachiopod
occurrences in hydrocarbon seep deposits (Campbell 2006;
Kiel and Peckmann 2008) related to wide dispersal of rhyn−chonellide Peregrinella few younger seep/vent localities yield
any brachiopod fauna. Moreover, there is no unequivocal re−port of brachiopod obligate association to chemosynthesis−−based communities in Recent hydrothermal vents and hydro−
carbon seeps. Herein we summarise published information onbrachiopod occurrences in the chemosynthesis−based associa−
tions, stressing that the Omagari seep site is the latest−known
occurrence of a rich brachiopod association but one that is
dominated by chlidonophorid terebratulidines rather thanrhynchonellides.
Acta Palaeontol. Pol. 55 (1): 73–84, 2010

Institutional abbreviations.—UMUT, University Museum,
University of Tokyo, Tokyo, Japan; ZPAL, Institute of Paleo−
biology, Polish Academy of Sciences, Warszawa, Poland.
Other abbreviations.—L, length; W, width; T, thickness.

Geological setting
All specimens described in this paper came from the Omagari
site which yields probably the best researched Cretaceous hy−
drocarbon seep deposits in Japan (Hikida et al. 2003). The
10−meter−wide seep carbonate body is located on a small islet
in the Abeshinai River near the mouth of its tributary Osou−shinai River (Fig. 1). The seep carbonate in Omagari was firstmentioned by Hashimoto et al. (1967) who reported a dense
assemblage of tube−like trace fossils, which were later identi−
fied (Hikida et al. 2003) as possible vestimentiferan worm
doi:10.4202/app.2009.0068

74

ACTA PALAEONTOLOGICA POLONICA 55 (1), 2010

by carbonate crusts, thus being difficult to clean. The numberof examined specimens is 21; however, additional samples arepresent at Nakagawa Museum of Natural History, Hokkaido,
Japan.
The internal morphology was investigated by transverse
serial sections of two specimens. The distance between thesections was variable (between 0.04 and 0.1 mm). Subse−
quently acetate peels have been prepared for each section.
For the study of shell ultrastructure, two specimens were em−
bedded in Araldite 2020 resin, cut and polished, then etchedwith 5% HCl before coating with platinum for observation
under a scanning electron microscope. The SEM micro−
graphs were taken in the SEM laboratory of ZPAL using a
Philips XL−20 scanning microscope.

Review of the brachiopod
occurrences in chemosynthesis
based associations

Fig. 1. Geological map of the Nakagawa area, Hokkaido, showing the loca−
tion of the Omagari site. Modified from Takahashi et al. (2003).

tubes. Hikida et al. (2003) identified some fossils from this lo−
cality. Subsequently Kiel et al. (2008) formally described bi−
valves and Kaim et al. (2008, 2009) gastropods. The islet at
Omagari is composed mainly from Campanian (Upper Creta−
ceous) rocks of two facies: (i) worm−tube boundstone faciesand (ii) brecciated facies (Hikida et al. 2003). The former fa−
cies consists of solid carbonates containing numerous silici−
fied worm tubes while the other fossils are uncommon. The
brecciated facies consists of a mixture of brecciated carbon−
ates and siliciclastics (mudstone to sandstone clasts). All typesof rocks in brecciated facies contain numerous fossils but they
are easiest to recover from mudstone and siltstone. For morecomprehensive description of the locality the reader is referred
to Hikida et al. (2003) and Kaim et al. (2009).

Material and methods
The investigated specimens were extracted by wet−sieving(mesh size 0.5 mm) of weakly consolidated mudstone andsiltstone clasts of the brecciated facies because of practicalreasons. We have not observed any brachiopods in the worm−
tube boundstone facies. Although all our specimens are articu−
lated, some are damaged and/or crushed, and most are covered

Brachiopods are present in the earliest−known chemosyn−
thesis−based associations containing metazoan animals. The
lingulate brachiopod Pyrodiscus reported from the Silurian(?Ludlow) of Yaman Kasy, Ural Mountains, Russia (Little etal. 1999b) is the sole inarticulate brachiopod described so far
from chemosynthesis−based associations (Table 1). Septa−
trypa lantenoisi is an atrypid brachiopod reported by Barbieri
et al. (2004) from Upper Silurian carbonate mound in Mo−
rocco. Barbieri et al. (2004) interpreted Septatrypa−bearing
rocks as hydrocarbon seep deposits, however, later authors(Buggisch and Krumm 2005; Himmler et al. 2008) argued thatthe reported isotope data indicate methanogenesis rather thanmethane oxidation which typifies seep environment. The firstrhynchonellide brachiopods in chemosynthesis−based associ−ations appear in the Devonian. The Devonian Dzieduszyckia isa very large, sturdily ribbed rhynchonellide with a wide but
disjunct worldwide distribution. It is known from Poland,
Morocco, western America, Russia, Kazakhstan, and China(Campbell and Bottjer 1995b; Sandy 1995; Baliński and
Biernat 2003). Campbell and Bottjer (1995b) and Sandy(1995) speculated that Dzieduszyckia apparently thrived inchemosynthesis−based associations. Baliński and Biernat(2003) challenged this speculation showing that isotope datafrom some Moroccan and Polish localities does not meet thecriteria for seep environments. However, Baliński and Biernat(2003) measured mainly shell material with admixture of sedi−ment, and—as already pointed out by Campbell (2006)—car−bon and oxygen isotopes measured from shell material fromhydrocarbon seeps are expected to fall within a range close tonormal marine conditions. A relation of Dzieduszyckia−bear−ing deposits from western Meseta of Morocco to hydrocarbon
seepage has been confirmed by Peckmann et al. (2007). Re−
maining localities await critical evaluation. Another rhyn−chonellide, a medium−sized, ribbed Ibergirhynchia has been

KAIM ET AL.—CRETACEOUS SEEP BRACHIOPOD FROM JAPAN75

Table 1. A list of brachiopods occurring in the chemosynthesis−based associations. (*) Buggisch and Krumm (2005) and Himmler et al. (2008) ar−gued that the reported isotope data from Septatrypa−bearing localities indicate methanogenesis rather than methane oxidation which typifies seep en−
vironment.
Taxon
Lingulida
Pyrodiscus lorrainae Little, Maslennikov,
Morris, and Gubanov, 1999
Atrypida: Septatrypidae
Septatrypa lantenoisi (Termier, 1936)
Rhynchonellida: Halorellidae
Dzieduszyckia kielcensis (Roemer, 1866)
Dzieduszyckia intermedia (Termier, 1936)
Dzieduszyckia crassicostata(Termier and Termier, 1948)
Dzieduszyckia tenuicostata (Termier, 1936)
Dzieduszyckia bashkirica(Tschernyschev, 1887)
Dzieduszyckia sonora
Noll, Dutro, and Beus, 1984
Rhynchonellida: Peregrinellidae

Reference

Environment

Locality

Age

Little et al. (1999b)

hot vent

Ural Mountains, Russia

Silurian

MoroccoUpper Silurian

Campbell and Bottjer (1995b), Barbieri?cold seep*
et al. (2004)Baliński and Biernat (2003)
Baliński and Biernat (2003)?cold seepcold seep

Holy Cross Mts., Poland
MoroccoFamennian
FamennianBaliński and Biernat (2003)

cold seep

MoroccoFamennianBaliński and Biernat (2003)

cold seep

FamennianBaliński and Biernat (2003)

uncertain

MoroccoUrals, Kazakhstan,Tadzhikistan, ChinaBaliński and Biernat (2003)

barite seep

Nevada and Mexico

FamennianAnarhynchia gabbi (Ager, 1968)

Manceñido and Dagys (1992); Sandy(1995, 2001); Stefanoff and Sandy(1998)?cold seep

California, USA

Lower Jurassic

Anarhynchia cf. gabbi (Ager, 1968)

Little et al. (1999a)hot vent

California, USA

Pliensbachian,Jurassic

Biernat (1957), Thieuloy (1972), Ageret al. (1972), Campbell and Bottjer(1995a, b); Kiel and Peckmann (2008)

cold seep

Peregrinella multicarinata(Lamarck, 1819)
Peregrinella silesica (Ascher, 1906)
Peregrinella whitneyi (Gabb, 1869)
Peregrinella chisania
Sandy and Blodgett, 1996
Peregrinella gongboxueensis Sun, 1986
Peregrinella cheboensis Sun, 1986
Peregrinella bifurcata Sun, 1986
Peregrinella baingoinensis Sun, 1986
Peregrinella dongqoensis Sun, 1986
Peregrinella sp.
Rhynchonellida: Dimerellidae
Ibergirhynchia contraria (Roemer, 1850)
Cooperrhynchia schucherti (Stanton, 1895)
Terebratulida: Beecheriidae
Beecheria sp.
Terebratulida: Zeilleriidae
Modestella jeletzkyi Sandy, 1990

FamennianFrance, Switzerland,
Poland, Czech Rep.,?Late
Germany, Romania,
Berriasian−Hauterivian
Italy, Crimea, Caucasus
Carpathians, Czech Rep.
Hauterivian

Ascher (1906)
Campbell et al. (1993), Campbell and
Bottjer (1995 a, b)?cold seepcold seep

California, USA

Hauterivian

Sandy and Blodgett (1996)?cold seep

Alaska, USA

Valanginian−Hauterivian

Sun (1986)
Sun (1986)
Sun (1986)
Sun (1986)
Sun (1986)
Ortiz−Hernández and Martínez−Reyes(1993)?cold seep?cold seep?cold seep?cold seep?cold seep

Tibet, China
Tibet, China
Tibet, China
Tibet, China
Tibet, China

Valanginian
Valanginian−Hauterivian
Valanginian−Hauterivian
Valanginian−Hauterivian
Valanginian−Hauterivian?cold seep

MexicoNeocomian−Aptian

Gischler et al. (2003)
Sandy and Campbell (1994)

cold seepcold seepHarz, Germany
California, USA

Carboniferous
Tithonian, Jurassic

Von Bitter et al. (1990, 1992)?hot vent

Newfoundland, Canada

Early Carboniferous

Sandy (1990, 1995), Beauchamp andSavard (1992)

cold seep

Arctic Canada?Albian, Lower
Cretaceousthis paper

cold seep

Hokkaido, Japan

Campanian, Upper
Cretaceous

Campbell (2006), Campbell et al.
(2008)

cold seep

New Zealand

MioceneKuroda (1931); Tanaka (1959)
Majima et al. (2003, 2005)

cold seepcold seep

Honsiu, Japan
Kyushu, Japan

MiocenePliocene

Terebratulida: Chlidonophoridae
Eucalathis methanophila Bitner sp. nov.
Terebratulida: Terebratulidae?Liothyrella sp.
Terebratulida: Terebrataliidae
Coptothyris sinanoensis Kuroda, 1931
Unidentified brachiopod

doi:10.4202/app.2009.0068

76

reported by Gischler et al. (2003) from the Early Carbonifer−ous of Germany. A relation of Ibergirhynchia−bearing deposit
to hydrocarbon seepage has been confirmed by Peckmann etal. (2001). There are no confirmed records of chemo−
synthesis−based associations from the Permian and Triassic
though Sandy (1995) suggests that some Triassic Halorella
and Carapezzia could be seep−related as they are strikingly
similar to Jurassic Anarhynchia and Cooperrhynchia (see be−low) respectively and usually occur in isolated carbonate
lenses emplaced in otherwise clastic sediments (Sandy 1995,
2001). The Early Jurassic Peregrinelloidea from Siberia(Dagys 1968) is very similar and occurs in monospecific asso−
ciations in carbonate lenses (Sandy 1995). The seep environ−ments for Halorella, Carapezzia, and Peregrinelloidea needcritical evaluation and a confirmation from isotope analyses.
The brachiopod Anarhynchia is known from Pliensbachian(Lower Jurassic) hydrothermal vent deposits of the Franciscan
Complex in California (Little et al. 1999a, 2004). The genus
has also been recorded from Oregon and Argentina (Man−ceñido and Dagys 1992; Sandy 1995, 2001; Stefanoff andSandy 1998) but these localities are not confirmed as seep/vent sites. Cooperrhynchia has been reported from Tithonian(Upper Jurassic) seep carbonates in California (Sandy andCampbell 1994).
The Early Cretaceous is a period of ubiquitous occur−
rences of the rhynchonellide Peregrinella in ancient hydro−
carbon seep deposits. This brachiopod, as with Dzieduszyc−
kia, is characterised by cosmopolitan but disjunct occur−
rences (Campbell and Bottjer 1995a, b; Sandy 1995; Sandy
and Blodgett 1996; Posenato and Morsilli 1999; Kiel andPeckmann 2008). It is known from Alaska (Sandy et al.
1995), US Pacific Coast (Campbell and Bottjer 1995a, b),
Mexico (Ortiz−Hernández and Martínez−Reyes 1993; Sandy
1995), Crimea (Kiel and Peckmann 2008), Tibet (Sun 1986),
and numerous localities in Europe (Ascher 1906; Biernat
1957; Thieuloy 1972; Posenato and Morsilli 1999; Lazăr etal. 2005). Campbell and Bottjer (1995b) have suggested that
Peregrinella is an end−member of vent−seep endemic lin−
eages of rhynchonellides (Dzieduszyckia–Ibergirhynchia–
Peregrinella), a hypothesis challenged by Baliński and Bier−nat (2003) who noted that morphological characters of thecrura suggest different familial placement.
Terebratulide brachiopods are much less common in theancient seep deposits. A short−looped brachiopod Beecheria
has been reported from alleged “low−temperature vent associ−ations” of Lower Carboniferous in Newfoundland (von Bitter
et al. 1990, 1992; Sandy 1995). However, the vent nature ofthese associations is still under debate (Campbell 2006). An−other unidentified terebratulide has been reported by Beau−champ and Savard (1992) from a Lower Cretaceous cold seep
carbonate in Arctic Canada. The species, named Modestella
jeletzkyi by Sandy (1990), is the sole occurrence of Modestellain seep associations as all the other species of this genus(known mostly from Europe) are not considered to be seep−re−lated (Sandy 1995). Campbell (2006) and Campbell et al.
(2008) mentioned a terebratulide ?Liothyrella from a possible

ACTA PALAEONTOLOGICA POLONICA 55 (1), 2010

Miocene seep deposit in New Zealand. This identification,
however, has not been critically evaluated in a taxonomic
work.
In Japan brachiopods are known from three hydrocarbon
seep localities. The Campanian species in this study has beenreported as a “terebratulid brachiopod” by Hikida et al. (2003).
A single occurrence of Coptothyris sinanoensis in a Miocene
carbonate in Nagano Prefecture, later identified as hydrocar−
bon seep deposit, was described by Kuroda (1931) and listed
by Tanaka (1959). Majima et al. (2003, 2005) reported an un−
identified brachiopod from a Pliocene locality at Kuge Shrine
on Kyushu Island. However, Majima et al. (2005: 89) consider
these “...Pliocene brachiopods associated with a cold seep as−
semblage ... not to be a chemosynthetic species because thebrachiopods are rare in the assemblage and have also been col−lected from the same horizon about 300 m away, where no ev−
idence of a seep is present.”
There are no extant brachiopods that are unequivocally as−sociated with vent/seep environments (Campbell 2006; Peck−mann et al. 2007). Some species are recorded in the vicinities(Zezina 2000; Lee et al. 2008) but there is no evidence they are
obligatory members of chemosynthesis−based communities.
Zezina (2000) reported a number of terebratulide brachiopods
living in the vicinity of hydrothermal vents. These include
Terebratulina kiiensis, Laqueus blanfordi, Macandrevia ame−ricana, Aneboconcha smithi, Nanacalathis atlantica, andAbyssothyris sp. Two other species (Platidia anomioides andP. concentrica) were found in a volcanic caldera in the Juan de
Fuca Ridge (Zezina 1990). Zezina (2000) noted that several
species (T. kiiensis, L. blanfordi, M. americana, Abyssothyrissp.) in these locations were represented exclusively by indi−viduals with no mature gonads, i.e., they could not breed there
while A. smithi is represented only by dwarfed individuals.
Lee et al. (2008) recently reported Melvicalathis macroctena(Zezina, 1981) attached to basalts and manganese crusts cov−
ering rocks in the Southeast Indian Ridge and southeast Pa−
cific but there is no obvious relation of the brachiopods to hy−
drothermal vents. Lee et al. (2008) suggest that Melvicalathis
macroctena is a primary opportunistic coloniser of available
hard substrates including fresh glassy basalt.

Systematic palaeontology(by M.A. Bitner)

Phylum Brachiopoda Duméril, 1806
Subphylum Rhynchonelliformea Williams, Carlson,
Brunton, Holmer, and Popov, 1996
Class Rhynchonellata Williams, Carlson, Brunton,
Holmer, and Popov, 1996
Order Terebratulida Waagen, 1883
Suborder Terebratulidina Waagen, 1883
Superfamily Cancellothyridoidea Thomson, 1926
Family Chlidonophoridae Muir−Wood, 1959

KAIM ET AL.—CRETACEOUS SEEP BRACHIOPOD FROM JAPAN

77

1 mm

A4

A1

A2

A3

1 mm

A5

1 mm

1 mm

B1

B3

B2

1 mm

D
B4

500 µm

F1

1 mm

1 mm

C

F2

500 µm

E

Fig. 2. Terebratulide brachiopod Eucalathis methanophila Bitner sp. nov., Campanian, Omagari, Japan. A. Holotype, UMUT MB30198, ventral (A1),
dorsal (A2), lateral (A3), and anterior (A4) views of complete specimen, enlargement (A5) of the posterior part. B. Paratype, UMUT MB30199, ventral (B1),
dorsal (B2), and lateral (B3) views of complete specimen, enlargement (B4) of shell surface to show nodular ornament. C–E. Paratypes, UMUT
MB30200−30202, dorsal views of complete specimens. F. UMUT MB30207, shell surface of ventral valve (F1) and enlargement (F2) to show details of
ornamentation. All SEM.

Subfamily Eucalathinae Muir−Wood, 1965
Genus Eucalathis Fischer and Oehlert, 1890
Type species: Terebratulina murrayi Davidson, 1878 by original desig−
nation of Fischer and Oehlert (1890: 72); Recent, 28°33’S, 177°50’W,
near Kermadec Islands, depth 1080 m (Davidson 1878).

Remarks.—Today Eucalathis has a worldwide distribution,
being known from the Caribbean, Atlantic, Indian, Pacific,
and Antarctic Oceans (Cooper 1973a, b, c, 1977, 1981a, b;
Foster 1974; Brunton and Curry 1979; Logan 1983, 1988,
1990, 1998, 2007; Zezina 1985, 1987, 2006; Hiller 1986,
doi:10.4202/app.2009.0068

78

ACTA PALAEONTOLOGICA POLONICA 55 (1), 2010

0.10

0.22

0.40

1.36

0.51

0.58

1.41

1.63

0.71

0.86

0.78

1.47

1.52

1.67

1.57

1.80

1.74

1.87

1.93

2.06

2.17

2.22

2.27

2.31

2.36

2.40

2.46

2.50

1 mm
2.65

2.70

Fig. 3. Transverse serial sections of terebratulide brachiopod Eucalathis methanophila Bitner sp. nov. through specimen UMUT MB30203, Campanian,Japan. Original dimensions of the specimens: L = 4.9 mm, W = 4.4. mm, T = 2.0 mm. Numbers indicate distance in mm from the tip of the ventral umbo.

1994; Dawson 1991; Laurin 1997; Gaspard 2003; Álvarez
and Emig 2005; Bitner 2006, 2008, 2009). The 13 Recent
species assigned to this genus exhibit a wide bathymetricrange from 185 to 3870 m (Logan 2007); one deep−water
species, E. macroctena, included to Eucalathis by Zezina(1981), has recently been transferred into a new genus Melvi−
calathis Lee, Lüter, and Zezina, 2008 (Lee et al. 2008).
Melvicalathis can be easily distinguished from Eucalathis by
its broad, smooth, triangular in cross−section ribs.
In the fossil record Eucalathis is very rare and has been so
far reported from the Eocene of Eastern Coast of America(Cooper 1988) and Miocene of Italy (Davidson 1870; Lee etal. 2006). Those fossil species are coarsely ribbed, and thus
differ strongly from the specimens described here. Apartfrom morphological differences, the great stratigraphical gap
between the hitherto described Eucalathis species and theLate Cretaceous species from Japan justifies the decision to

Fig. 4. Recontruction of internal structures of Eucalathis methanophila
Bitner sp. nov. based on transverse serial sections (Fig. 3) and 3D digital re−construction (see at http://app.pan.pl/SOM/app55-Kaim_etal_SOM.pdf).
Drawing by Ewa Widłak−Kaim.

KAIM ET AL.—CRETACEOUS SEEP BRACHIOPOD FROM JAPAN

A1

B1

50µm

100µm

79

50µm

A2

50µm

B2

Fig. 5. Terebratulide brachiopod Eucalathis methanophila Bitner sp. nov., Campanian, Omagari, Japan. A. UMUT MB30197, transverse section of ventral
valve, visible fused fibres (A1), visible boundary of primary and secondary layers (A2). B. UMUT MB30203, ventral valve, transverse sections of the wholeshell (B1), visible boundary between primary and secondary layers, and punctae (B2). All SEM.

erect a new species. The studied specimens represent not
only the first record of Eucalathis from Japan but also theoldest occurrence of the genus.

Eucalathis methanophila Bitner sp. nov.
Figs. 2–5.
Etymology: From methane and Latin suffix, phila, meaning to like. Re−
fers to its occurrence at the methane seep.
Type material: Holotype: UMUT MB30198 (Fig. 2A1–A5), articulatedshell; Paratypes: UMUT MB30199−30202 (Fig. 2B–E), articulatedshells.
Type locality: Omagari site, Nakagawa area, Northern Hokkaido, Japan.
Coordinates 44° 39’ 26’’N, 144° 2’ 25’’E.
Type horizon: Hydrocarbon seep deposits of Omagari Formation, Cam−panian, Upper Cretaceous.

Diagnosis.—Eucalathis of medium size with surface orna−
mented by numerous beaded ribs, high beak with large, oval
foramen, anterior commissure rectimarginate, loop wide with
subparallel descending branches narrowed anteriorly.
Measurements (in mm).—See Table 2.
Description.—External morphology: Shell small (maximum
length 5.7 mm), auriculate, variable in outline from widely
subtriangular to elongate oval, with maximum width usuallyat two thirds of shell length. Shell biconvex with ventral
valve slightly more convex. Shell surface covered with nu−
merous beaded, tuberculate ribs (20–30) which increase in
number by intercalations and bifurcations; growth lines dis−tinct (Fig. 2B4, F1, F2). Lateral commissures nearly straight;
anterior commissure rectimarginate to incipiently uniplicate.
Hinge line wide, slightly curved. Beak relatively high, erect
to suberect with sharp, distinct ridges. Foramen large, oval;
disjunct deltidial plates form narrow ridges.
Table 2. Measurements of Eucalathis methanophila Bitner sp. nov.
Abbreviations: L, length, W, width, T, thickness.
Specimen no.
UMUT MB30197
UMUT MB30198 (holotype)
UMUT MB30199 (paratype)
UMUT MB30200 (paratype)
UMUT MB30201 (paratype)
UMUT MB30202 (paratype)
UMUT MB30203
UMUT MB30204
UMUT MB30205
UMUT MB30206

L
5.7
4.1
4.3
4.0
4.4
5.2
4.9
5.3
3.2
3.2

W
4.9
3.8
3.7
3.2
4.1
4.3
4.4
4.9
3.0
3.1

T
?2.7
1.95
1.8
1.9
2.2
2.2
2.0
2.4
1.6
1.4

W/L
0.86
0.93
0.86
0.8
0.93
0.83
0.9
0.92
0.93
0.97

T/L
0.47
0.47
0.49
0.47
0.5
0.42
0.41
0.45
0.5
0.44

doi:10.4202/app.2009.0068

80Internal morphology: Two specimens were sectioned ofwhich the better preserved is figured (Fig. 3). Pedicle collar
present. Teeth long, smooth, nearly horizontally inserted
into large dental sockets. Inner socket ridges massive, long.
No hinge plates observed. Crura short, massive. Loop shortand wide with subparallel to slightly divergent descending
branches that narrow anteriorly (Fig. 4). Transverse band
not preserved in either of the sectioned specimens.
Shell microstructure: Two specimens were studied. The
shell is composed of two layers: a primary microgranularlayer and a secondary fibrous layer (Fig. 5). In both cases theshell is diagenetically modified, fragmentary fibres of thesecondary layer are completely fused (Fig. 5A1). The pri−mary layer is 27–53 μm thick in the ribs and 12–25 μm in
sulci. The secondary layer is 144–205 μm thick in the ribs
and 100–114 μm in sulci. The total thickness of the shell is
127 to 241 μm. The punctae are more densely distributed in
the rib regions (Fig. 5).
Remarks.—The investigated specimens display, both exter−
nally and in a loop character, typical features of the genus
Eucalathis. This genus was erected for the specimens simi−
lar to Terebratulina but having disjunct crural processes(Fischer and Oehler 1890); in Terebratulina crural pro−
cesses are united to form a ring. In the ornamentation of thenumerous fine ribs the specimens from Japan resemble
three living species, Indo−Pacific E. murrayi (Davidson,
1878), eastern Atlantic E. tuberata (Jeffreys, 1874), andCaribbean E. cubensis Cooper, 1977 but differ in having
higher beak and wider loop (Davidson 1878, 1886; Cooper
1977, 1981b; Zezina 1987). Additionally the studied speci−
mens are nearly twice as large as E. tuberata.
Stratigraphic and geographic range.—Campanian (Upper
Cretaceous) seep carbonate at Omagari (Hokkaido, Japan).

Discussion
Brachiopods are known from the chemosynthesis−based as−sociations since Silurian times (Little et al. 1999b) and they
were important members from Devonian up to Early Creta−
ceous constituting at some settings monospecific clusters ofnumerous individuals (Campbell and Bottjer 1995a, b; Little
et al. 1999a; Campbell 2006; Kiel and Peckmann 2008). No
Cainozoic fossil and Recent brachiopods are known to besustained by chemosymbiosis. However, some Devonian(Dzieduszyckia) and Early Cretaceous (Peregrinella) rhyn−chonellides attained large sizes at the hydrocarbon seeps as ischaracteristic of species from other phyla (e.g., molluscs)
hosting chemosymbiotic bacteria. Since the demise of Pere−grinella, brachiopods have become rare elements in the seep/vent environments and nowadays—although still found oc−
casionally nearby—there is no evidence that any brachiopod
is an obligatory member of such community. Therefore theoccurrence of Eucalathis methanophila in the Campanianhydrocarbon seep deposits is surprising. Firstly, it appears in

ACTA PALAEONTOLOGICA POLONICA 55 (1), 2010

relatively large numbers in the seep carbonate while it is un−
known from ambient strata and, secondly, the brachiopod be−longs to the Terebratulida and not to the Rhynchonellida (incontrast to the majority of older taxa). As outlined above,
terebratulides are known from occasional occurrences at ornear both ancient and modern chemosynthesis−based com−munities. Nevertheless all Recent examples are considered
as “normal” deep water taxa and early colonisers of the rocky(usually basaltic) substrate (Lee et al. 2008). It is worth
noting that at least two Recent eucalathines (Nanacalathis
atlantica and Melvicalathis macroctena) are reported fromthe vicinity of hydrothermal vents (Zezina 2000; Lee et al.
2008). Terebratulina tauriniensis described by Davidson(1870) from a Miocene locality at Gassino near Turin is ap−
parently another species of Eucalathis. Although this locality
has not been listed as hydrocarbon seep, several outcrops of“Calcari a Lucina” are known from this region. These car−bonate deposits have been interpreted by Taviani (1994) as
Miocene hydrocarbon seep deposits. Eucalathis tauriniensis
has been described by Davidson (1870) more than a century
before identification of the first hydrocarbon seep faunas andnow it is difficult to trace the exact Eucalathis−bearing local−
ity of Gassino. Otherwise eucalathines are unknown from thefossil record apart from two occurrences in the Eocene of
Eastern Coast of Northern America (Cooper 1988).
All articulate brachiopods are primary suspension feeders
collecting their food by a lophophore wedged between twoshells. They feed on suspended organic matter and do notpossess a functional anus (James et al. 1992). No endo−
symbiotic interactions have been observed for this group ofanimals. As mentioned above, some large rhynchonellide
brachiopods have been suspected of chemosynthetic metab−olism or at least as well adapted to environments typified byhigh concentrations of reduced chemicals (Campbell et al.
1993; Campbell and Bottjer 1995a, b). The ubiquitous occur−
rence of E. methanophila at Omagari suggests that it waswell adapted to a seep environment but its small size makes it
unlikely to have hosted any bacterial endosymbionts. Zezina(2000) suggested that most of the brachiopods in the vicinityof hydrothermal vents were immature and/or dwarfed. The
specimens of E. methanophila from Omagari are of average
size for eucalathines and most of them are seemingly adultspecimens. The size of the pedicle foramen suggests that it
was functional. Eucalathines, however, are known to possessa number of pedicle shapes. Lee et al. (2008) reported for
Melvicalathis macroctena short, branched, or brush−likepedicles while Bromley and Surlyk (1973) illustrated Euca−
lathis murrayi with the pedicle divided into rootlets immedi−
ately posterior to the pedicle opening. A brachiopod having ashort pedicle usually attaches to rocky substrate whilebrachiopods having branched or brush−like pedicles attach tosmall hard particles in the loose sediment. Thus E. methano−
phila may have been attached to exhumed seep carbonate, at−tached to worm tubes projecting above the seafloor, or an−
chored to sediment around the seep. The seep carbonate nor−mally precipitates subsurface within sediments (Ritger et al.

KAIM ET AL.—CRETACEOUS SEEP BRACHIOPOD FROM JAPAN

1987) and there is no direct indication that the carbonate waslater exhumed at Omagari seep. Nevertheless, the breccia−
tion events in Omagari seep described by Hikida et al. (2003)
may suggest that—at least in some periods—the carbonate
could be exhumed due to enhanced/explosive discharge ofthe methane. Moreover, some carbonate cements, e.g., iso−pachous rim cements, precipitated around worm tubes whichwere free of sediment inside (Takahashi et al. 2007). It sug−gests that the cements precipitated above the sea floor. Last
but not least the brachiopods could have attached to theworm tubes, which are extraordinarily common in Omagari
while relatively rare in other seep sites in Japan (Jenkins et al.
2007). The presence of these hard substrates could explain
why E. methanophila is common in Omagari while absent at
any other seep sites in the Upper Cretaceous of Hokkaido(AK and RGJ unpublished data).
Campbell and Bottjer (1995a) suggested that the rhyn−chonellide brachiopods were displaced in chemosynthesis−
based associations by chemosymbiotic bivalves at the end of
Early Cretaceous. Indeed demise of Peregrinella happened
around the same time as the onset of the modern, mollusc−
dominated seep fauna (Campbell and Bottjer 1995a; Kiel andLittle 2006; Kiel and Peckmann 2008). Kiel and Peckmann(2008) have demonstrated that the presence or absence ofdominant brachiopods is inversely correlated with the pres−ence or absence of dominant lucinids. They suggested, how−ever, that mass occurrences of thick−shelled brachiopods
made it difficult for lucinids and other infaunal bivalves tosurvive at seep sites (Kiel and Peckmann 2008: 757). We are
inclined rather to the suggestion of Campbell and Bottjer(1995a) that inversely, the increasing competition fromseep−adapted bivalves (including lucinids)—many of which
hosted chemosymbiotic bacteria—might be a plausible ex−planation of the demise of brachiopods from chemosynthesis
based associations. Such an explanation would parallel thegeneral trend in brachiopod decline during Mesozoic times.
Numerous authors (Gould and Calloway 1980; Thayer 1985,
1986; Ager 1986) have suggested that after the P−T crisis
brachiopods were outcompeted by bivalves in a majority ofenvironments. Still there are some Recent examples wherebrachiopods successfully coexist with bivalves (Lee 2008).
Large bivalves, especially lucinids are uncommon at
Omagari (Kiel et al. 2008) compared to gastropods, for ex−
ample (Kaim et al. 2009). In this respect the Omagari site isunusual among other Upper Cretaceous and younger seepsites. Scarcity of such bivalves at Omagari might explain theabundant presence of E. methanophila. However, it remains
unclear why the Omagari seep was settled by terebratulides
and not by rhynchonellides. Lee (2008) has shown thatrhynchonellides declined more rapidly than terebratulides af−
ter Middle Jurassic diversity zenith. By the Cretaceous rhyn−chonellides were much less common, less ecologically dif−ferentiated, and taxonomically diverse than terebratulides
and therefore immigration of the latter into Campanian seep
communities might be from purely stochastic reasons. Thishypothesis might be also supported by the lack of any reports

81

of rhynchonellides from the vicinity of Recent seep/vent lo−
calities. On the other hand, the oldest seep related sediments
reported so far from Japan came from Albian (Kaim andJenkins 2008, Kaim et al. 2009) while the youngest occur−
rence of Peregrinella is recorded from Neocomian–Aptianof Mexico (Ortiz−Hernández and Martínez−Reyes 1993),
therefore it remains unknown if Peregrinella was present in
the vicinity of present−day Japan during the Early Cretaceous
times. Geographically closest are the occurrences reported
by Sun (1986) from Tibet.

Conclusions
Eucalathis methanophila, as with some other brachiopods
associated with hydrocarbon seeps in the geological past,
constitutes a monospecific association with molluscs andworm tubes in the Campanian methane seep site at Omagari.
Although it still remains disputable if the species was an
obligatory member of hydrocarbon seep community, it may
have been adapted to thrive in such an extreme environment.
E. methanophila, unlike most of the earlier brachiopods as−sociated with hydrocarbon seeps, belongs to the Eucala−thinae, a subfamily within Terebratulida. Two species of thissubfamily are known from the vicinity of hydrothermal vents
although they are considered to be deep water opportunists.
The occurrence of E. methanophila in Omagari may suggestthat it was well adapted to environments typified by highconcentrations of reduced chemicals. If so it would be the lat−
est record of a brachiopod species closely connected to achemosynthesis−based association. The increasing impor−tance of chemosymbiont−bearing bivalves at Cretaceous hy−
drocarbon seeps might be a plausible explanation of the de−mise of brachiopods from these environments. The presenceof numerous brachiopods in Omagari might be related to rel−
atively low abundance of bivalves at this locality and/or pres−ence of numerous worm tubes serving as a source of hardsubstrate for attachment.

Acknowledgements
We thank Zbigniew Strąk (ZPAL) for help in preparing serial sections.
Thomas Kunze (University of Munich, Germany) prepared the digita−
lized 3−D reconstruction of the brachidium. Ewa Widłak−Kaim (Cele−
stynów, Poland) is thanked for the art reconstruction of the brachiopod
internal structures. Kazushige Tanabe (University of Tokyo, Japan) isheartily acknowledged for his support during this research. Daphne E.
Lee (University of Otago, Dunedin, New Zealand) and Jörn Peckmann(University of Bremen, Germany) are thanked for constructive peer re−views. The research of AK was partially supported by the Japan Society
for the Promotion of Science (JSPS) Postdoctoral Fellowship for For−eign Researchers and associated research grant number 17.05324 (pro−ject number 050500000614). The research of RGJ was financially sup−ported by the Nippon Foundation−HADal Environmental Science Edu−
cation Program (HADEEP).
doi:10.4202/app.2009.0068

82

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