Antiquity of the substrate choice among acmaeid limpets
from Late Cretaceous chemosynthesis−based communities
ROBERT G. JENKINS, ANDRZEJ KAIM, AND YOSHINORI HIKIDA
Jenkins, R.G., Kaim, A., and Hikida, Y. 2007. Antiquity of the substrate choice among acmaeid limpets from Late Creta−
ceous chemosynthesis−based communities. Acta Palaeontologica Polonica 52 (2): 369–373.
Two Campanian methane seep sites in the Nakagawa area of Hokkaido (northern Japan) yield fossils of the limpet genera
Serradonta and Bathyacmaea that appear to have had the same substrate preference as do their modern counterparts.
Serradonta cf. vestimentifericola was a species having an elongated and strongly compressed shell adapted to living on
vestimentiferan tubes, like its modern relatives. Bathyacmaea cf. subnipponica was an acmaeid with a relatively elon−
gated shell but with a more rounded aperture than Serradonta and thus apparently attached to small hard objects otherthan worm tubes. One Bathyacmaea specimen was found attached in situ to an ataphrid gastropod shell. The restrictedpresent−day distribution of Serradonta possibly reflects its spreading route exclusively through the hot vent and cold seep
communities settled by vestimentiferans.
Key wo r d s: Gastropoda, paleoecology, chemosynthetic community, methane−seep, Cretaceous, Hokkaido, Japan.
Robert Jenkins [robert@um.u−tokyo.ac.jp], University Museum and Department of Earth and Planetary Science, Uni−
versity of Tokyo, Tokyo 113−0033, Japan;
Andrzej Kaim [kaim@twarda.pan.pl], Institute of Paleobiology, Polish Academy of Sciences, PL−00−818 Warszawa,
Poland and Department of Earth and Planetary Science, University of Tokyo, Tokyo 113−0033, Japan;
Yoshinori Hikida [nmhikida@coral.ocn.ne.jp], Nakagawa Museum of Natural History, Hokkaido 068−0835, Japan.

Introduction
Since the surprising discovery of high biomass communities
of novel animals at hydrothermal vents on the Galápagos Rift
(Lonsdale 1977), studies of vent ecosystems have revealed
new modes of life in the animal kingdom. The trophic struc−ture of such communities is based on chemoautotrophic bac−teria that gain energy through the oxidation of reduced com−pounds, such as hydrogen sulfide or methane (for reviews
see Van Dover 2000; Campbell 2006). In the 30 years sincethose exciting discoveries similar chemosynthetic communi−ties have been found at various other sites in the world’s
oceans where bottom water is enriched with reduced com−pounds, e.g., cold−seeps, sunken whale carcasses, sunken
drift wood and even the rotting cargo of a sunken ship (Mar−shall 1988; Smith et al. 1989; Dando et al. 1992; Tunnicliffe
1992; Hasegawa 1997; Sibuet and Olu 1988; Van Dover
2000).
The fossil record of chemosynthesis−based communities
goes back to the Silurian (Barbieri et al. 2004; Little 2002;
Little and Vrijenhoek 2003; Campbell 2006). However, thefaunal composition of these communities underwent pro−found modifications since then. In the Palaeozoic brachio−
pods were predominant, but in the Mesozoic these werelargely replaced by molluscs (Campbell and Bottjer 1995;
Little and Vrijenhoek 2003). Molecular analyses of the mod−ern animals endemic to chemosynthetic communities also in−
dicate a late Mesozoic or even Cenozoic origin (for molecu−
Acta Palaeontol. Pol. 52 (2): 369–373, 2007

lar data see, e.g., Peek et al. 1997; Halanych et al. 1998;
McArthur and Koop 1999 and for discussion Van Dover et
al. 2002; Little and Vrijenhoek 2003; Kiel and Little 2006).
In order to elucidate the Mesozoic change in chemosynthesis
based faunas, it is important to trace autecology of specifictaxa. Herein we provide such a case study for two acmaeid
limpet gastropods, Bathyacmaea cf. subnipponica and Ser−radonta cf. vestimentifericola, found at two Late Cretaceous
methane−seep sites from the Nakagawa area of Hokkaido
in northern Japan. The Recent species of Serradonta andBathyacmaea are restricted to methane−seeps and hydrother−mal vents (Warén and Bouchet 2001) and herein we reportthat their attachment behavior was established as early as theLate Cretaceous.
Institutional abbreviations.—DEPUT, Department of Earthand Planetary Science, Faculty of Science, University ofTokyo, Japan; UMUT, University Museum, University ofTokyo, Tokyo, Japan.

Material and methods
The gastropods were found in the Omagari and Yasukawa
methane−seep sites (for the seep descriptions see Hikida et al.
2003 and Jenkins et al. 2007 respectively) located along theAbeshinai River, Nakagawa area, Hokkaido (Fig. 1). The lo−calities are Campanian (Late Cretaceous) seep carbonates
surrounded by siliciclastic sediments of the Omagari Forma−
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ACTA PALAEONTOLOGICA POLONICA 52 (2), 2007

tion in the Yezo Group (Takahashi et al. 2003; Jenkins et al.
2007).
The fossils were extracted by wet−sieving weakly consol−
idated muddy sediments (mesh size 0.5 mm). The samples
were first examined using a Keyence digital microscope
VHX−500 and detailed observations were carried out with a
Hitachi S−2400S scanning electron microscope at the
DEPUT. The latter required coating the specimens with plat−inum−palladium.
Carbonate isotopic signatures are expressed per mil rela−tive to the PDB standard.

Geological background
The Omagari methane−seep deposit consists of an approxi−
mately 10 m wide carbonate body having a very negative
carbonate carbon isotopic composition (as low as –45‰
versus PDB) (Hikida et al. 2003). The Yasukawa meth−ane−seep deposits are composed of several carbonate bod−ies, none of which exceeds 2 m maximum diameter. Thesealso have very negative carbon isotopic signatures (–43.5‰
versus PDB) (Jenkins et al. 2007). They have brittle andductile fabrics, probably caused by strong flow rates(Jenkins 2006). Fossils at the two methane−seep sites arepresent both within and near the carbonate bodies (see also
Hikida et al. 2003; Amano et al. in press; Jenkins et al.
2007). Some of the fossils closely resemble modern ventand seep taxa (compare e.g., Tunnicliffe 1991; Sibuet andOlu 1988; Warén and Bouchet 2001). The most striking ex−
amples are probable vestimentiferan worm tubes (Fig. 2D),
acmaeid limpets (Serradonta cf. vestimentifericola andBathyacmaea cf. subnipponica: Fig. 2A and B), lucinid,
thyasirid, solemyoid, and nuculid bivalves. However, thecomponent ratios of the fossils differ significantly between
the Yasukawa and Omagari sites. The epifaunal vesti−
mentiferans and acmaeid limpets flourished at the Omagari
methane−seep but are much less common at the Yasukawa
site. In contrast, infaunal thyasirid and solemyoid bivalves
were much more common at the Yasukawa methane−seep.
Detailed taxonomic treatment of these faunas will be pub−lished elsewhere.

Results
The fossil Serradonta cf. vestimentifericola specimens havean extremely tall and compressed shell (Fig. 2A) and strik−
ingly resemble the Recent S. vestimentifericola. The shellsurface has only growth lines which are curved towards the
apex on the anterior and posterior sides of the shell (Fig.
2A3). The aperture of Serradonta found at the Omagari and
Yasukawa sites neatly fits the worm tubes found at the same
localities. The mean width of Serradonta shells from theOmagari site is 1.85 mm (45 specimens measured) while theFig. 1. Geological map of the Nakagawa area, Hokkaido, showing the loca−
tion of the Yasukawa and Omagari sites. Modified from Takahashi et al.
(2003).

mean diameter of the worm tubes is 1.42 (68 specimens mea−
sured) but both samples were from slightly different areas ofthe outcrop and it cannot be excluded that the tubes wereslightly wider at the collection site of Serradonta. Indeed,
Hikida et al. (2003) reported tube diameters at the Omagari
site ranging from 1 to 4 mm.
Bathyacmaea cf. subnipponica also has an elongatedshell in the early stages (Fig. 2B), but its aperture expands
later in the ontogeny. The shell is ornamented by strong con−centric growth lines with small knobs. The aperture is weakly
concave dorsally and its outline becomes irregular. One
specimen of B. cf. subnipponica was found attached to an
ataphrid shell in a bulk sediment sample collected from the
Yasukawa methane−seep (Fig. 2C). Its aperture line fits per−fectly to the ataphrid gastropod shell surface and there is no
sediment between the two shells (Fig. 2C).

Discussion
A tall shell shape is rather unusual for limpets (Denny 2000)
and is observed mostly among those species utilizing otherorganisms of limited surface area as a substrate. For exam−
ple, some groups of Recent limpets have both tall and flat

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371

Fig. 2. Campanian (Late Cretaceous) molluscs and probable vestimentiferan fossils from the Yasukawa (A–C) and Omagari (D) sites (Hokkaido, Japan).
A. Serradonta cf. vestimentifericola UMUT MM29351. Apical (A1), lateral (A2) and anterior (A3) views. B. Bathyacmaea cf. subnipponica UMUT
MM29352. Apical (B2), lateral (B3) and anterior (B3) views. C. Acmaeid limpet Bathyacmaea cf. subnipponica attached to ataphrid gastropod (UMUT
MM29353). Lateral (C1, C2) and apical (C3) views; close up of the limpet: photograph (C4) and explanatory drawing (C5). D. Tube of ?vestimentiferanworm (UMUT MW29354).

species in the same genus, the shape of which depends
strictly on the substrate utilization. The Patelloida heroldigroup is a striking case, where P. conulus on living Batillaria
has a tall conical shell, but P. heroldi on rocks and P.
pygmaea on oyster beds show more depressed profiles likeother typical patellogastropod limpets (Sasaki and Okutani
1994, Nakano and Ozawa 2005; Nakai et al. 2006). In these
species, the difference in habitat selection has resulted in re−productive separation as revealed by allozyme and mito−
chondrial gene analyses (Nakano and Ozawa 2005; Nakai etal. 2006).
The shape of the fossil Serradonta cf. vestimentifericola
specimens, with their concave apertures and extremely tallshells (Fig. 2A), is just like that of the modern species, whichattaches to vestimentiferan tubes (Okutani et al. 1992; Sasaki
et al. 2003), and, indeed, fossil tubes, that are probably thoseof vestimentiferans occur abundantly in the Omagari meth−ane−seep carbonate body. Both Serradonta and vestimenti−
feran tubes are relatively rare at the Yasukawa methane−seepsite. These facts strongly suggest that S. cf. vestimentiferi−cola used the tubes of the vestimentiferans as a preferred sub−strate (Fig. 3). Moreover, it seems that the relatively small di−ameter of the worm tubes in the Campanian methane seeps
provided the selective pressure for these limpets to keep an
elongate and strongly compressed shell shape.
There are also other Recent limpets known to live on thevestimentiferan tubes, e.g., several species of Lepetodrilus.
The species L. fucensis living on the small sized tube worm

Ridgeia piscesae have elongated shells indeed (Desbruyères
et al. 2006) but the other small−sized species (e.g., L. ele−
vatus, L. ovalis, and L. pustulosus) have relatively flat shells
because they live on the large sized tubes of Riftia pachy−
ptila, a tube surface which would appear to be almost flat rel−
ative to the small−sized Lepetodrilus (compare Desbruyères
et al. 2006: 89–90).
Bathyacmaea cf. subnipponica probably lived on sub−strates of small size judging from its elongated shell shapeand supported by direct observation of the specimen still at−tached to the ataphrid shell (Fig. 2C). Living Bathyacmaea
were observed attached to bivalve shells, pieces of rock andeven crawling through the sandy sediment or browsing on
bacterial mats (Okutani et al. 1992; Hashimoto et al. 1995;
Sasaki et al. 2005). Shell elongation depending on the sub−strate type affects not only patellogastropods but also otherlimpet gastropods living in chemosynthesis−based communi−ties, e.g., a moderately elongated cocculiniform limpet
Pyropelta corymba lives attached to the gastropod Provanna
laevis (Warén and Bouchet 2001) but it is not known fromvestimentiferan tube worms (Desbruyères et al. 2006).
The Recent species of Serradonta are restricted exclu−
sively to the marine waters of the Japanese Archipelago (e.g.,
Sasaki et al. 2005) and its fossil record is also restricted to Ja−pan (Hikida et al. 2003; Jenkins et al. 2007; own unpublisheddata). This relatively weak dispersal over prolonged geologi−
cal time may suggest that species of Serradonta could flour−
ish only those chemosynthesis−based communities already
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ACTA PALAEONTOLOGICA POLONICA 52 (2), 2007

Fig. 3. Reconstruction of the Campanian (Late Cretaceous) Yasukawa methane−seep community. Abundant ataphrid and abyssochrysid gastropods and a
few vestimentiferans inhabited the carbonate mound that formed due to anaerobic oxidation of methane and was covered by sandy/silty sediments. The
bivalves Nucinella and Acharax lived in the peripheral zone of the seep. Bathyacmaea cf. subnipponica attached to an ataphrid shell and Serradonta cf.
vestimentifericola grazed on the surface of vestimentiferan tubes. 1, Ataphrid gastropod; 2, abyssochrysid gastropod; 3, Serradonta cf. vestimentifericola;
4, Bathyacmaea cf. subnipponica; 5, ?vestimentiferan tube; 6, Thyasira sp.; 7, Miltha sp.; 8, Nucinella sp.; 9, Acharax cretacea; 10, Leionucula formosa;11, unidentified decapod; 12, ammonoid. Soft part reconstructions based on Recent counterparts, ataphrid soft body reconstruction based on Recent turbinid
gastropods.

settled by vestimentiferan tube worms. This contrasts to
Bathyacmaea which could live on a variety of substrates andwas able to disperse from Japan (the only fossil occurrence)to other chemosynthesis−based communities in the Western
Pacific (Warén and Bouchet 2001; Sasaki et al. 2005 and ref−erences therein) including those not settled by vestimenti−
ferans (e.g., Beck 1996). It is worth noting that acmaeid lim−
pets were not found in the extensive fossil record of Ceno−
zoic seeps in Japan (e.g., Majima et al. 2005). This might berelated to the apparent absence of tubes worms at all but one(Kanie et al. 1992) of these sites. There is, however, little
knowledge about larval dispersal ability and food preferenceof Recent Serradonta and Bathyacmaea and these factors
may also influence the pattern of their distribution.

Conclusions
The mode of acmaeid limpet occurrences in ancient meth−ane−seeps of the Nakagawa area of Hokkaido suggests thatby the Campanian these gastropods had a mode of life similar

to that of their Recent counterparts. Serradonta already pre−ferred vestimentiferan tubes for attachment; Bathyacmaea
could utilize a wider variety of small substrates includingshells of other small gastropods, as do some other Recent tall
limpets. The difference in substrate preference is probably
responsible for the present−day distribution of these gastro−
pods. Serradonta has a more restricted distribution than
Bathyacmaea because it could spread only through those hotvents and cold seeps which were already settled by vesti−
mentiferans.

Acknowledgements
We are deeply indebted to Kazushige Tanabe (DEPUT) and Takenori
Sasaki (UMUT) for helpful discussion and critical reading of the manu−
script. The paper greatly benefited from peer reviews of Steffen Kiel
(University of Leeds, UK), James L. Goedert (Burke Museum, Univer−sity of Washington, USA), and an anonymous reviewer. We thankShin−ichi Fujiwara (DEPUT) for his helpful advice about the recon−
struction figure. The linguistic improvements of Maurice E. Jenkins(Tokyo, Japan) are highly appreciated. The study of RGJ and YH wasJENKINS ET AL.—ACMAEID LIMPETS FROM CRETACEOUS METHANE SEEPS IN JAPANfinancially supported by scientific research grants from the Fukada
Geological Institute, the Japan Science Society, the 21st Century COE
Program to the University of Tokyo (G3, Leader Toshio Yamagata),
and the Japanese Society for the Promotion of Science (nos. 18403013
and 18340165). The research of AK was supported by the Japan Soci−ety for the Promotion of Science (JSPS) Postdoctoral Fellowship for
Foreign Researchers, and JSPS research grant number 17.05324 (pro−ject number 050500000614).

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