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been made to this work since it was submitted for publication. The definitive version has been published in Palaeogeography, Palaeoclimatology,
Palaeoecology 166 (2001) 121-140 doi:10.1016/S0031-0182(00)00205-4
Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
www.elsevier.nl/locate/palaeo
Eocene gastropods of western Kamchatka Ð implications for
high-latitude north paci®c biostratigraphy and biogeography
A.E. Oleinik*
Department of Geography and Geology, Florida Atlantic University, 777 Glades Road, Physical Sciences Building 336,
Boca Raton, FL 33431, USA
Received 19 May 1999; accepted for publication 15 September 1999
Abstract
Fossiliferous rocks of the Snatolskaya and Kovachinskaya formations comprise a Middle and Late Eocene shallow-marine
record of the central part of western Kamchatka. Gastropod assemblages of these formations contain taxa that are conspeci®c
with those in Paleogene strata of western North America and Japan, as well as a large percentage of endemic species. Analysis
of the latitudinal ranges and worldwide occurrences of gastropod genera from these formations show the presence of three
biogeographic components: cosmopolitan, North Paci®c, and endemic. No Tethyan, or circumtropical genera are present in
these Kamchatkan Middle and Late Eocene gastropod faunas. Changes in the geographic distribution of North Paci®c gastropod
assemblages through the Middle and Late Eocene indicate that only eastern Paci®c Tethyan taxa were subjected to latitudinal
range reduction. The distribution of cosmopolitan and North Paci®c elements did not signi®cantly change from the Middle to
Late Eocene, which suggests a relatively stable environment and climate stability during that time. High-latitude Eocene
gastropod assemblages from western Kamchatka demonstrate a high level of endemism at the species level and a low-level
of endemism on the genus level. This pattern is thought to be a result of the unrestricted migration of cosmopolitan taxa
northward along the shallow-marine margin of the Paci®c rim. q 2001 Elsevier Science B.V. All rights reserved.
Keywords: Eocene; Kamchatka; gastropods; Tethyan; cosmopolitan; endemic; latitudinal ranges; biogeography
1. Introduction
Shallow-marine rocks of Middle and Late Eocene
age in western Kamchatka contain 86 species of
gastropods, in 53 genera. This diverse gastropod
assemblage remained very poorly studied until recent
years (Oleinik, 1987, 1988, 1994, Oleinik, 1996;
Sinelnikova et al., 1991). Fossil collections made
prior to the 1980s by Soviet ®eld geologists mainly
contain large and abundant bivalves (Krishtofovich,
1947). The combination of a remote location, harsh
* Fax: 11-561-2972985.
E-mail address: [email protected] (A.E. Oleinik).
climate, and lack of roads resulted in a very incomplete knowledge of the stratigraphy and paleontology
of this area. Specimens studied for the present work
were collected from 1984 to 1989 during extensive
®eld studies of the Kamchatka by expeditions of the
Soviet Academy of Sciences with participation of the
author. Much of this ®eldwork was focused primarily
on the stratigraphy and paleontology along the westcentral coast of the Kamchatka Peninsula (the Tigil
and Palana regions of local usage), adjacent to the
Gulf of Shelikhov in the Sea of Okhotsk (Fig. 1).
This region is characterized by a thick succession of
Cenozoic sedimentary rocks and is known as the
western Kamchatka Depression (Zinkevich and
0031-0182/01/$ - see front matter q 2001 Elsevier Science B.V. All rights reserved.
PII: S 0031-018 2(00)00205-4
2
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A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
Fig. 1. Geologic map of western Kamchatka with fossil localities.
Tsukanov, 1993). Paleogene marine deposits crop out
in the sea cliffs along the coast of the Gulf of Shelikhov for a distance of approximately 500 km, as well
as in river bluffs in the inland parts of the western
Kamchatka Depression. Middle Eocene to Miocene
rocks make up a large transgressive cycle, overlaying
a Late Paleocene±Early Eocene hiatus that can be
traced throughout western Kamchatka and the Sea
of Okhotsk region (Gladenkov et al., 1990). Middle
Eocene strata overlie Paleocene±Lower Eocene rocks
with angular unconformity and are conformably
overlain by the Uppermost Eocene±Lower Oligocene Amaninskaya and Gakhinskaya formations
(Gladenkov et al., 1991). The lithology, tectonic
history, and faunal composition of the Snatolskaya
and Kovachinskaya formations suggest that they
were deposited at upper subtidal to outer neritic
depths (10±500 m) in the back of an active volcanic arc (Oleinik, 1999). The existence of an active
volcanic arc is evident from the large volume of
Paleogene volcanic rocks, including those of
Middle to Late Eocene age, forming a Western
Kamchatka Volcanic Belt, also known as the
Koryak±West Kamchatka Volcanic Belt (Zonenshain
et al., 1990). Based on the K±Ar dating of the
volcanic rocks, the last episode of Paleogene
volcanism within the Western Kamchatka Volcanic
Belt occurred during the uppermost Bartonian±
lowermost Priabonian (46.5±37.4 my) (Gladenkov et
al., 1990).
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A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
2. Area description, materials and methods:
stratigraphy, age, and correlation of Middle and
Upper Eocene rocks of western Kamchatka
Fossiliferous Eocene shallow-marine rocks crop
out in sea-cliffs and river bluffs along the western
coast of Kamchatka. They are subdivided into the
Snatolskaya and Kovachinskaya formations that
comprise the basal part of a large transgressive
cycle, and are separated from the older Paleogene
rocks by a regional unconformity (Gladenkov et al.,
1991).
The most complete sections of the Snatolskaya and
Kovachinskaya formations in the Tigil and Palana
regions crop out in the sea cliffs along the southeastern coast of the Gulf of Shelikhov. This area is characterized by the Tigil and Lesnaya±Palana uplifts of
Neogene age, which produced a number of synclines
and anticlines of predominantly northeastern strike
that are incised by a shoreline running generally
perpendicular to strike. Poor exposures adjacent to
the seashore and inland areas necessitated making a
composite section from numerous outcrops in river
valleys. Nineteen stratigraphic sections of the
Snatolskaya and Kovachinskaya formations, ranging
in thickness from 40 to nearly 800 m, were measured
at 16 localities in Tigil and Palana regions of western
Kamchatka (Fig. 1).
The Snatolskaya Formation is represented mostly
by shallow-marine, medium- to ®ne-grained, laterally
continuous sandstones, with abundant lagoonal and
delta front deposits, containing a rich and diverse
fauna of benthic molluscs. Only its lowermost part
locally contains alluvial conglomerates and terrestrial
near-shore deposits, where the Snatolskaya Formation
directly overlay Cretaceous rocks. The thickness of
the Snatolskaya Formation in the Tigil and Palana
regions (a continuous and complete section with
observable lower and upper contacts) varies from 10
to 295 m. Rocks of the Snatolskaya Formation
become thinner and coarser-grained, and change
from ®ne-grained sandstones to medium- to coarsegrained sandstones and conglomerates, toward the
areas of exposure of Cretaceous rocks.
Benthic foraminifers in the Snatolskaya Formation
occur at two localities in the Tigil Region: the
Tochilinskaya Anticline and Mainach River sections
(Fig. 1). The lower and middle parts of the Snatolskaya
123
Formation in the Mainach River Section contain
Rhizammina indivisa, Bathysiphon nodosariaformis,
Silicosigmolinacf. californica, Haplophragmoides
glabratus, Asanospira cf. exavata, Asanospira cf.
akkeshiensis, Glomospira corona, and Recurvoides sp.
The upper part of the Snatolskaya Formation in the
southern ¯ank of the Tochilinskiy Section contains
Trochammina vitrea, Gyroidina kamtschatica,
Elphidium californicum, Reophax dif¯ugiformis,
Haplopharagmoides snatolensis, Cyclammina ezoensis,
Trochammina markini, Cyclammina tani, Dorothia
amakusaensis, Textularia imariensis, Guttulina irregularis, Gyroidina kamtschatica, Elphidium asagiensis,
Caucasina eocaenica kamtschatica, Globobulimina
paci®ca, and Bolivina jacksonensis (Gladenkov et al.,
1991b). These assemblages strongly imply a Middle
Eocene age for the Snatolskaya Formation (Gladenkov
et al., 1991a). Serova (1969), however, noted on the
possible presence of the Upper Eocene rocks within
this formation. The planktonic foraminiferal assemblage is poorly preserved and can be generally assigned
to the Globigerina boweri regional zone (Serova, 1969;
Krasheninnikov et al., 1988) and suggests a Lutetian to
Bartonian age for the Snatolskaya Formation. A number
of conspeci®c benthic and planktonic foraminifers in the
Snatolskaya Formation of western Kamchatka suggest
correlations with the Narizian Stage of California
(Mallory, 1959), the Middle Eocene (Lutetian±
Bartonian) Takisawa Formation (Honshu), and the
lowermost part of the Poronai Formation (Upper
Bartonian, Hokkaido) of Japan, which contain both
benthic foraminiferal assemblages and nannoplankton
(Saito et al., 1984).
The Kovachinskaya Formation conformably overlies the Snatolskaya Formation and consists of marine
siltstone, claystone, and sandstone. Rocks of the
Kovachinskaya Formation contain numerous tuff
horizons, which suggest an increase in volcanic activity during its deposition. The Kovachinskaya Formation in the Tigil and Palana regions varies in thickness
from 30 to 475 m and is conformably overlain by
Lower Oligocene rocks of the Amaninskya and
Gakhinskaya formations.
The Kovachinskaya Formation consists of a ®negrained clayey rocks, and reaches its maximum thickness in areas most distant from outcrops of Cretaceous
rocks. Rocks of this formation become progressively
enriched in coarser-grained clastics toward the
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A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
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A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
Cretaceous outcrops, where it reaches its minimal
thickness and consists mostly of sandstone and
conglomerate.
Because of its open-marine deposition, the
Kovachinskaya Formation contains more abundant
and diverse foraminiferal assemblages than does the
Snatolskaya Formation. Plectofrondicularia packardi
packardi, Alabamina californica, and Globocassidulina globosa among the most abundant forms of
benthic foraminifers in the Kovachinskaya Formation
on the southern ¯ank of the Tochilinskiy Section. The
Kovachinskaya Formation also contains a diverse
assemblage that is commonly found in most sections
of the Tigil and Palana regions: Gyrodina condoni,
Bulimina debilis, Bathysiphon eocenicus, Ammodiscus pennyi, Haplophragmoides obliquicameratus,
Budashevaella deserta, Cyclammina cushmani, C.
paci®ca, Ammobaculites kamtschaticus, A. akabiroensis, Ammomarginulina stephensoni, Robulus inornatus, R. alatolimbatus, Dentalina plamerae,
Alabamina californica, Cibicides pachecoensis, C.
beckii, Nonion durchani, N. sorachiense, Melonis
planatum, Bulimina debilis, and Globobulimina
paci®ca. In addition to these taxa mentioned above,
the Kovachinskaya Formation in the Mainach
Section, also contains Rhabdammina eocenica,
Reophax tappuensis, R. dentaliniformis, Ammodiscus
concinnus, Haplopharagmoides aff. laminatus, H. cf.
spadix, H. subimpressus, Budashevaella ex gr.
deserta, Guttulina irregularis, G. problema,
Globulina gibba, G. landesi, Gyrodina condoni,
Cibiscides metastersi, C. pachecoensis, C. becki, C.
eponidiformis, Melonis shimokinense, Epistominella
minuta, Caucasina eocenica kamtschatica, C. cf.
schwageri, Trifarina advena californica, Bolivina
kleinpelli, B. jacksonensis, and Globobulimina debilis
(Gladenkov et al., 1991b).
According to Serova (1985), the foraminiferal
assemblage of the Kovachinskaya Formation of the
Tochilinskiy Section contains over 150 species,
mostly benthic foraminifers. Planktonic taxa are rare
and represented only by species of Globigerina. The
most commonly found species in this assemblage are:
Gyroidina condoni, Cibicides hodgei, Plectofrondicu-
125
laria packardi packardi, P. packardi multilineata,
Caucasina eocaenica kamtschatica, C. schwageri,
Uvigerina garzaensis nudorobusta, Bulimina corrugata, B. sculptilis, and Anomalina californiensis
(Serova, 1985). The foraminiferal assemblage characteristic of the Kovachinskaya Formation also occurs
in the eastern Kamchatka, Japan and western North
America (Serova, 1985). Fotyanova and Serova
(1976) referred the Kovachinskaya Formation to the
Bulimina corrugata zone. Bulimina corrugata cooccurs with Globigerina frontosa in the Ilpinskiy
Peninsula of northeastern Kamchatka. The Globigerina frontosa zone of the Ilpinskiy Peninsula, as well as
the Bulimina corrugata zone in California, is correlative with the P14 zone of the standard planktonic
foraminiferal sequence and indicates a Bartonian
age (Krasheninnikov et al., 1988; Volobueva et al.,
1994). This correlation is also supported by the cooccurrence of Globigerina frontosa with calcareous
nannoplankton in the Ilpinskiy Peninsula, indicative
of the NP17 zone of Martini (1971) and the CP14 zone
of Bukry (1973) and Bukry (1975), which suggests a
late Bartonian age for at least the basal beds of the
Kovachinskaya Formation. Similar foraminiferal
assemblages in western North America, containing
Bulimina sculptilis, Plectofrondicularia packardi,
Uvegerina atwili, and Cyclammina clarki, have been
assigned to the Refugian Stage (Serova, 1985;
Krasheninnikov et al., 1988). Nannoplankton
biochronozone NP20 within the Refugian Stage
further suggests a Priabonian age for a large part
of the Kovachinskaya Formation. Analogs of the
Kovachinskaya Formation foraminiferal assemblage in Japan are found in the Okinoshiman
Stage (Kyushu), which is dated as late Middle to
Late Eocene based on the planktonic foraminifers,
and which contains a diverse benthic assemblage
(50 species and subspecies) of the Plectofrondicularia packardi±Bulimina ezoensis zone. This zone
is also found in the upper part of the Poronai
Formation of Hokkaido, where it is known as
the P. packardi±B. schwagerior B. schwageri±
Gyrodina yokoyamai zone (Gladenkov et al.,
1991) (Fig. 2).
Fig. 2. Correlation of the Snatolskaya and Kovachinskaya formations with Eocene formations of western North America and Japan, U.S. West
Coast Eocene molluscan stages (after Saul, 1983; Squires, 1984; Prothero and Armentrout, 1985; Berggren and Prothero, 1992; Nesbitt, 1995),
standard ages, planktonic foraminiferal zones, and nannoplankton zones after Berggren et al. (1995).
6
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A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
Fig. 3. Stratigraphic distribution of species ranging through the Snatolskaya and Kovachinskaya formations.
3. Results and analyses: biostratigraphy and
gastropod fauna of the Snatolskaya and
Kovachinskaya formations
Gastropod shells are moderately common in the
Snatolskaya and Kovachinskaya formations, but differ
in preservation from locality to locality. The best
preserved specimens (without signs of post-mortem
wear or secondary leaching of the shell material)
occur in a ®ne-grained sandstone of the lower member
of the Snatolskaya Formation, particularly at the
Snatol River locality (Fig. 1), or from hard carbonate
concretions that are scattered throughout the Eocene
section at numerous localities. Well-preserved specimens typically have intact protoconchs and delicate
ornamentation patterns. The dissolution and decortication of shell material occur rapidly when specimens
become exposed to weathering. Some traces of shell
leaching have been observed in both the Snatolskaya
and Kovachinskaya formations at the Uvuch
Mountain and Moroshka River localities (Fig. 1).
Thus, the breakage and abrasion of shells was caused
by diagenetic processes and not by reworking.
The Snatolskaya Formation contains a molluscan
assemblage recognizable over a large area of western
Kamchatka. Gastropods occur in all sections of the
Snatolskaya Formation. The biostratigraphic zonation
proposed herein for the Snatolskaya Formation is
based on the ®rst appearance of characteristic taxa,
in addition totaxa restricted to a particular interval.
Special consideration was given to a continuity, or
lateral stability of molluscan assemblages.
The individual stratigraphic ranges of 78 species
and 53 genera of gastropods in the Snatolskaya and
Kovachinskaya formations, plotted by each individual
measured section and locality in the studied region are
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A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
Fig. 4. Ranges of stratigraphically restricted species in the Snatolskaya and Kovachinskaya formations.
127
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A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
Fig. 5. Percentages and numbers (at the end of each bar) of genera and species present in the Snatolskaya and Kovachinskaya formations.
given in Figs. 3 and 4. Local molluscan units proposed
in the present paper are based on temporal changes in
the local stratigraphic ranges of gastropod species.
Fifty-three species and 34 genera occur in the
Snatolskaya Formation and 36 species and 32 genera,
in the Kovachinskaya Formation.
These two distinct, successive gastropod assemblages de®ne the Lower and Upper Snatolskaya
Formation (Figs. 3 and 4), as used in this paper. The
Lower Member contains 47 species and 23 genera of
gastropods. These two members do not necessarily
correspond to lithologic changes within the formations, but are based on a faunal assemblage that can
be recognized in the great majority of Snatolskaya
Formation sections in western Kamchatka. Twentyfour species and 11 genera are restricted to the
Lower Member, and four species and two genera to
the Upper Member of the Snatolskaya Formation. The
Lower and Upper members share 23 species and 19
genera, which suggests a strong facies- and climaterelated similarity. The relatively uniform lithologic
composition and similar molluscan assemblages
throughout the formation suggest uniform depositional and ecologic environments and stable climatic
conditions. The coef®cient of community (Jaccard,
1901) between the lower and upper members of the
Snatolskaya Formation is 0.45 at the species level and
0.59 at the genus level. Twenty-four species (30.8% of
the 78 total species) and 11 genera (20.7% of the 53
total genera) disappear at the Lower and Upper
Snatolskaya Formation boundary, six species (7.7%)
and three genera (5.7%) appear at the same boundary,
whereas 14 species (17.9%) and seven genera (13.2%)
range throughout both members (Fig. 5). There are
only two species that range from the underlying
Napanskaya Formation (Gladenkov et al., 1997) into
the overlying Snatolskaya Formation. One of them is
Calyptraea diegoana, which ranges through the
Snatolskaya into the Kovachinskaya Formation, and
the other is the Kamchatkan species Polinices (Polinices) snatolensis, which is morphologically similar
to the wide-ranging Polinices (Euspira) nuciformis
and Polinices (Euspira) hotsoni of western North
America.
A two-part subdivision of the Snatolskaya Formation has been proposed earlier, based on the entire
molluscan assemblage, known at that time, including
many bivalves (Krishtofovich, 1947; Dyakov, 1957).
Author's data on gastropod distribution do not support
the subdivision of the Snatolskaya Formation into
three `layers with molluscs', proposed by Gladenkov
et al. (1991). For example, the Plicacesta sameshimai±Solen tigilensis layer of Gladenkov et al.
(1991) was more or less clearly de®ned only in the
Pyatibratka River locality, and cannot be traced in
other sections in western Kamchatka. Molluscan
layers named for a particular taxon (or taxa) by
Gladenkov et al. (1991), following the practice usual
for microfossil zones is inappropriate because most
taxa used in these names are not restricted, or even
most abundant, in their nominate.
The Kovachinskaya Formation contains a gastropod assemblage that is distinctively different from
those of the Lower and Upper Snatolskaya Formation.
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A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
The Kovachinskaya fauna re¯ects deeper water and
generally softer substrates than the Snatolskaya fauna.
Twenty-®ve species (32%) and 18 genera (34%) are
known only in a single member within the Kovachinskaya Formation (Fig. 4). Two species (2.6%) and one
genus (1.9%) also occur in the Upper Snatolskaya
Formation. Eighteen species (23%) and 10 genera
(18.9%) disappear, and 25 species (32%) and 18
genera (34%), ®rst appear at the boundary between
the Snatolskaya and Kovachinskaya formations (Fig.
5). These numbers suggest that a larger faunal turnover occurred at the Snatolskaya/Kovachinskaya
boundary than at the Lower/Upper Snatolskaya
Formation boundary. The Jaccard coef®cients calculated for the total number of gastropod genera and
species in the Snatolskaya and Kovachinskaya formations are 0.14 for species and 0.25 for genera, which
are signi®cantly lower than coef®cients for the
Snatolskaya Formation. These coef®cients clearly
indicate that gastropod assemblages of the
Snatolskaya and Kovachinskaya formations are less
similar, than assemblages of the Lower and Upper
Snatolskaya Formation.
Thirteen species in the Snatolskaya Formation and
six species in the Kovachinskaya Formation are
conspeci®c with taxa of the western coast of North
America and Japan. Comparison of published
stratigraphic ranges of these species in the western
North America and Japan with the stratigraphic
ranges in Kamchatka suggest three main categories
of conspeci®c species for the North Paci®c.
3.1. Species with wide stratigraphic ranges
Lower member of the Snatolskaya Formation (Late
Middle Eocene of Kamchatka).
Calyptraea diegoana Conrad, 1855 occurs in the
Late Paleocene through Late Oligocene of
Washington, Oregon, and California (Squires,
1987, 1988).
Olivella matchewsonii umpquaensis Turner,
1938 ranges from the `Martinez' to `Tejon'
molluscan stages of California, and from the
Late Paleocene through Late Eocene in
Washington, Oregon, and California (Turner,
1938; Squires, 1984).
Polinices (Euspira) nuciformis Gabb, 1864
129
ranges from the Late Paleocene through
`Tejon,' in Washington, Oregon, and California
(Marincovich, 1977; Squires, 1987).
Scaphander (Mirascapha) costatus (Gabb, 1864)
ranges from the `Martinez' to `Transition' stages
(Late Paleocene±Middle Eocene) of California
Oregon, and western Washington (Squires,
1984).
Kovachinskaya Formation (Late Eocene of
Kamchatka).
Hataiella (Kotakiella) poronaiensis (Takeda,
1953) ranges from the Late Eocene through
early Late Oligocene in Japan (Poronai (Middle
to Late Eocene), Momijiyama (Late Eocene±
Early Oligocene), Omagari, Shitakara, Charo,
and Nuibetsu formations (Early to Late
Oligocene)) (Honda, 1989; Sinelnikova et al.,
1991; Titova, 1994).
Trominina dispar (Takeda, 1953) is also reported
from the latest Eocene±early Oligocene of
Sakhalin (Takaradai Formation) (Takeda, 1953)
and Early Oligocene (Charo and Nuibetsu
Formations) of Hokkaido (Gladenkov et al.,
1988; Honda, 1989; Titova, 1994).
3.2. Heterochronous species
Lower member of the Snatolskaya Formation (Late
Middle Eocene of Kamchatka).
Ficopsis meganosensis (Clark and Woodford,
1927). Known occurrences are in the upper part
of the `Meganos' stage (Clark and Vokes, 1936),
most probably ranging from Late Paleocene to
Early Eocene (Weaver, 1942).
Polinices (Euspira) lincolnensis (Weaver, 1916)
ranges from Late Eocene to Late Miocene in
Oregon, Washington and Vancouver Island
(British Columbia) (Marincovich, 1977).
Scaphander (Mirascapha) alaskensis Clark,
1932 is also known from the Early to Late
Oligocene parts of Poul Creek and Yakataga
formations of southern Alaska (Clark, 1932;
Kanno, 1971).
Turriola tokunagai (Yokoyama, 1924) also
occurs in the Early to Late Oligocene of the
northwestern Paci®c: the Ombetsu Group of
Hokkaido (Honda, 1989; Sinelnikova et al.,
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A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
1991; Titova, 1994) and Arakai Formation of
Sakhalin (Titova, 1994).
Exilia bentsonae Hickman, 1980 occurs in the
Late Eocene Keasey Formation of northwestern
Oregon (Hickman, 1980).
Upper Member of the Snatolskaya Formation
(Middle Eocene of Kamchatka).
Ancilla (Spirancilla) gabbiCossmann, 1899
occurs in the `Domengine' Stage of California
which is of Early to early Middle Eocene age
(Vokes, 1939; Squires, 1984)
Kovachinskaya Formation (Late Eocene of
Kamchatka).
Margarites (Pupillaria) ef®ngeri (Durham,
1944) occurs in the Echinophoria fax zone of
the Lincoln Creek Formation (Early Oligocene,
Washington) (Weaver, 1942; Prothero and
Armentrout, 1985)
Colus (Aulacofusus) asagaensisMakiyama,
1934, occurs in the Lower Oligocene Asagai
Formation of Kyushu and in the Matschigar
and Arakai formations of the Sakhalin Island
(Oyama et al., 1960; Gladenkov et al., 1988).
3.3. Species with narrow stratigraphic ranges
Lower Member of the Snatolskaya Formation (Late
Middle Eocene of Kamchatka).
Fusinus (Fusinus) willisi (Dickerson, 1915) from
the Cowlitz Formation (late Middle Eocene),
southwestern Washington (Weaver, 1942;
Nesbitt, 1995).
Odostomia (Doliella) hiltoni (Van Winkle, 1918)
from the Cowlitz Formation (late Middle
Eocene), southwestern Washington (Weaver,
1942; Nesbitt, 1995).
Kovachinskaya Formation (Late Eocene of
Kamchatka).
Turricula keaseyensis Hickman, 1976, from the
Late Eocene Keasey Formation of northwestern
Oregon (Hickman, 1980).
Neptunea altispirata (Nagao, 1928) from the
Late Eocene Doshi Formation of northern
Kyushu (Oyama et al., 1960; Gladenkov et al.,
1988; Sinelnikova et al., 1991).
Based on the species cited above, the last four species
are considered to be age-diagnostic for the
Snatolskaya and Kovachinskaya formations and
useful for biostratigraphic correlation.
4. Eocene paleogeography and paleocirculation of
the North Paci®c
The origin and distribution of Eocene faunas in the
northern Paci®c cannot be well understood without
insights into the general paleogeography of the Paci®c
rim and the inferred patterns of paleocirculation.
During the Eocene, the shorelines, total area, and
bathymetry of the Paci®c Ocean were quite different
from those existing today. One of the major paleogeographic elements in the northern part of the Paci®c
Ocean was the Bering Land Bridge, also known in
the literature as `Beringia', which connected Eurasia
and North America throughout most of the early and
middle Tertiary (Durham, 1967; Hopkins, 1967).
There are no reliable data to suggest the existence of
a `paleo-Bering Strait' during the Paleogene (Marincovich and Gladenkov, 1999). The similarity of land
mammals and plants suggests more or less unhindered
terrestrial migration between the Old and New Worlds
(Wolfe and Hopkins, 1967). The record of marine
molluscs clearly indicates an absence of faunal interchange between the Paci®c and Arctic basins during
the Paleogene (Scarlato and Kafanov, 1976; Kafanov,
1979; Marincovich, 1993). The western part of the
Beringian platform apparently extended up to the
Koryak Upland and eastern Kamchatka (Asano,
1963; Shantser, 1974; Vdovin, 1976). Paleogene
unconformities recorded in seismic data at the western
part o the Bering Sea (Goryachev, 1965) suggest the
existence of large areas of dry land from the Late
Cretaceous (Maastrichtian) through Middle Eocene.
Some authors suggest that extensive dry land existed
at the present-day location of the Sea of Okhotsk and
the Sea of Japan (Asano, 1963; Alexandrov, 1973;
Shantser, 1974; Sergeev, 1976).
Relatively small sedimentary basins, with normal
marine to restricted circulation, existed in the areas of
modern Sakhalin and Hokkaido (Iijima, 1964;
Zhidkova et al., 1974; Matsumoto, 1964; Honda,
1991). The Kamchatka was represented by an island
archipelago, which consisted of islands of various size
and formed a volcanic arc. The distribution of
11
A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
131
Fig. 6. Eocene paleogeography, inferred circulation, and distribution of the major biogeographic groups of gastropods in the North Paci®c.
shallow-marine deposits and invertebrate faunas
along the western coast of North America suggests
a continuous shallow-marine habitat extending
from northern Mexico to Alaska (Snavely and
Wagner, 1963; Nilsen and McKee, 1979). This
North±South coastline provided a submeridional
dispersal route for shallow-marine molluscs during
the Paleogene (Fig. 6).
The circulation of the world ocean in the Eocene
was dominated by the westward-¯owing, circumtropical Tethys Current, which resulted in the widespread dispersal and largely cosmopolitan nature of
Paleogene marine faunas (Davies, 1971). The exact
position of seaways connecting the Atlantic and
Paci®c oceans during the Paleocene and Eocene is
not known, and a summary of the Eocene Atlantic±
Paci®c connection is given in Squires (1987). The
main evidence for a marine connection between the
Atlantic and Paci®c is the presence of many Old
World Tethyan taxa in Eocene faunas of western
North America, particularly California. Some species
are strikingly similar to those occurring in Europe and
Africa (Squires, 1984, 1987). Circulation since the
Paleocene outside of the equatorial northern Paci®c,
has been well documented in recent years. It had a
pattern similar to the modern one in the North Paci®c,
with an anticyclonic subtropical gyre and a cyclonic
subarctic gyre as the main features (Kennett, 1982)
(Fig. 6). However, the latitudinal extent and intensity
of North Paci®c oceanic circulation during Paleogene,
especially local shallow-water patterns, are still
poorly understood.
5. Discussion: gastropod faunas and biogeography
The gastropod fauna of Kamchatka, except for a
few specimens from Alaska (Marincovich, 1988), is
the northernmost record of Paleogene molluscs in the
North Paci®c. Two biogeographic components of
Eocene faunas have previously been identi®ed in
lower latitudes of the western and eastern North
Paci®c (the western coast of North America and
Japan), a Tethyan or Tethyan±Indo-Paci®c element
and a northern Paci®c element (Oyama et al., 1960;
Hickman, 1980; Squires, 1984; Squires and
Demetrion, 1992; Honda, 1991; Honda, 1994). The
Tethyan (Tethyan±Indo-Paci®c) element included
genera of presumed warm-water af®nities with a
circumtropical distribution. These genera were
12
132
A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
Table 1
Worldwide occurrences of gastropod genera present in the Snatolskaya and Kovachinskaya formations. Asterisks next to genus name indicate
taxa found only in the North Paci®c region (Kamchatka, Sakhalin, Japan, and West Coast of North America). North Paci®c occurrences are
excluded. Worldwide data from Davies (1971) and Wenz (1933±1944). European data from: Bellardi (1872±1888), Cossmann and Pissarro
(1910±1913), Cossmann (1895±1925), Edwards (1849), Furon and Soyer (1947), and Klyushnikov (1958) North American data (excluding the
West Coast) from Garvie (1996), Dockery (1977), McNeil and Dockery (1984), Richards and Palmer (1953), Toumlin (1977), Palmer (1937),
Palmer and Brann (1965±1966) and Gardner (1923, 1927, 1939) South American data from Steimann and Wilckens (1908), Woods et al.
(1922), Von Ihering (1905±1907) and Zinsmeister (1981) African data from Newton (1922) and Eames (1957). Asian data (excluding Sakhalin
and Japan) from Alexeev (1963), Ilyina (1955), Eames (1952), Shuto (1980, 1984), Vredenburg (1923a, 1923b, 1928), Douville (1928) and Cox
(1930). Australian data from Long (1981). New Zealand data from Beu and Maxwell (1990), Marwick (1931), Maxwell (1992) and Powell
(1942). Antarctic data from Wilckens (1911, 1924), Stilwell and Zinsmeister (1992) and Zinsmeister and Camacho (1982)
Genus
Europe
North America
Acmaea s. l.
Acteon
Ancilla
Antimelatoma p
Apiotoma
Beringius p
Bonellitia
Calyptraea
Cancellaria
Clivoluturris p
Colus p
Comitas
Conominolia
Conus
Creonella p
Crepidula
Diodora
Epitonium s. l.
Exilia
Ficopsis
Ficus
Fulgoraria p
Fusinus
Gibbula
Hataiella p
Homalopoma
Lirosoma
Lishkeia p
Makiyamaia
Margarites p
Marshallena
Mipus p
Molopophorus p
Nekewis p
Neptunea p
Nihonia p
Notoacmaea p
Odostomia
Olivella
Polinices
Problacmaea p
Pseudoliomesus p
Puncturella p
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£
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£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
£
South America
£
£
Africa
£
£
£
£
£
£
Asia
Australia
New Zealand
Antarctica
£
£
£
£
£
£
£
£
£
£
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£
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13
A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
133
Table 1 (continued)
Genus
Europe
North America
South America
Scaphander
Siphonalia
Snatolia p
Solariella
Trominina p
Turcicula p
Turricula
Turrinosyrinx p
Turriola p
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£
£
£
£
£
£
£
Africa
Asia
Australia
New Zealand
Antarctica
£
£
£
£
£
£
ranging south of 508N in the Eocene. The northern
Paci®c element was represented by genera that were
widespread in the high and mid-latitude North Paci®c
in the Paleogene, including central to northern Japan,
Sakhalin, Kamchatka, Alaska, Washington, Oregon,
and California. The majority of these genera are still
living in the northern Paci®c today. Some northern
Paci®c genera penetrated into the subtropical Paci®c
Ocean, but did not attain circumtropical distribution
within the Tethyan (sensu Kauffman, 1973; Sohl,
1987) realm. The worldwide occurrences of gastropod
genera recorded from the Eocene of Kamchatka (Table
1) and their latitudinal ranges in the high-latitude North
Paci®c (Fig. 7), suggest that Kamchatkan gastropod
assemblages were largely composed of cosmopolitan
genera. The high percentage of cosmopolitan taxa in
the Eocene faunas was a product of low latitudinal
temperature gradients (Addicott, 1970) that allowed
warm-water taxa to dwell in high latitudes.
Taxa with at least one occurrence outside of both
the North Paci®c and the Tethyan (tropical) realms
(Table 1) are here considered to be cosmopolitan.
Genera found only in the Eocene of Kamchatka are
considered to be endemic. Genera that occur elsewhere in the North Paci®c realm are cited as North
Paci®c elements (Fig. 7). No Tethyan genera have
been found in Middle or Upper Eocene rocks of
Kamchatka.
The cosmopolitan element remained high throughout the Middle Eocene, with 20 genera (62.5%) in the
Lower Snatolskaya and 14 genera (60.9%) in the
Upper Snatolskaya Formation, but declined to 17
genera (53.1%) in the Upper Eocene Kovachinskaya
Formation. This decline was accompanied by an
increase of the North Paci®c component, with seven
£
£
(21.9%), ®ve (21.7%), and 11 (34.4%) genera in the
Lower Snatolskaya, Upper Snatolskaya, and Kovachinskaya formations, respectively. The endemic
component remained stable at four to ®ve species,
which corresponded to 15.6% of the Lower
Snatolskaya, 17.4% of the Upper Snatolskaya, and
12.5% of the Kovachinskaya Formation faunas (Fig.
8). The in¯uence of the eastern and western Paci®c
faunas on Kamchatka can be evaluated by tallying the
number of species in common with Eocene of western
North America and Japan. The Lower Snatolskaya
Formation contains 11 species (23.4%), the Upper
Snatolskaya has eight species (27.6%), and the
Kovachinskaya Formation has six species (16.6%)
that also occur in western North America (Fig. 8).
Four such species: Calyptraea diegoana, Crepidula
pileum, Odostomia (Doliella) hiltoni, and Scaphander
(Mirascapha) alaskensis, range throughout the
Snatolskaya and Kovachinskaya formations. In
addition, Ficopsis meganosensis, Fusinus (Fusinus)
willisi, Polinices (Euspira) nuciformis, and Scaphander (Mirascapha) costatus occur only in the Lower
member of the Snatolskaya Formation, while Exilia
bentsonae, Olivella matchewsonii umpquaensis, and
Polinices (Euspira) lincolnensis occur in both the
Lower and Upper Snatolskaya Formation, and
Margarites (Pupillaria) ef®ngeri and Turricula
keaseyensis occur in the Kovachinskaya Formation.
Only one species known in Japan, Turriola tokunagai,
occurs in the Lower and Upper Snatolskaya Formation, and also ranges into the Kovachinskaya Formation. The Kovachinskaya Formation contains four
species that also occur in the Paleogene of Japan
and Sakhalin: Colus (Aulacofusus) asagaensis, Trominina dispar, Hataiella (Kotakiella) poronaiensis, and
14
134
A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
Neptunea altispirata. All of these species belong to
the North Paci®c biogeographic component (Figs. 7
and 8).
This pattern of decreasing numbers of eastern
Paci®c and increasing numbers of western Paci®c
species may indicate a weaker eastern Paci®c in¯uence on high-latitude northern Paci®c faunas towards
the end of the Eocene. However, this pattern may
result, at least partly, from a poor preservation and
incomplete knowledge of the Middle Eocene faunas
of Japan.
Temporal changes in the geographic distribution of
characteristic gastropod assemblages in the northern
Paci®c, from Middle to Late Eocene (Fig. 9), clearly
indicate a latitudinal range reduction for Tethyan
genera, particularly in the eastern Paci®c. Tethyan
genera that had dwelled in the northeastern Paci®c
as far North as western Washington during the middle
Eocene, become restricted to southern California
during the Late Eocene. The latitudinal ranges of
Tethyan genera in the western North Paci®c remained
the same, being con®ned in their northernmost penetration to Kyushu, in the Taiwan±South Japan
Province of Honda (1991, 1994). There was no significant change in the latitudinal ranges of cosmopolitan
or North Paci®c genera from the Middle to Late
Eocene (Fig. 9). This pattern suggests that high
latitudes were not severely affected by global climate
changes, at least during the Late Eocene. On the other
hand, minor changes in global climate and/or
circulation patterns may have limited the latitudinal
migration of the more temperature-sensitive circumequatorial genera, into the northern high latitudes.
There are few quantitative data on which to base
speculation about high-latitude Eocene biotic
provinces in the North Paci®c. Provinces can be
recognized on the basis of their unique taxonomic
compositions. Such taxa are commonly species, but
Fig. 7. Latitudinal ranges of gastropod genera of the Snatolskaya
and Kovachinskaya formations in the northern Paci®c. Latitudinal
range data from museum specimens and the following literature
sources: Anderson and Hanna, 1925; Clark, 1938; Clark and Anderson, 1938; Devjatilova and Volobueva, 1981; Dickerson, 1914,
1915, 1916; Hanna, 1927; Hickman, 1976, 1980; Honda, 1991,
Honda, 1994; Krishtofovich and Ilyina, 1954; Marincovich, 1977;
Oyama et al., 1960; Squires, 1984, 1985, 1987, 1988; Squires and
Goedert, 1994; Squires and Groves, 1992; Squires et al., 1992;
Turner, 1938; Vokes, 1939; Weaver, 1916, 1942.
15
A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
135
Fig. 8. Diversity and changes in the biogeographic composition of gastropod assemblages in the Snatolskaya and Kovachinskaya formations.
Fig. 9. Temporal changes in the latitudinal ranges of characteristic Eocene gastropod assemblages in the northern Paci®c. An asterisk next to the
genus name indicates that the record for the genus is probably incomplete.
16
136
A.E. Oleinik / Palaeogeography, Palaeoclimatology, Palaeoecology 166 (2001) 121±140
may be higher taxa, and it is common for paleontologists to use genera in reconstructing provincial
boundaries. Biogeographic patterns de®ned at the
species-level can also sometimes be recognized at
the genus- and family-levels (Campbell and Valentine,
1977; Smith, 1989). One of the most frequently
quoted rules that have been used to identify provincial
regions is that at least one-half of the species living in
the region must be endemic. However, there is no
special reason to employ any particular arbitrary
level of endemism to de®ne a province. The generic
level of endemism which can be considered satisfactory to de®ne a province, have to be lower than the
species level by the virtue of higher taxonomic rank.
Kauffman (1973) assigned a 10±25% of endemic
genera for a subprovince, 25±50% for a province,
and 50±75% for a regions. These numbers were
proposed based on study of distribution of Cretaceous
bivalves is not universally accepted for all ages and
other groups of organisms.
Taking into account the warm and equable climatic
conditions, and the linearity of shallow-marine
habitats around the North Paci®c rim in the Paleogene
(Fig. 6). Endemism at the genus-level may have been
low, while species-level endemism could still have
been high, owing to local conditions. This pattern of
high species-level endemism is present in the Middle
and Late Eocene gastropod fauna of the western
Kamchatka, ranged from highs of 65.5% in the
Upper Snatolskaya and 72.3% in the Lower
Snatolskaya Formation, to lows in genus-level
endemism of 12.5% in the Kovachinskaya and
17.4% in the Upper Snatolskaya Formation. It is
possible to propose a separate high-latitude province
in the northwestern Paci®c, using a species-level
gastropod data, but matter becomes questionable
when based on genus-level data. High latitude North
Paci®c biogeography and the extent of Paleogene
biogeographic provinces must be based on a more
quantitative approach, utilizing zoogeographic coef®cients for more than one group of benthic molluscs.
Acknowledgements
The author wishes to thank Valentina Sinelnikova
of the Russian Academy of Sciences, Geological
Institute, for providing part of the material used in
this study and for numerous stimulating discussions.
The author would also like to extend his deep gratitude to Richard Squires of California State University
at Northridge, for sharing a wealth of information on
Eocene faunas of the US. West Coast and for his
insightful comments on this study. Special thanks to
Louie Marincovich of California Academy of
Sciences and David T. Dockery III of the Mississippi
Department of Environmental Quality, Of®ce of
Geology, for a very helpful comments and critical
review of the early version of the manuscript.
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