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Paleontological Society
Genus Arctomelon (Gastropoda, Volutidae) in the Tertiary of the Northwestern Pacific:
Evolution and Adaptations
Author(s): Anton E. Oleinik
Reviewed work(s):
Source:
Journal of Paleontology,
Vol. 70, No. 2 (Mar., 1996), pp. 236-246
Published by: Paleontological Society
Stable URL: http://www.jstor.org/stable/1306387 .
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J. Paleont.,
70(2), 1996,
pp. 236-246
Copyright
? 1996,
The Paleontological Society
0022-3360/96/0070-0236$03.00
GENUS ARCTOMELON
(GASTROPODA,
VOLUTIDAE)
IN THE
TERTIARY OF THE NORTHWESTERN PACIFIC:
EVOLUTION
AND ADAPTATIONS
ANTON E. OLEINIK
Department
of Earth and Atmospheric Sciences,
Purdue
University,
West Lafayette,
Indiana 47907
ABSTRACT-The fossil record of the volutid gastropod
Arctomelon is represented
by three
species
from
the Koryak
Upland
and the
Kamchatka Peninsula
(Russian
northeast), ranging
in age from late middle Eocene to early
Miocene.
Although
Recent Arctomelon
stearnsii
(Dall, 1872) from the northeastern Pacific is a cold-water
species,
paleontological
data point to the origin
of the genus
in
shallow,
warm-water
conditions that existed
in high latitudes of the northwestern Pacific
during Paleogene
time. The fossil record
also indicates survival
of the genus during
the climatic
coolings through
the Cenozoic.
It is apparent
that Arctomelon
successfully
adapted
to the new cooler and deeper-water
conditions.
Despite this major change
of life habit, shell morphology
has remained
fairly unchanged
since the Eocene.
Two new species
of the genus
Arctomelon,
A. harasewychi
sp. nov. and A. rateginense
sp. nov.,
are described.
INTRODUCTION
THE GENUS Arctomelon includes the northernmost known
species of volute. The type species of the genus, Arctomelon
stearnsii (Dall, 1872) (Figures 1.1, 1.2, 1.3), has been dredged
from 73 to 180 m off the coast of southwestern Alaska (type
locality: Shumagin Islands, westward from Captains Bay, about
55?N) in bottom water temperatures of 2.8 to 5.0?C, and also
from the Bering Sea northward to the limit of floating ice in
winter in depths of 111 to 640 m (Weaver and DuPont, 1970;
Habe and Ito, 1965; Smith, 1942). More recent data show a
bathymetric range of 120 to 800 m for Arctomelon stearnsii,
with the vast majority of records between 200 and 500 m. The
shallower records are concentrated in the Bering Sea, whereas
the deepest records are off British Columbia (M. G. Harasewych,
personal commun.), which makes this species a good example
of latitudinal submergence. A second species, Arctomelon ben-
thalis (Dall, 1896), is known only from the material dredged by
the U.S.S. Albatross from the Gulf of Panama in a depth ex-
ceeding 3,000 m (Dall, 1896; Smith, 1942; Weaver and DuPont,
1970). One additional species that probably belongs to the genus
Arctomelon, Boromelon tamikoae Kosuge, 1970 (see synonymy
below), was recorded from about 120 m depth in the South
China Sea (Kosuge, 1970).
Arctomelon stearnsii is well known in high-latitude North
Pacific waters, but the fossil record of the genus is still poorly
known. Only one fossil species, Arctomelon lautenschlageri (Vo-
lobueva in Devjatilova and Volobueva, 1981), has been de-
scribed previously, from Oligocene and lower Miocene strata
of the Koryak Upland. Recent discoveries from the Russian
Northeast extend the fossil record of Arctomelon back to the
late middle Eocene. These new data allow insights into the or-
igin, evolution, and distribution of this high-latitude volutid
genus.
Material used for the present paper was collected during the
1984 through 1990 field seasons by expeditions of the Russian
Academy of Sciences, Institute of Geology, and by the Russian
Ministry of Geology, "Aerogeology" field parties.
Abbreviations used: CGMSP, Central Museum of Geology,
Ministry of Geology, St. Petersburg, Russia; USNM, National
Museum of Natural History, Smithsonian Institution; IGM,
Russian Academy of Sciences, Institute of Geology, Moscow,
Russia.
The holotype of the single previously described fossil species,
Arctomelon lautenschlageri (Volobueva in Devjatilova and Vo-
lobueva, 1981), was designated without a specific number (num-
ber of the paleontological collection in the CGMSP is 11292).
Therefore, in the present paper, it will be cited as a particular
plate and figure from the "Atlas of the Paleogene and Neogene
fauna of the U.S.S.R. northeast" by Devjatilova and Volobueva,
1981.
STRATIGRAPHIC SETTING AND DESCRIPTION OF
FOSSIL LOCALITIES
Cenozoic marine sediments are widespread and fairly well
exposed along the eastern and western coasts of the Kamchatka
Peninsula and in the Koryak Upland (Figure 2). They are rep-
resented by a series of alternating marine, transitional, and non-
marine clastic sediments ranging in age from the Late Paleo-
cene(?) to Late Pliocene and assigned to various formations.
The molluscan fossils described here occur in middle Eocene
through upper Miocene shallow to deep marine sediments crop-
ping out in the sea cliffs and tidal flats on the western coast of
the Kamchatka Peninsula and in bluffs along river valleys in
the Koryak Upland (Figure 2).
Remains of the oldest known species of Arctomelon were
discovered in upper middle Eocene deposits of the Koryak Up-
land in the Mayn River basin area (Figure 2). They occur in the
uppermost part of about 450 meters of shallow marine sand-
stones and conglomerates assigned to the Aglikitschian For-
mation (Figure 3). Age determinations of these strata are based
primarily on molluscan fossils. Common Aglikitschian bivalves
and gastropods similar to those from the upper middle Eocene
deposits (Snatolian and Kovachian Formations) of Western
Kamchatka include: Papyridea utcholokensis Slodkevitsch, 1938;
Cyclocardia (Cyclocardia) akagii (Kanehara, 1937); Glycymeris
(Glycymeris) aglikitschensis Devjatilova in Devjatilova and Vo-
lobueva, 1981; Ficopsis tuberculata Oleinik in Sinelnikova et
al., 1991; and Margarites cf: M. uvutschensis Oleinik in Sinel-
nikova et al., 1991. Early Oligocene through Early Miocene
specimens of the single previously described fossil species, Arc-
tomelon lautenschlageri (Volobueva in Devjatilova and Volob-
ueva, 1981), occur in the Khatyrka River basin and the Koryak
Upland (Figures 2, 3), and are known from the Ionaian and
Khaiidinian Formations.
The lower Oligocene Ionaian Formation consists of more than
1,000 m of gray marine clays and claystones with separate layers
of sandstone and conglomerate. Large carbonate concretions are
scattered throughout the section (Figure 3). The low species
236
OLEINIK- TERTIARY GASTROPOD
3
1 2
FIGURE
1-1-3, Arctomelon stearnsii
(Dall, 1872), paratype
USNM 91352, Aleutian
Islands, x 1 (courtesy
of M. G. Harasewych).
1, apertural
view; 2, abapertural
view; 3, portion
of radula of Arctomelon stearnsii
(Dall, 1872) (from
Pilsbry
and Olsson, 1954),
USNM 22504, Shumagin
Islands,
Alaska.
diversity of the molluscan assemblage and the predominantly
fine-grained sediments suggest an outer shelf or bathyal envi-
ronment during most of Ionaian deposition. An exception is the
lowermost part of the formation, where several tens of meters
of sandstone and conglomerate with Ostrea are probably indic-
ative of a shallower environment. Remains of Arctomelon lau-
tenschlageri are known only from the deep marine part of the
section (Figure 3). Mollusks such as Trominina (Trominina)
bicordata (Hatai and Koike, 1957); Trominina (Trominina) ish-
ikariensis (Hayasaka and Matsui, 1951); Fulgoraria (Musashia)
olutorskiensis (L. Krishtofovich, 1973); Fulgoraria (Musashia)
cordata Oleinik, 1993; Papyridea harrimani Dall, 1909; Sole-
mya bosoana (Hatai & Koike, 1957); Yoldia (Portlandella) wat-
sei (Kanehara, 1937), and Yoldia (Portlandia) nitida (Slodkev-
itsch, 1938) provide good correlation with lower Oligocene
bathyal deposits of Kamchatka and shallower deposits of Japan.
The Khaiidinian Formation, which has a total thickness of more
than 2,500 m, and consists of interbedded marine sandstone,
siltstone, and claystone in the lower 900 meters of the section
and 450 meters of sandstone and conglomerate in the upper
part, conformably overlies the Ionaian Formation (Figure 3).
The age of the Khaiidinian Formation has been determined as
late Oligocene to early Miocene, based on molluscan and fo-
raminiferal data (Devjatilova and Volobueva, 1981; Gladenkov
et al., 1985). Type of sediments and changes in diversity of
molluscan and foraminiferal assemblages within the section of
the Khaiidinian Formation could be interpreted as indicating a
regressive sequence, from relatively deep water outer shelf and
bathyal environments in the lower part of the formation towards
shallower inner shelf facies in the upper part. Arctomelon lau-
tenschlageri has been recorded only from the lower and deeper
water part of this formation.
Arctomelon is also known in the northwestern and western
parts of the Kamchatka Peninsula, from the upper part of the
Rategian Formation of Podkagernaya Bay and the Kuluvenian
Formation of the Mainatsch River mouth section (Figure 2).
The Rategian Formation consists of about 200 meters of shallow
marine sandstones and conglomerates containing abundant
molluscan remains (Figure 3). The age of the formation is ques-
tionable; Serova (1985) used benthic foraminifera to assign an
Oligocene age to the Rategian Formation, whereas more recent
analyses of molluscan fossils shows species-level correlation with
upper Eocene deposits of the North Pacific region. The age of
the Rategian Formation is not certain, but herein is considered
to be latest Eocene to possibly earliest Oligocene.
The Kuluvenian Formation consists of tuffaceous siltstones,
sandstones, claystones and cherty rocks, which together with
the occurrence of Yoldia, Neancistrolepis, Oenopota, Suavod-
rillia, Periploma, and Thyasira are indicative of a relatively deep
marine, probably outer shelf environment. Total thickness of
237
I%
JOURNAL OF PALEONTOLOGY, V. 70, NO. 2, 1996
FIGURE
2-Geologic sketch-map
of the Kamchatka
Peninsula
showing
the localities mentioned
in the text. Fossil localities:
A, Mainatsch
River mouth (Kamchatka Peninsula);
B, Podkagernaya Bay (Kam-
chatka
Peninsula);
C, Mayn
River
Basin,
Aglikitsch
River
valley (Ko-
ryak
Upland);
C, Khatyrka
River Basin (Koryak
Upland). Legend:
1-Quaternary alluvial sediments, 2-Quaternary basalts, 3-Neo-
gene sediments,
4-Paleogene sediments,
5-Volcanic rocks of var-
ious age,
6-Late Cretaceous
sediments,
7-Mesozoic (Triassic-Early
Cretaceous)
sediments,
8-Paleozoic sediments,
9-Major faults,
10-
Fossil localities.
the formation is about 300 m (Figure 3). Based on the molluscan
and foraminiferal data, the formation is confidently inferred to
be early Miocene in age (Bratseva et al., 1984). Specimens of
Arctomelon lautenschlageri (Volobueva in Devjatilova and Vo-
lobueva, 1981) are known from the uppermost beds of the for-
mation (Gladenkov and Sinelnikova, 1990).
SYSTEMATIC PALEONTOLOGY
Phylum MOLLUSCA
Linnaeus, 1758
Class GASTROPODA
Cuvier, 1797
Family VOLUTIDAE
Rafinesque, 1815
Subfamily ZIDONINAE
H. and A. Adams, 1853
Genus ARCTOMELON
Dall, 1915
Fulgoraria
(Arctomelon)
DALL, 1915, p. 57.
Fulgoraria
(Boromelon) DALL, 1918, p. 138.
K7il 1j =2 3
I,jj
3 34 1 5
rrr r _____
6
6 rr
I 7 8 - 19
FIGURE
3-Location of the genus
Arctomelon
through
the formations
mentioned
in the
text.
Legend:
1
-sandstones, 2- siltstones,
3-mud-
stones,
4-conglomerates, 5-carbonate concretions, 6--volcaniclas-
tic and
cherty
rocks,
7-volcanic rocks,
8-coal, 9-fossil occurrences
of the genus
Arctomelon.
Boreomelon
Dall. SMITH,
1942, p. 59.
Arctomelon Dall. PILSBRY
AND
OLSSON, 1954, p. 287.
Type species. - Voluta (Scaphella) stearnsii Dall, 1872; by
original designation; Recent, northeast Pacific, Bering Sea (Fig-
ures 1.1, 1.2).
Diagnosis. -"Shells are of moderate size (105 to 125 mm in
length). They are solid and fusiform or ovate-fusiform with a
high, blunt spire. The protoconch of Arctomelon stearnsii is
small, white and mammilate. Teleoconch is smooth (except for
growth lines) with whorls that are slightly convex to roundly
convex. Suture is calloused or appressed. Siphonal notch and
fasciole are present. Columella has several plaits. Periostracum
and operculum are absent. The radula is uniserial with tricuspid
teeth" (Weaver and Du Pont, 1970, p. 119).
Remarks. -A dark-colored form with low spire and oval shape
from the Bering Sea has been named A. stearnsii ryosukei (Habe
and Ito, 1965). Recent observations, however, demonstrate that
there are no significant differences in color between A. stearnsii
and A. stearnsii ryosukei, although the two forms are very dis-
tinct in shape (M. G. Harasewych, personal commun.).
Discussion. -The shell characteristics of the genus Arctome-
lon closely resemble those of a number of volutid genera and
subgenera, not only from the subfamily Zidoninae, but also
Scaphellinae and Odontocymbiolinae. Therefore, it seems ap-
propriate to include a brief comparison of the genera morpho-
logically close to the Arctomelon. Ericusa (Ericusa) H. & A.
Adams, 1858 (subfamily Zidoninae) (Darragh, 1988), type spe-
cies E. (E.)fulgetra (Sowerby, 1825) (Figure 4.1), differs in hav-
ing a much less well developed fasciole, smoother shell surface,
less elongated spire, more numerous and smaller columellar
plaits, large papillary protoconch, distinct shoulder and thinner
callus. Similarly, subgenus Ericusa (Mesericusa) Iredale, 1929,
type species E. (M.) sowerbyi (Kiener, 1839) (Figure 4.2), has,
238
OLEINIK- TERTIAR Y GASTROPOD
in general, a less elongated spire, less developed fasciole, four
columellar plaits, smoother shell surface and thinner and broad-
er callus. Genus Iredalina Finlay, 1926 (subfamily Zidoninae)
(Darragh, 1988), type species I. mirabilis Finlay, 1926 (Figure
4.3), differs in having a very smooth glossy shell surface, a
smaller conical protoconch, and in lacking a fasciole and col-
umellar plaits. Guivillea Watson, 1886 (subfamily Zidoninae)
(Knudsen, 1973), type species G. alabastrina Watson, 1882 (Fig-
ure 4.4), is similar to Arctomelon in shape, but differs in having
a very thin and smooth shell, irregular mammilate protoconch,
wider aperture, lack of fasciole and siphonal notch, and absence
of true columellar plaits (abrupt break along the midpoint may
look like a columellar plait). Genus Teremelon Marwick, 1926
(subfamily Zidoninae), type species T. tumidior (Finlay, 1926)
(Figure 4.5), can be distinguished easily from Arctomelon by
having a smaller and more slender shell with a smooth surface
and a narrow aperture. The columella of Teremelon normally
carries four relatively strong columellar plaits. Another species
of this genus, T. knoxi Dell, 1956 (Figure 4.6) approaches Arc-
tomelon in shell morphology very closely, but differs in having
four strong columellar plaits and a more slender and smaller
shell with a narrower aperture. Pachymelon (Palomelon) Finlay,
1926 (subfamily Zidoninae), type species P. (P.) lutea (Watson,
1882) (Figure 4.7), although somewhat similar to Arctomelon
in pattern of surface sculpture, has a more convex shell, broader
aperture and larger protoconch. Adelomelon (Adelomelon) Dall,
1906 (subfamily Zidoninae), type species A. ancilla (Lightfoot,
1786) (Figure 4.8), differs in having a much more slender,
smoother, and larger shell, narrower aperture and a protoconch
often sculptured with a calcarella. Scaphella (Scaphella) Swain-
son, 1832 (subfamily Scaphellinae H. and A. Adams, 1858),
type species S. (S.) junonia (Lamarck, 1804) (Figure 4.9), has a
more slender shell and more elongated last whorl, narrower
aperture, and protoconch typically sculptured with a calcarella.
Notopeplum Finlay, 1927 (subfamily Zidoninae) (Wilson, 1972;
Darragh, 1988), type species N. politum (Tate, 1889) (Figure
4.10), differs in having a smaller and smoother shell, narrower
aperture, more numerous and stronger columellar plaits and
large convex low domal protoconch. Odontocymbiola Clench
and Turner, 1964 (subfamily Odontocymbiolinae Clench and
Turner, 1964), type species 0. magellanica (Gmelin, 1791) (Fig-
ure 4.11), has many shell characteristics common with Arcto-
melon. It differs, however, in its generally larger size, broadening
anteriorly aperture, more convex shell, lower spire and in having
a thin glazing callus.
ARCTOMELON HARASEWYCHI n. sp.
Figures 5, 6.1, 6.2
Diagnosis. -Shell convex, solid, ovate-fusiform; low spire with
distinct narrow apex; four anterior oblique columellar plaits
with the uppermost weakest.
Description. -Shell ovate-fusiform, solid, with relatively low
spire; last whorl comprises approximately three-quarters of shell
length; teleoconch has five whorls, slightly concave just below
the suture, then moderately convex; suture appressed; sculpture
of irregular oblique axial ribs parallel to the thin growth lines;
aperture semiovate, narrowing posteriorly; outer lip thin and
simple; siphonal notch deep and narrow; fasciole strong and
broad; parietal canal indistinct; columella slightly arched with
four oblique anterior plaits; upper two plaits very close to each
other with the uppermost weakest; callus moderately thin and
narrow, covering columella and parietal area; protoconch, al-
though slightly eroded, is relatively small, homeostrophic with
2.5 to possibly 3 smooth whorls, forming distinctive raised nar-
row apex.
Dimensions. -Holotype USNM 483409: length (incomplete),
66.7 mm; maximum diameter, 38.9 mm; length of the body
whorl (incomplete), 55.9 mm. Paratype IGM 1894/4: length
(incomplete) 59.3 mm; maximum diameter, 27.5 mm; length
of the body whorl (incomplete), 47.2 mm.
Type locality.-Koryak Upland, Mayn River Basin, Black
River Valley.
Material. -Holotype and paratype from the type locality.
Age. -Aglikitschian Formation, late middle Eocene.
Geographic distribution. -Known only from the type locality.
Repository. -United States National Museum, Smithsonian
Institution, Washington, D.C.
Discussion. -Although this species morphologically ap-
proaches Arctomelon stearnsii and A. stearnsii ryosukei, it differs
by its very low spire, highly elevated, narrow blunt apex and
by having four oblique columellar plaits that are stronger than
those ofA. stearnsii, with the upper one the weakest. Arctomelon
stearnsii and A. stearnsii ryosukei, in contrast, do not normally
have an elevated apex, but have a small to medium sized pro-
toconch with about three and a half smooth whorls and two or
three weak, strongly oblique columellar plaits. The new species
also differs from A. lautenschlageri by having more convex
whorls, an elevated apex, four stronger columellar plaits, and a
broader aperture.
Etymology. -This species is named for Dr. M. G. Harasew-
ych, Curator of Mollusks, U.S. National Museum of Natural
History, Smithsonian Institution, Washington D.C.
ARCTOMELON
RATEGINENSE
n. sp.
Figure 6.3-6.6
Diagnosis. -Large body whorl; aperture semi-ovate to half-
moon; columella has four oblique anterior plaits of equal strength;
growth lines almost straight.
Description. -Shell elongate-fusiform, rather solid, with 4 or
5 lightly convex whorls. Last whorl comprises approximately
four-fifths of shell length; suture appressed; surface smooth, ex-
cept for thin and almost straight growth lines; aperture semi-
ovate, narrowing both anteriorly and posteriorly; siphonal notch
deep and fairly narrow; fasciole strong and narrow;
parietal canal
narrow, weak, and almost indistinct; columella almost straight,
with four oblique prominent anterior plaits of equal strength;
outer lip thin and simple; callus thin and narrow, covering col-
umella and part of the parietal area.
Dimensions. -Holotype USNM 483410: length (incomplete),
45.4 mm; maximal diameter, 24.1 mm; length of the body whorl
(incomplete), 41.6 mm. Paratype USNM 483411: length (in-
complete), 61.05 mm; maximum diameter, 31.7 mm; length of
the body whorl 56.3 mm.
Type locality. -Northwestern Kamchatka Peninsula, Pod-
kagernaya Bay, near the mouth of the Podkagernaya River,
outcrop exposed at low tide.
Material. -Holotype and paratype from the type locality.
Age. -Rategian Formation, late Eocene-(?) early Oligocene.
Geographic distribution. -Known only from the type locality.
Repository. -United States National Museum, Smithsonian
Institution, Washington D.C.
Discussion. -This species closely approaches Arctomelon lau-
tenschlageri, but can be distinguished by a slightly more narrow
and elongated shell, narrower fasciole, almost straight growth
lines, and four oblique columellar plaits of equal strength that
are stronger and narrower than those of A. lautenschlageri. It
also differs from A. stearnsii by having a narrower aperture,
narrower siphonal canal, and four columellar folds that are less
oblique and stronger than those of A. stearnsii. Because both
available specimens were collected from the outcrop subjected
239
JOURNAL OF PALEONTOLOGY, V. 70, NO. 2, 1996
A
2
1
5 6
10
3
7
'11
11
8
240
9
OLEINIK- TERTIARY GASTROPOD
to the tidal and wave action, the surface of both shells could
have been worn smooth by wave erosion.
Etymology. -The name of this species is from the Rategian
Formation, Podkagernaya Bay, northwestern Kamchatka.
ARCTOMELON LAUTENSCHLAGERI
(Volobueva in Devjatilova and Volobueva, 1981)
Figure 6.7, 6.8, 6.11, 6.12
Scaphella
lautenschlager
VOLOBUEVA
IN
DEVJATILOVA
AND
VOLOBUEVA,
1981, p. 130, Pl. 33, figs.
4 a, b, 5.
Arctomelon
cf. lautenschlageri
(Volobueva).
GLADENKOV
AND
SINELNK-
OVA,
1990, p. 139, P1. 1, figs. 5 a, b.
Diagnosis. -Shell of medium size, length 50 to 85 mm, solid,
moderately fusiform, with low spire and three equal strength
anterior oblique columellar plaits.
Description.--Shell elongate-fusiform, solid with relatively
short spire; last whorl comprising slightly more than two-thirds
of shell length; suture appressed; teleoconch of five rounded
whorls; aperture serr iovate, narrowing posteriorly; outer lip thin
and simple; columetla slightly concave, carrying three oblique
weak equally developed anterior plaits; fasciolar fold flattened;
distance between columellar plaits increases posteriorly; si-
phonal notch narrow, moderately deep, fasciole present; pos-
terior notch indistinct; callus thin and narrow, covering colu-
mella and part of the anterior end; shell surface smooth except
for growth lines.
Dimensions. -Holotype CGMSP, No. (of paleontologic col-
lection) 11292; Devjatilova and Volobueva, 1981, P1. 33, fig. 4
a, b: length, 64.1 mm; maximum diameter, 32.6 mm; length of
the body whorl, 50.7 mm. Paratypes CGMSP: length, 65.0-83.8
mm; maximum diameter, 33.2-43.2 mm; length of body whorl
48.2-64.0 mm. Hypotype USNM 483411: length, 59.1 mm;
maximum diameter, 39.6 mm; length of body whorl, 51.2 mm.
Type locality. -Koryak Upland, Khatyrka River basin, Vych-
ginai Mountains, Khaiidinian Formation.
Material. -Holotype, paratypes, hypotype, and several spec-
imens of fair and poor preservation.
Age. -Koryak upland, Khatyrka River Basin: Vychginai
mountains, Khaiidinian Formation, latest Oligocene-earliest
Miocene, Imyneiveem river, Ionaian Formation, early Oligo-
cene; Western Kamchatka: sea cliffs south from the mouth of
the Mainatsch River, Kuluvenian Formation, early Miocene.
Geographic distribution. -Koryak Upland and western Kam-
chatka.
Discussion. -Shells of this species show very few morpho-
logical differences from Recent Arctomelon stearnsii (Dall, 1872).
Shells ofA. stearnsii vary in shape from elongated fusiform with
moderately high spire of typical stearnsii, to rather ovate and
convex with a low spire, such as the Bering Sea subspecies A.
stearnsii ryosukei Habe and Ito (1965). Arctomelon lautensch-
lageri is intermediate in spire height and shell width between
A. stearnsii and A. stearnsii ryosukei. However, it differs by
having a generally smaller shell and less oblique and slightly
stronger columellar plaits that are located closer to the anterior
end of the shell than those of A. stearnsii and A. stearnsii ry-
FIGURE
5--Arctomelon
harasewychi
sp. nov. Reconstructed
drawing
of
the protoconch.
Scale bar - 2.5 mm.
osukei. The aperture ofA. lautenschlageri seems to be relatively
narrow at the posterior end. The siphonal notch appears to be
narrower and deeper than those of both A. stearnsii and A.
stearnsii ryosukei.
ARCTOMELON
sp.
Figure 6.9, 6.10
Description. -Shell ovate-fusiform, rather slender with 5-6
teleoconch whorls; last whorl comprises approximately two-
thirds of shell length; suture appressed; surface smooth, with
thin and almost straight growth lines; aperture semiovate, nar-
rowing posteriorly; siphonal notch broad and shallow; fasciole
rather weak; outer lip thin and simple; columella slightly arched
with two oblique anterior plaits.
Dimensions.-Hypotype IGM 13/159: length, 62.8 mm;
maximum diameter, 33.5 mm; length of body whorl, 53.1 mm.
Material. -1 poorly preserved specimen.
Age. -Western Kamchatka, to the south of the mouth of the
Mainatsch River, Kuluvenian Formation, late early Miocene.
Geographic distribution.
-Western Kamchatka.
Discussion. -This specimen closely approaches A. lautensch-
lageri in morphology. However, a more convex last whorl, al-
most straight growth lines, and two more oblique columellar
plaits distinguish the present species. It also differs from A.
stearnsii by having a narrower aperture, lower spire, straight
and fine growth lines and generally smaller size. It differs from
A. stearnsii ryosukei in having a narrower aperture, smoother
surface with almost straight growth lines and less convex whorls.
Poor preservation of the single specimen does not allow deter-
mination of its exact taxonomic position.
DISCUSSION
The Recent North Pacific species Arctomelon stearnsii is a
cold, relatively deep-water (mainly bathyal) taxon. However,
its ancestral Paleogene species lived in warm, shallow-marine
FIGURE
4-1-11, Line drawings (new herein) of the volutid genera
that morphologically
approach
Arctomelon.
1, Ericusa
(Ericusa)
fulgetra
(Sowerby,
1825), Zidoninae,
Recent,
Australia
(x '/3). 2, Ericusa
(Mesericusa) sowerbyi
(Kiener, 1839), Zidoninae, Recent,
Australia
(x '/3).
3,
Iredalina
mirabilis
Finlay, 1926, Zidoninae,
Recent, New Zealand
(x3/4). 4, Guivillea alabastrina
Watson, 1882, Zidoninae,
Recent, South
Indian
Ocean
(x 1/2).
5, Teremelon
tumidior
(Finlay,
1926),
Zidoninae,
early
Miocene
(Duntroonian-Altonian),
New Zealand
(x 2). 6, Teremelon
knoxi Dell, 1956, Zidoninae,
Recent,
New Zealand
(x 1). 7, Pachymelon
(Palomelon)
lutea (Watson,
1882),
Zidoninae, Recent,
New Zealand
(x /2).
8, Adelomelon
(Adelomelon)
ancilla
(Lightfoot,
1786),
Zidoninae,
Recent,
eastern South America
(x '/3).
9, Scaphella
(Scaphella) junonia
(Lamarck,
1804), Scaphellinae,
Recent, Florida
(x /2). 10, Notopeplum politum (Tate, 1889),
Zidoninae,
middle Miocene,
Australia
(x 2). 11,
Odontocymbiola
magellanica
(Gmelin, 1791), Odontocymbiolinae,
Recent,
South America
(x 1/3).
241
242 JOURNAL OF PALEONTOLOGY, V. 70, NO. 2, 1996
3 4
1 2
5 6
7 8
10 11 12
FIGURE
6-1, 2, Arctomelon harasewychi sp. nov., holotype USNM 483409, x 1. 1, abapei lural view, 2, apertural view. 3, 4, Arctomelon rateginense
sp. nov., holotype USNM 483410, x 1. 3, abapertural view, 4, apertural view. 5, 6, Arctomelon rateginense sp. nov., paratype USNM 483411,
9
OLEINIK- TERTIARY
GASTROPOD
habitats
that existed during
the Eocene
along
the Northwestern
Pacific (63?-65?N). Existence of these mild warm climatic con-
ditions during
the Eocene is clearly
supported by a variety of
data provided
by marine
invertebrates,
plant remains,
and ox-
ygen
isotope studies.
Surface
water
temperatures
at 60?N
in the
Pacific Ocean
during
the Eocene
probably
exceeded
15?C
(Frakes
and
Kemp, 1973).
The distribution
of plant
fossils indicates
that
broad-leaved
evergreen
forests
extended north of 60?N and that
the mean annual
range
of temperatures during
the middle Eo-
cene must have been at least 21?C
(Wolfe 1971, 1978). This
type
of relatively
warm and
equable
climate
with low latitudinal
temperature gradients
is known
as "reduced
subtropical,"
"sub-
tropical-warm-temperate"
(Kafanov, 1982) or "paratropical"
(Wolfe, 1972) and has no analogs
in the modern world.
The molluscan
assemblage
of the upper
Middle
Eocene
Agli-
kitschian
Formation
of the Koryak
Upland (about 62?-63?N),
which includes
Arctomelon
harasewychi
n. sp., the oldest
known
species
of the genus, contains
a diverse warm-water
molluscan
assemblage,
characterized
by temperate
and subtropical
genera:
Glycymeris,
Kelletia, Trochita, Fulgoraria, Ficopsis, Ancilla
(Weaver,
1942;
Oyama,
et al., 1960;
Oleinik, 1994).
Arctomelon
has not been recorded
previously from either the Eocene of
Oregon,
Washington,
California,
and Alaska or from
Japan
and
Sakhalin.
Since
numerous
paleontological
studies
have been done
on Eocene mollusks of these areas, it is more likely that its
absence
is real,
rather
than related
to the incompleteness
of the
fossil record,
and indicates that the initial origin of the genus
was in the high latitude
northwestern
Pacific or that migration
of Arctomelon
or its ancestral
taxon to these latitudes
occurred
during
the Late Cretaceous-Early
Tertiary.
Possible southward
expansion of the genus during
the late
Eocene is suggested
by occurrence
of Arctomelon
rateginense
n.
sp., in the latest Eocene-(?) earliest Oligocene Rategian For-
mation
of Podkagernaya
Bay,
Northwestern
Kamchatka,
about
60?N
(Figure
7), together
with a diverse and abundant fauna
of
bivalves and gastropods:
Acesta,
Periploma, Thyasira,
Ostrea,
Fulgoraria,
Hataiella, Trominina,
Epitonium,
and Cancellaria.
This assemblage suggests
that warm climatic conditions per-
sisted until
the very end of the Eocene.
It is also notable for one
of the first occurrences
of a significant
number of gastropod
genera
of Turridae and
Buccinidae:
Aforia,
Pseudoliomesus,
An-
cistrolepis,
Colus
(Aulacofusus)
and
Beringius
(Gladenkov
et al.,
1988;
Oleinik, 1988;
Titova, 1993),
which
compose
a significant
percentage
of the Recent boreal molluscan assemblage
in the
Northern
Pacific
(Golikov, 1980; Goryachev, 1978; Kafanov,
1982).
Occurrences
of these taxa
also suggest
their
initial warm-
water
origin
and later
adaptation
to colder conditions.
Deterioration
of the warm
equable
climate started sometime
during
the Oligocene,
with apparently
more severe
temperature
decline in higher
latitudes.
Oxygen
isotope determinations
for
the equatorial
Pacific,
for instance, show gradual
reduction
of
the surface
water
temperatures
from 25?C
(11-12?C
bottom) in
the Middle
Eocene to 17?C
(5? bottom)
in the middle Oligocene
(Douglas
and Savin, 1971). Evidence
from fossil floras
in high
latitudes shows a major decline in mean annual temperatures
at the Eocene/Oligocene
boundary
in the northeastern
Pacific
from about 12-13?C at 60?N latitude
to about 10-11?C at 45?N
latitude
(Wolfe, 1971, 1978). Based on the study of the fossil
floras
of Hokkaido,
Tanai (1970) suggested
mean annual
early
Oligocene
temperatures
of 16-200C.
Fossil floras allowed doc-
umentation of a "cold"
period in the Northwestern Pacific
by
Kobayashi
and Shikama
(1961) for
Japan
and
by Vlasov
(1964),
Fotyanova (1977), and Akhmetiev (1976) for Sakhalin and
Kamchatka.
Estimates based on molluscan fossils have been
made for
the Northern
Pacific
mainly using
data from the west-
ern coast of North America. These data yield a range
of water
temperatures
in the early
Oligocene
of 5-8'C at bathyal
depth,
Blakeley Formation, Washington (Moore, 1984) to 10-12?C
(summer months) in outer shelf depths ("Narrow Cape For-
mation" of Sitkinak
Island,
Alaska) (Allison
and Marincovich,
1981). Oxygen isotope determinations based on Cyclocardia
shells from
the lower
Oligocene bathyal
deposits
of the Ilpinian
Peninsula,
Northeastern Kamchatka
(65-70?N),
showed bottom
temperatures
ranging
from 3? to 50C at bathyal
depths, which
can be compared
to those found between 32?N and 45?N at
present.
These data suggest
that the climatic conditions in the
high
latitudes at the beginning
of the Oligocene
were colder than
in the Eocene but still warmer than those of the Holocene.
Existence
of warmer
than present-day
climatic conditions in
this high-latitude
area
(60?N)
during
at least
the early Oligocene
can probably
explain
a wide distribution of fulgorariid
volutes
around the Northern Pacific
at that time (Oleinik, 1990, 1993).
The record of Arctomelon lautenschlageri
(Volobueva in
Devjatilova
and Volobueva, 1981) from the Oligocene
through
early Miocene of the Koryak
Upland (Figure
7) shows its ex-
istence in outer shelf to bathyal
environments at that time to-
gether
with other
gastropod genera
such as Trominina,
Fulgo-
raria,
Bathybembix,
Buccinum,
Beringius,
and Colus (Aulaco-
fusus). This first bathyal occurrence coincides with the most
severe phase of the Oligocene cooling. Such alteration or ex-
pansion
of the habitat
may suggest
two principal
mechanisms:
climatic
changes
and competition.
Assuming
that fossil species
of Arctomelon
were carnivores
(based on Recent A. stearnsii,
which have a uniserial radula with tricuspid rachidian teeth
(Pilsbry
and Olsson, 1954;
Weaver and Du Pont, 1970) (Figure
1.3), they probably
competed with common and diverse Ful-
gorariinae.
These two groups
even may have had close, if not
identical,
food preferences.
High
levels of competition
with oth-
er more diverse and successful
Fulgorariinae,
therefore,
may
have caused retreat of Arctomelon offshore.
Based on the few
known fossil specimens,
arctomelons were not very
common or
numerous
during
the Tertiary,
especially
before the Oligocene.
Their successful survival and wide distribution
in cold waters
today was probably
a result
of their replacement
of the Fulgo-
rariinae,
a group that apparently
was unable to adapt to the
onset of cold conditions and retreated
southward
during the
final cooling of the late Cenozoic. Reduction of competition
pressure
for the genus probably
caused an increase
in species
diversity
in the Holocene. The fossil record
indicates that
Arc-
tomelon was not a very successful
taxon during most of the
Cenozoic and only cooling of climate in the late Tertiary
and
Quaternary
gave Arctomelon an evolutionary
opportunity
not
only for successful survival and diversification
in the reduced
competition
of the cold waters
of the northern
Pacific,
but also
the wider
geographic
distribution of the genus seen today. The
low latitude occurrence
ofArctomelon benthalis at abyssal depths
of the Eastern Pacific and Arctomelon tamikoae
at lower sub-
littoral depths in the South China Sea can be viewed as an
x 1. 5, abapertural view; 6, apertural
view. 7, 8, Arctomelon
lautenschlageri (Volobueva, 1981), drawing
(new herein)
of the holotype from
Devjatilova
and Volobueva, 1981, pl. XXXIII, figs.
4 a, b, x 1. 7, abapertural view; 8, apertural
view. 9, 10, Arctomelon
sp., hypotype
IGM
13/159. 9, abapertural
view, x 1; 10, apertural view, x 1.2. 11, 12, Arctomelon
lautenschlageri (Volobueva,
1981), hypotype
USNM 483412,
x 1. 11, abapertural
view; 12, apertural
view.
243
JOURNAL OF PALEONTOLOGY,
V. 70, NO. 2, 1996
FIGURE 7-The stratigraphic
distribution and
types
of habitats of the
species
of Arctomelon
in the
Northwestern Pacific
plotted
against
the
paleoclimatic
curve. Habitats:
1-bathyal,
2-outer shelf,
3-inner
shelf.
Climates:
A-High boreal-Arctic,
CT-cool temperate,
WT-
warm
temperate, P-paratropical.
attempt
toward
further
southward
expansion
of the genus
in the
Quaternary.
Relatively cold conditions existed in high latitudes
through
the Oligocene
and for most of the early
Miocene.
The first
Neo-
gene climatic
warming
that is well documented and recognized
both on a global
scale and for the North Pacific
(Chinzei, 1978;
Masuda, 1978;
Gladenkov and
Sinelnikova, 1990;
Margolis
and
Kennett, 1970) began
in the late early Miocene and continued
during
the early
middle Miocene.
The southernmost
known fossil occurrence ofArctomelon lau-
tenschlageri
and Arctomelon
sp. from the upper
lower
Miocene
Kuluvenian Formation of the central part of Western Kam-
chatka
(about 56?-57?N)
coincides with the first Neogene cli-
matic warming.
Evidence from this area shows that, regardless
of its southward
migration,
and rising
of the temperature
of the
seawater
during
the early middle Miocene climatic optimum,
Arctomelon
kept
inhabiting
outer shelf to possibly upper
bathyal
environments
(Figure 7).
Evidence from the fossil record shows that Arctomelon did
not
undergo any
significant
changes
in external
shell
morphology
during
the Cenozoic, and especially
since the early Oligocene.
There
are, however,
a few morphologic
features that allow sep-
aration of the Recent Arctomelon
stearnsii
from Oligocene-
early
Miocene
Arctomelon
lautenschlageri.
Comparison
of shell
morphologies
of A. stearnsii
and A. lautenschlageri
indicates
a
general
trend
towards a more
slender
shell shape
and narrowing
of the aperture.
The two Eocene species, on the other hand,
seem to be slightly
more variable
in shape, outline of the ap-
erture,
number
and strength
of the columellar
plaits, and ex-
ternal
shell
sculpture.
However,
even these features have a fairly
low degree
of variation. It can be generally
concluded from the
comparison
of the shell
morphology
of fossil taxa that all major
changes
in shell morphology
occurred
during
the Eocene
when
the genus inhabited
shallow and warm marine environments.
Even after
major
changes
in climatic
conditions and habitat
at
the beginning
of the Oligocene (Figure 7), shell morphology
appears
to have remained
quite conservative,
except for an in-
crease
in size. Most
Recent
adult
specimens
ofA. stearnsii
range
in length
from 100 to 125 mm, whereas
fossil species
vary
from
45 to 80 mm in length.
The reason
for such morphologic
con-
servatism
is unclear.
Conservatism
in morphology may be an
indication
of environmental
stability
or a lack of competitors.
This phenomenon
may also be related
to an isolation
and, as a
consequence,
restriction of gene flow, or to adaptation
to a cer-
tain ecologic
niche. Both cases are known
and documented
for
marine
gastropods (Batten, 1984;
Hickman, 1976, 1984;
Houb-
rick, 1984; Stanley, 1984). Conservative shell morphology
of
Arctomelon
through
the Cenozoic
can probably
be attributed to
geographic
isolation of the genus in the high latitudes of the
northwestern
Pacific
since early Tertiary.
Since the Oligocene,
it can also be related to the existence of the genus in a deeper,
colder,
and more stable
environment.
Arctomelon
ancestry
is not quite clear
from the fossil record.
Occurrence
of the Zidoninae
in the northern
Pacific is unusual.
Zidoninae are common mostly in the Southern
Hemisphere
(Australia,
New Zealand, and South America). The northern
part of the Pacific, on the other hand, is well known for the
diverse Fulgorariinae,
which constitutes
a unique component
of both Recent and Tertiary
molluscan faunas of this region.
Both Fulgorariinae
and Zidoninae are restricted
in their distri-
bution due to lack of planktonic larvae. "Bipolarity"
in the
distribution of the Zidoninae
may suggest
an origin
of the com-
mon ancestor of the Zidoninae in the lower latitudes of the
Pacific
region
and later
dispersal
in the northern
and the south-
ern Pacific.
The
Cretaceous
and
early
Paleogene
fossil
record
in the north-
western
Pacific does not have any known
taxon which can pro-
vide
evidence of direct or
indirect
ancestry
to Arctomelon.
Among
Cretaceous
mollusks
of the eastern
margin
of the Pacific
Ocean,
the volutid genus Varens Saul and Popenoe, 1993, from the
Turonian
of California,
although
referred
to the subfamily
Vol-
utoderminae
Pilsbry
and
Olsson,
1954
(Saul
and
Popenoe,
1993),
can be viewed as a possible ancestral
taxon for the subfamily
Zidoninae in the Pacific,
based on the similarity
of the external
shell morphology
with most members of Zidoninae.
The evolutionary
history
of the genus
can probably
be inter-
preted
as gradual
evolution from the middle Eocene A. hara-
sewychi
to the Recent
A. stearnsii.
Together
with
this
conclusion,
one must
keep
in mind that
if we accept
an origin
of A. stearnsii
from A. lautenschlageri,
we are
left with a gap
of about 16.3 my
with no records
of the genus (Figure 7). That gap may contain
at least one undiscovered
species
from middle-late
Miocene
or
Pliocene. It is not surprising
that no Pleistocene
species
of Arc-
tomelon have yet been reported.
This fact can be explained
by
a lack of deep-water
Pleistocene
deposits in the North Pacific
realm.
ACKNOWLEDGMENTS
The author
gratefully
acknowledges
V. N. Sinelnikova
of the
Russian
Academy
of Sciences
Institute
of Geology
for
providing
several
specimens
for the study, W. J. Zinsmeister
of the De-
partment
of Earth
and Atmospheric
Sciences,
Purdue Univer-
sity,
for discussions
on the text
of the manuscript,
critical review
of the manuscript
and helpful
comments,
M. G. Harasewych
of
the Division of Mollusks,
National
Museum of Natural
History,
Smithsonian
Institution,
for
providing photographs
of the para-
244
OLEINIK- TERTIAR Y GASTROPOD
type of Arctomelon stearnsii from the Smithsonian collections
and valuable critique and suggestions for improvement of the
manuscript. This paper was considerably improved by a critical
review by L. N. Marincovich of the United States Geological
Survey, Menlo Park, California. The author is also grateful to
W. D. Allmon of the Paleontological Research Institution, Ith-
aca, New York, and C. R. Givens of the Department of Earth
Science, Nicholls State University, Thibodaux, Louisiana for
review and comments on the manuscript. This work would not
have been possible without the support of the Department of
Earth and Atmospheric Sciences, Purdue University, which pro-
vided laboratory facilities.
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