Palaeogeography, Palaeoclimatology, Palaeoecology 267 (2008) 138–146

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Palaeogeography, Palaeoclimatology, Palaeoecology

j o u r n a l h o m e p a g e : w w w. e l s e v i e r. c o m / l o c a t e / p a l a e o

Paleobiogeography of the pectinid bivalve Neithea, and its pattern of step-wise

demise in the Albian Northwest Pacific
Yasuhiro Iba a,⁎, Shin-ichi Sano b
a
Department of Earth and Planetary Science, Graduate School of Science, University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan
b
Fukui Prefectural Dinosaur Museum, Katsuyama, Fukui 911-8601, Japan

a r t i c l e i n f o a b s t r a c t

Article history: The pectinid bivalve genus Neithea is one of the most important indicators for understanding the
Received 4 December 2007 biogeographic relationships between the Tethyan Realm and North Pacific Province during the Cretaceous
Received in revised form 22 June 2008 Period. Changes in temporal species diversity, endemic/widespread species composition, and origination
Accepted 4 July 2008
and demise ratios of Neithea at each Cretaceous stage boundary in the Northwest Pacific were analyzed from
a biogeographic perspective. Neithea is continuously present in the Northwest Pacific during the Berriasian
Keywords:
Mid-Cretaceous
to late Albian time interval. Species diversity reached its maximum in the late Aptian, being correlated with
Greenhouse period the global warming phase. Step-wise demise of Neithea in the Northwest Pacific during the Albian is
Neithea subdivided into three stages: the late Aptian/early Albian, early Albian/middle-late Albian, and late Albian/
Paleobiogeography early Cenomanian. Thereafter, Neithea disappeared in the Northwest Pacific and never reappeared. This
North Pacific pattern is the reverse of the Albian diversification of Neithea in the Mediterranean, and also contrary to the
Tethys Mid-Cretaceous global warming trend. Demise of Neithea in the Northwest Pacific occurred simultaneously
Diversity with the step-wise demise of Mesogean taxa (e.g., rudists) which strongly supports the idea that the
Northwest Pacific gradually became independent from the Tethyan Realm during the Albian. It also suggests
a long-term deterioration of the faunal interchange between the North Pacific Province and Tethyan Realm
throughout the Late Cretaceous. This biogeographic change was possibly caused by Albian “cooling” and
changes in oceanic flow/heat transport in the Northwest Pacific.
© 2008 Elsevier B.V. All rights reserved.

1. Introduction Realm. The North Pacific Province was clearly distinguishable during
the Late Cretaceous Epoch. Late Cretaceous bivalve faunas in the
Mid-Cretaceous is a well-documented greenhouse period of global Northwest Pacific contain many endemic taxa, which first appeared in
importance during the Earth's history (Johnson et al., 1996; Clarke and the Albian (e.g., Hayami and Yoshida, 1991; Tashiro, 2000). Thus it is
Jenkyns, 1999; Wilson and Norris, 2001; Huber et al., 2002; Steuber et expected that remarkable biotic changes occurred in the mid-
al., 2005). The typical Tethyan biota (Mesogean taxa in the sense of Cretaceous Pacific, already at that time the world's largest aquatic
Masse, 1992) (e.g., rudists and orbitolinid foraminifers), extensively reservoir.
flourished within tropical shallow marine settings in the world's In addition to Mesogean taxa, some bivalves (e.g., Neithea and
oceans throughout the Cretaceous, and therefore are regarded as Chondrodonta), for which a term “Tethyan non-rudist bivalves” was
essential indicators of tropical realm and climate (Masse, 1992). coined (Dhondt, 1992; Dhondt and Dieni, 1992), inhabited warm
Recently, Iba and Sano (2007) summarized the Cretaceous record of shallow marine environments, together with Mesogean taxa. For this
Mesogean taxa (sensu Masse, 1992) mainly from clastic sequences of reason, they are also considered as a good indicator of the Cretaceous
the Northwest Pacific, and described their demise during latest Tethyan Realm and warm climatic environment. The pectinid bivalve
Aptian–middle Albian. Iba and Sano (2007) explained this bio-event Neithea, has often been used for Cretaceous biogeographical studies
by means of vicariance, which led to the establishment of the North in Europe, the Mediterranean, Western Interior Seaway, and South
Pacific Province (Jeletzky, 1971) being independent from the Tethyan Atlantic (e.g., Dhondt, 1981, 1985, 1992; Dhondt and Dieni, 1991, 1992;
Kauffman et al., 1993; Andrade et al., 2004). This genus commonly
occurs in the Cretaceous shallow marine calcareous deposits in the
Northwest Pacific, and has the most abundant and continuous record
⁎ Corresponding author. Tel.: +81 3 5841 4072; fax: +81 3 5841 4555. among the Tethyan non-rudist bivalves in this region (e.g., Hayami,
E-mail address: [email protected] (Y. Iba). 1975; Hayami and Noda, 1977; Iba and Sano, 2007). There are many

0031-0182/$ – see front matter © 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2008.07.002
 Y. Iba, S. Sano / Palaeogeography, Palaeoclimatology, Palaeoecology 267 (2008) 138–146 139

taxonomic, stratigraphic, and paleontological studies of Neithea in the by Hayami (1965) as junior synonyms of European species. Sub-
Northwest Pacific, and therefore, we can easily compare its spatio- sequently, Hayami (1975) and Hayami and Noda (1977), with reference
temporal distribution pattern in the Northwest Pacific with other to Dhondt's (1973) interpretation, revised the taxonomy of Japanese and
regions. Although a mid-Cretaceous “local extinction” of Neithea in the Taiwanese species and described eight species from this region
Northwest Pacific has been recognized (Hayami, 1989; Hayami and (N. aketoensis, N. atava, N. ficalhoi, N. kochiensis, N. matsumotoi,
Yoshida, 1991) as an important biotic change in the Cretaceous Pacific, N. nipponica, N. notabilis, N. syriaca amanoi). Subsequently, Tashiro and
its detailed process, timing, and paleobiogeographic significance Kozai (1986) described one new species (N. hanourensis). Species of
however remain unknown. The present study analyses statistically Neithea from Northwest Pacific are characterized by two well-developed
all the available data on Neithea in the Northwest Pacific, in order to secondary ribs between each two tripartite principal ribs, and have been
elucidate its spatiotemporal occurrence pattern. Furthermore, we identified as the Mediterranean species N. ficalhoi (Choffat, 1888)
compare the spatiotemporal distribution patterns of Neithea in the (Hayami, 1965; Hayami and Noda, 1977; Tashiro and Kozai, 1986; Tanaka
Northwest Pacific with those in Mediterranean region, and other et al., 1999, 2002; Kawano et al., 2002), which recently has been
contemporaneous biotic changes in the Northwest Pacific. Finally, we synonimized with N. alpina (d'Orbigny, 1847) (Andrade et al., 2004).
discuss the mid-Cretaceous paleobiogeographic changes of marine Herein we followed the interpretation of Andrade et al. (2004).
biota in the Northwest Pacific and its possible causes. The taxonomic status of N. kochiensis and N. aketoensis described
from the Aptian deposits of Japan by Hayami (1965) and Hayami
2. Note on taxonomy of Neithea in the Northwest Pacific and Kawasawa (1967) remains unclear due to scarce and poorly pre-
served material. N. kochiensis was proposed by Hayami and Kawasawa
Cretaceous shallow marine deposits are widely distributed in the (1967) based on a poorly preserved inner mould of a specimen
Northwest Pacific margin (Taiwan–Japanese Islands) which occupied that possess no prominent secondary ribs. The feature of secondary
the eastern margin of the Asian Continent during this period. These ribs of N. kochiensis is possibly a misinterpretation due to its poor
yield numerous well-preserved macro- and microfossils from various preservation. Well-preserved specimens of alleged N. kochiensis
horizons. Since the first report of Neithea from the Northwest Pacific were reported by Tanaka et al. (1996), but the rib morphology and
by Yabe et al. (1926), Neithea has been reported from many localities distribution pattern of these specimens resembles that of N. atava
in this region (Fig. 1 and Table 1). Although Cretaceous marine de- (Roemer, 1839), a species which displays worldwide distribution
posits are distributed in Northeast China and Far East Russia inclusive of the Japanese Islands. N. aketoensis Hayami (1965) was
(Sikhote-Alin and Kamchatka), there is no documented occurrence based on a single specimen from the upper Aptian of the Hiraiga
of Neithea in these regions. Several species of Neithea were reported Formation on the Pacific coast of the Northeast Honshu (Loc. 10 in
from the mid- to Upper Cretaceous in Tibet and Tarim Basin, western Fig. 1). We re-examined the type specimen, and concluded that the rib
China (e.g., Wen, 1999). However, since these seas were not morphology and its distribution pattern both on the inner mould and
connected directly to the Northwest Pacific, but to the Tethys Sea external shell surface resembled those of N. nipponica Hayami (1965).
(e.g., Chen, 1987; Wen, 1999), these occurrences are not discussed in Therefore, N. aketoensis should be considered as a junior synonym
this paper. of N. nipponica. Taking into account the discussion above we excluded
Detailed taxonomic studies of Neithea species from the Northwest N. kochiensis and N. aketoensis from further consideration in this
Pacific have been done by Hayami (1965) and Hayami and Kawasawa paper. Detailed taxonomic revision of these two species will be pro-
(1967). Dhondt (1973) regarded several endemic species described vided elsewhere.

3. Material and methods

Eight species of Neithea (N. alta, N. atava, N. hanourensis, N. notabilis,

N. matsumotoi, N. alpina, N. syriaca amanoi, N. nipponica) reported
from more than 60 publications in Taiwan–Japanese Islands (see
Appendix) are considered in the present study. We have not taken into
account the species of Neithea left in open nomenclature. The objec-
tives are to clarify temporal diversity changes, demise and origination
ratios, and endemic/widespread species compositions in the surveyed
region. The Aptian–Albian time interval is a crucial period for marine
paleobiogeography in the Northwest Pacific (e.g., Iba and Sano, 2006,
2007) and so the interval is analyzed to the substage level. However,
because of difficulty in recognizing the middle Albian stage in the all
circum-North Pacific regions due to the paucity of index fossils, we
treated middle and late Albian jointly.
We calculated demise and origination ratios at each stage and/or
substage boundary, and then attributed the biogeographic-type of
species (i.e., endemic or widespread species) for each stage and sub-
stage. Demise ratio (DR) and origination ratio (OR) are defined as
follows: DR = (number of preexisting species absent above each
boundary)/(total number of species below each boundary), OR =
(number of successor species not present below each boundary)/
(total number of species above each boundary). The ratios of endemic
and widespread species were examined based on previous biostrati-
graphic, biogeographic and taxonomic studies of each species in the
Europe, Mediterranean, Caribbean–Western Interior Seaway, and
Fig. 1. Occurrences of Neithea in the Cretaceous of Northwest Pacific. All locality numbers
Atlantic (Dhondt, 1973, 1981, 1982, 1992; Dhondt and Dieni, 1991,
are compatible with those from Table 1. HK; Hokkaido Island, HO; Honshu Island, SH; 1992; Kauffman et al., 1993; Bogdanova and Yanin, 1995; Kues, 1997;
Shikoku Island, KY; Kyushu Island, to; Tohoku area, ka; Kanto area, ki; Kinki area. Andrade et al., 2004). Endemic species are defined here as species that
140 Y. Iba, S. Sano / Palaeogeography, Palaeoclimatology, Palaeoecology 267 (2008) 138–146

Table 1
Neithea species reported from the Cretaceous of the Northwest Pacific

Species Original Biogeographic Stratigraphic range in Northwest Pacific Formation

designation type of species (in Mediterranean region) (locality and loc. no.)
N. alta Hayami in Hayami and Endemic Berriasian Ayukawa Formation (Tohoku, 1)
Noda (1977)
N. atava Roemer (1839) Widespread Hauterivian–late Aptian Kimigahama (Kanto, 2), Ishido (Kanto, 3),
(Beriasian–Albian?) Idaira (Kanto, 4), Arida (Kinki, 5), Lower Hanoura,
Hanoura (eastern Shikoku,6), Monobe,
Lower Monobe (central Shikoku), 7),
Haidateyama, Osaka (eastern Kyushu, 8),
Sanpozan, Hachiryuzan,
Hinagu (western Kyushu, 9) formations
N. hanourensis Tashiro and Kozai (1986) Endemic Barremian Lower Hanoura Formation (eastern Shikoku, 6)
N. notabilis Von Münster in Widespread Barremian–late Aptian Ashikajima, Kimigahama (Kanto, 2),
Goldfuss (1833) (Neocomian–Turonian) Ishido (Kanto, 3), idaira (Kanto, 7), Hiraiga
(Tohoku, 10) formations
N. matsumotoi Hayami (1965) Endemic Barremian–late Albian “Sebayashi” (Kanto, 3), Doganaro
(central Shikoku, 7), Sukubo, Haidateyama
(eastern Kyushu, 8) Hachiryuzan, Kesado,
(western Kyushu, 9) formations and Upper Aptian
of Peikang area (Taiwan, 11)
N. alpina d'Orbigny (1847) Widespread Early–late Aptian Hibihara (central Shikoku, 7),
(Albian–Maastrichtian) Osaka (eastern Kyushu, 8), Tomochi, Imaizumigawa
(western Kyushu, 9), Hiragia (Tohoku, 10),
Shuparogawa? (central Hokkaido, 12) formations
N. syriaca amanoi Hayami (1965) Widespread Early Aptian–early Albian Ashikajima, Kimigahama (Kanto, 2),
(see text for detail) (Barremian–Cenomanian = range of Bunjo, Hagino (central Shikoku, 7),
N. syriaca syriaca) Tamarimizu, Osaka (eastern Kyushu, 8),
Kesado (western Kyushu, 9),
Kamiji (northern Hokkaido, 13) formations
N. nipponica Hayami (1965) Endemic Late Aptian–early Albian Hiraiga and Aketo formations (Tohoku, 10)
N. “kochiensis” Hayami in Hayami and Uncertain taxonomic position Doganaro (central Shikoku, 7) and
Kawasawa (1967) (see text for detail) Osaka (eastern Kyushu, 8) formations
N. “aketoenis” Hayami (1965) Uncertain taxonomic position Aketo Formation (Tohoku, 10)
(see text for detail)

See Appendix for references in each area. All localities are comparable to Fig. 1.

are known exclusively in the Northwest Pacific region, whereas 4. Results: spatiotemporal changes in Neithea species in the
widespread species are those that have been recorded from other Cretaceous Northwest Pacific
regions as well. In addition, the subspecies (N. syriaca amanoi) is
considered here to be a widespread species. Neithea is known from 32 formations in the Northwest Pacific
(Fig. 1 and Table 1). Stratigraphic distribution of each species is shown
in Fig. 2. The earliest record of Neithea in this region is known from the
Berriasian, and it occurs almost continuously up into the upper Albian
(Fig. 2). We could not obtain any specimens from Valanginian strata
because shallow marine deposits of this age have a very restricted
distribution in the Northwest Pacific.
Neithea alta, N. hanourensis, N. nipponica, and N. matsumotoi (Fig. 3)
are all endemic species, whereas N. atava, N. notabilis, N. alpina, and
N. syriaca are widespread species (Table 1). Species diversity clearly
increased during the Berriasian to late Aptian interval, when it
reached a maximum of six species (Fig. 4A). Subsequently, the diversity
gradually decreased during the early to late Albian (Fig. 4A). There is
no record of Neithea in the post-late Albian Cretaceous. Low demise
ratios were obtained for the Hauterivian/Barremian (0), Barremian/
early Aptian (0.25), and early Aptian/late Aptian (0) boundaries (Fig. 4B).
Thereafter demise ratios increased during the late Aptian/early Albian
to the middle–late Albian/Cenomanian (0.50, 0.66 and 1 for the
late Aptian/early Albian, early Albian/middle–late Albian and middle-
late Albian/early Cenomanian, respectively) (Fig. 4B). Origination
ratios gradually decreased during the Hauterivian/Barremian to early
Aptian/late Aptian (0.75, 0.40, and 0.16 for the Hauterivian/Barremian,
Barremian/early Aptian and early Aptian/late Aptian, respectively). In
addition, no origination occurred in the late Aptian/early Albian, early
Albian/middle-late Albian, and middle-late Albian/early Cenomanian
(Fig. 4B). In particular, high demise ratios in the absence of origination
are characteristic of the late Aptian/early Albian, early Albian/middle-
late Albian, and middle-late Albian/early Cenomanian boundaries
(Fig. 4B). The ratio of widespread species gradually decreased during
Fig. 2. Stratigraphic ranges of Neithea species in the Cretaceous of Northwest Pacific. early Aptian to early Albian (80, 66, and 33% for early Aptian, late Aptian,
 Y. Iba, S. Sano / Palaeogeography, Palaeoclimatology, Palaeoecology 267 (2008) 138–146 141

Fig. 3. Endemic species of Neithea in the Northwest Pacific. A) N. alta. B) N. hanourensis C) N. matsumotoi. D) N. nipponica. A is the holotype, B–D are plaster models of holotypes. All
specimens are right valves. Scale-bars are 1 cm.

Fig. 4. Spatiotemporal changes of Neithea in the Northwest Pacific. A) species diversity changes. B) ratios of demise (DR) and origination (OR) at each stage and/or substage boundary.
C) ratios of endemic and widespread species for each stage and/or substage. D) important features of Neithea spatiotemporal changes.

and early Albian, respectively), and widespread species were absent in phase, the Aptian Greenhouse Earth II period determined by Weissert
middle-late Albian (Fig. 4C). and Lini (1991). The maximum diversity of Neithea is most probably
related to this Late Aptian warming phase.
5. Discussion
5.2. Step-wise demise of Neithea in the Albian of the Northwest Pacific
5.1. Diversification phase of Neithea during the Early Cretaceous
The Albian demise of Neithea in the Northwest Pacific can be sub-
Neithea originated in the Berriasian, thrived in the mid-Cretaceous, divided into three stages (Fig. 4D); Stage 1 (late Aptian–early Albian
then decreased in diversity and finally became extinct in the terminal interval), Stage 2 (early Albian to middle–late Albian interval), and
Cretaceous (Dhondt, 1981, 1992). The first appearance of Neithea in the Stage 3 (late Albian–early Cenomanian interval). In Stage 1, three wide-
Northwest Pacific is virtually synchronous with the earliest occur- spread species, N. atava, N. notabilis, and N. alpina disappeared. Species
rences (Berriasian) of Neithea in the Tethyan region (Dhondt, 1973). diversity started to decrease coincident with no origination and high
Since its first appearance in the Berriasian, Neithea was continuously demise ratios from this stage (Fig. 4). In Stage 2, widespread species
present until the late Albian in the Northwest Pacific. Iba and Sano disappeared completely and only one species N. matsumotoi could
(2006, 2007) concluded that tropical–subtropical conditions prevailed survive. In Stage 3, Neithea completely disappeared in the Northwest
in the Northwest Pacific during the Berriasian to Albian, based on Pacific and never reappeared (Figs. 2 and 4). Although Upper Cretaceous
spatiotemporal distribution patterns of Mesogean taxa. The continu- shallow marine deposits containing abundant molluscan fossils such
ing presence of Neithea in the Northwest Pacific during this interval as trigonid and ostreid bivalves are distributed widely in the Northwest
strongly supports Iba and Sano's (2006, 2007) conclusions. After a Pacific margin (e.g., Komatsu, 1999; Komatsu and Maeda, 2005; Ando,
period of gradual increase in diversity of Neithea species in the 2003), there have been no reports of Neithea in this period. This suggests
Northwest Pacific during the Hauterivian to late Aptian, they reached that the demise of Neithea was not the result of facies change and/or lack
their maximum diversity in the late Aptian. This coincides with late of Upper Cretaceous marine strata.
Aptian northward development of large carbonate platforms, inhab- In the Northwest Pacific three widespread species, N. atava,
ited by diverse Mesogean taxa (Sano, 1995; Iba and Sano, 2006, 2007; N. notabilis, and N. alpina, disappeared in Stage 1 and N. syriaca in
Takashima et al., 2007). This northward expansion of carbonate plat- Stage 2, however, these taxa are known to have been in Tethys sea
form distribution in the Northwest Pacific is interpreted by Takashima (e.g., Mediterranean region) until the Albian, Turonian, Maastrichtian,
et al. (2007) as being a consequence of the Late Aptian global warming and Cenomanian, respectively (Table 1) (Dhondt, 1973). These lines of
142 Y. Iba, S. Sano / Palaeogeography, Palaeoclimatology, Palaeoecology 267 (2008) 138–146

evidence illustrate the different stratigraphic ranges of the same species deed in any other regions of Tethys. Therefore, the biotic change of
between the Northwest Pacific and Tethyan regions and should Neithea in the Northwest Pacific was a unique bio-event in this area.
therefore been considered as a demise of Neithea in the Northwest
Pacific, and not the extinction of the species. It indicates that some 5.3. Paleobiogeographic implications of Neithea demise
profound paleoceanographic changes caused the demise of Neithea in
the Northwest Pacific. Contemporaneous and profound long-term biotic changes took
In order to investigate the disparity in temporal change of Neithea place in the Northwest Pacific during the mid-Cretaceous. Iba and
species diversity between the Northwest Pacific and the Mediterranean Sano (2006, 2007) have analyzed the gradual demise pattern of
region (Fig. 5), available data from Dhondt (1973, 1982, 1985), Dhondt Mesogean taxa (sensu Masse, 1992) in the mid-Cretaceous North-
and Dieni (1993), and Perrilliat et al. (2006) is compiled in this study. west Pacific and revealed that Mesogean key reference taxa (rudists
The data reveals that in the Mediterranean region Neithea gradually and dasyclads) and some Mesogean indicators (hermatypic corals
diversified during the Berriasian to Cenomanian, at which point it and stromatoporoids) disappeared in the latest Aptian to early
reached maximum diversity (15 species), and thereafter species diver- Albian interval. Iba and Sano (2007) coined the term “Mesogean
sity decreased until its extinction in the terminal Cretaceous (Fig. 5). key taxa demise event” to describe the simultaneous and gradual
Mid-Cretaceous times are known to be a typical greenhouse period, and disappearance of several taxa. That event was followed by the final
a significant warming trend during Late Aptian to Turonian has been disappearance of all Mesogean indicators in the early Albian to
reconstructed from oxygen isotopic records (Fig. 5) (e.g., Clarke and middle Albian interval in the Northwest Pacific (“Mesogean indi-
Jenkyns, 1999; Wilson and Norris, 2001; Huber et al., 2002; Steuber cators demise event” of Iba and Sano, 2007). This step-wise demise
et al., 2005). The Albian diversification of Neithea in the Mediterranean of Mesogean taxa clearly indicates that the Northwest Pacific
region is consistent with the mid-Cretaceous global warming trend became independent from Tethyan Realm during latest Aptian to
and sea-level rise (Fig. 5). The step-wise demise of Neithea during middle Albian, and led to the establishment of the North Pacific
the Albian in the Northwest Pacific is opposite and counterpart to the Province (Iba and Sano, 2007) (Fig. 6).
diversification trend seen in the Mediterranean, and contrary to the Stages 1 and 2 of Neithea demise are simultaneous with the
mid-Cretaceous global warming trend (Fig. 5). This gradual demise of Mesogean key taxa demise event and Mesogean indicator demise
Neithea and its subsequent long-term absence throughout the Late event, respectively. This shows that the step-wise demise of biota in the
Cretaceous has not been recorded in Mediterranean, Caribbean, or in- Northwest Pacific can be recognized not only in Mesogean taxa but

Fig. 5. Graph of Neithea species diversity in the Northwest Pacific and the Mediterranean region plotted against Cretaceous global changes in sea-level and climate. Species diversity
of Neithea in the Mediterranean region compiled from Dhondt (1973, 1982, 1985), Dhondt and Dieni (1993), and Perrilliat et al. (2006). Climatic changes based on Clarke and Jenkyns
(1999) (Exmouth Plateau) and Steuber et al. (2005) (Mediterranean). Global sea-level changes based on Hardenbol et al. (1998). Cretaceous time scale based on Ogg et al. (2004).
 Y. Iba, S. Sano / Palaeogeography, Palaeoclimatology, Palaeoecology 267 (2008) 138–146 143

Fig. 6. Illustrating the changes in distribution of Neithea and Mesogean taxa, and changing northern limit of the Tethyan Realm in the Cretaceous. A) Early Cretaceous, B) Late
Cretaceous. Map for the Early (120 Ma) and Late Cretaceous (80 Ma) based on Barron et al. (1981). Northern limit of the Tethyan Realm in Mediterranean region based on Masse (1992)
and Voigt et al. (1999).

also in the Tethyan non-rudist bivalve Neithea, though the Albian de- factor for the distribution and immigration of tropical biota in the
mise of the Neithea (three stages) was more protracted than the demise Northwest Pacific (e.g., Iba and Sano, 2007) (Fig. 6).
of Mesogean taxa (two stages). These profound Albian “cool” conditions and changes in oceanic
Demise of widespread species of Neithea in Stages 1 and 2 indicate circulation/heat transport would affect the demise of Tethyan biota
start of a weakening in the faunal connection between the Northwest (i.e., Mesogean taxa and Neithea) in the Northwest Pacific and also
Pacific and Tethys, whilst the complete demise of Neithea in Stage 3 restrict faunal immigration into the Northwest Pacific from the Tethys
suggests that the faunal connection between the two oceans deterio- Sea (Fig. 6). This new hypothesis of paleobiogeography and paleocea-
rated significantly in this stage (Fig. 6). During the mid-Cretaceous nography in the mid-Cretaceous Northwest Pacific warrants verifica-
warming period, widespread species that disappeared in the Northwest tion and further research.
Pacific in Stages 1 and 2 (e.g., N. atava, N. notabilis), and other wide-
spread species (e.g., N. hispanica, N. sexangularis, N. reguraris) were 6. Conclusions
widely distributed in other oceans (Dhondt, 1973, 1982; Dhondt and
Dieni, 1993; Andrade et al., 2004; Perrilliat et al., 2006). However, these 1. Neithea was continuously distributed in the Northwest Pacific
widespread species never penetrated and/or re-immigrated into the during the Berriasian to late Albian, suggesting that the Northwest
Northwest Pacific (Fig. 6). Long-term absence of Neithea in the Late Pacific was not distinct biogeographically from the Tethyan Realm
Cretaceous of the Northwest Pacific could have resulted from restricted during this time interval. During the Hauterivian to late Aptian, the
faunal interchange between Northwest Pacific and other oceans during Northwest Pacific species diversity of Neithea increased gradually
that time. with a low demise rate and a high origination rate, reaching a
maximum in the late Aptian. This most probably occurred in con`-
5.4. Possible causes of Neithea demise in the Northwest Pacific junction with the global warming phase at that time.
2. The step-wise demise of Neithea in the Northwest Pacific is re-
Recently, the early Albian “cooling” episode is recognized in the cognized and subdivided into three stages: at the late Aptian–early
Northwest Pacific (Iba, in press). A typical Arctic-type ammonite Albian interval (Stage 1), early Albian to middle–late Albian interval
Arcthoplites (Subarcthoplites) sp. was discovered from the lower Albian (Stage 2), and in the late Albian–early Cenomanian (Stage 3).
of northern Hokkaido, northern Japan (Loc. 13 in Fig. 1). Iba (in press) Following the Albian, Neithea disappeared in the Northwest Pacific
considered this southward distribution of Arctic-type ammonite as and never reappeared. This pattern is unique to the Northwest
the appearance of a distinct “cooling” episode in the early Albian Pacific as it has not been recorded from any other regions of
Northwest Pacific. Tethyan Realm. This pattern is the reverse of the Albian diversifica-
Results of some recent climatic model simulations can explain tion of Neithea in the Mediterranean region and Mid-Cretaceous
this Albian “cooling” episode in the Northwest Pacific. The Aptian– global warming trend.
Albian interval should be paid attention as a time of major changes in 3. Stages 1 and 2 of Neithea demise coincide in timing with the
paleogeographic/paleoceanographic settings, due to the formation of Mesogean key taxa demise event and Mesogean indicator demise
new oceanic gateways. It has been postulated that the opening of the event (sensu Iba and Sano, 2007), respectively. Gradual demise of
equatorial Atlantic gateway (EAG) changed not only the adjacent ocean Neithea during the Albian and its subsequent absence in the
current system, but also those in the Pacific (Poulsen et al., 2003; Northwest Pacific strongly supports the idea that the independence
Poulsen and Huynh, 2006). Recently, Poulsen and Huynh (2006) of the North Pacific Province from the Tethyan Realm was gradual,
numerically simulated the conditions of oceanic circulation in the and possibly caused by long-term deterioration of the faunal inter-
Albian Pacific, paying special attention to the increase in atmospheric change. This deterioration of faunal interchange could be explained
CO2 and opening of the EAG. As a consequence of the EAG opening, the by Albian “cooling” conditions and changes in oceanic circulation/
simulation suggested the generation of a relatively cold water mass in heat transport in the Northwest Pacific.
the Pacific, contrasting with the Atlantic and Indian Oceans. In addition,
an increase in atmospheric CO2 would strongly weaken the paleo- Acknowledgements
Kuroshio warm current, a clockwise current flows from south to north
along the Northwest Pacific margin (Poulsen and Huynh, 2006). The We thank to K. Tanabe (Univ. of Tokyo), A. Kaim (Polish Science
paleo-Kuroshio warm current was the most important controlling Academy) for the critical reading of the manuscript and to C.J.
144 Y. Iba, S. Sano / Palaeogeography, Palaeoclimatology, Palaeoecology 267 (2008) 138–146

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