Laguna (2022) 29: 11-25

ISSN 2185-2995

Short Article
Modern benthic foraminiferal fauna (Rhizaria)
in Miyazu Bay, central Japan
Hiroyuki Takata1, Satoshi Tanaka2, Hideki Sawada3, Keita W. Suzuki3
Abstract: We investigated modern benthic foraminifera in Miyazu Bay, central Japan, basedon 20 surface sediment samples in April 2019, in order to consider ecological characteristics ofmodern benthic foraminifera. Horizontal distributions of the major species of living (stained)
benthic foraminifera in Miyazu Bay are divided into the several groups that seem to be generally
affected by (1) the water depths and the substrates (grain size) and probably by (2) the influence ofthe pelagic water on the bay. These spatial occurrences of modern benthic foraminifera in Miyazu
Bay seem to support a hypothesis that enhanced winter monsoons may explain changes in theabundance of opportunistic benthic foraminifera in the coastal areas along Wakasa Bay.

Key words: Wakasa Bay; distribution of benthic foraminifera; Aso-kai lagoon; dispersal of benthic
foraminifera

north and west coasts of the Japanese Islands are closely

Introduction

related to the East Asian winter monsoon (e.g., Suzuki

Seasonal contrast of climates in the Japanese Islands is

et al., 2020), in addition to precipitation in the summer
monsoon.

strongly affected by East Asian summer/winter monsoon(e.g., Nakagawa et al., 2009). In particular, the north andA number of small inlets and lagoons are present

west coasts of the Japanese Islands are characterized byalong the shoreline of Wakasa Bay which is locatedabundant snow falls are a result of the intense seasonal

paleoclimatic studies have been carried out based on pasthigh precipitation in winter due to heavy snow fall. The

along the north coast of central Japan (Fig. 1). Several

northwesterly winds driven by the East Asian wintersediment records. For example, studies of sediment coresmonsoon which carry large amounts of moisture fromfrom Lake Suigetsu have provided high quality proxy

the surface ocean under the influence of the Tsushima

data on paleoclimatic conditions during the Holocene

Warm Current. Thus, variations in climate, regional(e.g., Fukusawa, 1997; Suzuki et al., 2016). Takata ethydrology and behavior of marine organism along theal. (2014) investigated fossil benthic foraminifera from1

BK21 School of Earth and Environmental Systems, Pusan National University, 2 Busandaehak-ro 63 beon-gil, Busan 46241, Korea.

2

Faculty of Education, Kyoto University of Education, 1 Fukakusa-Fujimori, Kyoto 612-8522, Japan.

3

Maizuru Fisheries Research Station, Field Science Education and Research Center, Kyoto University, Nagahama, Maizuru 625-0086, Japan.

受付日：2021 年 11 月 12 日，受理日：2022 年 1 月 16 日，WEB 掲載日：2022 年 2 月 10 日

11

Hiroyuki Takata・Satoshi Tanaka・Hideki Sawada・Keita W. Suzuki

a sediment core in Aso-kai lagoon along Wakasa Bay

130˚(Fig. 1). They recognized high abundances of benthic

Figure1_takata
Single-column width

140˚

150˚

(a)

foraminifera in AD ~1070, ~1400 and ~1600, andsuggested that these events resulted from an enhanced
East Asian winter monsoon through passive dispersal of

40˚

benthic foraminifera into the lagoon. In addition, Takata

(b)

Korea

et al. (2019) investigated modern benthic foraminifera

Japan

in Maizuru Bay (~20 km ESE-direction from Aso-

kai lagoon). They found individuals of several species

Pacific Oceanof benthic foraminifera in the water column, based on

30˚plankton tow sampling. According to these observations,they confirmed the passive dispersal potential of some(b)benthic foraminifera in the coasts of Wakasa Bay.

WakasaBay

This insight about the dispersal potential of benthic

foraminifera would aid the interpretation of core-based

1-20

N35.6ºE135.2ºstudy of Aso-kai lagoon, and improve understandingof past variation of the East Asian winter monsoon. In

foraminifera in Miyazu Bay (Fig. 1) which sits between

Aso-kailagoonAso-kai lagoon and the open ocean. Information about

4-20

4-15
4-10

5-15
6-20
6-10

an opportunity for a more precise paleoenvironmental

study of Aso-kai lagoon. In this study, we investigate

6-15

9-15

10-20

8-10
8-15
15 m

the horizontal distribution of living (stained) benthic
foraminifera from Miyazu Bay, based on 20 surface

11-20
20 m

5-20

modern benthic foraminifera in this bay would provide

25 m

13-25
12-25

3-20

contrast, there have been no studies of modern benthic

Miyazu Bay

2-20

7-15
7-10

4 km

sediments, in order to describe some of the ecologicalFig. 1 Locality map of the study area (bathymetric datacharacteristics of modern benthic foraminifera in thissetting.

referred from the Japan Hydrographic Association)

Materials and methods

temperatures at that depth range from ~11 ºC to ~28 ºC.

Miyazu Bay is an elongate inlet along the coast of

Dissolved oxygen content at a depth of 10 m is higher

Wakasa Bay (central Japan) (Fig. 1) with a long axis in athan 5 mg l-1 throughout the year (Fig. 2). The salinity ofnortheast to southwest direction. The deepest part of thethe waters on April 23, 2019 in the bay ranged between

bay is approximately 30 m deep. Small rivers discharge

31.05 and 34.35 (mean 33.90 throughout the water

into the Miyazu Bay, such as the Ohte River (in thecolumn among the 6 stations) (Sawada and Suzuki,

southern bay). In addition, Aso-kai lagoon, a brackish

unpublished data). In particular, there is also no marked

lagoon, is located in the southwest part of the bay (Fig.

lateral salinity gradient in the hypolimnetic water (>34)

1). It is largely separated from Miyazu Bay by sand bars

from the inner (station 6-20) to outer parts (station 1-20)(“Amano-hashidate” and “Sho-tenkyo”), and only connectedof the bay (Sawada and Suzuki, unpublished data). The

by a narrow channel (~4 m deep), which branches into two

substrate in the bay generally consists of sandy sediment,in the southern area of the lagoon (Fig. 1).

whereas substrates at greater depths (>20 m water

Seasonal variations of water temperature and dissolved

depth) sometimes consist of muddy sediments (Table

oxygen content at a depth of 10 m in Miyazu Bay are1). In addition, surface sediments at some stations along

shown in Fig. 2 (Kyoto Prefecture; http://www.pref.

the east coast of the bay consist of various grain size

kyoto.jp/suishitu/jyoujikanshi/wakasawan.html). Water

sediments ranging from gravel to mud. This seems to be12Modern benthic foraminiferal fauna (Rhizaria) in Miyazu Bay, central Japan
30
25

Water
temperature
(ºC)

20
15
10
15

Dissolved
oxygencontent(mg l-1)

Ejiri (10 m)
Shimazaki (10 m)

10
5
0

2015

2016

2017

2018

2019

Fig. 2 Seasonal variation of water temperature and dissolved oxygen content
at 10 m water depth at Stations Ejiri and Shimazaki (near our stations 11-20
and 6-20, respectively) in Miyazu Bay during 2015-2019 (data from Kyoto
Prefecture; http://www.pref.kyoto.jp/suishitu/jyoujikanshi/wakasawan.html).

Table1_takata
Double-column width

Table 1 Locations, water depths and substrates of ourdue to the complicated and steep geomorphology along
the east coast of the bay, compared to the west coast of

study stations in Miyazu Bay.

the bay (Fig. 1).

Station13-25*
13-20
12-25*
12-20
11-20*
11-15
10-20*
10-15
10-10
9-15*
9-10
9-5
8-15*
8-10*
8-5
7-15*
7-10*
7-5
6-20*
6-15*
6-10
6-5
5-20*
5-15*
5-10
5-5
4-center
4-20*
4-15*
4-10*
4-5
3-center
3-20*
3-15
3-10
3-5
2-20*
2-15
2-10
2-5
1-20*
1-15
1-10

We used a Smith-Macintyre type sampler to collectsurface-sediment samples from 43 stations along 13transects of Miyazu Bay (Fig. 1; Table 1) on April 23,

2019, using a R/V Ryokuyo-maru (Kyoto University).
Sediment samples with a thickness of two centimeters

were collected from a sub-core that had a diameter of 5
cm for foraminiferal analysis (39.3 cm3).

The foraminiferal subsamples were washed througha 250-mesh sieve (63-μm openings). The residues were

stained with rose Bengal solution (0.5 g l–1) for 24 hours,
after which they were washed with warm water toremove excess dye (Oda, 1978). The residues were driedat 40ºC and split into 1/2 to 1/32 aliquots. Living (stained)

foraminiferal specimens were collected from the 20

samples and were identified under a stereo-binocular
microscope. Taxonomic assignments followed Matoba(1970), Nomura and Seto (1992), Oki (1989) and Akimoto
et al. (2002). Generic classification was based on

Loeblich and Tappan (1987). The number of specimens
per unit volume (# cm–3) was calculated in each sample.

Latitude35 35.688
35 35.202
35 35.498
35 35.224
35 34.771
35 34.785
35 34.477
35 34.246
35 34.172
35 33.883
35 33.969
35 33.945
35 33.449
35 33.455
35 33.451
35 32.907
35 32.584
35 32.516
35 33.271
35 32.790
35 33.076
35 33.006
35 33.728
35 33.884
35 33.971
35 34.026
35 34.651
35 34.722
35 34.375
35 34.345
35 34.325
35 35.015
35 35.027
35 34.916
35 34.977
35 34.958
35 35.605
35 35.773
35 35.844
35 35.971
35 36.050
35 36.192
35 36.062

Longitude
135 14.362
135 14.205
135 13.969
135 13.935
135 13.322
135 13.607
135 13.321
135 13.546
135 13.522
135 12.746
135 12.878
135 12.890
135 12.136
135 12.199
135 12.219
135 12.106
135 12.401
135 12.423
135 11.738
135 11.543
135 11.384
135 11.346
135 11.794
135 11.461
135 11.487
135 11.472
135 12.454
135 12.435
135 11.842
135 11.768
135 11.710
135 12.808
135 12.713
135 12.477
135 12.466
135 12.408
135 13.590
135 13.672
135 13.347
135 13.235
135 14.070
135 13.955
135 13.673

Depth (m)
25.0
21.9
24.3
24.1
21.2
20.4
19.7
15.8
13.8
15.3
10.2
4.2
14.8
12.8
8.9
14.6
9.2
4.6
16.8
14.9
14.8
10.4
18.6
13.0
13.2
6.0
20.8
20.4
15.4
10.9
4.1
21.6
20.6
15.9
7.8
5.5
20.3
14.5
8.9
5.5
17.1
13.6
10.1

* shows samples that we studied benthic foraminiferal fauna

13

Substrate
mud
mud
mud
mud
mud
mud
mud
sandy mudgravelly-sandy mud
mud
gravelly medium sandgravelly medium sandmudsandy mudfine-medium sand
mud
mud
mud
(no description)(no description)(no description)(no description)mudfine sandmudfine sand(no description)(no description)(no description)(no description)(no description)
mud
mud
mud
fine sandfine sand(no description)(no description)(no description)(no description)
sandy mudsandfine sandHiroyuki Takata・Satoshi Tanaka・Hideki Sawada・Keita W. Suzuki

Fig. 3 Light micrographs of benthic foraminifera in Miyazu Bay.

1a, b, 2a, b. Ammomarginulina sp. A; 3a, b. Nouria sp.; 4a, b. Textularia foliacea; 5a,b. Eggerelloides advenus; 6a, b. Haplophragmoides sphaeriloculum; 7. Textularia
earlandi. Scale bar=100μm.

and Nonionellina sp. A. Agglutinated species, such

Results and discussion

as Ammomarginulina sp. A and Haplophragmoides

Living (stained) specimens of benthic foraminifera

spaeriloculum, are also similar, although they are

genera and 73 species were recognized in this study

the outer portion. Preservation of benthic foraminifera

were present at all 20 stations in Miyazu Bay. Fifty-threesometimes more common in Miyazu Bay, especially in(Appendix 1). Ammomarginulina sp. A, Eggerelloides

was generally good with no sign of strong destruction
or abrasion, suggesting no marked transportation ofadvenus, Haplophragmoides spaeriloculum, Noninella

stella, Nonionellina sp. A and Pseudononion japonicum

foraminiferal tests (Fig. 3). Although we cannot totallywere com mon constituents ( Fig. 3). T his faunal

dismiss possible post-mortem transport of foraminiferal

association is similar to that of Maizuru Bay (Takata

tests, these features imply that this benthic foraminiferal

et al., 2019), for example the common Noninella stellaassemblage was unlikely to be biased by significant
14Modern benthic foraminiferal fauna (Rhizaria) in Miyazu Bay, central Japan

Fig. 3 Continued.

8a-c. Trochammina pacifica; 9a, b. Triloculina rotunda; 10. Brizalina striatula; 11.

Buliminella elegantissima; 12a, b. Uvigerinella glabra; 13a-c. Ammonia sp. A; 14.

Pseudoparrella tamana; 15a-c. Nonionella stella; 16a-c. Nonionellina sp. A.; 17a, b.
Elphidium excavatum forma excavata; 18a-c. Pseudononion japonicum. Scale bar=100 µm.

transportation from another area, such as the outside area

P. japonicum is present in the outer part of the bay.

Horizontal distributions of the major species of modern

the two stations along the east coast of the bay. (d)of Miyazu Bay.

(c) Eggerelloides advenus is specifically common at

benthic foraminifera within Miyazu Bay show different

Ammomarginulina sp. A is common in the outer part ofpatterns (Fig. 4). (a) Nonionella stella is generally

the bay. (e) Finally, Haplophragmoides spaeriloculum is

sp. A and Pseudononion japonicum are common alongof the bay. Thus, these horizontal distributions imply thatabundant in the inner part of the bay. (b) Nonionellina

generally common at the several stations along the coasts

the west coast of the bay. Nonionellina sp. A tends tothese major species occupy specific habitats.

be more common in the inner part of the bay, whereas

Some species of benthic foraminifera show specific15Hiroyuki Takata・Satoshi Tanaka・Hideki Sawada・Keita W. Suzuki

Figure4_takata
Double-column width

Abundance(# cm-3)

5

25

50

4 km
Eggerelloides
advenusAmmomarginulina sp. A

0% 2% 10% 20%

0% 2% 10% 20%

4 km

4 km

Haplophragmoides spaeriloculum

Nonionella stella

0% 2% 10% 20%

0% 5% 25% 50%

4 km

4 km
Nonionellina sp. A

Pseudononion japonicum

0% 2% 10% 20%

0% 2% 10% 20%

4 km

4 km

Fig. 4 Distribution of the abundance of living (stained) benthic
foraminifera per unit volume (# cm–3) and the relative abundances of six

selected species of living (stained) benthic foraminifera in Miyazu Bay.

trends with variations of water depth and/or medianno marked tendency with water depth. This species is(Fig. 5). Ammomarginulina sp. A tends to be common

and Bowser, 1999) and is common in the seasonallygrain size (Sawada and Suzuki, unpublished data)

known as a low oxygen tolerant species (e.g., Bernhard

in deeper (20–25 m water depth) and fine-grained

oxygen-poor hypolimnetic water of Maizuru Bay (Takata

substrates in the stations off the east coasts. In contrast,

et al., 2019).

H. sphaeriloculum tends to be common in the shallower

Neritic benthic foraminifera commonly show specificdepths (<20 m water depth). In addition, N. stella isdistributions in relation to horizontal salinity gradientsbasically abundant in the fine-grained substrates, despite

established by coastal water/pelagic water mixing
16

Figure5_takata
Double-column width

Modern benthic foraminiferal fauna (Rhizaria) in Miyazu Bay, central Japan

Depth (m)

10

10

15

20

Abundance(# cm-3)25

Depth (m)

1.0

1.5

0

20

10
15

30

30

40 20

50

1.0

10

15

15

20

20

Ammomarginulina
sp. A (%)
1.0

1.5

2.0

15

15

20

20

Eggerelloides
advenus (%)
1.5

2.0

1.5

2.5

2.0

2.5

Pseudononion
japonicum (%)

25

2.5

2.0

Nonionellina
sp. A (%)
1.0

10

1.0

1.5

25

2.5

10

25

Nonionella
stella (%)

25

2.5

10

25

Depth (m)

2.0

10

1.0

1.5

2.0

2.5

log (median grain size)

10

Depth (m)

0
10

15

20
20

30

Haplophragmoides
sphaeriloculum (%).

25
1.0

1.5

2.0

Area
off west coasts
off east coasts
inner part2.5

log (median grain size)

Fig. 5 Crossplots between the median grain size and water depth in the Miyazu
Bay. The circle symbols represent the abundances of benthic foraminifera perunit volume and the relative abundance of major six species. The stations in thethree areas are as follows: off the west shore: transects 1 to 5; off the east shore:transects 8 to 13; inner bay: transects 6 and 7.

17

Hiroyuki Takata・Satoshi Tanaka・Hideki Sawada・Keita W. Suzuki(Matoba, 1970; Kosugi et al., 1991; Nomura and Seto,

major benthic foraminifera, including opportunistic

modern benthos and noted a general tendency in thetimes. Further study is highly recommended focusing on

1992). In Miyazu Bay, Habe and Yamazi (1955) studied

species, are quite variable at one location at different

distribution of benthos that is related to the inflow ofseasonal monitoring of benthic foraminifera, in order tothe pelagic water from the outer sea (Wakasa Bay)evaluate the possibility that passive dispersal of benthic

into Miyazu Bay, seen generally along the west coast.

foraminifera is due to intense wind mixing of the water

The common occurrences of Noninellina sp. A andcolumn driven by the East Asian winter monsoon. SuchP. japonicum along the western shoreline of the bay

studies will require additional seasonal surveys, withseem to be consistent with the influence of the anti-

plankton net towing and sediment trap mooring (e.g.,clockwise inflow of pelagic water to the bay. In contrast,

Nomura and Seto, 2004; Nomura et al., 2010).

the common E. advenus has been reported from neritic

eutrophic environments (Tsujimoto et al., 2006).

Acknowledgements

Common E. advenus at the two stations off the east
coast of Miyazu Bay is also consistent with this kind of

We thank Mr. Yoshihito Ogura (Captain of R/Vhydrology in Miyazu Bay (i.e., weaker influence of the

Ryokuyo-maru, Kyoto University) and undergraduate

in Miyazu Bay usually (Sawada and Suzuki, unpublishedassistances in our field sampling in Miyazu Bay. We

pelagic water). Thus, despite no marked salinity gradientstudents of Kyoto University of Education for their kinddata), the inflow of the pelagic water into the bay seems

appreciate Dr. Akira Tsujimoto (Shimane University)

to affect the spatial occurrences of several species.

for his assistance in taking light micrographs. We are

In summary, the distribution of modern benthic

indebted to Dr. Yoshiki Saito (Shimane University) andforaminifera in Miyazu Bay seem to be generally

two anonymous reviewers for constructive comments

to improve the manuscript. We also thank Dr. David L.

affected by (1) the water depths and the substrates (grainsize) (Ammomarginulina sp. A, H. sphaeriloculum andDettman (University of Arizona/Shimane University) for

N. stella) and probably by (2) the influence of the pelagic

his English editing.

water on the bay (Nonionellina sp. A, P. japonicum and E.
advenus).

Faunal reference list

Takata et al. (2014) reported high abundances ofbenthic foraminifera in Aso-kai lagoon (core ASC2) atAmmobaculites exiguus Cushman and Brönnimann,

the following times: ca. AD 1070, 1400 and 1600. The

1948, Contribution from the Cushman Laboratory forincreased abundance of benthic foraminifera at theseForaminiferal Research, 24, p. 38, pl. 7, figs. 7, 8.

dates were largely due to the abundance of E. advenus.

Ammonia “beccarii” (Linné) forma 1 Matoba, 1970,

Alve and Goldstein (2010) suggested that Eggerelloides

Science Reports of Tohoku University, 2nd ser.

scaber (which has a similar test morphology to E.

(Geology), 42 (1), pl. 5, figs. 8, 9.

advenus) might spread through propagule dispersal.

Ammonia japonica (Hada) = Rotalia japonica Hada,

events of E. advenus in Aso-kai lagoon, as proposed

University, ser. 4, Biology, 6 (3), p. 137, text-figs. 93.

Enhanced wind mixing may have driven past dispersal

1931, Science Repor ts of the Tohoku Imperial

by Takata et al. (2014). In addition, Takata et al. (2014)

Bolivina humilis Cushman and McCulloch = Bolivina

into Aso-kai lagoon from the outer sea (Wakasa Bay),

McCulloch, 1942, Allan Hancock Pacific Expeditions,hypothesized that E. advenus may have dispersed

seminuda Cushman var. humilis Cushman and6, p. 211, pl. 26, figs. 1–6.

whereas this study indicates that the species is more

likely to have dispersed from the inner-middle portion

Bolivina robusta (Brady) = Bulimina (Bolivina) robusta

support the hypothesis that enhanced winter monsoons

Results of the Voyage of the H. M. S. Challengerof Miyazu Bay to the lagoon (Fig. 4). These observations

Brady, 1881; Brady, 1884, Report on the Scientific

may explain changes in the abundance of opportunistic

during the years 1873–1876, Zoology, 9, p. 421, pl. 53,

benthic foraminifera in the Aso-kai lagoon. In contrast,

figs. 7–9.

Takata et al. (2019) demonstrated that occurrences of

Bolivina tokiokai Uchio, 1962, Publications of the Seto
18

Modern benthic foraminiferal fauna (Rhizaria) in Miyazu Bay, central Japan

Marine Biological Laboratory, 10, p. 245, pl. XVIII,

pl. 1, 17–20.

figs. 5a, b.

Elphidium reticulosum Cushman, 1933, Bulletin United

Cushman, 1922, Department of Marine Biology, 311, p.

Elphidium somaense Takayanagi, 1955, Contributions

Brizalina striatula (Cushman) = Bolivina striatula

States National Museum, p.51, pl.12, figs. 5a, b.

27, pl. 3, fig. 10.

from the Institute of Geology and Paleontology,

Buccella frigida (Cushman) = Pulvinulina frigida

Tohoku University, 45, p. 43, 52, text-figs. 28a, b.

Cushman, 1922a, Contributions to Canadian Biology, p. 12.

F u r s e n k o i n a c o m p a c t i f o r m i s ( M c C u l l o c h) =Mem. Volume, p. 234, text-figs. 2a-c, pl. 1, figs.

University of Southern California, p. 248, pl. 104, fig. 25.

Buccella makiyamai Chiji, 1961, Prof. Jiro Makiyama

Neobuliminoides compactiformis McCulloch, 1977,

13a–14b.

Geminospira simaensis Makiyama and Nakagawa, 1941,

Sciences Naturelles, 7, p. 269, pl.12, figs. 10-12.

Goessella iizukae Takayanagi, 1955, Contributions from

Bulimina marginata d’Orbigny, 1826, Annales desJournal of the Geological Society of Japan, 48, p. 241, figs. 3–5.

Bulimina nojimaensis Asano, 1950, Illustrated Catalogue

the Institute of Geology and Paleontology, Tohoku

of Japanese Tertiary Smaller Foraminifera, pt. 2, p. 4,

University, 45, p. 41, text-fig. 25.

text-figs. 15, 16.

Hanzawaia nipponica Asano 1944, Journal of the

Buliminella elegantissima (d'Orbigny) = Bulimina

Geological Society of Japan, 41, p. 99, pl. 4, figs. 1a, b,

el ega n t i s s i m a d' O r big ny, 1839, Voyage d a n s

2a, b.

l’Amérique méridionale–Foraminifères, t. 5, pt. 5, p.

Haplophragmoides canariensis (d’Orbigny) = Nonionina

Cancris auriculus (Fichtel and Moll) = Nautilus

Canaries, in Barker-Webb and Berthelot, Hist. Nat. des51, pl. 7, figs. 13, 14.

canariensis d’Orbigny, 1839, Foraminiferes des Iles

auriculus var. α Fichtel and Moll, 1798, Testacea

Iles Canaries 2, p. 128, pl. 2, figs. 33, 34.

m i c r o s c o p i c a , a l i a q u e m i n u t a ex g e n e r i b u s

Haplophragmoides sphaeriloculum Cushman, 1917,

Argonauta et Nautilus, ad naturam picta et descripta

Proceedings of the United States National Museum,(Microscopische und andere klein Schalthiere aus den

51, p. 652.

geschlechtern Argonaute und Schiffer, p. 108, pl. 20,

Haynesina sp. A = Haynesina sp., Kosugi et al., 1991,

Cibicides lobatulus (Walker and Jacob) = Nautilus

Islandiella algida (Cushman) = Cassidulina algida

Adam’s Essays on the Microscope. In, Kanmmacher, E.

Laboratory for Foraminiferal Research, (12), p. 35, pl.

figs. a-c.

Fossil (51), p.53, figs. 5, 10-11.

lobatulus Walker and Jacob in Kanmacher, 1798,

Cushman, 1944, Special Publications Cushmaned., the Second Edition, with Considerable Additions

4, figs. 24a–c.

and Improvements. Chapter 11, p. 642, pl. 14, fig. 36.

Murrayinella minuta (Takayanagi) = Rotalia? minuta

Cushman, 1922, Contributions to Canadian Biology, 9, p. 9.

Geology and Paleontology, Tohoku University, 45, p.

Eggerelloides advenus (Cushman) = Verneuilina advena

Takayanagi, 1955, Contributions from the Institute ofElphidium advenum (Cushman) = Polystomella advena

52, text-figs. 29a–c.

Cushman, 1922, Department of Marine Biology, 311, p.

Nonion manpukuziensis Otuka, 1932, Journal of theElphidium excavatum (Terquem) forma clavata =Nonionella stella Cushman and Moyer = Nonionella

Tappan, 1953, Smithsonian Miscellaneous Collections,

1930, Contribution from the Cushman Laboratory for56, pl. 9, figs. 11, 12.

Geological Society Tokyo, 39, p. 654, fig. 1.

Elphidium clavatum Cushman, emend, Loeblich andmiocenica Cushman var. stella Cushman and Moyer,

121 (7), p. 98, 101, 102, pl. 19, figs. 8a, b, 9, 10.

Foraminiferal Research, 6, p. 56, pl. 7, fig. 17.

Elphidium excavatum (Terquem) forma excavata =Oolina hexagona (Williamson) = Entosolenia squamosaclassement des animaux qui vivent sur la plage et dansand Magazine of Natural History, ser. 2, p. 20, pl. 2,

Polystomella excavata Terquem, 1875, Essai sur le(Montagu) var. hexagona Williamson, 1848, Annals

les environs de Dunqerque, Fasc. 2, p. 20, pl. 2, figs. 2a, b.

fig. 23.

Elph i d iu m k a n eh a ra i Ish iwad a, 1958, Rep or t s,

Pseudononion japonicum Asano, 1936, Journal of theGeological Survey of Japan, 180, p. 18, text-figs. 2a, b;

Geological Society of Japan, 43 (512), p. 347, text-figs. a-c.

19

Hiroyuki Takata・Satoshi Tanaka・Hideki Sawada・Keita W. Suzuki

Pseudoparrella naraensis Kuwano, 1950, Journal of thefrom the Cushman Laboratory for Foraminiferal

Pseudoparrella tamana Kuwano, 1950, Journal of theUvigerina proboscidea Schwager, 1866, Reise derGeological Society of Japan, 56 (657), p. 317, text-fig. 6.

Research, 1, p. 39, pl. 6, fig. 3.

Geological Society of Japan, 56, p. 317, text-figs. 5a-c.

Österreichischen Fregatte Novara um die Erde in den

Pseudorotalia gaimardii compressiuscula (Brady) =Jahren, p. 250, pl. 7, fig. 96.

Rotalia papillosa Brady compressiuscula Brady, 1884,

Uvigerinella glabra (Millett) = Uvigerina auberiana

M. S. Challenger during the years 1873–1876, Zoology,

Royal Microscopical Society, p. 268, pl. 5, figs. 8a, b.

Report on the Scientific Results of the Voyage of the H.

d’Orbigny var. glabra Millett, 1903, Journal of the9, p. 708, pl. 107, fig. 1; p. 108, fig. 1.

Virgulinella fragilis Grindell and Collin, 1976, Rev.

Q u i n q u e l o c u l i n a a k n e r i a n a d’O r b ig ny, 18 4 6,Espanola Micropaleont., 8 (2), p. 274, 277, 278, pl. 1,

Foraminifères fossiles du Bassin Tertiaire de Vienne

figs. 1-8, text-fig. 2.

(Autriche), p. 290, pl. 18, figs. 16-21.

Quinqueloculina elongata Natland, 1938, Bulletin of theReferences

Scripps Institution of Oceanography of the Universityof California, Technical Series, Tech. Ser. 4, (5), p. 141,

Akimoto, K., Matsui, C., Shimokawa, A. and Furukawa,

Quinqueloculina lamarckiana d’Orbigny, 1839, in de laof Shimabara Bay, Kyushu, Southwest Japan. The

pl. 4, fig. 5.

K., (2002) Atlas of Holocene benthic foraminifers

Sagra R., Histoire physique, politique et naturelle deKagoshima University Museum Monographs, 2:1–112.

l'ile de Cuba, p. 189, pl. 11, figs. 14–15.

Alve, E. and Goldstein, S.T. (2010) Dispersal, survival

Quin queloculin a sem inulu m ( Li n né) = Serpulaa nd delaye d g row t h of b ent h ic fora m i n i fera lseminulum Linné, 1758, Systema Naturae, p. 786.

propagules. Journal of Sea Research, 63:36–51.

Rectobolivina raphana (Parker and Jones) = Uvigerina

Ber nha rd, J.M. and Bowser, S. S. (1999) Benthic

Philosophical Transactions of the Royal Society, 155, p.

sequestration and functional morphology. Earth-(S agr i n a) ra p h a n u s Pa rke r a nd Jones, 1865,

foraminifera of dysoxic sediments: chloroplast

364, pl. 18, figs. 16a-b, 17.

Science Reviews 46:149–165.

Reussella aculeata Cushman, 1954, Contribution from

Fukusawa, H. (1997) High-resolution reconstruction of

the Cushman Laboratory for Foraminiferal Research,paleoenvironmental informations detected by using

21, p. 41, pl. 7, figs 10–11.

lake sediment. Journal of Environmental Information

Sagrinella convallaria (Millett) = Bolivina convallaria

Science, 26:42–53. (in Japanese)

Millett, 1900, Journal of the Royal MicroscopicalHabe, T. and Yamazi, I. (1955) Bio-oceanographical

Stainforthia f usiformis (Williamson) = Bulimina

Biogeographical Society of Japan. vols. 16–19:399–

Society, p. 544, pl. 4, fig. 7a, b.

characteristics of the Miyazu-wan. Bulletin of the

pupoides d’Orbigny var. fusiformis Williamson, 1858,

406. (in Japanese)On the Recent foraminifera of Great Britain, p. 63, pl.

Kosugi, M., Kataoka, H. and Hasegawa, S. (1991)

Textularia earlandi Parker, 1952, Bulletin of the Museumof environments in an inner bay and its application to5, figs. 129, 130.

Classification of foraminifer communities as indicatorsof Comparative Zoology, 106 (10), p. 458.

reconstruction paleoenvironment. Fossils, 50:37–55. (inTextularia foliacea Heron-Allen and Earland, 1915,

Japanese with English abstract)

Transactions of the Zoological Society of London, 20,

Loeblich, A.R. Jr. and Tappan, H. (1987) Foraminiferal

p. 628, pl. 47, figs. 17–20.

genera and their classification. Van Nostrand Reinhold,

Triloculina rotunda d’Orbigny, 1826, Annales des

New York, 970p. and its plates (1–847).

Sciences Naturelles, p. 298.

Matoba, Y. (1970) Distribution of recent shallow water

Trochammina cf. japonica Ishiwada = Cf. Trochammina

foraminifera of Matsushima Bay, Miyagi Prefecture,japonica Ishiwada, 1950, Bulletin of the Geological

Nor t hea st Japa n. Scienc e Rep or t s of Tohok uSurvey of Japan, 1 (4), p. 190, pl., figs. 2a-c.

University, ser. 2 (Geology) 42:4–85.

Trochammina pacifica Cushman, 1925, Contribution

Nakagawa, T., Okuda, M., Yonenobu, H., Miyoshi, N.,
20

Modern benthic foraminiferal fauna (Rhizaria) in Miyazu Bay, central Japan

Fujiki, T., Gotanda, K., Tarasov, P.E., Morita, Y.,

and Khim, B.-K. (2019) Dispersal potential of

monsoon climate by two different orbital rhythms andof the Japanese Islands and its application to

Takemura, K. and Horie, S. (2009) Regulation of theneritic benthic foraminifera in the coastal areas

forcing mechanisms. Geology, 36:491–494.

paleoenvironmental studies. Estuarine, Costal and

Nomura, R. and Seto, K. (1992) Benthic foraminifera

Shelf Science, 227:Article 106288.

from brack ish La ke Na kaum i, San-in distr ict,Tsujimoto, A., Nomura, R., Yasuhara, M. and Yoshikawa,Japanese Micropaleontology, (eds.) Ishizaki, K., Saito,

Bay, southwestern Japan: faunal changes over the lastsouthwester n Honshu, Japan. In: Centenar y ofS. (2006) Benthic foraminiferal assemblages in OsakaT., pp. 227–240, Terra Scientific Publishing Company,

50 years. Paleontological Research, 10:141–161.

Tokyo.

Nomura, R. and Seto, K. (2004) Water conditions during

the time of red tide occurrence in spring, 2003, andthe associated benthic foraminifera in Lake Nakaumi.
LAGUNA, 11:125–130. (in Japanese with English
abstract)

Nomura, R., Seto, K. and Tsujimoto, A. (2010) Hightide dispersion of marine benthic foraminifera into
brackish waters: implications for dispersion processes
during sea-level rise. LAGUNA, 17:15–21.

Oda, M. (1978) I Foraminifera and Ostracoda, In:
Manual of Microfossil studies, (ed.) Takayanagi, Y.,
pp. 33–46, Asakura Books, Tokyo. (in Japanese)

Ok i, K. (1989) Ecological analysis of benthonicforaminifera in Kagoshima Bay, south Kyushu, Kapan.
South Pacific Study, 10:1–191.

Suzuki K.W., Fuji, T., Kasai, A., Itoh, S., Kimura, S. andYamashita, Y. (2020) Winter monsoon promotes thetransport of Japanese temperate bass Lateolabrax

japonicus eggs and larvae toward the innermost part of
Tango Bay, the Sea of Japan. Fisheries Oceanography,
29:66–83.

Suzuki, Y., Tada, R., Yamada, K., Irino, T., Nagashima,K., Na kagawa, T. a nd Omor i, T. (2016) Massaccumulation rate of detrital materials in Lake
Suigetsu as a potential proxy for heavy precipitation:
a comparison of the observational precipitation andsedimentary record. Progress in Earth and Planetary
Science, 3：5 https://doi.org/10.1186/s40645-0160081-x.

Takata, H., Tanaka, S., Seto, K., Sakai, S., Takayasu, K.

and Khim, B.-K. (2014) Biotic response of benthic
foraminifera in Aso-kai lagoon, central Japan, tochanges in terrestrial climate and ocean conditions(~AD 700–1600). Journal of Paleolimnology, 51:421–
435.

Ta kata, H., Nomura, R., Sa mpei, Y., Ta na ka, S.
21

Table2_takata
Double-column width
Hiroyuki Takata・Satoshi Tanaka・Hideki Sawada・Keita W. Suzuki

Appendix 1 Faunal list of living (stained) benthic foraminifera in Miyazu Bay.
StationAmmobaculites exiguus
Ammobaculites sp. A
Ammomarginulina sp. A
Eggerelloides advenus
Gaudryina sp.
Goessella iizukae
Haplophragmoides canariensis
Haplophragmoides sphaeriloculum
Haplophragmoides sp. indet.
Lagenammina sp.
Leptohalysis sp.
Nouria ? sp.
Reophax sp.
Textularia earlandi
Textularia foliacea
Textularia sp. indet.
Trochammina cf. japonica
Trochammina pacifica
Agglutinated Foram. gen. et sp. indet.
Massilina ? sp.
Miliolinella sp.
Quinqueloculina akneriana
Quinqueloculina elongata
Quinqueloculina lamarckiana
Quinqueloculina seminulum
Sigmoillina sp.
Siphonaperta sp.
Spiroloculina sp.
Triloculina rotunda
Calc. Porc. Foram. gen. et sp. indet.
Ammonia "beccarii" forma 1
Ammonia japonica
Ammonia sp. A
Amphicoryna sp.
Bolivina cf. decussata
Bolivina humilis
Bolivina robusta
Bolivina tokiokai
Bolivina sp. indet.
Brizalina striatula
Buccella frigida
Buccella makiyamai
Bulimina marginata
Bulimina nojimaensis
Buliminella elegantissima

1-20

2-20

6
7
1

1

3-20

4-10

4-20

5-15

5-20

6-10

6-15

4
6
3
1

4-15
1

2
1

2

8
3

7
3

10
8

3
3

8

2

2

8

1

17

5
7

4

3

2

10

4

5
1
5

3

2

5
11
5

7

1
1

2
1

2
6

4

1

1

6

6

2

21

3

17
2
2
1
2

14

5

1

2

1

3
1

3
2
1
1

1
1

3

2

3

1

3
3

1

2
1

3
1

3

6
3

1

1

1
1
1

1

2

2

1
3
6
4
2

2

1

1
1

1

9

1

1
1

2

2

3

1
1

1

1

2

3
1

3

5

2
2

2

1

1

1
1

1

22

5

3

2

2
2

1

1

1

1

3

4
1

3
1
1
1

1

Modern benthic foraminiferal fauna (Rhizaria) in Miyazu Bay, central Japan

Appendix 1 Continued.
StationCancris auriculus
Cibicides lobatulus
Elphidium advenum
Elphidium excavatum forma clavata
Elphidium excavatum forma excavata
Elphidium kaneharai
Elphidium reticulosum
Elphidium somaense
Fissurina sp.
Fursenkoina compactiformis
Fursenkoina sp. A
Geminospira simaensis
Globocassidulina sp.
Guttulina sp.
Hanzawaia nipponica
Haynesina sp. A
Islandiella algida
Murraynella minuta
Nonion manpukuziensis
Nonionella stella
Nonionellina sp. A
Oolina hexagona
Pseudononion japonicum
Pseudoparrella naraensis
Pseudoparrella tamana
Pseudorotalia gaimardii compressiuscula
Rectobolivina raphana
Reussella aculeata
Rosalina sp.
Sagrinella convallaria
Stainforthia fusiformis
Uvigerina proboscidea
Uvigerinella glabra
Virgulinella fragilis
Calc. Hyaline Foram. gen. et sp. indet.
Total
Split

1-20

2-20

3-20

1

2

1

1

1

1

4-10

3
1

1
2

4-20

5-15

1

1

5-20

6-10

6-15

3

1

1

1

1
3

3

1

1

2

4-15

1

1
1

2

1

1
1
2
1

1

15
6
12
1
1
1
2

18
6
1
10

1

10
4

10
22

3
11

2
12

12

6

1

6

6

4

3
2

2
2

2
1
1

1

1
1
114
16

1
73
16

48
5.3

23

3
77
8

1

2

72
16

1
66
5.3

1
95
10.7

1
94
33

1
3

2
3

1

1

29
22
1
1
1

3

3
3
1
193
4

2
3
57
32

128
8

Table2_takata
Double-column width
Hiroyuki Takata・Satoshi Tanaka・Hideki Sawada・Keita W. Suzuki

Appendix 1 Continued.
StationAmmobaculites exiguus
Ammobaculites sp. A
Ammomarginulina sp. A
Eggerelloides advenus
Gaudryina sp.
Goessella iizukae
Haplophragmoides canariensis
Haplophragmoides sphaeriloculum
Haplophragmoides sp. indet.
Lagenammina sp.
Leptohalysis sp.
Nouria ? sp.
Reophax sp.
Textularia earlandi
Textularia foliacea
Textularia sp. indet.
Trochammina cf. japonica
Trochammina pacifica
Agglutinated Foram. gen. et sp. indet.
Massilina ? sp.
Miliolinella sp.
Quinqueloculina akneriana
Quinqueloculina elongata
Quinqueloculina lamarckiana
Quinqueloculina seminulum
Sigmoillina sp.
Siphonaperta sp.
Spiroloculina sp.
Triloculina rotunda
Calc. Porc. Foram. gen. et sp. indet.
Ammonia "beccarii" forma 1
Ammonia japonica
Ammonia sp. A
Amphicoryna sp.
Bolivina cf. decussata
Bolivina humilis
Bolivina robusta
Bolivina tokiokai
Bolivina sp. indet.
Brizalina striatula
Buccella frigida
Buccella makiyamai
Bulimina marginata
Bulimina nojimaensis
Buliminella elegantissima

6-20

7-10

5
2

3

13

3
17

2
5
2
1

7-15
1

8-10

8-15

13
29

5
11

3
8

8
8

1

1
2
12

1
1
1
2
8
3

4

11
1

2
1

4
4

5

6
1
2
3

9-15 10-20 11-20 12-25 13-25

5
7
4
2
1

1

19
4

8

3

1

2

11

3

9
1

2

4

20
5
1

1
2

14

3

3

3

1
2

1

1

1

21

1
2

2
2
3

2

4

27
29

2

6

1
2

8

2
1

2
8

2

1

2
1

3

1

1

6

1

3
2

2

24

1
8

3

4

1
2

2

2
1

2
1

4

1

2

1

4

2

3

1

3

2

Modern benthic foraminiferal fauna (Rhizaria) in Miyazu Bay, central Japan

Appendix 1 Continued.
StationCancris auriculus
Cibicides lobatulus
Elphidium advenum
Elphidium excavatum forma clavata
Elphidium excavatum forma excavata
Elphidium kaneharai
Elphidium reticulosum
Elphidium somaense
Fissurina sp.
Fursenkoina compactiformis
Fursenkoina sp. A
Geminospira simaensis
Globocassidulina sp.
Guttulina sp.
Hanzawaia nipponica
Haynesina sp. A
Islandiella algida
Murraynella minuta
Nonion manpukuziensis
Nonionella stella
Nonionellina sp. A
Oolina hexagona
Pseudononion japonicum
Pseudoparrella naraensis
Pseudoparrella tamana
Pseudorotalia gaimardii compressiuscula
Rectobolivina raphana
Reussella aculeata
Rosalina sp.
Sagrinella convallaria
Stainforthia fusiformis
Uvigerina proboscidea
Uvigerinella glabra
Virgulinella fragilis
Calc. Hyaline Foram. gen. et sp. indet.
Total
Split

6-20

7-10

7-15

1
4

8-10

1

2

2

2

9

2

1

1

8-15

9-15 10-20 11-20 12-25 13-25
3

4
3

1

1

1

9

28

1

6

1

5
1

11

1
1
1
1
1
47
25

2
2

12

1

1
1
82
13

92
8

88
17

5
4

35
23

7
4

9
4

1
3
4
1

8
1
1

2

4

5

4
2
1

1

6

1

1

5
4

1

1

1

6

1

1

1

3

2

3
2

128
8

52
5.3

104
2

25

4

6

7

3

1
3

1

1
183
8

162
4

219
8

1
119
4

139
4

61
8

1
1

1

87
8