Timing of megafaunal extinction in the late Late Pleistocene on the Japanese Archipelago

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Quaternary International 255 (2012) 114e124

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Quaternary International

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Timing of megafaunal extinction in the late Late Pleistocene on the JapaneseArchipelago

Akira Iwase a,*, Jun Hashizume b, Masami Izuho a, Keiichi Takahashi c, Hiroyuki Sato d

aArchaeology Laboratory, Faculty of Social Sciences and Humanities, Tokyo Metropolitan University, 1-1 Minami Osawa, Hachioji City, Tokyo 192-0397, JapanbMeiji University Center for Obsidian and Lithic Studies, Nagawa, Nagano 386-0601, Japanc Lake Biwa Museum, Kusatsu, Shiga 525-0001, JapandDepartment of Archaeology, University of Tokyo, Bunkyo-ku, Tokyo 113-0033, Japan

a r t i c l e i n f o

Article history:Available online 2 April 2011

* Corresponding author.E-mail addresses: [email protected] (A. Iwa

(J. Hashizume), [email protected] (M. Izuho), [email protected] (H. Sato).

1040-6182/$ e see front matter � 2011 Elsevier Ltd adoi:10.1016/j.quaint.2011.03.029

a b s t r a c t

In the late Late Pleistocene (lLP), Japanese terrestrial large mammals consisted of two main groups; thePalaeoloxodon-Sinomegaceroides complex and the mammoth fauna. The former inhabited temperateforests and the latter were adapted to patches of taiga and grassland in cold environments. Among thetwo groups, almost all large mammals became extinct in the Late Quaternary. The lLP extinction is one ofthe most interesting topics currently debated in Japan.

This paper evaluates previously reported radiocarbon dates of mammal fossils to determine the timingof lLP megafaunal extinctions on the Japanese Archipelago. Unreliable specimens which were dated byconventional 14C decay counting, samples obtained from poorly preserved fossils, samples inconsistentwith geological context, and samples dated by combining bone fragments of several species and whoseexact provenances are unknown are rejected. The timing of extinctions was compared with the vege-tational changes. As a result, the present paper indicates that the extinction of large mammals in thePalaeoloxodon-Sinomegaceroides complex roughly coincided with the onset of the last glacial maximum(LGM: from ca. 25,000 BP to 16,000 BP) and subsequent domination by subarctic conifers. In contrast, themammoth fauna survived the LGM and became extinct or migrated northward when the climate startedto ameliorate. The lLP extinction on the Japanese Islands occurred in two pulses. These results imply thatthe main causes of lLP extinction on the Japanese Archipelago were changes of the ecosystem driven byclimatic changes rather than “overkill” by human hunters.

� 2011 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction

Geographically, Japan is a long chain of islands situated between45�N and 24�N along the northwestern Pacific Rim comprising fourmain islands; Hokkaido, Honshu, Shikoku, Kyusyu, and smallerislands such as Ryukyu. They extend from north to south alonga northeastesouthwest axis. With the Japan Sea to the west, thearchipelago stretches parallel to the Asian mainland (Fig. 1). Due toglobal glacioeustatic sea level drop during the late Late Pleistocene(lLP: MIS3 and MIS2, roughly 50,000 to 10,000 BP), the four mainislands were combined into two land masses; the Palaeo-Sakhalin/

se), [email protected]@lbm.go.jp (K. Takahashi),

nd INQUA. All rights reserved.

Hokkaido/Kurile Peninsula (Palaeo-SHK Peninsula), which wasformed by connecting Hokkaido, Sakhalin, a part of the KurileIslands, and the Russian Far East (Ono, 1990, 1991), and Palaeo-Honshu Island formed by combining Honshu, Shikoku, and Kyu-syu (Ota and Yonekura, 1987) (Fig. 2).

The lLP terrestrial mammals are divided into two main faunalgroups (Takahashi, 2007); one group with Naumann’s elephant(Palaeoloxodon naumanni) and giant deer (Sinomegaceros yabei), andanother group with woolly mammoth (Mammuthus primigenius)and bison (Bison priscus). The former group mainly consisted ofanimals that inhabited temperate forests and had a high percentageof endemic species (Hasegawa, 1972; Kawamura, 1994). Hasegawa(1972) called this group the Palaeoloxodon-Sinomegaceroidescomplex, which was composed of Naumann’s elephant, giant deer,extinct cervid (Cervus praenipponicus), Sika deer (Cervus nippon),brown bear (Ursus arctos), marten (Martes melampus), raccoon dog(Nyctereutes procyonoides), wolf (Canis lupus), fox (Vulpes vulpes),

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Fig. 1. Map of the Japanese Archipelago and locations of sites from which the 14C-dated fossils were recovered. The numbers correspond with those in Table 1.

A. Iwase et al. / Quaternary International 255 (2012) 114e124 115

and Japanese monkey (Macaca Fuscata) (Hasegawa, 1972). Thisgroup inhabited mainly Palaeo-Honshu Island and probablymigrated to the southern part of Palaeo-SHK Peninsula (Hokkaido)when the temperate forest expanded (Takahashi et al., 2004, 2006).

The second group is called the mammoth fauna (Kawamura,1994, 2007; Takahashi, 2007) adapted to patches of taiga andgrassland in cold environment. In the Asian mainland, this groupconsisted of woolly mammoth, steppe bison, red deer (Cervus ela-phus), giant deer (Megaroceros giganteus), musk ox (Ovibosmoschatus), woolly rhinoceros (Coelodonta antiquitatis), wolverine(Gulo gulo), arctic fox (Alopex lagopus), reindeer (Raingifer tarandus),horse (Equus caballus), brown bear and moose (Alces alces) (Lister

and Bahn, 2007). Only a few species of the mammoth fauna (e.g.,mammoth, bison and moose) have been found in Japan (e.g.Kawamura, 1994). This group inhabited mainly Palaeo-SHK Penin-sula, and probably expanded its habitat onto Palaeo-Honshu Islandduring colder periods (Kawamura and Taruno, 2000; Kawamura,1991, 1994, 1998, 2007).

On the Japanese Archipelago, almost all large mammals of thetwo groups, except for a few species such as U. arctos, becameextinct in the Late Quaternary (e.g. Kawamura, 1991, 1994, 2007).The lLP extinction of terrestrial large mammals is one of the mostinteresting topics currently debated in Japan (e.g. Kawamura, 1991,1994, 2007; Norton et al., 2010; Takahashi, 2007; Kitagawa et al.,

Fig. 2. Topography and vegetation zones of the Japanese Archipelago and surroundingregions during the LGM.

A. Iwase et al. / Quaternary International 255 (2012) 114e124116

2009; Iwase et al., 2010). This paper reconstructs a reliable AMS 14Cchronology of lLP terrestrial mammals to determine the timing oflLP extinctions on the Japanese Archipelago. The causes of extinc-tion will also be discussed. Because of inconsistencies betweenvarious calibration programs (e.g. Reimer et al., 2009; Weningerand Jöris, 2008) and the difficulty calibrating samples older than25,000 14C BP, dates are given in uncalibrated radiocarbon BPexcept for some cases.

2. Methods, materials and criteria for the evaluation ofexisting dates

2.1. Methods

Recent studiesutilize adatabaseofwell-dated fossils todepict theaccurate chronology of Quaternary extinctions across theworld (e.g.,Stuart et al., 2002, 2004; MacPhee et al., 2002; Kuzmin and Orlova,2004; Guthrie, 2006). In light of these studies, the reported radio-carbon dates of mammal fossils were used to reconstruct the chro-nology of extinction on the Japanese Archipelago. In Japan, althoughmany previously published dates for lLP mammal fossils exist, theyinevitably include problematic specimens. Here, the existing radio-carbon dates are evaluated, and unreliable specimens rejected.

2.2. Materials

This study is based on 89 published radiocarbon dates ofterrestrial mammal fossils recovered across the Japanese Archi-pelago (Fig. 1). These specimens include 58 dates for P. naumanni,8 for S. yabei, 12 for M. primigenius, 1 for Bison sp., and 11 for otherspecies (Table 1).

2.3. Criteria for the evaluation

From these collected radiocarbon dates, obviously problematicspecimens were rejected according to the following five criteria.

2.3.1. Dating methodFirst, 14C dates made on bone can potentially have several

problems (e.g., Taylor, 1997; Stafford et al., 1987). A fossil bone canlose most of its original organic material and frequently containscontaminants. Thus, for bone dating it is advisable to use theaccelerator mass spectrometry (AMS) method, which utilizesmilligram-sized amounts of purified compounds rather thanconventional 14C decay counting (e.g. Stafford et al., 1987; Sawadaet al., 1992). In this paper, specimens dated by the conventionalmethod are rejected.

2.3.2. Bone preservationSecond, to obtain a more accurate date for a fossil bone, XAD

and/or ultrafiltration are considered more reliable (Stafford andDuhamel, 1982; Stafford, 1987, 1988; Brown et al., 1988; BronkRamsey et al., 2004), but gelatin-extraction techniques weregenerally applied in Japan (Longin, 1971; Arita et al., 1990;Nakamura et al., 1996). The collagen extracted from bone is sepa-rated into two types: one soluble in acid (solution collagen), andanother insoluble (gelatin collagen) (Arita et al., 1990; Nakamuraet al., 1996). Although gelatin collagen is more reliable for datingthan is solution collagen, a fossil which contains less than 0.7%gelatin collagen in weight and has a C/N ratio higher than 4.0 in thecollagen is considered poorly preserved and will date younger thanwell-preserved bones of comparable age (Deniro, 1985; Sawadaet al., 1992; Nakai et al., 1992). The well-preserved fossils (withgelatin collagen content more than 0.7%) from Lake Nojiri, forexample, dated the same as associated wood samples, but poorlypreserved specimens (less than 0.7%) were several thousands yearsyounger than the associated wood (Arita et al., 1990; Sawada et al.,1992; Nakai et al., 1992). Thus, samples which are dated by AMS,contain sufficient gelatin collagen (more than 0.7%), and have C/Nratios lower than 4.0 are accepted.

2.3.3. Consistency with the geological contextThird, 14C dates that are inconsistent with their geological

context were rejected. For example, the 14C date of 34,800� 800 BPobtained from the tusk of P. naumanni (Sakura specimen) recoveredfrom the Chiba Prefecture has a discrepancy with the age of itsassociated stratigraphic layer (Narita formations: MIS5) (Akiyamaand Nakai, 1988) (Table 1).

2.3.4. Combined bone specimenFourth, single dates obtained from combined bone fragments of

several animals are also excluded.

2.3.5. ProvenanceLast, fossils whose exact provenances are unknown are also

rejected. For instance, Takahashi et al. (2006) pointed out that it isproper to exclude the youngest mammoth, the Yubari specimen(Table 1), from consideration because its exact provenance isunknown.

3. Results and AMS 14C chronologies

3.1. P. naumanni

Fifty eight specimens of P. naumanni are directly 14C dated. Asingle date was obtained from a specimen recovered in Hokkaido,and 57 dates were from Honshu, Shikoku, and Kyusyu. The range of

Table 1Previously Reported 14C Dates for Terrestrial Mammal Fossils in Japan. The Numbers Correspond Fig. 1.

Site Region No. Lab Number 14C Age (BP) Fossil Element Collagen(%)

C/N ratio d13C(&)

Datingmethod

Remarks Evaluation Reference

Yubari ? Unknown 7 Beta-187606 16,250 � 90 Mammuthusprimigenius

Molar �20.8 AMS Provenanceunknown

Reject Takahashiet al., 2005

Ogoshi Hokkaido 5 Beta-188519 19,530 � 80 Mammuthusprimigenius

Molar �21.7 AMS Accept Takahashiet al., 2005

Off Notsukezaki Hokkaido 2 20,243 � 670 Mammuthusprimigenius

Molar AMS Sample A Accept Akiyama et al., 1989;Takahashi et al., 2005

Off Notsukezaki Hokkaido 2 Beta-184269 20,700 � 120 Mammuthusprimigenius

Molar �20.6 AMS Re-datingof sample A

Accept Takahashi et al., 2005

Off Onsentsu Honshu (Shimane) 24 NUTA-633 23,680 � 880 Mammuthusprimigenius

Molar 2.7 AMS Accept Akiyama et al., 1989;Takahashi 2007

Off Rausu Hokkaido 1 23,816 � 884 Mammuthusprimigenius

Molar AMS Sample B Accept Nakai et al., 1991;Takahashi et al., 2005

Off Rausu Hokkaido 1 Beta-185830 25,010 � 120 Mammuthusprimigenius

Molar �24.1 AMS Re-datingof sample B

Accept Nakai et al., 1991Takahashi et al., 2005

Yuni Hokkaido 6 IAAA-32223 37,400 � 250 Mammuthusprimigenius

Molar �25.6 AMS Accept Takahashi et al., 2005

Off Rausu Hokkaido 1 Beta-85090 38,920 � 760 Mammuthusprimigenius

Molar �23.4 AMS Accept Yamada et al., 1996

Churui Hokkaido 4 Beta-241409 42,850 � 510 Mammuthusprimigenius


Off Notsukezaki Hokkaido 2 Beta-184935 ＞42,980 Mammuthusprimigenius

Molar �17.7 AMS Accept Takahashi et al., 2005Takahashi et al. 2006

Yuni Hokkaido 6 IAAA-32222 45,110 � 480 Mammuthusprimigenius


Funkawan Bay,Yakumo cho

Hokkaido 8 Beta-87674 17,900 � 90 Bison sp. Horn �22.9 AMS Accept Akamatsu et al., 1999

Anikawa River Honshu(Yamanashi)

15 NUTA-2598 15,780 � 380 Palaeoloxodonnaumanni

Unknown 0.06(G)0.06(S)

AMS G-collagenþ S-collagen

Reject Nakamura 1995

Kumaishi-doCave F4

Honshu (Gifu) 18 Gak-7007 16,720 � 880 Palaeoloxodonnaumanniþ Sinomegacerosyabeiþ Cervuspraenipponicus

mix of bonefragments

Beta Reject Okumura et al., 1982

Hanaizumi site Honshu (Iwate) 12 GaK-15893 21,430 � 1260 Palaeoloxodonnaumanni

Molar Beta Reject Hanaizumi hakkutsuchosadan 1993

Shikkari No.4 Honshu(Aomori)

9 IAAA-53431 23,600 � 130 Palaeoloxodonnaumanni

Molar �26.81 AMS accept Takahashi et al., 2006

Kumaishi-do cave Honshu (Gifu) 18 23,960 � 200 Palaeoloxodonnaumanni

Molar AMS Sample C Reject Yasui et al., 2004

Off Islands ofTowa machi

Honshu(Yamaguchi)

28 24,280 � 190 Palaeoloxodonnaumanni

Molar AMS Sample D Reject Kitagawa et al., 2006

Off Natori-Kajitanihana

Honshu (Ehime) 27 NUTA-5723 24,880 � 580 Palaeoloxodonnaumanni

Molar 1.22 3.2 �20.2 AMS S-collagenof sample E

Reject Nakamura et al.,1998; Minami andNakamura 1999

Shikkari No.4 Honshu (Aomori) 9 IAAA-52430 25,780 � 120 Palaeoloxodonnaumanni




Molar 3 �20 AMS Re-dating ofsample E byXAD-2 method

Accept Minami andNakamura 1999

Off Shimotsu’i Honshu(Okayama)


Tusk 0.039(G)0.365(S)

�24.5 AMS G-collagenþ S-collagen

Reject Otsuka et al., 2008

Off Hinomisaki Honshu(Shimane)


Tusk AMS Accept Akiyama et al., 1988Akiyama et al. 1989

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Table 1 (continued )

Site Region No. Lab Number 14C Age (BP) Fossil Element Collagen(%)

C/N ratio d13C(&)

Datingmethod

Remarks Evaluation Reference

Serikawa River Honshu (Shiga) 22 PLD-11230 29,030 � 130 Palaeoloxodonnaumanni

Molar 0.34 2.82 �19.12 � 0.21 AMS Reject Kitagawa et al., 2009



Molar 1.22 3.2 �20.2 AMS G-collagenof sample E

Accept Nakamura et al., 1998Minami andNakamura 1999

Yubetsu Hokkaido 3 Beta-134800 30,520 � 220 Palaeoloxodonnaumanni


Aikawa River Honshu(Yamanashi)


Unknown AMS? Inconsistent withgeological context

Reject Nakai et al., 1991

Shikkari No.4 Honshu(Aomori)

9 IAAA-52429 31,270 � 160 Palaeoloxodonnaumanni


Terayamano-Ana Cave

Honshu(Yamaguchi)

29 PLD-11227 31,350 � 160 Palaeoloxodonnaumanni

Molar 0.65 2.84 �20.13 � 0.22 AMS Reject Kitagawa et al., 2009

Lake Nojiri Honshu(Nagano)


Molar 0.37 4.24 �20.88 AMS Reject Sawada et al., 1992

Lake Nojiri Honshu (Nagano) 17 NUTA-631 33,620 � 810 Palaeoloxodonnaumanni

Tusk AMS Accept Nakai et al., 1991

Serikawa River Honshu (Shiga) 22 PLD-11229 34,010 � 180 Palaeoloxodonnaumanni

Molar 2.31 2.84 �20.74 � 0.22 AMS Accept Kitagawa et al., 2009


Molar 0.77 3.63 �20.6 AMS Accept Nakai et al., 1991Nakai et al. 1992Sawada et al., 1992

Sakura Honshu (Chiba) 14 NUTA-505 34,800 � 800 Palaeoloxodonnaumanni

Tusk AMS Inconsistent withgeological context

Reject Akiyama andNakai 1988


Tusk 0.21(G)0.15(S)

5.16(G)5.30(S)

AMS Reject Nakai et al., 1991

Terayamano-Ana Cave

Honshu(Yamaguchi)


Molar 5.53 2.79 �18.48 � 0.22 AMS Accept Kitagawa et al., 2009

Off Hinomisaki Honshu (Shimane) 23 Gak-12898 35,560 � 1300 Palaeoloxodonnaumanni

Tusk Beta Reject Hoshimi andMorioka 1987


Tusk 0.15 AMS Reject Nakai et al., 1991


Honshu(Yamaguchi)


Molar 0.48 2.66 �17.76 � 0.20 AMS Re-datingof sample D

Reject Kitagawa et al., 2009


Tusk 0.29(G) 4.32(G) �22.48(G) AMS G-collagenof sample F



Tusk AMS Accept Nakai et al., 1991

Kumaishi-do Cave

Honshu (Gifu) 18 PLD-11226 37,990 � 250 Palaeoloxodonnaumanni

Molar 1.25 2.89 �20.53 � 0.15 AMS Re-datingof sample C

Accept Yasui et al., 2004Kitagawa et al., 2009


Honshu(Yamaguchi)


Molar 1.93 2.99 �18.82 � 0.16 AMS Re-datingof sample D

Accept Kitagawa et al., 2006Takahashi et al., 2005Kitagawa et al., 2009

Lake Nojiri Honshu (Nagano) 17 NUTA-1262 38,310 � 1400(G) Palaeoloxodonnaumanni

Molar 2.14(G)0.52(S)

3.65(G)3,12(S)

�20.3(G) AMS Accept Nakai et al., 1991

Off Hinomisaki Honshu (Shimane) 23 NUTA-478 38,500 � 600 Palaeoloxodonnaumanni

Tusk AMS Accept Akiyama et al., 1989


Molar 0.98(G)0.26(S)

3.75(G)3.21(S)



Molar 0.55(G)0.27(S)

3.89(G) �23.12(G) AMS Reject Nakai et al., 1991

Off Onsentsu Honshu (Shimane) 24 PDL-10879 39,680 � 290 Palaeoloxodonnaumanni

Tusk 15.2 2.7 �22.33 � 0.17 AMS Accept Kitagawa et al., 2009

Lake Nojiri Honshu (Nagano) 17 39,944 � 1043(S) Palaeoloxodonnaumanni

Tusk 0.63(S) 4.42(S) �22.5 AMS S-collagenof sample F

Reject Sawada et al., 1992

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Lake Nojiri Honshu (Nagano) 17 40,130 � 1080 Palaeoloxodonnaumanni

Molar 1.2(G)2.41(S)

3.79(G) �22.05 AMS Accept Sawada et al., 1992


Molar 2.53(G)0.64(S)

3.55(G)3.04(S)

AMS Accept Nakai et al., 1991

Lake Nojiri Honshu (Nagano) 17 NUTA-1231 40,860 � 1165(S) Palaeoloxodonnaumanni

Molar 1.47(S) 3.82(S) �21.54(S) AMS S-collagen Reject Nakai et al., 1991


Molar 1.36(G)0.31(S)

3.49(G)3.00(S)


Lake Nojiri Honshu (Nagano) 17 41.566 � 927 Palaeoloxodonnaumanni

Molar 1.22(S) 3.92(S) �21.78(S) AMS S-collagenof sample G



Molar 1.54(G)1.26(S)



Molar 0.73(G)0.3(S)



Molar 2.06(G) 3.82(G) �21.8(G) AMS G-collagenof sample G

Accept Nakai et al., 1991


Molar 4.22(G)1.35(S)

3.43(G)2.94(S)



Molar 1.3(G)0.85(S)



Molar 1.96(G)1.8(S)

4.10(G)4.24(S)



Molar 2.22(G)0.43(S)

3.48(G)2.99(S)


Lake Nojiri Honshu (Nagano) 17 43,351 � 1164(G) Palaeoloxodonnaumanni

Molar 1.78(G)1.68(S)

3.64(G) �22.23(G) AMS Accept Sawada et al., 1992


Molar 0.58(G)0.4(S)

3.92(G) AMS Reject Sawada et al., 1992


Molar ＜3.54(G) 3.52(G) �20.95(G) AMS Accept Sawada et al., 1992


Molar 2.92(G)1.12(S)

3.73(G)3.79(S)

�21.42(G)�21.41(S)


Lake Nojiri Honshu(Nagano)

17 NUTA-1267 45,120 � 1350(G) Palaeoloxodonnaumanni

Molar 2.97(G)0.54(S)

3.62(G)3.10(S)



Molar 2.88(G)1.26(S)

3.52(G)3.02(S)



Molar 1.7(G)1.24(S)



Molar 0.7(G)0.29(S)

3.68(G)3.15(S)


Off Mishima Honshu (Yamaguchi) 30 NUTA-1078 48,870 � 1770 Palaeoloxodonnaumanni

Tusk AMS Accept Akiyama et al. 1992

Lake Nojiri Honshu (Nagano) 17 NUTA-1194 30,580 � 1290(G) Sinomegacerosyabei


5.62(G) AMS Reject Nakai et al., 1991


Antler 0.22(G) 3.99(G) AMS G-collagen ofsample H


Lake Nojiri Honshu (Nagano) 17 34,355 � 1345 Sinomegacerosyabei

Antler 0.33(S) 4.96(S) AMS S-collagenof sample H




4.46(G)5.72(S)

AMS Reject Nakai et al., 1991



3.61(G)3,10(S)


Lake Nojiri Honshu (Nagano) 17 41,250 � 1190 Sinomegacerosyabei

Antler 3.72(G) AMS Accept Sawada et al., 1992

Yuni Hokkaido 6 49,250 � 740 Sinomegacerosyabei

Antler AMS Accept Takahashi 2007

Negata Cave Honshu (Shizuoka) 19 Beta-29709 13,970 � 90 Cervus sp. Metapodial AMS Accept Kondo andMatsu’ura 2005

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Table

1(con

tinu

ed)

Site

Reg

ion

No.

LabNumbe

r14CAge

(BP)

Fossil

Elem

ent

Collage

n(%)

C/N

ratiod1

3C

(&)

Dating

method

Rem

arks

Evaluation

Referen

ce

Suse

Quarry

(eastfissure)

Hon

shu(A

ichi)

21Gak

-114

9514

,710

�67

0So

mekind

oflarge

mam

mal

bonefrag

men

tBeta

Reject

Kaw

amura

etal.,19

90

Neg

ataCav

eHon

shu(Shizuok

a)19

Beta-94

983

17,910

�70

Panthe

racf.

pardus

Phalan

xAMS

Accep

tKon

doan

dMatsu

’ura

2005

Yag

eQuarry

No.5

Hon

shu(Shizuok

a)20

Gak

-114

9418

,040

�99

0Cervu

ssp

.þ

some

kindsof

mam

mal

mix

ofbo

ne

frag

men

tsBeta

Reject

Kaw

amura

and

Matuhashi1

989

Kaza-an

aCav

eHon

shu(Iwate)

11Beta-11

7392

18,140

�60

Elep

han

tidae

Femur

�20.4

AMS

G-collage

nAccep

tMatsu

’ura

&Kon

do20

03b

Han

aizu

mis

ite

Hon

shu(Iwate)

12GaK

-157

9818

,470

�66

0Bov

idea

gen.

etsp

.Inden

tMolar

Beta

Reject

Han

aizu

mih

akku

tsu

chosad

an19

93Kuch

imino-se

Ryu

kyu

32NUTA

-314

425

,750

�22

0Elap

hulussp

.Fe

mur

4.38

(G)

1.21

(S)

�14.7

AMS

Accep

tNak

amura

etal.,19

96

Kumaish

i-doCav

eHon

shu(G

ifu)

1831

,010

�32

0Ca

nislupu

sUlm

aAMS

Accep

tYasuia

nd

Matsu

oka20

02Obu

roCav

eHon

shu

(Hiroshim

a)25

Beta-18

7550

31,640

�90

0Ce

rvus

sp.

Tooth

�21.1

AMS

Accep

tKaw

amura

2009

Seiryu

kutsu

Cav

eHon

shu(Fuku

oka)

31TE

RRA12

1905

0845

,500

�22

00Ce

rvus

sp.?

Limb

0.86

3.27

�23.8

AMS

Accep

tNak

agaw

aet

al.,20

07

Aba

kuch

iCav

eHon

shu(Iwate)

10Beta-11

7393

48,260

�11

00Ursus

sp.

Metacau

pus

�17.9

AMS

G-collage

nAccep

tMatsu

’ura

&Kon

do20

03a


existing radiocarbon dates is between ca. 49,000 BP and ca.16,000 BP (Fig. 3). Among the existing radiocarbon dates thenumber of unreliable samples is 22, including two conventional14C-dated specimens, 17 poorly preserved examples, two incon-sistent with their geological contexts, and one from mixed bonefragments (Table 1). For example, the youngest specimen ofP. naumanni was recovered from a bank of Anikawa River in theYamanashi Prefecture, which represented an AMS 14C date of15,780 � 380 BP (Nakamura, 1995). The gelatin collagen, however,formed only 0.06% of weight. Moreover, a piece of wood from abovethe fossil layer was dated at >36,860 BP and wood below the bonewas >47,210 BP (Nakamura, 1995). These results suggest that theAnikawa specimen is poorly preserved and dated considerablyyounger than its actual age. Although the second youngestP. naumanni example, the Kumaishido (F4) specimen, was dated to16,720 � 880 BP, this date was obtained from combined bonefragments of P. naumanni, S. yabei, and C. praenipponicus, not fromP. naumanni only (Okumura et al., 1982).

The remaining 36 P. naumanni specimens are considered morereliable with dates from 48,870 � 1770 BP (Mishima specimen) to23,600 � 130 BP (Shikkari No.4 specimen) (Fig. 4).

3.2. S. yabei

Eight specimens of S. yabei are directly 14C-dated. A single datewas obtained from a specimen recovered from Hokkaido and theothers from Honshu specimens. The range of existing radiocarbondates is between ca. 49,000 BP and ca. 17,000 BP (Fig. 3). Theyoungest specimen, which dated to 16,720 � 880 BP, was obtainedfrom combined bone fragments including other mammal species(Okumura et al., 1982). Another four specimens with low gelatincollagen content and high C/N ratios are also rejected (Table 1). Thenumber of reliable S. yabei specimens then is three, which showa range of 14C dates is from 49,250 � 740 BP (Yuni specimen) to40,560 � 1500 BP (Nojiriko specimen) (Fig. 4).

3.3. M. primigenius

Twelve M. primigenius specimens are directly dated by the AMSmethod. A single date was obtained from a sample recovered fromthe bottom of the Sea of Japan and the others from Hokkaido. Therange of existing 14C dates is from ca. 45,000 BP to ca. 16,000 BP(Fig. 3). As mentioned above, the youngest date, the Yubari spec-imen, is rejected because its exact provenance is unknown (Kamei,1990; Takahashi et al., 2006) (Table 1). The number of remaining

Fig. 3. Plot of all dates of terrestrial mammal fossils on the Japanese Archipelago.

Fig. 4. Plot of the accepted dates and the range of the LGM.


acceptableM. primigenius specimens is eleven, which represent 14Cdates that range from 45,110 � 480 BP (Yuni specimen) to19,530 � 80 BP (Ogoshi specimen) (Fig. 4).

3.4. Bison sp.

Only a single specimen (Funkawan Bay) is directly dated by theAMS method. The specimen recovered from Hokkaido was dated at17,900 � 90 BP (Akamatsu et al., 1999) (Fig. 4). From the Hanaizumisite in Iwate Prefecture, northern Honshu, the molar of a bovid,which was considered B. priscus dated at 18,470 � 660 BP(Hanaizumi iseki hakkutsu chosadan,1993), was rejected because itwas measured by the conventional method (Table 1).

3.5. Other species

Ten specimens are directly radiocarbon dated from the remainsof several other species: C. lupus, Cervus sp., Ursus sp., Panthera cf.pardus, elephantidae gen. et sp. indent., and other unidentifiedmammals. A single specimen originated from Kyusyu and theothers from Honshu. The existing radiocarbon dates range from ca.48,000 BP to ca. 14,000 BP (Fig. 3). Three specimens, obtained frommixed bone fragments and analyzed by the conventional method,are rejected (Table 1). The ranges of the remaining 14C dates are31,010� 320 BP for C. lupus, three dates between 45,500� 2200 BPand 13,970� 90 BP for Cervus sp., a single date of 48,260 � 1100 BPfor Ursus sp., a date of 17,910 � 70 BP for Panthera cf. pardus, anda date of 18,140 � 60 BP for elephantidae gen. et sp. indent (Fig. 4).The femur of elephantidae recovered from Kaza-ana Cave in IwatePrefecture is important because it suggests that proboscideasurvived until 18ka in northern Honshu.

A single specimen of Elaphulus sp. recovered from the seabottom at a depth of 122m around the Ryukyu Islands was dated at

25,750 � 220 BP (Kuchimino-se specimen) (Nakamura et al., 1996)(Fig. 1; Table 1). This specimen may play an important role in thediscussion of land-bridge formation in the East China Sea aroundthe LGM (Nakamura et al., 1996).

4. Discussion

Pleistocene megafaunal extinction is one of the most debatedsubjects in Quaternary research throughout the world (e.g., Martin,1984, 2005; Grayson, 2001; Grayson and Melzer, 2002; Stuart et al.,2002, 2004; Steadman and Martin, 2003; Barnosky et al., 2004;Meltzer, 2004; Burney and Flannery, 2005; Surovell et al., 2005;Koch and Barnosky, 2006; Guthrie, 2006; Wroe and Field, 2006;Ugan and Byers, 2007, 2008; Surovell and Waguespack, 2008;Pushkina and Raia, 2008). Recent reviews for worldwide extinc-tions reached the conclusions that 1) the timing varied betweeneach region, 2) it is not possible to explain the causes of extinctionsolely by environmental changes, and 3) direct and/or indirecthuman impacts possibly precipitated the extinctions across theworld (Barnosky et al., 2004; Burney and Flannery, 2005; Koch andBarnosky, 2006). However, in northern Europe, Siberia, Alaska andYukon extinctions occurred in two pulses coincident with climaticchanges: extinction of warm-adapted animals at the end of the LastInterglacial and LGM, and cold-adapted animals during the Pleis-toceneeHolocene transition (Guthrie, 2003, 2004, 2006; MacPheeet al., 2002, 2005; Stuart et al., 2002, 2004; Kuzmin and Orlova,2004; Barnosky et al., 2004; Koch and Barnosky, 2006). Thefollowing section discusses the range of reliable 14C dates of fourspecies: P. naumanni, S. yabei, M. primigenius and Bison sp., whichrepresent the large mammals of the Palaeoloxodon-Sinomegacer-oides complex and mammoth fauna, respectively, in Japan, whichare adequate for the discussion of extinctions of warm-adapted andcold-adapted animals.

4.1. Palaeoloxodon-Sinomegaceroides complex: timing of theextinction and its relationship with vegetation changes

The reliable dates of P. naumanni range from 49 ka to 23 ka.Specimens younger than 23 ka are unacceptable (Table 1; Fig. 3) asalready pointed out by Takahashi (2007) and Kitagawa et al. (2009).This indicates that P. naumanni was nearly extinct by around 23 ka,although the possibility of their survival later in some refugia (suchas Kaza-ana Cave in northern Honshu) is not rejected. As thenumber of reliable dates of S. yabei is small, it is difficult toreconstruct details of its chronology. P. naumanni and S. yabei,however, are considered to be adapted to temperate forests(Hasegawa, 1972; Kawamura, 1991, 1994), indicating that bothspecies were affected when the climate became colder.

The analysis of sediment cores from the sea bottom, includingthe Oki Ridge in the Sea of Japan, suggests that the LGM started atca. 30,000 cal BP and ended at ca. 19,000 cal BP (Yokoyama et al.,2007; Lambeck et al., 2002). The results of analysis on pollen,organic carbon and nitrogen from sediment cores in Lake Nojiri,central Honshu (Kumon et al., 2009, 2003) showed that from ca.62,000 cal BP to ca. 29,000 cal BP (roughly corresponding to MIS3)deciduous broad-leaved trees were dominant, indicating a rela-tively warmer period. Then the climate became coldest from ca.29,000 cal BP to ca. 18,000 cal BP (early MIS2), which correspondsto the LGM. During this period, the vegetation was dominated bysubarctic conifers such as Picea and Abies. The results of sedimentcore analysis suggest that the LGM started at ca. 30,000e29,000 calBP (ca. 25,000 14C BP), and ended at ca. 19,000e18,000 cal BP (ca.16,000 14C BP). It appears that the timing of the P. naumanniextinction, whose principal habitat was in temperate climates, was


roughly coincident with the onset of the LGM and the subsequentvegetation change (Fig. 4).

4.2. Mammoth fauna: timing of the extinction and its relationshipto vegetation changes

The range of reliable 14C dates of M. primigenius is divided intotwo clusters: one from 45 ka to 37 ka and another from 25 ka to 20ka (Fig. 4). In Hokkaido, between 60e35 ka and 25e10 ka thevegetation included open-forest taiga composed mainly of Larixgmelinii, Picea pumila, and Picea jezoensis with grassy plains(Igarashi et al., 1989, 1990; Igarashi, 1993). Such vegetation, similarto that of northernmost Sakhalin at present, provided a suitableenvironment for M. primigenius. Between 34 ka and 26 ka, theclimate slightly ameliorated in Hokkaido and the vegetation of theIshikari Lowland inwestern Hokkaidowas dominated by deciduousbroadleaf trees (Igarashi and Kumano, 1981). The lack ofM. primigenius dates from 37 ka to 25 ka indicates that they shiftedtheir habitats to the north in response to vegetational change assuggested by Takahashi et al. (2004, 2006). The radiocarbon date ofBison sp., at 17,900 � 90 BP implies that the mammoth faunasurvived at least to ca. 18 ka. This implication is supported by pollenanalysis, which shows that steppe with open-larch forests devel-oped in the Kenbuchi Basin, northern Hokkaido, under cold/dryclimates during the LGM (25-16 ka) (Igarashi et al., 1993). Tosummarize, woolly mammoth on the Japanese Archipelago becameextinct or migrated northward after the LGM when the climateameliorated (Fig. 4).

4.3. Timing and causes of lLP megafaunal extinction on theJapanese Archipelago

Several hypotheses about the timing and causes of lLP mega-faunal extinction have already been proposed for Japan. Kawamura(1991, 1994, 2007) suggested that the combination of climatechange and human impacts from 20 ka to10 ka led to the extinctionof large animals including P. naumanni, S. yabei, C. praenipponicus,M. primigenius, and B. priscus. Norton et al. (2010) also proposedthat the extinctions of P. naumanni, S. yabei andM. primigeniuswereinfluenced by direct and/or indirect anthropogenic impacts at theMIS 3-2 transition (30e20 ka). Akazawa (2005) reviewed studies ofJapanese Pleistocene mammalian fauna and postulated thatP. naumanni and S. yabeiwent extinct at 10,000 BP, but these studiesare not founded on the direct evidence of 14C dates. While hiscompilation was incomplete, Takahashi (2007) examined the 14Cdates of themain fossils on the Japanese Archipelago and suggestedthat P. naumanni became extinct by around 23 ka. Based on the newand previously published radiocarbon dates of P. naumanni andM. primigenius specimens from Hokkaido, Takahashi et al. (2004,2006) also indicated that the boundary of these two species thatmigrated back and forth north-south was matched with the rangeof vegetation type, implying that climatic and vegetation changeshad significant roles in their migrations and extinctions. Kitagawaet al. (2009) dated seven specimens of P. naumanni recoveredfrom western Japan, and pointed out that this species disappearedby ca. 30,000 cal BP.

Evaluation of the radiocarbon dates and the examination ofrelationships between the vegetational change and AMS 14C chro-nology of terrestrial mammals indicate that megafauna comprisingof the Palaeoloxodon-Sinomegaceroides complex were extinct by theonset of LGM (ca. 23,000 BP), and large mammals of the mammothfauna survived during the LGM and probably became extinct ormigrated northward only after the end of LGM. This suggests thatthe two species of Proboscidea, P. naumanni and M. primigenius,went extinct at different times according to the disappearance of

their preferable environments. Thus, lLP extinction on the JapaneseArchipelago could occur in “two pulses” similar to northern Europe,Siberia, Alaska and Yukon (Guthrie, 2003, 2004, 2006; MacPheeet al., 2002, 2005; Stuart et al., 2002, 2004; Kuzmin and Orlova,2004; Barnosky et al., 2004; Koch and Barnosky, 2006).

It is also certain that humans have inhabited the JapaneseArchipelago since 40e35 ka, the onset of Early Upper Palaeolithic(Izuho, 2010; Izuho and Sato, 2008), considerably earlier than theoccurrence of megafaunal extinction. This implies that the lLPmegafauna coexisted with human hunters for at least 10,000 years.

Although the possibility that direct or indirect human impacts(e.g. Norton et al., 2010) may have partly influenced the extinctionof some large herbivores is not denied, the process of Japanesemegafaunal extinction can be explained without consideration ofhuman influence. It is important to separate the “main” cause ofextinction from other “minor” factors. The result of this studysuggests that the “main” cause of lLP extinction on the JapaneseArchipelago was climate-induced ecosystem changes. The possiblerole of human hunters in these extinctions, however, has not beensufficiently discussed in Japan. Thus, in future studies it will benecessary to examine details of human impacts on both ecosystemsand large mammals.

The evidence of lLP extinction on the Japanese Archipelago hasyet to be adequately clarified. The dates of S. yabei and B. priscus areespecially limited and several species have never been dated (e.g.,A. alces). This topic will need to be discussed in more detail in thefuture when additional evidence has accumulated.

5. Conclusion

This paper reconstructed the reliable AMS 14C chronology of lLPterrestrial mammals and determined the timing of lLP extinctionson the Japanese Archipelago. The 14C extinction chronology wascompared with lLP vegetation changes in Honshu and Hokkaido.The demise of warm-adapted animals (the Palaeoloxodon-Sinome-gaceroides complex) occurred around the LGM and cold-adaptedanimals (the mammoth fauna) disappeared after the LGM whenthe climate started to warm, suggesting that the lLP extinction onthe Japanese Archipelago could have occurred in “two pulses”similar to the Eurasian continent and North America. The similarityof extinction patterns between the Japanese Archipelago andseveral other regions of the Northern Hemisphere, and the corre-spondence between the extinction timing and the vegetationchange, imply that the main causes of lLP extinction wereecosystem changes rather than direct human impacts such as“overkill” on the Japanese Archipelago.

Acknowledgements

An earlier version of this paper was presented at the symposium“The World of Mammoths: Vth International Conference onMammoths and their Relatives” in Le Puy-en-Velay, France. Wewould like to thank Koh Hamaguchi for helping us to collect 14Cdates of mammal fossils, and Takuya Yamaoka, Yoshinori Oda, andTsubasa Kamei for reading an earlier version of the manuscript. Wedeeply thank also Ian Buvit for correcting the English. Errors, biases,and shortcomings in the paper, if any, are our responsibility.

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Timing of megafaunal extinction in the late Late Pleistocene on the Japanese Archipelago

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