

後期更新世から完新世の初めにかけて、世界の大型動物(通常、体重が 44 キログラム(97 ポンド)を超える動物種と定義される)の大半が絶滅し、[1]、その結果、地球全体で動物の密度と多様性が崩壊した。[2]後期更新世の絶滅は、大型動物への極端なサイズの偏り(小型動物はほとんど影響を受けなかった)、これらの絶滅した大型動物種に代わる生態学的継承の広範な欠如、 [3]、および結果として以前に確立された動物相の関係と生息地の体制シフトによって、以前の絶滅と区別される。絶滅の時期と深刻さは地域によって異なり、人為的要因と気候的要因のさまざまな組み合わせによって引き起こされたと考えられている。[3]大型動物の個体群に対する人為的影響は、狩猟(「過剰」)、 [4] [5]およびおそらく環境の変化によって引き起こされたと考えられている。 [6]絶滅における人為的要因と気候的要因の相対的な重要性は、長年にわたる論争の対象となっている。[3]
大規模な絶滅は、オーストラリア・ニューギニア(サフル)で約5万年前に始まり、アメリカ大陸では約13,000年前に発生し、これらの地域への初期の人類の移住と時期を同じくしています。 [7]ユーラシア北部での絶滅は、5万年前から1万年前までの数万年にわたって段階的に発生しましたが、[2]アメリカ大陸での絶滅は事実上同時に発生し、最大でも3000年しか続きませんでした。[4] [8]全体として、後期更新世には世界中の大型動物種の約65%が絶滅し、[9]北米では72%、南米では83%、オーストラリアでは88%にまで増加し、[10]オーストラリアとアメリカ大陸では体重1,000キログラム(2,200ポンド)を超えるすべての哺乳類が絶滅し、[11]世界全体では約80%が絶滅しました。[12]アフリカ、南アジア、東南アジアでは他の地域よりも穏やかな絶滅が見られました。[10]
生物地理学的領域による絶滅
まとめ
導入
_in_each_TDWG_country_during_the_last_132_000_years,_only_counting_extinctions_earlier_than_1000_years_BP.jpg/500px-thumbnail.jpg)
後期更新世には体重40キログラム(88ポンド)以上の哺乳類が多数絶滅し、その中には体重1トン以上の哺乳類の約80%が含まれていた。大型動物の絶滅の割合は、アフリカからの人間の移住距離が遠くなるにつれて次第に大きくなり、オーストラリア、北米、南米で絶滅率が最も高かった。[12]
絶滅の拡大は、現代人の移住パターンを反映している。アフリカから遠く離れれば離れるほど、人間がその地域に居住したのが最近であるほど、その環境(大型動物を含む)が人間に慣れるのに要した時間は短くなる(そしてその逆もまた同様)。
この絶滅を説明する主な仮説は 2 つあります。
現在入手可能なデータと先史時代の過剰絶滅仮説の間には矛盾がいくつかある。例えば、オーストラリアの大型動物の絶滅の時期については曖昧な点がある。[14]先史時代の過剰絶滅仮説を裏付ける証拠には、大陸に生息していた同族が姿を消した後も、いくつかの島で大型動物が数千年にわたって生息し続けたことがある。例えば、地上性ナマケモノは南北アメリカの地上性ナマケモノが絶滅した後もアンティル諸島で生き延び、ケナガマンモスは本土で絶滅してから6000年後に遠く離れたウランゲル島で絶滅し、ステラーカイギュウは北太平洋の大陸岸から姿を消した後も、孤立した無人のコマンドル諸島沖で数千年にわたって生息し続けた。[15]これらの島の種のその後の絶滅は、これらの島々への後の人類の入植と相関している。
大型動物絶滅の主な原因が人類の到来時期か気候変動かという当初の議論は、必然的に古生物学的証拠と地質学的年代測定技術の組み合わせに基づいていた。最近、生き残った大型動物集団の遺伝子分析によって新たな証拠がもたらされ、「特に過去 75,000 年間の大型動物の個体数の減少を気候が予測できないということは、この時期の大型動物の動態の主な要因は人間の影響であったことを示唆している」という結論に至った。[16]
最近の研究では、それぞれの種が環境の変化に対して異なる反応を示しており、単一の要因だけでは多様な絶滅を説明できないことが示されています。原因には、気候変動、種間の競争、不安定な個体群動態、人間の捕食の相互作用が関係している可能性があります。[17]
アフリカ
アフリカは最も影響を受けていない地域の一つであるが、それでも絶滅に見舞われ、特に後期更新世から完新世への移行期には顕著であった。これらの絶滅は主に草原の生息地の変化による気候的要因によるものと考えられる。[18]
- 有蹄類
- 偶蹄類
- イノシシ科
- メトリディオコエルス (亜種)
- コルポコエルス(亜種)
- ウシ科(ウシ、カモシカ)
- 巨大な水牛( Syncerus antiquus )
- メガロトラガス
- ルシンゴリクス
- ミナミスプリングボック( Antidorcas australis )
- ボンドスプリングボック( Antidorcas bondi )
- ダマリスクス・ヒプソドン
- ダマリスクス・ニロ
- 大西洋ガゼル( Gazella atlantica )
- ガゼラ・ティンギタナ
- カプリナエ
- シカ科
- メガセロイデス・アルジェリクス(北アフリカ)
- イノシシ科
- 奇蹄類
- 偶蹄類
- 長鼻類
- ゾウ科(ゾウ)
- Palaeoloxodon iolensis ? (他の著者は、この分類群は中期更新世の終わりに絶滅したと示唆している)
- ゾウ科(ゾウ)
- 齧歯類
- Paraethomys filfilae ?
南アジアと東南アジア


インド亜大陸における絶滅の時期は、信頼できる年代測定がないため不明である。[20]中国の遺跡でも同様の報告があるが、その地域で大型動物の分類群が完新世まで生き延びたという証拠はない。[21]東南アジアと中国南部での絶滅は、開けた森林生息地から閉じた森林生息地への環境の変化の結果であると提案されている。[22]
- 有蹄類
- フォリドタ
- オオセンザンコウ ( Manis palaeojavanica )
- 食肉類
- イヌ目
- ヒキガエル科
- クマ
- パンダ(ジャイアントパンダの祖先)
- クマ
- ヒキガエル科
- イヌ目
- アフロテリア
- 鳥類
- 飛べないカモ(シリヤネッタハセガワ)[27]
- レプトプティロス・ロブストゥス
- 爬虫類
- 霊長類
- いくつかのサル(Simiiformes)属。
- ポンゴ(オランウータン)
- ポンゴ・ウェイデンレイチ(中国南部)
- さまざまなホモ属(旧人類)
- ホモ・エレクトス・ソロエンシス(ジャワ)
- ホモ・フローレシエンシス(フローレス)
- ホモ・ルゾネンシス(フィリピン、ルソン島)
- デニソワ人(ホモ属)
- ポンゴ(オランウータン)
- いくつかのサル(Simiiformes)属。
ヨーロッパ、北アジア、東アジア




RhinocérosEnFuite.jpg/500px-16_PanneauDesLions(CentreGauche)RhinocérosEnFuite.jpg)
.jpg/500px-Lions_painting,_Chauvet_Cave_(museum_replica).jpg)




旧北区はヨーロッパ大陸全体に広がり、北アジア、コーカサス、中央アジアを経て中国北部、シベリア、ベーリング地峡にまで広がっている。絶滅はアフリカや南アジア、東南アジアよりも北ユーラシアで深刻だった。これらの絶滅は数万年にわたって段階的に起こり、約5万年前から1万年前までの間に発生し、温帯に適応したハタハタゾウやナナフシサイなどの種は、寒帯に適応したマンモスやケナガサイなどの種よりも一般的に早く絶滅した。気候変動は絶滅の大きな要因であると考えられており、おそらく人間の狩猟と相まってそうである。[2]
- 有蹄類
- 偶蹄類哺乳類
- ウシ科
の各種
- 草原バイソン( Bison priscus )
- バイカルヤク ( Bos baikalensis ) [28]
- ヨーロッパスイギュウ( Bubalus murrensis )
- ヨーロッパタール( Hemitragus cedrensis ) [29] [30]
- オオジャコウウシ( Praeovibos priscus ) [31]
- サイガ(サイガボレアリス)[32]
- ねじれた角を持つカモシカ(Spirocerus kiakhtensis)[33] [34]
- ヤギツネレイヨウ(パラブバリス・カプリコルニス)[33] [34]
- Bubalus wansijocki(中国北部原産の絶滅した水牛)
- 様々なシカ(Cervidae)種。
- オオジカ/ アイルランドヘラジカ ( Megaloceros giganteus )
- クレタ島鹿( Candiacervus spp.)
- ハプロイドセロス・メディテラネウス[35] [36]
- シノメガセロス属(日本ではシノメガセロス・ヤベイ、中国ではシノメガセロス・オルドシアヌス、おそらくシノメガセロス・パキオステウスを含む)。 [37]
- ドワーフリュウクジカ( Cervus astylodon )
- 在来種のカバ類すべて[38]
- Camelus knocklochi [39]およびその他のCamelus属
- ウシ科
の各種
- 奇蹄類哺乳類
- 様々なエクウス属
- さまざまな野生馬の亜種 (例: Equus c. gallicus、[40] [41] Equus c. latipes、[34] [40] [42] Equus c. uralensis [40] )
- エクウス・ダリアンエンシス(中国北部に生息する野生馬の一種)
- ヨーロッパノロバ( Equus hydruntinus ) (アナトリアの避難所で完新世後期まで生き延びた)
- Equus ovodovi(完新世後期まで中国北部の避難所で生き延びた)
- すべての在来サイ(Rhinocerosidae)種。
- 様々なエクウス属
- 偶蹄類哺乳類
- 食肉類
- イヌ目
- イヌ科
- ヒキガエル科
- 各種のUrsus属。
- ステップヒグマ( Ursus arctos " priscus ") [44]
- ガムスルゼンホラアナグマ( Ursus ingressus ) [45]
- 更新世の小型洞窟クマ( Ursus rossicus )
- 洞窟クマ( Ursus spelaeus )
- 巨大なホッキョクグマ( Ursus maritimus tyrannus )
- イタチ科
- イタチ科
- いくつかのカワウソ科(Lutrinae)属。
- 頑強な更新世のヨーロッパカワウソ( Cyrnaonyx )
- アルガロルトラ
- サルデーニャオオカワウソ( Megalenhydris barbaricina )
- サルデーニャコビトカワウソ( Sardolutra )
- クレタカワウソ( Lutrogale cretensis )
- いくつかのカワウソ科(Lutrinae)属。
- イタチ科
- 各種のUrsus属。
- フェリフォルミア
- ネコ科のさまざまな種。
- ホモテリウム・ラティデンス(シミター歯猫とも呼ばれる)
- 洞窟オオヤマネコ( Lynx pardinus spelaeus ) [46]
- イソワールオオヤマネコ( Lynx issiodorensis )
- パンサー属
- ヨーロッパホラアナライオン( Panthera spelaea )
- ヨーロッパ氷河期ヒョウ( Panthera pardus spelaea )
- ハイエナ科
- 洞窟ハイエナ( Crocuta crocuta spelaeaおよびCrocuta crocuta ultima )
- 「ハイエナ」プリスカ
- ネコ科のさまざまな種。
- イヌ目
- すべての在来ゾウ(ゾウ科)種。
- マンモス
- ケナガマンモス( Mammuthus primigenius )
- ドワーフサルデーニャマンモス( Mammuthus lamarmorai )
- インドゾウ( Palaeoloxodon antiquus ) (ヨーロッパ)
- ナウマンニ古生代化石(日本、おそらく韓国と中国北部にも生息)
- パレオロクソドン・フアイホエンシス(中国)
- ドワーフゾウ
- パレオロクソドン・クロイツブルギ(クレタ島)
- キプロスゾウ( Palaeoloxodon cypriotes )
- パレオロクソドン・ムナイドリエンシス(シチリア島)
- マンモス
- げっ歯類
- アロクリケトゥス
- Cricetus major(別名、 Cricetus cricetus major)
- Dicrostonyx gulielmi (北極レミングの祖先)
- オオヤマアラシ( Hystrix refossa )
- レイシア属(マルタとシチリアのオオヤマネ) [47]
- マルモタ パレオカウカシカ
- ミクロタス・グラフィ
- ミモミス属
- M.ピレナイカス
- M. シャンドレンシス
- プリオミス・レンキ
- スペルモフィルス・シテロイデス
- スペルモフィルス・セヴェルスケンシス
- スペルモフィルス・スーパーシリオサス
- Trogontherium cuvieri(大型ビーバー)
- ウサギ目
- Lepus tanaiticus (別名Lepus timidus tanaiticus )
- ナキウサギ( Ochotona ) 属など
- オオナキウサギ( Ochotona whartoni )
- トノモチョタ属
- T. カサネンシス
- T.シコタナ
- T.メジャー
- 鳥類
- 霊長類
- 爬虫類
- Solitudo sicula ;約 12,500 年前までシチリア島で生き残っていました。
- ラケルタ シクリメリテンシス;マルタから。
北米
北米における絶滅は、現在からおよそ13,800~11,400年前、後期更新世の終わりに集中しており、これはヤンガードリアス寒冷期の始まりと狩猟採集民のクローヴィス文化の出現と一致している。北米の絶滅における人間と気候要因の相対的な重要性は、大きな論争の的となっている。絶滅した種は、約35属に上る。[4]アラスカ・ユーコン地域以南の北米の放射性炭素記録は、信頼できる年表を構築するには「不十分」であるとされている。[48]
_1_(15257877377).jpg/500px-Bison_latifrons_fossil_buffalo_(Pleistocene;_North_America)_1_(15257877377).jpg)






















北米の絶滅(草食動物 (H)または肉食動物(C)として記載)には以下が含まれます。
- 有蹄類
- 偶蹄類哺乳類
- ウシ科
の各種
- 更新世のバイソンのほとんどの形態(北米のバイソンとユーラシアの
バイソン・ボナサスのみが生き残った)
- 古代バイソン( Bison antiquus ) ( H )
- オオバイソン( Bison latifrons ) ( H )
- 草原バイソン( Bison priscus ) ( H )
- バイソン オクシデンタリス( H )
- Caprinae属のいくつかの種(ジャコウウシは生き残った)
- オオジャコウウシ( Praeovibos priscus ) ( H )
- シュラブオックス( Euceratherium collinum ) ( H )
- ハーランジャコウウシ( Bootherium bombifrons ) ( H )
- セルゲルの牛( Soergelia mayfieldi ) ( H )
- ハリントンシロイワヤギ( Oreamnos harringtoni ;現存する近縁種よりも小型で、分布は南方である) ( H )
- サイガ(Saiga tatarica ; 絶滅)(H)
- 更新世のバイソンのほとんどの形態(北米のバイソンとユーラシアの
バイソン・ボナサスのみが生き残った)
- 鹿
- クワガタヘラジカ( Cervalces scotti ) ( H )
- アメリカシカ( Odocoileus lucasi ) ( H )
- トロントセロス ヒプノゲオス( H )
- 様々なAntilocapridae属(プロングホーンは生き残った)
- カプロメリクス( H )
- ストコセロス( H )
- テトラメリクス( H )
- パシフィックプロングホーン( Antilocapra pacifica ) ( H )
- いくつかのペッカリー(Tayassuidae)種。
- フラットヘッドペッカリー( Platygonus ) ( H )
- ナガバペッカリー( Mylohyus ) ( H )
- クビワペッカリー( Dicotyles tajacu ; 絶滅、分布域は半再定着) ( H ) ( Muknalia minimusは新参シノニム)
- ラクダ科の様々な種
- ウシ科
の各種
- 奇蹄類哺乳類
- ウマ科の在来種すべて
- 北アメリカの後期更新世に生息していたカバル系の馬(Equus cf. ferus )は、歴史的にはEquus fraternus、Equus scotti、Equus lambeiなど多くの異なる種に分類されてきたが、これらの馬の分類学は不明瞭であり、これらの種の多くは互いに同義語である可能性があり、おそらく単一の種のみを表している可能性がある。[49] [50] [51]
- 竹馬( Haringtonhippus francisci / Equus francisci ; ( H )
- バク ( Tapirus ; 3種)
- ウマ科の在来種すべて
- †ノトゥングラタ目
- ミクソトキソドン[52] [53] ( H )
- 偶蹄類哺乳類
- 食肉類
- フェリフォルミア
- いくつかのネコ科の種。
- サーベルタイガー ( †マカイロドン亜科)
- スミロドン・ファタリス(サーベルタイガー) ( C )
- ホモテリウム血清シミター歯ネコ( C)
- アメリカチーター( Miracinonyx trumani ; 真のチーターではない)
- クーガー(Puma concolor ;北米から絶滅した大型動物の生態形態、南米の個体群が以前の生息域に再定着)( C)
- ジャガランディ( Herpailurus yagouaroundi ; 絶滅、範囲が半再定着) ( C )
- マーゲイ( Leopardus weidii ; 絶滅) ( C )
- オセロット(Leopardus pardalis ;絶滅したが、生息域はわずかに再定着した)(C)
- ジャガー
- 更新世の北米ジャガー(Panthera onca augusta ; 分布域は他の亜種によって部分的に再植民化されている)(C)
- 北米ジャガー
- Panthera balamoides (疑わしいが、短頭熊類Arctotheriumの同義語ではないかと示唆されている)
- ライオンズ
- サーベルタイガー ( †マカイロドン亜科)
- いくつかのネコ科の種。
- イヌ目
- イヌ科
- ヒキガエル科
- イタチ科
- メフィティダエ
- スズメバチ(Brachyprotoma obtusata)[55](C)
- イタチ科
- ステップケナガイタチ(Mustela eversmanii ; 絶滅)[56](C)
- メフィティダエ
- クマ科の
各種
- アルクトドゥス・シムス(C)
- フロリダメガネグマ( Tremarctos floridanus ) ( C )
- 南米ヒメグマ(Arctotherium wingei)[57] [54] ( C )
- 巨大なホッキョクグマ( Ursus maritimus tyrannus ; 生息者の可能性あり) ( C )
- イタチ科
- フェリフォルミア
- アフロテリア
- エウアルコントグリレス
- コウモリ
- ストックス吸血コウモリ( Desmodus stocki ) ( C )
- 原始のヒゲコウモリ( Pteronotus ( Phylodia ) pristinus ) ( C )
- げっ歯類
- ジャイアントビーバー( Castoroides ) 属
- オヒオエンシス( H )
- カストロイデス・レイセヨラム( H)
- クラインヤマアラシ ( Erethizon kleini ) [59] ( H )
- オオシロイバネズミ( Peromyscus nesodytes ) ( C )
- ネオコエルス属など
- ピンクニーカピバラ( Neochoerus pinckneyi ) ( H )
- ネオコエルス・アエソピ( H)
- ネオトマ・フィンドレイ
- ネオトマ・ピグマエア
- シナプトミス・オーストラリス
- すべてのジャイアント フーティア (Heptaxodontidae) 属。
- 鈍歯オオヒメグマ( Amblyrhiza inundata ;アメリカクロクマほどの大きさに成長する) ( H )
- 板歯巨大フティア( Elasmodontomys obliquus ) ( H )
- ねじれ歯ネズミ( Quemisia gravis ) ( H )
- オズボーンズキーマウス( Clidomys osborn's ) ( H )
- サイマカ フルボプルビス( H )
- ジャイアントビーバー( Castoroides ) 属
- ウサギ目
- コウモリ
- エウリポティフラ
- 異節動物
- 鳥類
- 水鳥
- アヒル
- バミューダ飛べないカモ( Anas pachyscelus ) ( H )
- カリフォルニア飛べない海ガモ( Chendytes lawi ) ( C )
- メキシコオオカモ( Oxyura zapatima ) [57] ( H )
- ネオチェンバルバディアナ( H )
- アヒル
- トルコ( Meleagris ) 属
- カリフォルニアターキー( Meleagris californica ) ( H )
- メレアグリス・クラッシペス[57] ( H )
- ツル目魚類各種
- すべての洞窟クイナ(Nesotrochis)属
- アンティル洞窟レール( Nesotrochis debooyi ) ( C )
- バルバドスクイナ( Incertae sedis ) ( C )
- キューバ飛べないヅル( Antigone cubensis ) ( H )
- ラ ブレア クレーン( Grus pagei ) ( H )
- すべての洞窟クイナ(Nesotrochis)属
- フラミンゴ(Phoenicopteridae)の各種種。
- ドウツボシドリ( Fratercula dowi ) ( C )
- 更新世のメキシコダイバー属
- プリオリンバス・バリオステウス( C )
- ポディセプス属
- ポディセプス・パルヴス[57] ( C )
- コウノトリ
- ラ・ブレア/アスファルトコウ(Ciconia maltha)[57](C)
- ウェットモアコウノトリ(Mycteria wetmorei)[57](C)
- 更新世メキシコウ属( Phalacrocorax属)[57]
- ゴレテンシス( C )
- ファラクロコラックス・チャパレンシス( C )
- 残りのテラトルニ科(Teratornithidae)属
すべて。
- アイオロニス・インクレディビリス( C )
- ニシキヘビ( C)
- オスカラビス・オルソニ( C )
- テラトルニス・メリアミ( C)
- テラトルニス・ウッドバーネンシス( C )
- いくつかの新世界ハゲワシ類(Cathartidae)種。
- 更新世クロコンドル(Coragyps occidentalis ssp.)(C)
- 大型動物相 カリフォルニアコンドル( Gymnogyps amplus ) ( C )
- クラークコンドル( Breagyps clarki ) ( C )
- キューバコンドル( Gymnogyps varonai ) ( C )
- いくつかのタカ科の種。
- アメリカハゲワシ( Neophrontops americanus ) [57] [70] ( C )
- アメリカワシ( Amplibuteo woodwardi ) ( C )
- キューバオオタカ( Buteogallus borrasi ) ( C )
- ダゲットワシ( Buteogallus daggetti ) ( C )
- ワシ( Buteogallus fragilis ) ( C )
- キューバオオタカ(Gigantohierax suarezi)[71] [72](C)
- エララントワシ( Neogyps errans ) ( C )
- グリンネルズクレステッドワシ ( Spizaetus grinnelli ) [57] ( C )
- ウィレットクマタカ(Spizaetus willetti)[57](C)
- カリブタイタンホーク(Titanohierax)(C)
- いくつかのフクロウ(Strigiformes)属の種。
- ブレアミニチュアフクロウ( Asphaltoglaux ) ( C )
- クロチキンのコビトフクロウ( Glaucidium kurochkini ) ( C )
- ブレアフクロウ( Oraristix brea ) ( C )
- キューバオオフクロウ(オルニメガロニクス) ( C )
- バミューダキツツキ( Colaptes oceanicus ) ( C )
- いくつかのカラカラ(Caracarinae)属。
- バハマ陸生カラカラ ( Caracara sp.) ( C )
- プエルトリコ陸生カラカラ ( Caracara sp.) ( C )
- ジャマイカ カラカラ( Carcara Tellustris ) ( C )
- キューバカラカラ ( Milvago sp.) ( C )
- イスパニオラン カラカラ ( Milvago sp.) ( C )
- オウムガイ
- オウム目
- メキシコオオハシインコ(Rhynchoppsitta phillipsi)[57](H)
- オウム目
- 水鳥
- いくつかの巨大なカメの種。
- ヘスペロテストゥド( H )
- ゴフェラス属
- ゴフェラス・ドンラロイ( H )
- ケロノイディス属
- ケロノイディス・マルカノイ( H )
- ケロノイディス・アルブリオルム( H )
生き残った種も、ある意味では絶滅した種と同じくらい重要です。バイソン( H )、ハイイロオオカミ( C )、オオヤマネコ( C )、ハイイログマ( C )、アメリカクロクマ( C )、シカ(例:カリブー、ヘラジカ、ワピチ (ヘラジカ)、Odocoileus spp.) ( H )、プロングホーン( H )、メジロペッカリー( H )、ジャコウウシ( H )、オオツノヒツジ( H )、シロイワヤギ( H ) です。生き残った種のリストには、第四紀の絶滅イベントで絶滅したが、中期完新世の間に南米の残存個体群から少なくとも生息域の一部に再定着した種も含まれています。たとえば、クーガー( C )、ジャガー( C )、オオアリクイ( C )、クビワペッカリー( H )、オセロット( C )、ジャガランディ( C ) などです。プロングホーンとオオアリクイを除くすべての動物は、人間の捕食者とともに進化したアジアの祖先の子孫である。[73]プロングホーンは(チーターに次いで)2番目に速い陸上哺乳類であり、それがハンターの目を逃れるのに役立ったのかもしれない。過剰殺害という文脈で説明するのがより難しいのは、バイソンの生き残りである。なぜなら、これらの動物が最初に北米に現れたのは24万年未満であり、したがってかなりの期間、地理的に人間の捕食者から隔離されていたからである。[74] [75] [76]古代のバイソンが現生のバイソンに進化したため、[ 77] [78]更新世の終わりに大陸全体でバイソンが絶滅することはなかった(ただし、この属は多くの地域で地域的に絶滅した)。したがって、完新世および近年までバイソンが生き残ったことは、過剰殺害のシナリオと矛盾している。[要出典]更新世の終わり、人類が初めて北米大陸に到達したとき、これらの大型動物は 20 万年以上にわたって人間の集中的な狩猟から地理的に隔離されていました。この膨大な地質学的時間を考慮すると、バイソンは北米原産の大型哺乳類と同じくらい無知であったことはほぼ間違いありません。[[要出典]
北米における絶滅の波と関連づけられている文化は、大型動物を殺すために槍投げ器を使用していたと考えられているクローヴィス人(参照) に関連する古アメリカ文化です。「先史時代の過剰殺戮仮説」に対する主な批判は、当時の人類の人口が少なすぎたり、地理的に十分に広まっていなかったりしたため、生態学的に重大な影響を与えることはできなかったというものです。この批判は、絶滅を説明する気候変動シナリオが自動的にデフォルトで優先されることを意味するものではありませんが、気候変動の議論の弱点が過剰殺戮を支持するものとみなされる可能性があるのと同じです。両方の要因が何らかの形で組み合わさっている可能性はあり得ますが、気候変動によってすでにストレスを受けている人口では、過剰殺戮の方が大規模な絶滅を達成するのがはるかに簡単でしょう。
南アメリカ
.jpg/500px-Macrauchenia_(reconstruction).jpg)

)_(Aenocyon_dirus).jpg/500px-The_American_Museum_journal_(c1900-(1918))_(Aenocyon_dirus).jpg)





南米は大陸の中で最も深刻な損失を被り、大型動物の約83%が絶滅した。[10]これらの絶滅は、約15,000年前に南米に現生人類が到着した後のことである。様々な著者によって、人為的要因と気候的要因の両方が絶滅の要因であるとされている。[79]一部の大型動物は歴史的に、放射性炭素年代に基づいて完新世初期まで生き延びたと示唆されてきたが、これは汚染による年代測定の誤りの結果である可能性がある。[80]これらの絶滅は、南極の寒冷逆転(北半球のヤンガードリアスよりも早く、より軽度の寒冷期)の終焉と、南米全土に広まった魚の尾のような尖頭器官の出現と一致している。魚の尾のような尖頭器は大型動物の狩猟に使われたと考えられているが、人間が絶滅した大型動物を搾取したという直接的な証拠はまれである。 [79]ただし、大型動物の搾取は多くの場所で記録されている。[80] [81]魚の尾のような尖頭器は大型動物の絶滅後に急速に姿を消し、より小型の獲物を狩るのに適した他の形状のものに取って代わられた。[79]一部の著者は「壊れたジグザグ」モデルを提唱しており、人間による狩猟と気候変動により大型動物が好む開けた生息地が減少したことが、南米における大型動物の絶滅の相乗要因であったとしている。[82]
- 有蹄類
- 霊長類
- 食肉類
- フェリフォルミア
- いくつかのネコ科の種。
- サーベルタイガー(スミロドン)属[93]
- スミロドン・ファタリス(南アメリカ北西部)
- スミロドン・ポピュレーター(南アメリカ東部および南部)
- パタゴニアジャガー(Panthera onca mesembrina )(一部の研究者は、これらの遺骨は実際にはアメリカライオンのものであると示唆している[94])
- サーベルタイガー(スミロドン)属[93]
- いくつかのネコ科の種。
- イヌ目
- イヌ科
- ダイアウルフ ( Aenocyon dirus )
- ネーリングのオオカミ ( Canis nehringi )
- プロトキオン[95]
- 更新世ブッシュドッグ ( Speothos pacivorus )
- クマ科
- 南米のコガネグマ(Arctotherium属)
- アルクトテリウム・ボナイレンセ
- アルクトテリウム・タリイエンセ
- アルクトテリウム・ウィンゲイ
- 南米のコガネグマ(Arctotherium属)
- イヌ科
- フェリフォルミア
- げっ歯類
- コウモリ
- オオ吸血コウモリ ( Desmodus draculae )
- 長鼻類(ゾウとその近縁種)
- 異節族
- 残りの地上ナマケモノ属
- 残りのGlyptodontinae属すべての種。
- ドエディクロス[96] [97] [98] [99]
- グリプトドン/クラミドテリウム[100]
- ヘテログリプトドン[101]
- ホプロフォロス
- ロマフォラス
- ネオスクレロカリプタス
- ネウリュルス[100] [102]
- パノクトゥス
- パラパノクトゥス[100] [103]
- プラクシャプルス
- スクレロカリプタス
- いくつかのDasypodidae属。
- 美しいアルマジロ ( Dasypus bellus )
- ユータトゥス
- パキアルマテリウム
- プロパオプス[38] [86]
- Pampatheriidae科のすべての種。
- 鳥類
- カラカリナエ亜科
の各種
- ベネズエラのカラカラ (カラカラ メジャー) [107]
- シーモアのカラカラ (カラカラ セイモウリ) [108]
- ペルーのカラカラ (ミルバゴ・ブロドコルビ) [109]
- ニシキヘビ科の各種
- 各種のTadorninae属
- プシロプテルス(小型の恐怖鳥の化石は後期更新世に遡るが、 [110] [111]議論の余地がある) [112]
- カラカリナエ亜科
の各種
- 爬虫類
- クロックス&ゲイターズ
- テッチュディン
- Chelonoidis lutzae(アルゼンチン)
- ペルトセファルス・マチュリン[113]
サフル(オーストラリア・ニューギニア)と太平洋




確実に年代測定された大型動物の骨の堆積物が不足しているため、特定の地域での大型動物の絶滅のタイムラインを構築することが困難であり、大型動物の種がいつ、どのように絶滅したかについての研究者の間で分裂が生じています。[114] [115]
オーストラリアの大型動物の絶滅に関しては、少なくとも 3 つの仮説があります。
- 彼らはオーストラリア先住民が大陸に到着した際に絶滅した。
- 自然の気候変動により絶滅したと考えられています。
.png/500px-Marsupial_lion_(Thylacoleo_carnifex).png)
この理論は、大型動物が4万年前まで生き残っていたという証拠、つまりホモ・サピエンスが初めてオーストラリアに上陸してから実に3万年も経った後まで生き残っていたという証拠、つまり2つのグループが長きにわたって共存していたという証拠に基づいています。これらの動物が当時存在していたという証拠は、化石記録と海洋堆積物から得られています。まず、オーストラリア南西部のインド洋で掘削された堆積物コアは、植物食哺乳類の糞で生き延びたスポロルミエラと呼ばれる菌類の存在を示しています。4万5000年前より前の堆積物にこれらの胞子が豊富に存在していたことは、その時点まで南西オーストラリアの地形に多くの大型哺乳類が存在していたことを示しています。堆積物データはまた、大型動物の個体数が数千年以内、つまり4万5000年前頃に激減したことを示しており、急速な絶滅イベントがあったことを示唆しています。[116]さらに、オーストラリア北部で最も新しい大型動物の遺跡であるサウスウォーカークリークで発見された化石は、少なくとも16種の大型動物が4万年前までそこに生息していたことを示しています。さらに、 4万年前にホモサピエンスがサウスウォーカークリークに生息していたという確固たる証拠はないため、これらの大型動物の絶滅は人間が原因ではないと言えます。しかし、4万年前にサウスウォータークリークで大きな環境悪化の証拠があり、それが絶滅を引き起こした可能性があります。これらの変化には、火災の増加、草原の減少、淡水の喪失が含まれます。[117]当時のオーストラリア全土で同様の環境悪化が見られ、気候変動の議論をさらに強化しています。当時のオーストラリアの気候は、降水量の減少により地形が全体的に乾燥し、その結果、淡水の利用可能性が低下し、干ばつ状態が増えたと表現するのが最も適切です。全体として、これは植生の変化、火災の増加、草原の全体的な減少、そしてすでに不足している淡水をめぐる競争の激化につながりました。[118]これらの環境変化はオーストラリアの大型動物にとって対処するには大きすぎることが判明し、大型動物種の90%が絶滅しました。
- 一部の科学者が共有している3番目の仮説は、人間の影響と自然の気候変動がオーストラリアの大型動物の絶滅につながったというものです。オーストラリアの約75%は半乾燥または乾燥地帯であるため、大型動物種が人間と同じ淡水資源を使用していたことは理にかなっています。この競争により、大型動物の狩猟が増えた可能性があります。[119]さらに、ホモサピエンスは火農法[要説明]を使用して、通行不能[要説明]な土地を焼き払いました。これにより、すでに消滅しつつあった草地がさらに減少しました。草食性の大型動物の主要な食物成分である植物が含まれていました。これについては科学的なコンセンサスはありませんが、ホモサピエンスと自然の気候変動が複合的に影響を及ぼしたと考えられます。全体として、人間が原因であることを示す証拠は多数ありますが、オーストラリアの大型動物絶滅の原因として気候変動を完全に排除することで、全体像を把握することはできません。 45,000年前のオーストラリアの気候変動により生態系が不安定になり、人間による狩猟や火を使った農業に対して特に脆弱になりました。これがオーストラリアの大型動物の絶滅につながったと考えられます。
いくつかの研究は、気候変動がオーストラリアの更新世に大型動物相の絶滅を引き起こしたという証拠を示しています。ある研究者グループは、オーストラリア南東部のカディスプリングスで発見された化石の歯を分析しました。酸素同位体を分析することで乾燥度を測定し、炭素同位体と歯の微細摩耗組織分析を分析することで、大型動物相の食事と植生を評価しました。中期更新世の間、南東部オーストラリアはC4植物を食べる動物相を含む草食動物が主流でした。後期更新世までに、C4植物の食事成分はかなり減少しました。この変化は、食事制限を引き起こした可能性のあるますます乾燥した状況によって引き起こされた可能性があります。卵殻とウォンバットの歯の他の同位体分析も、45 Ka以降のC4植生の衰退を示しています。このC4植生の衰退は、乾燥度の上昇と一致しています。後期更新世におけるオーストラリア南東部のますます乾燥した気候は、大型動物にストレスを与え、その減少に寄与した可能性がある。[120]



サフル(かつてオーストラリア大陸とニューギニアで構成されていた)では、世界の他の地域よりも早く、突然の大規模な絶滅の波が起こった。[121] [122] [123] [124]ほとんどの証拠は、紀元前63,000年頃の人類の到着から20,000年間を示していますが、[125]正確な日付の範囲については科学的な議論が続いています。[126]太平洋の残りの地域(ニューカレドニアやオセアニアなどの他のオーストラリア諸島)では、いくつかの点ではるかに後ではあるものの、固有種の動物相も、後期更新世から前期完新世に人類が到着するとすぐに絶滅するのが一般的でした。
- 有袋類
- ディプロトドン科の様々な種
- パロルケステス(「有袋類のバク」)
- ボンバチ科の様々な種
- Lasiorhinus angustidens (巨大ウォンバット)
- ファスコロヌス(巨大ウォンバット)
- ラマサイア・マグナ(巨大ウォンバット)
- Vombatus hacketti(ハケットウォンバット)
- ワレンジャ・ウェイクフィールディ(ドワーフ・ウォンバット)
- セドファスコロミス(巨大ウォンバット)
- コアラ(巨大コアラ)
- 有袋類ライオン( Thylacoleo carnifex)
- ボルンガブーディー(巨大ポトルー)
- カンガルー科のさまざまな種(カンガルー、ワラビーなど)
- サルコフィラス(タスマニアデビル)
のさまざまな形態
- サルコフィラス・ラニアリウス(現生種より25%大きいが、現生のタスマニアデビルとは別種であるかどうかは不明[131])
- サルコフィラス・ムルナエンシス
- 単孔類:卵を産む哺乳類。
- 鳥類
- コヒクイドリ ( Casuarius lydekkeri )
- ゲニオルニス(体高2メートル(6.6フィート)のドロモルニス類)
- オオツチハシ( Progura gallinacea )
- クリプトジプス ラセルトサス
- ダイナトエトゥス・ガファエ
- いくつかのPhoenicopteridae属。
- Xenorhynchopsis属(オーストラリアフラミンゴ) [132]
- ゼノリンクシス・マイナー
- ゼノリンクシス・ティビアリス
- 爬虫類
原因
研究の歴史
大型動物の絶滅は、19世紀にはすでに一部の科学者によって独自の現象として認識されていました。[133] [134]
[南アメリカ] の変わり果てた状態を振り返ると、深い驚きを感じずにはいられません。かつてはアフリカ南部のように巨大な怪物が群がっていたに違いありませんが、今ではバク、グアナコ、アルマジロ、カピバラしか見られません。先祖の種族に比べれば、単なる小動物です... 彼らがいなくなってから、この国の自然に大きな物理的変化は起きていません。では、これほど多くの生き物を絶滅させたものは何なのでしょうか?
— チャールズ・ダーウィン『ビーグル号航海記』(1834年)
したがって、私たちは今、地球の歴史においてまったく例外的な時期にいるのは明らかです。私たちは動物学的に貧しい世界に住んでおり、最近になって最も巨大で、最も獰猛で、最も奇妙な形態の動物がすべて姿を消しました。そして、それらがいなくなった今、私たちにとって世界ははるかに良いものになっていることは間違いありません。しかし、これほど多くの大型哺乳類が、一箇所だけではなく、地球の陸地の半分以上で突然死滅したことは、確かに驚くべき事実であり、十分に論じられていない事実です。この大きな変化には何らかの物理的原因があったに違いないと考えざるを得ません。そして、それは地球の表面の大部分にほぼ同時に作用する原因であり、少なくとも第三紀に関する限り、例外的な性質のものでした。
— アルフレッド・ラッセル・ウォレス『動物の地理的分布、現存動物と絶滅動物の関係の研究による地球表面の過去の変化の解明』(1876年)
この問題に関する議論は20世紀に広まり、特に1960年代にポール・シュルツ・マーティンが「過剰狩猟仮説」を提唱して以降、さらに広まった。20世紀末までに、この問題に関して2つの「陣営」の研究者が出現し、1つは気候変動を支持し、もう1つは絶滅の主な原因として人間の狩猟を支持した。[134]
狩猟
The hunting hypothesis suggests that humans hunted megaherbivores to extinction, which in turn caused the extinction of carnivores and scavengers which had preyed upon those animals.[135][136][137] This hypothesis holds Pleistocene humans responsible for the megafaunal extinction. One variant, known as blitzkrieg, portrays this process as relatively quick. Some of the direct evidence for this includes: fossils of some megafauna found in conjunction with human remains, embedded arrows and tool cut marks found in megafaunal bones, and European cave paintings that depict such hunting. Biogeographical evidence is also suggestive: the areas of the world where humans evolved currently have more of their Pleistocene megafaunal diversity (the elephants and rhinos of Asia and Africa) compared to other areas such as Australia, the Americas, Madagascar and New Zealand without the earliest humans. The overkill hypothesis, a variant of the hunting hypothesis, was proposed in 1966 by Paul S. Martin,[138] Professor of Geosciences Emeritus at the Desert Laboratory of the University of Arizona.[139]


Circumstantially, the close correlation in time between the appearance of humans in an area and extinction there provides weight for this scenario.[140][9][3] Radiocarbon dating has supported the plausibility of this correlation being reflective of causation.[141] The megafaunal extinctions covered a vast period of time and highly variable climatic situations. The earliest extinctions in Australia were complete approximately 50,000 BP, well before the Last Glacial Maximum and before rises in temperature. The most recent extinction in New Zealand was complete no earlier than 500 BP and during a period of cooling. In between these extremes megafaunal extinctions have occurred progressively in such places as North America, South America and Madagascar with no climatic commonality. The only common factor that can be ascertained is the arrival of humans.[142][143] This phenomenon appears even within regions. The mammal extinction wave in Australia about 50,000 years ago coincides not with known climatic changes, but with the arrival of humans. In addition, large mammal species like the giant kangaroo Protemnodon appear to have succumbed sooner on the Australian mainland than on Tasmania, which was colonised by humans a few thousand years later.[144][145] A study published in 2015 supported the hypothesis further by running several thousand scenarios that correlated the time windows in which each species is known to have become extinct with the arrival of humans on different continents or islands. This was compared against climate reconstructions for the last 90,000 years. The researchers found correlations of human spread and species extinction indicating that the human impact was the main cause of the extinction, while climate change exacerbated the frequency of extinctions. The study, however, found an apparently low extinction rate in the fossil record of mainland Asia.[146][147] A 2020 study published in Science Advances found that human population size and/or specific human activities, not climate change, caused rapidly rising global mammal extinction rates during the past 126,000 years. Around 96% of all mammalian extinctions over this time period are attributable to human impacts. According to Tobias Andermann, lead author of the study, "these extinctions did not happen continuously and at constant pace. Instead, bursts of extinctions are detected across different continents at times when humans first reached them. More recently, the magnitude of human driven extinctions has picked up the pace again, this time on a global scale."[148][149] On a related note, the population declines of still extant megafauna during the Pleistocene have also been shown to correlate with human expansion rather than climate change.[16]
The extinction's extreme bias towards larger animals further supports a relationship with human activity rather than climate change.[150] There is evidence that the average size of mammalian fauna declined over the course of the Quaternary,[151] a phenomenon that was likely linked to disproportionate hunting of large animals by humans.[5]
Extinction through human hunting has been supported by archaeological finds of mammoths with projectile points embedded in their skeletons, by observations of modern naive animals allowing hunters to approach easily[152][153][154] and by computer models by Mosimann and Martin,[155] and Whittington and Dyke,[156] and most recently by Alroy.[157]

Major objections have been raised regarding the hunting hypothesis. Notable among them is the sparsity of evidence of human hunting of megafauna.[158][159][160] There is no archeological evidence that in North America megafauna other than mammoths, mastodons, gomphotheres and bison were hunted, despite the fact that, for example, camels and horses are very frequently reported in fossil history.[161] Overkill proponents, however, say this is due to the fast extinction process in North America and the low probability of animals with signs of butchery to be preserved.[162] The majority of North American taxa have too sparse a fossil record to accurately assess the frequency of human hunting of them.[10] A study by Surovell and Grund concluded "archaeological sites dating to the time of the coexistence of humans and extinct fauna are rare. Those that preserve bone are considerably more rare, and of those, only a very few show unambiguous evidence of human hunting of any type of prey whatsoever."[163] Eugene S. Hunn points out that the birthrate in hunter-gatherer societies is generally too low, that too much effort is involved in the bringing down of a large animal by a hunting party, and that in order for hunter-gatherers to have brought about the extinction of megafauna simply by hunting them to death, an extraordinary amount of meat would have had to have been wasted.[164]
Second-order predation


The Second-Order Predation Hypothesis says that as humans entered the New World they continued their policy of killing predators, which had been successful in the Old World but because they were more efficient and because the fauna, both herbivores and carnivores, were more naive, they killed off enough carnivores to upset the ecological balance of the continent, causing overpopulation, environmental exhaustion, and environmental collapse. The hypothesis accounts for changes in animal, plant, and human populations.
The scenario is as follows:
- After the arrival of H. sapiens in the New World, existing predators must share the prey populations with this new predator. Because of this competition, populations of original, or first-order, predators cannot find enough food; they are in direct competition with humans.
- Second-order predation begins as humans begin to kill predators.
- Prey populations are no longer well controlled by predation. Killing of nonhuman predators by H. sapiens reduces their numbers to a point where these predators no longer regulate the size of the prey populations.
- Lack of regulation by first-order predators triggers boom-and-bust cycles in prey populations. Prey populations expand and consequently overgraze and over-browse the land. Soon the environment is no longer able to support them. As a result, many herbivores starve. Species that rely on the slowest recruiting food become extinct, followed by species that cannot extract the maximum benefit from every bit of their food.
- Boom-bust cycles in herbivore populations change the nature of the vegetative environment, with consequent climatic impacts on relative humidity and continentality. Through overgrazing and overbrowsing, mixed parkland becomes grassland, and climatic continentality increases.
The second-order predation hypothesis has been supported by a computer model, the Pleistocene extinction model (PEM), which, using the same assumptions and values for all variables (herbivore population, herbivore recruitment rates, food needed per human, herbivore hunting rates, etc.) other than those for hunting of predators. It compares the overkill hypothesis (predator hunting = 0) with second-order predation (predator hunting varied between 0.01 and 0.05 for different runs). The findings are that second-order predation is more consistent with extinction than is overkill[165][166] (results graph at left). The Pleistocene extinction model is the only test of multiple hypotheses and is the only model to specifically test combination hypotheses by artificially introducing sufficient climate change to cause extinction. When overkill and climate change are combined they balance each other out. Climate change reduces the number of plants, overkill removes animals, therefore fewer plants are eaten. Second-order predation combined with climate change exacerbates the effect of climate change.[167] (results graph at right). The second-order predation hypothesis is further supported by the observation above that there was a massive increase in bison populations.[168]
However, this hypothesis has been criticised on the grounds that the multispecies model produces a mass extinction through indirect competition between herbivore species: small species with high reproductive rates subsidize predation on large species with low reproductive rates.[157] All prey species are lumped in the Pleistocene extinction model. Also, the control of population sizes by predators is not fully supported by observations of modern ecosystems.[169] The hypothesis further assumes decreases in vegetation due to climate change, but deglaciation doubled the habitable area of North America. Any vegetational changes that did occur failed to cause almost any extinctions of small vertebrates, and they are more narrowly distributed on average, which detractors cite as evidence against the hypothesis.
Competition for water
In southeastern Australia, the scarcity of water during the interval in which humans arrived in Australia suggests that human competition with megafauna for precious water sources may have played a role in the extinction of the latter.[119]
Landscape alteration
One consequence of the colonisation by humans of lands previously uninhabited by them may have been the introduction of new fire regimes because of extensive fire use by humans.[7] There is evidence that anthropogenic fire use had major impacts on the local environments in both Australia[6] and North America.[170]
Climate change
At the end of the 19th and beginning of the 20th centuries, when scientists first realized that there had been glacial and interglacial ages, and that they were somehow associated with the prevalence or disappearance of certain animals, they surmised that the termination of the Pleistocene ice age might be an explanation for the extinctions.
The most obvious change associated with the termination of an ice age is the increase in temperature. Between 15,000 BP and 10,000 BP, a 6 °C increase in global mean annual temperatures occurred. This was generally thought to be the cause of the extinctions. According to this hypothesis, a temperature increase sufficient to melt the Wisconsin ice sheet could have placed enough thermal stress on cold-adapted mammals to cause them to die. Their heavy fur, which helps conserve body heat in the glacial cold, might have prevented the dumping of excess heat, causing the mammals to die of heat exhaustion. Large mammals, with their reduced surface area-to-volume ratio, would have fared worse than small mammals. A study covering the past 56,000 years indicates that rapid warming events with temperature changes of up to 16 °C (29 °F) had an important impact on the extinction of megafauna. Ancient DNA and radiocarbon data indicates that local genetic populations were replaced by others within the same species or by others within the same genus. Survival of populations was dependent on the existence of refugia and long distance dispersals, which may have been disrupted by human hunters.[171]
Other scientists have proposed that increasingly extreme weather—hotter summers and colder winters—referred to as "continentality", or related changes in rainfall caused the extinctions. It has been shown that vegetation changed from mixed woodland-parkland to separate prairie and woodland.[172][173][174] This may have affected the kinds of food available. Shorter growing seasons may have caused the extinction of large herbivores and the dwarfing of many others. In this case, as observed, bison and other large ruminants would have fared better than horses, elephants and other monogastrics, because ruminants are able to extract more nutrition from limited quantities of high-fiber food and better able to deal with anti-herbivory toxins.[175][176][177] So, in general, when vegetation becomes more specialized, herbivores with less diet flexibility may be less able to find the mix of vegetation they need to sustain life and reproduce, within a given area. Increased continentality resulted in reduced and less predictable rainfall limiting the availability of plants necessary for energy and nutrition.[178][179][180] It has been suggested that this change in rainfall restricted the amount of time favorable for reproduction.[181][182] This could disproportionately harm large animals, since they have longer, more inflexible mating periods, and so may have produced young at unfavorable seasons (i.e., when sufficient food, water, or shelter was unavailable because of shifts in the growing season). In contrast, small mammals, with their shorter life cycles, shorter reproductive cycles, and shorter gestation periods, could have adjusted to the increased unpredictability of the climate, both as individuals and as species which allowed them to synchronize their reproductive efforts with conditions favorable for offspring survival. If so, smaller mammals would have lost fewer offspring and would have been better able to repeat the reproductive effort when circumstances once more favored offspring survival.[183] A study looking at the environmental conditions across Europe, Siberia and the Americas from 25,000 to 10,000 YBP found that prolonged warming events leading to deglaciation and maximum rainfall occurred just prior to the transformation of the rangelands that supported megaherbivores into widespread wetlands that supported herbivore-resistant plants. The study proposes that moisture-driven environmental change led to the megafaunal extinctions and that Africa's trans-equatorial position allowed rangeland to continue to exist between the deserts and the central forests, therefore fewer megafauna species became extinct there.[171]
Evidence in Southeast Asia, in contrast to Europe, Australia, and the Americas, suggests that climate change and an increasing sea level were significant factors in the extinction of several herbivorous species. Alterations in vegetation growth and new access routes for early humans and mammals to previously isolated, localized ecosystems were detrimental to select groups of fauna.[184]
Some evidence from Europe also suggests climatic changes were responsible for extinctions there, as the individuals extinctions tended to occur during times of environmental change and did not correlate particularly well with human migrations.[2]
In Australia, some studies have suggested that extinctions of megafauna began before the peopling of the continent, favouring climate change as the driver.[185]
In Beringia, megafauna may have gone extinct because of particularly intense paludification and because the land connection between Eurasia and North America flooded before the Cordilleran Ice Sheet retreated far enough to reopen the corridor between Beringia and the remainder of North America.[186] Woolly mammoths became extirpated from Beringia because of climatic factors, although human activity also played a synergistic role in their decline.[187] In North America, a Radiocarbon-dated Event-Count (REC) modelling study found that megafaunal declines in North America correlated with climatic changes instead of human population expansion.[188]
In the North American Great Lakes region, the population declines of mastodons and mammoths have been found to correlate with climatic fluctuations during the Younger Dryas rather than human activity.[189]
In the Argentine Pampas, the flooding of vast swathes of the once much larger Pampas grasslands may have played a role in the extinctions of its megafaunal assemblages.[8]
Critics object that since there were multiple glacial advances and withdrawals in the evolutionary history of many of the megafauna, it is rather implausible that only after the last glacial maximum would there be such extinctions. Proponents of climate change as the extinction event's cause like David J. Meltzer suggest that the last deglaciation may have been markedly different from previous ones.[190] Also, one study suggests that the Pleistocene megafaunal composition may have differed markedly from that of earlier interglacials, making the Pleistocene populations particularly vulnerable to changes in their environment.[191]
Studies propose that the annual mean temperature of the current interglacial that we have seen for the last 10,000 years is no higher than that of previous interglacials, yet most of the same large mammals survived similar temperature increases.[192][193][194] In addition, numerous species such as mammoths on Wrangel Island and St. Paul Island survived in human-free refugia despite changes in climate.[195] This would not be expected if climate change were responsible (unless their maritime climates offered some protection against climate change not afforded to coastal populations on the mainland). Under normal ecological assumptions island populations should be more vulnerable to extinction due to climate change because of small populations and an inability to migrate to more favorable climes.[citation needed]
Critics have also identified a number of problems with the continentality hypotheses. Megaherbivores have prospered at other times of continental climate. For example, megaherbivores thrived in Pleistocene Siberia, which had and has a more continental climate than Pleistocene or modern (post-Pleistocene, interglacial) North America.[196][197][198] The animals that became extinct actually should have prospered during the shift from mixed woodland-parkland to prairie, because their primary food source, grass, was increasing rather than decreasing.[199][198][200] Although the vegetation did become more spatially specialized, the amount of prairie and grass available increased, which would have been good for horses and for mammoths, and yet they became extinct. This criticism ignores the increased abundance and broad geographic extent of Pleistocene bison at the end of the Pleistocene, which would have increased competition for these resources in a manner not seen in any earlier interglacials.[191] Although horses became extinct in the New World, they were successfully reintroduced by the Spanish in the 16th century—into a modern post-Pleistocene, interglacial climate. Today there are feral horses still living in those same environments. They find a sufficient mix of food to avoid toxins, they extract enough nutrition from forage to reproduce effectively and the timing of their gestation is not an issue. Of course, this criticism ignores the obvious fact that present-day horses are not competing for resources with ground sloths, mammoths, mastodons, camels, llamas, and bison. Similarly, mammoths survived the Pleistocene Holocene transition on isolated, uninhabited islands in the Mediterranean Sea until 4,000 to 7,000 years ago,[201] as well as on Wrangel Island in the Siberian Arctic.[202] Additionally, large mammals should have been able to migrate, permanently or seasonally, if they found the temperature too extreme, the breeding season too short, or the rainfall too sparse or unpredictable.[203] Seasons vary geographically. By migrating away from the equator, herbivores could have found areas with growing seasons more favorable for finding food and breeding successfully. Modern-day African elephants migrate during periods of drought to places where there is apt to be water.[204] Large animals also store more fat in their bodies than do medium-sized animals and this should have allowed them to compensate for extreme seasonal fluctuations in food availability.[205]
Some evidence weighs against climate change as a valid hypothesis as applied to Australia. It has been shown that the prevailing climate at the time of extinction (40,000–50,000 BP) was similar to that of today, and that the extinct animals were strongly adapted to an arid climate. The evidence indicates that all of the extinctions took place in the same short time period, which was the time when humans entered the landscape. The main mechanism for extinction was probably fire (started by humans) in a then much less fire-adapted landscape. Isotopic evidence shows sudden changes in the diet of surviving species, which could correspond to the stress they experienced before extinction.[206][207][208]
Some evidence obtained from analysis of the tusks of mastodons from the American Great Lakes region appears inconsistent with the climate change hypothesis. Over a span of several thousand years prior to their extinction in the area, the mastodons show a trend of declining age at maturation. This is the opposite of what one would expect if they were experiencing stresses from deteriorating environmental conditions, but is consistent with a reduction in intraspecific competition that would result from a population being reduced by human hunting.[209]
It may be observed that neither the overkill nor the climate change hypotheses can fully explain events: browsers, mixed feeders and non-ruminant grazer species suffered most, while relatively more ruminant grazers survived.[210] However, a broader variation of the overkill hypothesis may predict this, because changes in vegetation wrought by either Second Order Predation (see below)[167][211] or anthropogenic fire preferentially selects against browse species.[citation needed]
Disease
The hyperdisease hypothesis, as advanced by Ross D. E. MacFee and Preston A. Marx, attributes the extinction of large mammals during the late Pleistocene to indirect effects of the newly arrived aboriginal humans.[212][213][214] In more recent times, disease has driven many vulnerable species to extinction; the introduction of avian malaria and avipoxvirus, for example, has greatly decreased the populations of the endemic birds of Hawaii, with some going extinct.[215] The hyperdisease hypothesis proposes that humans or animals traveling with them (e.g., chickens or domestic dogs) introduced one or more highly virulent diseases into vulnerable populations of native mammals, eventually causing extinctions. The extinction was biased toward larger-sized species because smaller species have greater resilience because of their life history traits (e.g., shorter gestation time, greater population sizes, etc.). Humans are thought to be the cause because other earlier immigrations of mammals into North America from Eurasia did not cause extinctions.[212] A similar suggestion is that pathogens were transmitted by the expanding humans via the domesticated dogs they brought with them.[216] A related theory proposes that a highly contagious prion disease similar to chronic wasting disease or scrapie that was capable of infecting a large number of species was the culprit. Animals weakened by this "superprion" would also have easily become reservoirs of viral and bacterial diseases as they succumbed to neurological degeneration from the prion, causing a cascade of different diseases to spread among various mammal species. This theory could potentially explain the prevalence of heterozygosity at codon 129 of the prion protein gene in humans, which has been speculated to be the result of natural selection against homozygous genotypes that were more susceptible to prion disease and thus potentially a tell-tale of a major prion pandemic that affected humans of or younger than reproductive age far in the past and disproportionately killed before they could reproduce those with homozygous genotypes at codon 129.[217]
If a disease was indeed responsible for the end-Pleistocene extinctions, then there are several criteria it must satisfy (see Table 7.3 in MacPhee & Marx 1997). First, the pathogen must have a stable carrier state in a reservoir species. That is, it must be able to sustain itself in the environment when there are no susceptible hosts available to infect. Second, the pathogen must have a high infection rate, such that it is able to infect virtually all individuals of all ages and sexes encountered. Third, it must be extremely lethal, with a mortality rate of c. 50–75%. Finally, it must have the ability to infect multiple host species without posing a serious threat to humans. Humans may be infected, but the disease must not be highly lethal or able to cause an epidemic.[citation needed]
As with other hypotheses, a number of counterarguments to the hyperdisease hypothesis have been put forth. Generally speaking, disease has to be very virulent to kill off all the individuals in a genus or species. Even such a virulent disease as West Nile fever is unlikely to have caused extinction.[218] The disease would need to be implausibly selective while being simultaneously implausibly broad. Such a disease needs to be capable of killing off wolves such as Canis dirus or goats such as Oreamnos harringtoni while leaving other very similar species (Canis lupus and Oreamnos americanus, respectively) unaffected. It would need to be capable of killing off flightless birds while leaving closely related flighted species unaffected. Yet while remaining sufficiently selective to afflict only individual species within genera it must be capable of fatally infecting across such clades as birds, marsupials, placentals, testudines, and crocodilians. No disease with such a broad scope of fatal infectivity is known, much less one that remains simultaneously incapable of infecting numerous closely related species within those disparate clades. On the other hand, this objection does not account for the possibility of a variety of different diseases being introduced around the same era.[citation needed] Numerous species including wolves, mammoths, camelids, and horses had emigrated continually between Asia and North America over the past 100,000 years. For the disease hypothesis to be applicable there it would require that the population remain immunologically naive despite this constant transmission of genetic and pathogenic material.[citation needed] The dog-specific hypothesis in particular cannot account for several major extinction events, notably the Americas (for reasons already covered) and Australia. Dogs did not arrive in Australia until approximately 35,000 years after the first humans arrived there, and approximately 30,000 years after the Australian megafaunal extinction was complete.[citation needed]
Extraterrestrial impact
An extraterrestrial impact, which has occasionally been proposed as a cause of the Younger Dryas,[219] has been suggested by some authors as a potential cause of the extinction of North America's megafauna due to the temporal proximity between a proposed date for such an impact and the following megafaunal extinctions.[220][4] However, the Younger Dryas impact hypothesis lacks widespread support among scholars due to various inconsistencies in the hypothesis,[221][222] and has been comprehensively refuted.[223]
Geomagnetic field weakening
Around 41,500 years ago, the Earth's magnetic field weakened in an event known as the Laschamp event. This weakening may have caused increased flux of UV-B radiation and has been suggested by a few authors as a cause of megafaunal extinctions in the Late Quaternary.[224] The full effects of such events on the biosphere are poorly understood, however these explanations have been criticized as they do not account for the population bottlenecks seen in many megafaunal species and nor is there evidence for extreme radio-isotopic changes during the event. Considering these factors, causation is unlikely.[225][226]
Effects
The extinction of the megafauna has been argued by some authors to be disappearance of the mammoth steppe rather than the other way around. Alaska now has low nutrient soil unable to support bison, mammoths, and horses. R. Dale Guthrie has claimed this as a cause of the extinction of the megafauna there; however, he may be interpreting it backwards. The loss of large herbivores to break up the permafrost allows the cold soils that are unable to support large herbivores today. Today, in the arctic, where trucks have broken the permafrost grasses and diverse flora and fauna can be supported.[227][228] In addition, Chapin (Chapin 1980) showed that simply adding fertilizer to the soil in Alaska could make grasses grow again like they did in the era of the mammoth steppe. Possibly, the extinction of the megafauna and the corresponding loss of dung is what led to low nutrient levels in modern-day soil and therefore is why the landscape can no longer support megafauna.
However, more recent authors have viewed it as more likely that the collapse of the mammoth steppe was driven by climatic warming, which in turn impacted the megafauna, rather than the other way around.[229]
Megafauna play a significant role in the lateral transport of mineral nutrients in an ecosystem, tending to translocate them from areas of high to those of lower abundance. They do so by their movement between the time they consume the nutrient and the time they release it through elimination (or, to a much lesser extent, through decomposition after death).[230] In South America's Amazon Basin, it is estimated that such lateral diffusion was reduced over 98% following the megafaunal extinctions that occurred roughly 12,500 years ago.[231][232] Given that phosphorus availability is thought to limit productivity in much of the region, the decrease in its transport from the western part of the basin and from floodplains (both of which derive their supply from the uplift of the Andes) to other areas is thought to have significantly impacted the region's ecology, and the effects may not yet have reached their limits.[232] The extinction of the mammoths allowed grasslands they had maintained through grazing habits to become birch forests.[233] The new forest and the resulting forest fires may have induced climate change.[233] Such disappearances might be the result of the proliferation of modern humans.[234][235]
Large populations of megaherbivores have the potential to contribute greatly to the atmospheric concentration of methane, which is an important greenhouse gas. Modern ruminant herbivores produce methane as a byproduct of foregut fermentation in digestion, and release it through belching or flatulence. Today, around 20% of annual methane emissions come from livestock methane release. In the Mesozoic, it has been estimated that sauropods could have emitted 520 million tons of methane to the atmosphere annually,[236] contributing to the warmer climate of the time (up to 10 °C warmer than at present).[236][237] This large emission follows from the enormous estimated biomass of sauropods, and because methane production of individual herbivores is believed to be almost proportional to their mass.[236]
Recent studies have indicated that the extinction of megafaunal herbivores may have caused a reduction in atmospheric methane. One study examined the methane emissions from the bison that occupied the Great Plains of North America before contact with European settlers. The study estimated that the removal of the bison caused a decrease of as much as 2.2 million tons per year.[238] Another study examined the change in the methane concentration in the atmosphere at the end of the Pleistocene epoch after the extinction of megafauna in the Americas. After early humans migrated to the Americas about 13,000 BP, their hunting and other associated ecological impacts led to the extinction of many megafaunal species there. Calculations suggest that this extinction decreased methane production by about 9.6 million tons per year. This suggests that the absence of megafaunal methane emissions may have contributed to the abrupt climatic cooling at the onset of the Younger Dryas. The decrease in atmospheric methane that occurred at that time, as recorded in ice cores, was 2–4 times more rapid than any other decrease in the last half million years, suggesting that an unusual mechanism was at work.[239]
The extermination of megafauna left many niches vacant, which has been cited as an explanation for the vulnerability and fragility of many ecosystems to destruction in the later Holocene extinction. The comparative lack of megafauna in modern ecosystems has reduced high-order interactions among surviving species, reducing ecological complexity.[240] This depauperate, post-megafaunal ecological state has been associated with diminished ecological resilience to stressors.[241] Many extant species of plants have adaptations that were advantageous in the presence of megafauna but are now useless in their absence.[242] The demise of megafaunal ecosystem engineers in the Arctic that maintained open grassland environments has been highly detrimental to shorebirds of the genus Numenius.[243]
Relationship to later extinctions
There is no general agreement on where the Quaternary extinction event ends, and the Holocene, or anthropogenic, extinction begins, or if they should be considered separate events at all.[244][245] Some authors have argued that the activities of earlier archaic humans have also resulted in extinctions, though the evidence for this is equivocal.[246]
This hypothesis is supported by rapid megafaunal extinction following recent human colonisation in Australia, New Zealand and Madagascar,[247] in a similar way that any large, adaptable predator moving into a new ecosystem would. In many cases, it is suggested even minimal hunting pressure was enough to wipe out large fauna, particularly on geographically isolated islands.[248][249] Only during the most recent parts of the extinction have plants also suffered large losses.[250]
See also
- Australian megafauna – Large animals in Australia, past and present era
- Holocene extinction – Ongoing extinction event caused by human activity
- Late Quaternary prehistoric birds – Extinct bird species
- List of Ice Age species preserved as permafrost mummies
- List of quaternary mammalian fauna of China
- Megafauna – Large animals
- Pleistocene rewilding – Ecological practice
- Toba catastrophe theory – Volcanic supereruption 74,000 years ago in Indonesia
References
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External links
Hyperdisease hypothesis
- MacFee RD, Marx PA (1998). "Lightning Strikes Twice: Blitzkrieg, Hyperdisease, and Global Explanations of the Late Quaternary Catastrophic Extinctions". American Museum of Natural History. Archived from the original on 3 September 2011.
- J.H. Brown. "Was a hyperdisease responsible?" (PDF). Archived from the original (PDF) on 27 May 2006.
Second-order predation
- Elin Whitney-Smith. "Quaternary.Net". Archived from the original on 3 December 2020. Retrieved 9 August 2008.
Other links
- "Ice Age Bay Area". Archived from the original on 26 December 2008.
- "The Extinct Late Pleistocene Mammals of North America". PBS.
- Peter Tyson. "End of the Big Beasts". PBS. Archived from the original on 21 December 2012. Retrieved 3 September 2017.
- S. Kathleen Lyons, Felisa A. Smith, James H. Brown (2004). "Of mice, mastodons and men: human-mediated extinctions on four continents" (PDF). Evolutionary Ecology Research. 6: 339–358.
- "Return to the Ice Age: The La Brea Exploration Guide". Archived from the original on 12 August 2011.

