志留紀—泥盆紀陸地革命

志留紀—泥盆紀陸地革命英語Silurian-Devonian Terrestrial Revolution),也稱泥盆紀植物爆發Devonian Plant Explosion,簡稱DePE[1]泥盆紀大爆發Devonian explosion),是顯生宙古生代中期有胚植物(特別是維管植物)和真菌開始在陸地上出現快速定殖多樣化輻射的時期,發生在距今4.28至3.59億年前的志留紀泥盆紀期間[2][3][4],其中最關鍵的階段發生在兩紀之交[5]。這個時期的植物相主要是通過孢子繁殖的蕨類植物,與現今主流的被子植物無論在外形、結構和繁殖策略上都有很大不同,而且大多在後來的泥盆紀後期滅絕事件石炭紀雨林崩潰事件二疊紀末大滅絕滅絕事件消亡[6][7],其生態位大多被後出現的種子植物裸子植物和被子植物)所取代。

泥盆紀沼澤的藝術想像畫,源自《The World Before the Deluge》(1872)一書

陸生自營生物的出現和不斷繁盛對地球表面的自然環境產生了很大的衝擊,特別是通過不斷光合作用改變了地球大氣層的成分,使得氧氣濃度首次超過了10%。陸生植物日漸強健的根系也不斷侵蝕岩石層,所產生的細砂與各種有機物碎屑混合後形成了土壤,最終在地殼上形成了可以保水並且富含腐殖質土壤層,為之後動物的登陸演化創造了棲息條件[8]。植物對地表風化作用的加速,也顯著影響了地球的水圈(特別是淡水),使得大量礦物質被從岩石中釋放並隨著地表徑流進入水體,最終也逐漸提高了海洋鹽度。植物葉部蒸騰作用產生的水蒸氣也影響了水循環,加上固碳移除二氧化碳降低了溫室效應,使得全球氣候受到影響,部分造成了晚古生代大冰期

陸地生命

最先登陸並定殖陸地的複雜生物其實是真菌,其中一些與綠藻藍綠菌發生共生演化成了早期的地衣,另一些(特別是球囊菌)甚至發展成了直徑達1公尺(3.3英尺)、高度則可達8.8公尺(29英尺)的原杉菌Prototaxites),是當時陸地上最大的生物,直到石炭紀早期才在尺寸上被植物超過。植物很可能沿著並利用真菌的菌絲擴展根系形成了共生的菌根,並且在植物根際之間的土壤內形成了可以交換養分菌根網絡[9]。最早由真菌與維管植物共生產生的菌根化石出現於泥盆紀早期[10]

最早可以在陸地環境存活的綠色質體生物植物綠藻的共同祖先)很可能在奧陶紀就已出現[11]。最早登陸的有胚植物非維管苔蘚植物,出現在4.7億年前的奧陶紀中後期[12][13][14],其中在波羅的大陸達瑞威爾期就已經存在了頗具多樣性的植物群[15]汞同位素調查(∆199Hg和∆200Hg)發現陸生植物在志留紀早期就已經擴散至陸地上很大區域[16]。在侯默期冰期早古生代大冰期英語Early Paleozoic Icehouse的一部分)結束後出現了一個間冰期暖化,三縫孢的植物也首次出現了大規模的多樣化事件,但之後在盧德福德期中期又再次因冰期出現大規模海退,在引發勞階滅絕事件的同時也讓出大量新暴露的陸地供植物和藍綠菌菌毯定殖,使得多孢植物開始繁盛[17]並在之後普里道利世的溫暖期開始進一步多樣化[18]。最早的維管植物化石以孢子體的形態出現在文洛克世[19],而最早的石松門物種——巴氏石松Baragwanathia)出現在隨後的羅德洛世[20][21]孢粉學證據表明志留紀的陸生植物一致性很強,而不向現代植物那樣會因為區域不同而呈現物種多樣性[22]。志留紀的植物多樣化得到了瑞亞克洋中數個快速變化的小型火山島相助,其環境多變性使得演化得以加速[23],但當時的植物尺寸很小,其中一個物種(Tichavekia grandis)達到13公分(5.1英寸)的高度已經算作巨型植物[24]

泥盆紀的地表則出現了廣泛綠化[25],許多現代維管植物的演化支都出現在這個時期。真葉植物基群出現在泥盆紀早期的化石中[26],而石松也首次出現了演化輻射[11]。和志留紀一樣,泥盆紀早期的植物群落無論地處哪裡都十分相似[27],只有極少數具有特有性.[28]。真葉植物的多樣性在泥盆紀中期繼續增加[29],高度超過8公尺(26英尺)的真正意義上的樹林也首次出現[30],最早的森林化石出現在艾菲爾期[31],最古老的樹木是枝蕨綱[32]。泥盆紀的樹沼森林主要充斥著巨型木賊真蕨類羊齒植物和可以長到40公尺(130英尺)高的鱗木科石松[6],而種子蕨前裸子植物也在這段時期興盛[33][34],其中古蕨屬發展出了複雜的根系系統,可以應對乾燥環境[35]。因為蕨類植物和前裸子植物的迅速輻射,植物在泥盆紀晚期的多樣化最為迅速[36],而枝蕨綱繼續成為森林生態系統的主力.[32]。在泥盆紀末期,真正意義上的種子植物開始出現[37]。與現代植物外形相似的泥盆紀植物是真蕨類),雖然其中許多物種被懷疑是附生植物。真正的裸子植物(銀杏蘇鐵)則是在石炭紀才真正出現[6]

各植物之間為了爭奪單位面積內總量有限的日光照射、土壤養分和生長空間,其表型的多樣性在志留紀和泥盆紀期間增加的幅度不亞於動物在寒武紀大爆發中的程度[38]。這期間維管植物因為演化出了木質素更能縱向生長,使得高聳茂密的樹冠開始成為常態,並在之後的石炭紀形成了廣袤的煤炭森林沼澤植被在地表上產生的遮掩也為各類陸生動物(主要是六足類蛛形類多足類節肢動物真肺類腹足綱軟體動物四足類脊椎動物寡毛類環節動物)提供了庇護環境,同時創造了植食這一種新的覓食策略。植物和動物之間的互動使其共同演化出了各種反捕適應共生關係[39],其中一些新演化出的裸子植物甚至開始依賴動物(主要是昆蟲)協助其授粉和傳播種子

對大氣、氣候和土壤的影響

根系更深的維管植物對土壤大氣海洋中的氧氣含量都造成了深遠影響,因此誕生了用來解釋生物地質形態變化的「泥盆紀植物假說」(Devonian Plant Hypothesis)[8]。泥盆紀的陸生植物擴張改變了土壤成分,並增加了矽酸鹽的風化[40][41]。因為陸地植物所受的日光照射更充足,光合固碳使得大氣中的二氧化碳含量從6300 ppmv降至2100 ppmv。雖然植被可以明顯減少了陸地的日光反照率,但當時分解木質食碎屑動物微生物尚未演化成熟,這使得大量碳元素被轉換為截存在淤泥河岩層中而不是回歸碳循環,因此大大降低了溫室效應[42]。這加上當時岡瓦納大陸構造抬升暴露出的玄武岩也會吸收大量二氧化碳[43],引發了泥盆紀晚期開始的晚古生代大冰期[44][45][46]。但學術界也有另外的假說認為陸地植物的演化其實會減少矽酸鹽風化[47]

泥盆紀植物興盛的另一個結果是大氣層中的氧氣含量劇增,使其在氧氣地質歷史上首次上升超過大氣成分的10%(現今水平的一半),是地球第三次大規模的氧化事件[42](前兩次分別是太古宙末期因藍綠菌產氧引發的大氧化事件、以及元古宙後期因質體藻類繁盛導致的新元古代氧化事件)。伴隨氧氣一同劇增的是野火的頻率[48],而地球的氧氣濃度在普里道利世首次達到了可以產生林火的水平,在化石證據中留下了最早的木炭痕跡[49],但泥盆紀早期和中期的大部分時期大氣層的含氧量還不足以引發頻繁的火災[50]。到了法門期晚期,大規模野火已經是常見現象[51][52]

陸生植物的繁盛使得內陸水體中的細砂沉積物淤泥)總量劇增,這些沉積物在沖積平原的積累造成了河流曲流交織的複雜度,並形成了大量濕地池塘湖泊以及經常被洪澇河岸帶林地,這使得各類節肢動物(包括水生和陸生)、淡水軟體動物(包括淡水螺貝類)、淡水魚食蟲四足動物都得到了更豐富的棲息條件[53]

泥盆紀的植物爆發所造成的風化加速在全球範圍造成了淡水水體和海洋的富養化也加速了沉積物的循環,很可能因此導致赤潮造成海洋低氧現象也引發了泥盆紀末大滅絕[54][55][56]。植物根系對岩石的侵蝕釋放出了大量礦物質(特別是磷酸鹽),而這些礦物質和土壤中的腐殖質降水大量沖刷到地表水體中後會造成養分污染,很可能會引發大範圍的藻華,從而使得當地水體缺氧[57]。這種水體缺氧對水生動物——特別是體型較大的自游動物(比如當時處於霸權生態位的盾皮魚)——尤其致命,其造成的極端選擇壓力也使得淡水生態系統中的一部分四足形類肉鰭魚因為愈加依賴用與魚鰾同源的原始呼吸空氣而向著陸生方向演化,最終在之後的石炭紀成為了四足動物並在二疊紀演化出了徹底擺脫水生的羊膜動物

陸地植物物質在沼澤中沉積的增加也可以解釋泥盆紀岩層煤礦沼氣石油的產生[6]

另見

參考

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