食木蛤科Xylophagaidae的成員是一類專性棲息于深海沉木的雙殼綱生物,與其“近親”船蛆科由同一祖先演化而來。這兩科是雙殼綱“唯二”的專性木食類群,也是海洋中最主要的木材降解生物類群。但與船蛆不同,食木蛤的內(nèi)臟完全在殼瓣之內(nèi),僅水管伸出,貝殼沒有殼內(nèi)柱。與棲息于溫帶與熱帶淺海浮木和潮間帶木材內(nèi)的船蛆不同,食木蛤生活于深水沉木(最深可達7250米)和高緯度地區(qū)淺海沉木中,極少數(shù)會在風(fēng)暴沖刷上岸的沉木中被發(fā)現(xiàn)。因此,船蛆能耐受鹽度與溫度的劇烈波動,通過分泌鈣質(zhì)蛀道內(nèi)壁和用鎧板封閉孔口抵御在潮間帶暴露于空氣中時的干燥環(huán)境;而深海食木蛤不需要耐受干燥環(huán)境,因此沒有用于封閉蛀道口的鎧板,僅Xyloredos屬的兩個物種具有在蛀道末端分泌鈣質(zhì)蛀道內(nèi)壁的能力。
Xyloredo teramachii,2023年2月拖網(wǎng)采集于東海1956海區(qū)4小區(qū)370~420米深 / 作者拍攝自有標本
食木蛤能將沉木能量和養(yǎng)分輸送到深海環(huán)境,是連接陸地與深海生態(tài)系統(tǒng)的橋梁。其通過特化的帶齒脊的殼前緣將木材銼磨成碎屑,并將碎屑儲存在胃盲囊中,利用共生的γ-變形菌分解纖維素。作為深海木材降解的主要生物類群,食木蛤能在數(shù)月內(nèi)形成高密度種群。例如,Xylophaga alexisi用7個月就能在木材表面形成170孔/厘米2的鉆孔密度,Xylophaga depalmai在云杉中的密度6個月就能達(1484.5±259.8)個/分米3。一些物種在半年內(nèi)即可將木材蛀至可用手壓碎的程度。例如,Xylophaga washingtona僅用2.4個月就能讓木材被用手壓碎,而Xylonora atlantica用8~10個月即可使木材崩解,Xylophaga depalmai在6個月內(nèi)可降解51.88%的木材。
Xylonora atlantica。 PAS. 后閉殼肌痕;PR. 縮足肌痕;SR. 水管收縮肌痕 / Romano et al,2020
纖維素分解會形成硫化環(huán)境,船蛆可通過鈣質(zhì)蛀道內(nèi)壁抵御有毒的硫環(huán)境,而食木蛤則演化出多種方式,如水管的角質(zhì)層、角質(zhì)保護錐,或用黏液混合糞便填滿水管與蛀道之間的空隙,隔絕水管與環(huán)境中的硫。Xylophaga dorsalis支系成員更進一步,會在木材表面形成糞便堆積的煙囪狀結(jié)構(gòu)?!凹S煙囪”富含細菌,能誘導(dǎo)新幼蟲定殖并減弱水流,加之食木蛤的高密度集群,導(dǎo)致局部含氧量下降。因此,被Xylophaga dorsalis支系成員蛀食的木材生物多樣性普遍較低,僅有耐受低氧環(huán)境的生物如擁有血紅蛋白的Xylophaga dorsalis存活。Xylophaga dorsalis支系成員還能通過濾食輔助獲取能量,這些特征使其成為非常強大的競爭者。例如,胡安·德富卡海脊(奮進段)[Juan de Fuca Ridge (Endeavour)]的樣本最初以廣布性及高豐度高度特化于新基質(zhì)快速定殖的Xylophaga microchira為主,但在24個月后則以Xylophaga dorsalis支系的Xylophaga oregona為主。而另一項研究的同支系的Xylophaga indica則占該樣本個體總數(shù)的95.7%。【待續(xù)】
Xylophaga dorsalis。A. 帶軟體的生物體背視圖;B. 帶軟體部的生物體側(cè)視圖;C. 正視圖;D. 貝殼內(nèi)面 / Romano et al,2014
參考文獻及圖片來源
Ablett J, Brown C, Gallichan J, et al, 2019. Mollusca Types in Great Britain[EB/OL]. Amgueddfa Cymru-National Museum Wales / Natural History Museum. https://gbmolluscatypes.ac.uk [Accessed: 16 July 2025].
Amon D J, Daniel S, Farah A, et al, 2015. Burrow forms, growth rates and feeding rates of wood-boring xylophagaidae bivalves revealed by micro-computed tomography[J]. Frontiers in Marine Science, 2(10): 00010.
Gaudron S M, Haga T, Wang H, et al, 2016. Plasticity in reproduction and nutrition in wood-boring bivalves (Xylophaga atlantica) from the Mid-Atlantic Ridge[J]. Marine biology, 163(10): 213.
Haga T, Kase T, 2008. Redescription of the deep-sea wood borerNeoxylophaga teramachiitaki & habe, 1950 and its assignment to the genusXyloredo(Bivalvia: Myoida: Pholadoidea) with comments on fossil photadoidae[J]. The Veliger, 50: 107-119.
Haga T, Kase T, 2013. Progenetic dwarf males in the deep-sea wood-boring genusXylophaga(Bivalvia: Pholadoidea)[J]. Journal of Molluscan Studies, 79(1): 90-94.
Harvey R, 1996. Deep water Xylophagaidae (Pelecypoda: Pholadacea) from the North Atlantic with descriptions of three new species[J]. Journal of Conchology, 35(6): 473-481.
Jayachandran P R , Velásquez M, Jima M, 2022. A new species of wood-boring bivalve (Mollusca: Xylophagaidae) from the Eastern Arabian Sea[J]. Marine Biodiversity, 52(1): 1-9.
Judge J, 2014. Testing relationships of a new form of boring bivalve: the interesting kind[J]. Friday Harbor Laboratories, http://hdl.handle.net/1773/27289.
Knudsen J, 1961. The bathyal and abyssalXylophaga(Pholadidae, Bivalvia)[J]. Galathea Reports. 5: 163-209.
Lai K Y, 2023. A new species of Xylophaga (Bivalvia: Xylophagaidae) from Taiwan[J]. 貝類學(xué)報, (6): 101-105.
Li Y, Altamia M A, Shipway J R, et al, 2022. Contrasting modes of mitochondrial genome evolution in sister taxa of wood-eating marine bivalves (Teredinidae and Xylophagaidae)[J]. Genome Biology and Evolution, 14(6): evac089.
Li Y, Halanych K M, 2017. Comparative genomics reveal symbiont-host evolution of deep-sea tubeworms (siboglinidae, annelida) and wood-boring bivalves (xylophagaidae, mollusca)[C]. 6th International Symposium on Chemosynthesis-Based Ecosystems. Woods Hole, Mass. USA. Augustus 2017.
MacIntosh H, Voight J R, 2021. Deep-sea wood-boring bivalves (Xylophagaidae) from southeast Australia, with a new species described[J]. Molluscan Research, 41(1): 16-25.
Romano C, Nunes-Jorge A, Le Bris N, et al, 2020. Wooden stepping stones: Diversity and biogeography of deep-sea wood boring Xylophagaidae (Mollusca: Bivalvia) in the North-East Atlantic Ocean, with the description of a new genus[J]. Frontiers in Marine Science, 7: 579959.
Romano C, Voight J R, Pérez-Portela R, et al, 2014. Morphological and genetic diversity of the wood-boringXylophaga(Mollusca, Bivalvia): new species and records from deep-sea Iberian Canyons[J]. PLoS ONE, 9(7): e102887.
Santhakumaran L N, 1980. Two new species ofXylophagafrom trondheimsfjorden, Western Norway (Mollusca, Pelecypoda)[J]. Sarsia North Atlantic Marine Science, 65(3): 269-272.
Turner R D, 2002. On the subfamily Xylophagainae[J]. Bulletin of the Museum of Comparative Zoology, 157(4): 223-307.
Voight J R, 2007. Experimental deep-sea deployments reveal diverse Northeast Pacific wood-boring bivalves of Xylophagainae (Myoida: Pholadidae)[J]. Journal of Molluscan Studies,73(4): 377-391.
Voight J R, 2008. Deep-sea wood-boring bivalves ofXylophaga(Myoida: Pholadidae) on the continental shelf: a new species described[J]. Journal of the Marine Biological Association of the United Kingdom, 88(7): 1459-1464.
Voight J R, 2009. Diversity and reproduction of near-shore vs offshore wood-boring bivalves (Pholadidae: Xylophagainae) of the deep eastern Pacific ocean, with three new species[J]. Journal of Molluscan Studies, 75(2): 167-174.
Voight J R, 2024. Three new species of wood-boring bivalves (Mollusca: Xylophagaidae) from the deep Northwest Atlantic Ocean[J]. Zootaxa, 5493(3): 275-290.
Voight J R, Heck P R, Du C, et al, 2022. Competition in the deep sea: phylogeny determines destructive impact of wood-boring xylophagaids (Mollusca: Bivalvia)[J]. Marine Biodiversity, 53: 1.
Voight J R, Marshall B A, Judge J, et al, 2019. Life in wood: preliminary phylogeny of deep-sea wood-boring bivalves (Xylophagaidae), with descriptions of three new genera and one new species[J]. Journal of Molluscan Studies, 85(2): 232-243.
Voight J R, Segonzac M, 2012. At the bottom of the deep blue sea: a new wood-boring bivalve (Mollusca, Pholadidae, Xylophaga) from the Cape Verde Abyssal Plain (subtropical Atlantic)[J]. Zoosystema, 34(1): 171-180.
文中部分圖片版權(quán)來源不詳,無法與著作權(quán)人一一取得聯(lián)系,未能及時支付稿酬,在此表示由衷的歉意。有關(guān)著作權(quán)人可與我們聯(lián)系。
特別聲明:以上內(nèi)容(如有圖片或視頻亦包括在內(nèi))為自媒體平臺“網(wǎng)易號”用戶上傳并發(fā)布,本平臺僅提供信息存儲服務(wù)。
Notice: The content above (including the pictures and videos if any) is uploaded and posted by a user of NetEase Hao, which is a social media platform and only provides information storage services.