植物与昆虫是当今陆地生态系统的重要组成部分，两者之间的相互关系为认识陆地生态系统结构和功能提供独一无二的信息。古生代是植物和昆虫起源和早期演化的关键地质时期，然而由于缺乏对该时期化石材料的系统研究，植物和昆虫之间的相互关系及其协同演化过程还存在许多未解之谜。对二叠纪晚期（约2.53亿年前）植物茎干中甲虫蛀孔的详细解剖学研究，证实当时蛀木甲虫利用真菌实现个体发育过程中食性的转变，而且蛀木甲虫可能存在初步的社会分工，并生活在复杂的生态网络中；进一步研究显示晚二叠世蛀木甲虫或对松柏类植物引起严重的虫灾。当前研究表明，渗矿化植物化石标本在探索地质历史时期动植物相互作用方面具有巨大研究潜力，可为深时陆地生态系统的重建和演化提供宝贵信息。
Plants and insects are the most diverse groups among macroscopic organisms on land today; their interactions play a pivotal role in the comprehension of the structure and the function of terrestrial ecosystems. The Paleozoic Era is a critical time-interval during the evolutionary history of both plants and insects. However, because of the lack of systematic investigations of the fossil records, the relationships between plants and insects and their co-evolution are still shrouded in mystery to date. Based on the systematic study of beetle's borings preserved in the late Permian (some 253 million-years ago) fossil conifer wood, this paper presents the earliest-known evidence for wood-boring beetles with changed diets during their developments by utilizing fungi, and living in a complicated ecological network. Statistical analysis indicates that the wood-boring beetles are probably the cause of a significant insect plague onto conifers during the late Permian. This study shows that the permineralized plant fossils are precious materials for a better understanding of the deeptime terrestrial ecosystems.
[1] Chapin Ⅲ F S, Matson P A, Vitousek P M. Principles of terrestrial ecosystem ecology[M]. 2nd ed. New York:Springer-Verlag, 2011.
[2] Schoonhoven L M, Van Loon J J A, Dicke M. Insect-plant biology[M]. 2nd ed. Oxford:Oxford University Press, 2005.
[3] Feng Z, Su T, Yang J Y, et al. Evidence for insect-mediated skeletonization on an extant fern family from the Upper Triassic of China[J]. Geology, 2014, 42(5):407-410.
[4] Feng Z, Wang J, Liu L J. First report of oribatid mite (arthropod) borings and coprolites in Permian woods from the Helan Mountains of northern China[J]. Palaeogeography, Palaeoclimatology, Palaeoecology, 2010, 288(1/4):54-61.
[5] Falcon-Lang H J, Labandeira C, Kirk R. Herbivorous and detritivorous arthropod trace fossils associated with subhumid vegetation in the middle Pennsylvanian of southern Britain[J]. Palaios, 2015, 30:192-206.
[6] Labandeira C C, Phillips T L. Stem borings and petiole galls from Pennsylvanian tree ferns of Illinois, USA:Implications for the origin of the borer and galling functional-feeding-groups and holometabolous insects[J]. Palaeontographica Abt. A, 2002, 264:1-84.
[7] Naugolnykh S V, Ponomarenko A G. Possible traces of feeding by beetles in coniferophyte wood from the Kazanian of the Kama River Basin[J]. Paleontological Journal, 2010, 44:468-474.
[8] Tapanila L, Roberts E M. The earliest evidence of holometabolan insect pupation in conifer wood[J]. PLoS One, 2012, 7:e31668.
[9] Gullan P J, Cranston P S. The insects:An outline of entomology (5th Edition)[M]. Wiley-Blackwell, 2014.
[10] Labandeira C C. A paleobiologic perspective on plant-insect interactions[J]. Current Opinion in Plant Biology, 2013, 16:414-421.
[11] Feng Z, Wang J, Rößler R, et al. Late Permian wood-borings reveal an intricate network of ecological relationships[J]. Nature Communications, 2017, 8:556.
[12] Feng Z, Wang J, Rößler R. A unique gymnosperm from the latest Permian of China, and its ecophysiological implications[J]. Review of Palaeobotany and Palynology, 2011, 165:27-40.
[13] Feng Z. Ningxiaites specialis, a new woody gymnosperm from the uppermost Permian of China[J]. Review of Palaeobotany and Palynology, 2012, 181:34-46.
[14] Feng Z. Late Palaeozoic plants[J]. Current Biology, 2017, 27(17):R905-R909.
[15] Feng Z, Wang J, Rößler R, et al. Complete tylosis formation in a latest Permian conifer stem[J]. Annuals of Botany, 2013, 111:1075-1081.
[16] Wei H B, Feng Z, Yang J Y, et al. Specialised emission pattern of leaf traces in a late Permian (253 million-years old) conifer[J]. Scientific Reports, 2015, 5:12405.
[17] Yang J Y, Shen J J, Chen Y X, et al. The bark anatomy of Ningxiaites specialis from the Permian of China[J]. Review of Palaeobotany and Palynology, 2017, 240:11-21.
[18] Kirkendall L R, Kent D S, Raffa K F. Interactions among males, females and offspring in bark and ambrosia beetles:The significance of living in tunnels for the evolution of social behavior[M]//Chloe J C, Crespi B J. The Evolution of Social Behavior in Insects and Arachnids. Cambridge University Press, 1997:181-215.
[19] Villari C, Herms D A, Whitehill J G, et al. Progress and gaps in understanding mechanisms of ash tree resistance to emerald ash borer, a model for wood-boring insects that kill angiosperms[J]. New Phytologist, 2016, 209(1):63-79.
[20] Liebhold A M, Brockerhoff E G, Kalisz S, et al. Biological invasions in forest ecosystems[J]. Biological Invasions, 2017, 19(11):3437-3458.
[21] Taylor T N, Taylor E L, Krings M. Paleobotany:The biology and evolution of fossil plants[M]. 2nd ed. Salt Lake City:Academic Press, 2009.
[22] Feng Z, Wei H B, Wang C L, et al. Wood decay of Xenoxylon yunnanensis Feng sp. nov. from the Middle Jurassic of Yunnan Province, China[J]. Palaeogeography, Palaeoclimatology, Palaeoecology, 2015, 433:60-70.
[23] Wan M, Yang W, Liu L J, et al. Plant-arthropod and plantfungus interactions in late Permian gymnospermous woods from the Bogda Mountains, Xinjiang, northwestern China[J]. Review of Palaeobotany and Palynology, 2016, 235:120-128.
