Traces of marine nematodes from 470 million years old Early Ordovician rocks in China

in Nematology
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Cylindrical, mostly horizontal, burrows of 20-60 μm diam. and sinusoidal course, found in the middle part of the Early Ordovician (early Floian) Fenxiang Formation in the Hubei Province of China, represent the oldest record of activity by marine nematodes, preceding known nematode body fossils by 70 million years. The burrows are filled with secondarily oxidised pyrite framboids and clay mineral flakes, indicating low oxygen content in the mud and proving that the animals lined their burrows with organic matter, being bacteriovores and mud-eaters. The marine bottom environment enabling such a mode of life originated no earlier than the mid Early Cambrian (approximately 535 million years ago) owing to peristaltic bioturbation, mostly by nemathelminthans of priapulid affinities. Before the so-called ‘Agricultural Revolution’, the bottoms of shallow seas were covered with microbial mats preventing within-sediment animal life. This event imposes the lower time limit on the possible date of origin of nematodes.


International Journal of Fundamental and Applied Nematological Research



ArduiniP.PinnaG.TeruzziG. (1983). Eophasma jurassicum n. g. n. sp., a new fossil nematode of the Sinemurian of Osteno in Lombardy. Atti della Società Italiana di Scienze Naturali a del Museo Civico di Storia Naturale, Milano 124, 61-64.

BalińskiA.SunY.DzikJ. (2012). 470-Million-years-old black corals from China. Naturwissenschaften 99, 645-653.

BottingJ.P.MuirL.A.Van RoyP.BatesD.UptonC. (2012). Diverse middle Ordovician palaeoscolecidan worms from the Builth-Llandrindod inlier of central Wales. Palaeontology 55, 501-528.

BoucotA.J.PoinarG.O.Jr (2010). Predation and feeding behaviors. In: BoucotA.J.PoinarG.O. Fossil behavior compendium. Boca Raton, FL, USA, CRC Press, pp.  79-118.

Conway MorrisS. (1997). The cuticular structure of the 495-Myr-old type species of the fossil worm Palaeoscolex, P. piscatorum (?Priapulida). Zoological Journal of the Linnaean Society 119, 69-82.

CrimesT.P.AndersonM.M. (1985). Trace fossils from late Precambrian-Early Cambrian strata of southeastern Newfoundland (Canada): temporal and environmental implications. Journal of Paleontology 59, 310-343.

DzikJ. (2005). Behavioral and anatomical unity of the earliest burrowing animals and the cause of the ‘Cambrian explosion’. Paleobiology 31, 507-525.

GabbottS.E.Xiang-GuangH.NorryM.J.SiveterD.J. (2004). Preservation of Early Cambrian animals of the Chengjiang biota. Geology 32, 901-904.

GłuszekA. (1995). Invertebrate trace fossils in the continental deposits of an Upper Carboniferous coal-bearing succession, Upper Silesia, Poland. Studia Geologica Polonica 108, 171-202.

GoodayA.J.CedhagenT.KamenskayaO.E.CorneliusN. (2007). The biodiversity and biogeography of komokiaceans and other enigmatic foraminiferan-like protists in the deep Southern Ocean. Deep-Sea Research Part II 54, 1691-1719.

HaspelG.O’DonovanM.J.HartA.C. (2010). Motoneurons dedicated to either forward or backward locomotion in the nematode Caenorhabditis elegans. The Journal of Neuroscience 30, 11151-11156.

HeipC.VincxM.VrankenG. (1985). The ecology of marine nematodes. Oceanography and Marine Biology Annual Reviews 23, 399-489.

HerbertR.B.BennerS.G.PrattA.R.BlowesD.W. (1998). Surface chemistry and morphology of poorly crystalline iron sulfides precipitated in media containing sulfate-reducing bacteria. Chemical Geology 144, 87-97.

HoltermanM.van der WurffA.van den ElsenS.van MegenH.BongersT.HolovachovO.BakkerJ.HelderJ. (2006). Phylum-wide analysis of SSU rDNA reveals deep phylogenetic relationships among nematodes and accelerated evolution toward crown clade. Molecular Biology and Evolution 23, 1792-1800.

HopeW.D.MurphyD.G. (1969). Syringonornus typicus new genus, new species (Enoplida: Leptosomatidae) a marine nematode inhabiting arenaceous tubes. Proceedings of the Biological Society of Washington 82, 511-518.

JensenP. (1996). Burrows of marine nematodes as centres for microbial growth. Nematologica 42, 320-329.

KnaustD. (2010). Remarkably preserved benthic organisms and their traces from a Middle Triassic (Muschelkalk) mud flat. Lethaia 43, 344-356.

LeeD.L.BiggsW.D. (1990). Two- and three-dimensional locomotion of the nematode Nippostrongylus brasiliensis. Parasitology 101, 301-308.

MacLeanL.C.TyliszczakT.GilbertP.U.ZhouD.PrayT.J.OnstottT.C.SouthamG. (2008). A high-resolution chemical and structural study of framboidal pyrite formed within a low-temperature bacterial biofilm. Geobiology 6, 471-480.

MetzR. (1987). Sinusoidal trail formed by a recent biting midge (Family Ceratopogonidae): trace fossil implications. Journal of Paleontology 61, 312-314.

MoodleyL.ChenG.HeipC.H.R.VincxM. (2000). Vertical distribution of meiofauna in sediments from contrasting sites in the Adriatic Sea: clues to the role of abiotic versus biotic control. Ophelia 53, 203-212.

MooreR.A.LiebermanB.S. (2009). Preservation of early and Middle Cambrian soft-bodied arthropods from the Pioche Shale, Nevada, USA. Palaeogeography, Palaeoclimatology, Palaeoecology 277, 57-62.

MoussaM.T. (1970). Reviewed source: nematode fossil trails from the Green River Formation (Eocene) in the Uinta Basin, Utah. Journal of Paleontology 44, 304-307.

NehringS.JensenP.LorenzenS. (1990). Tube-dwelling nematodes: tube construction and possible ecological effects on sediment-water interfaces. Marine Ecology Progress Series 64, 123-128.

OrłowskiS. (1990). Trace fossils in the Lower Cambrian sequence in the Świętokrzyskie Mountains, Central Poland. Acta Palaeontologica Polonica 34, 211-231.

PikeJ.BernhardJ.M.MoretonS.G.ButlerI.B. (2001). Microbioirrigation of marine sediments in dysoxic environments: implications for early sediment fabric formation and diagenetic processes. Geology 29, 923-926.

PoinarG.O.Jr (2011). The evolutionary history of nematodes: as revealed in stone, amber and mummies. Nematology Monographs & Perspectives 9 (Series Editors: HuntD.J.PerryR.N.). Leiden, The Netherlands, Brill.

PoinarG.O.JrKerpH.HassH. (2008). Palaeonema phyticum gen. n., sp. n. (Nematoda: Palaeonematidae fam. n.), a Devonian nematode associated with early land plants. Nematology 10, 9-14.

RobinsonA.F.PerryR.N. (2006). Behaviour and sensory perception. In: PerryR.N.MoensM. (Eds). Plant nematology. Wallingford, UK, CABI Publishing, pp.  210-233.

SchramF.R. (1973). Pseudocoelomates and a nemertine from the Illinois Pennsylvanian. Journal of Paleontology 47, 985-989.

SchramF.R. (1979). Worms of the Mississippian Bear Gulch Limestone of central Montana, USA. Transactions of the San Diego Society of Natural History 19, 107-120.

SeilacherA. (1999). Biomat-related lifestyles in the Precambrian. Palaios 14, 86-93.

SharmaJ.BluhmB.A. (2011). Diversity of larger free-living nematodes from macrobenthos (>250 μm) in the Arctic deep-sea Canada Basin. Marine Biodiversity 41, 455-465.

StørmerL. (1963). Gigantoscorpio willsi, a new scorpion from the Lower Carboniferous of Scotland and its associated preying microorganisms. Skrifter utgitt av Det Norske Videnskaps-Akademi i Oslo I. Mat.-Naturv. Klasse Ny Serie 8, 1-171.

UchmanA.KazakauskasV.GaigalasA. (2009). Trace fossils from Late Pleistocene varved lacustrine sediments in eastern Lithuania. Palaeogeography, Palaeoclimatology, Palaeoecology 272, 199-211.

WallaceH.R. (1968). The dynamics of nematode movement. Annual Review of Phytopathology 6, 91-114.

WhittardW.F. (1953). Palaeoscolex piscatorum gen. et sp. nov., a worm from the Tremadocian of Shropshire. Quarterly Journal of the Geological Society of London 109, 125-135.

ZhanR.JinJ. (2007). Ordovician-Early Silurian (Llandovery) stratigraphy and palaeontology of the Upper Yangtze Platform, South China. Beijing, China, Science Press.


  • Location of the Tianjialing section in the Three Gorge area of Hubei Province, southern China. A: Geological map; B: Rock column of the Fenxiang Formation with the source level of the sinusoidal traces indicated (modified after Baliński et al., 2012).

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  • Sinusoidal tracks in the shale intercalation within the Fenxiang Formation collected at the Tianjialing section. A: The longest linear track crossing with another one under angle. Note that the pyritised (secondarily oxidised) infill of the burrows is preserved three-dimensionally. The rusty coloration of the rock nearby is a result of weathering; B: More typical arrangement of tracks; C: Burrow crossing sediment laminae and with vertical plane of sinusoidal undulation (arrows point to parts of the burrow covered with sediment; see also the seemingly straight burrow in B); D, E: Intensely bioturbated portion of the shale. This figure is published in colour in the online edition of this journal, which can be accessed via

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  • SEM images and EDS analysis of burrows from the Early Ordovician Fenxiang Formation, China. A: Images with Back-scatter Electron Detector of a portion of rock slab with sinusoidal burrow; B: Enlarged infill of the burrow with pyritic framboids; C: Oblique view of transverse fracture of the burrow showing its three-dimensional aspect and low degree of compaction; D-I: SEM picture and element maps of a piece of burrow; width of the burrow marked with arrows; I: EDS spot analysis spectrum revealing relative contents of elements in a framboid. Note the Fe peak, total absence of S, and presence of Si, Al, Mg, and K connected with the matrix clay minerals; specimen coated with carbon. This figure is published in colour in the online edition of this journal, which can be accessed via

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