Abstract
Large, actively swimming suspension feeders evolved several times in Earth’s history, arising independently from groups as diverse as sharks, rays and stem teleost fishes1, and in mysticete whales2. However, animals occupying this niche have not been identified from the early Palaeozoic era. Anomalocarids, a group of stem arthropods that were the largest nektonic animals of the Cambrian and Ordovician periods, are generally thought to have been apex predators3,4,5. Here we describe new material from Tamisiocaris borealis6, an anomalocarid from the Early Cambrian (Series 2) Sirius Passet Fauna of North Greenland, and propose that its frontal appendage is specialized for suspension feeding. The appendage bears long, slender and equally spaced ventral spines furnished with dense rows of long and fine auxiliary spines. This suggests that T. borealis was a microphagous suspension feeder, using its appendages for sweep-net capture of food items down to 0.5 mm, within the size range of mesozooplankton such as copepods. Our observations demonstrate that large, nektonic suspension feeders first evolved during the Cambrian explosion, as part of an adaptive radiation of anomalocarids. The presence of nektonic suspension feeders in the Early Cambrian, together with evidence for a diverse pelagic community containing phytoplankton7,8 and mesozooplankton7,9,10, indicate the existence of a complex pelagic ecosystem11 supported by high primary productivity and nutrient flux12,13. Cambrian pelagic ecosystems seem to have been more modern than previously believed.
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Acknowledgements
Our expeditions to North Greenland were financed by Geocenter Denmark, the Agouron Institute and the Carlsberg Foundation. We are grateful for discussions with members of the Bristol Palaeobiology group, E. Sperling, C. Hull, M. Matz and M. Friedman. M.S. was supported by the Carlsberg Foundation. We thank POLOG for logistic support. A. T. Nielsen and M. P. Smith assisted in the field in 2009. S. Powell assisted with figures. S. L. Jakobsen and A. T. Nielsen facilitated work and curation of the collected material at the Statens Naturhistoriske Museum, Copenhagen.
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J.V., M.S. and N.R.L. designed, analysed and performed research. D.A.T.H. obtained funding for the fieldwork. J.V., M.S., N.R.L. and D.A.T.H. wrote and discussed the paper.
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Extended data figures and tables
Extended Data Figure 1 T. borealis MGUH 30500, frontal appendage.
a, Part, photographed in low-angle lighting coated with MgO. b, Camera lucida drawing with spines indicated (s1–s15). Bs, spines broken at the base. c, Detail of spine preserving auxiliary spines in relief (arrowed).
Extended Data Figure 2 T. borealis MGUH 30500, frontal appendage.
a, Part, photographed submerged in water and with high-angle illumination. b, Counterpart, displaying articulating membranes across the appendage, as indicated by their relatively lower reflectivity. c, Detail of b, and the articulating membranes (Am) and articles (Art) along the mid-section of the appendage. d, Detail of broken spine in b, displaying auxiliary spines.
Extended Data Figure 3 T. borealis MGUH 30501 frontal appendage with well-preserved auxiliary spines.
a, Part. b, Detail of auxiliary spines in a. c, Schematic drawing of MGUH 30501, from a combination of part and counterpart. d, Counterpart. e, Detail of d showing regular arrangement of auxiliary spines.
Extended Data Figure 4 MGUH 30502 frontal appendages and head shield assemblage, lateral view.
a, Part. b, Camera lucida drawing of the part indicating the head shield (Hs), left frontal appendage (Lfa) and right frontal appendage (Rfa). Partially superimposed on the specimen is the arthropod Buenaspis (Ba). c, Detail of distal section of frontal appendages in counterpart. d, Detail of head shield.
Extended Data Figure 5 Modern crustacean suspension feeders.
a, The Northern krill, Meganyctiphanes norvegica (image reproduced with permission from Wikipedia/Ø. Paulsen). Insert shows a reconstruction of the thoracic region of the krill, Euphausia suberba, reproduced from ref. 41 with permission of Verlag J. Neuman-Neudamm. b, Proximal elements of the thoracopods in E. suberba (image reproduced with permission from U. Kils). c, Distal elements of the thoracopods in E. suberba (image reproduced with permission from U. Kils). d, The filter basket in an undetermined mysid (image reproduced with permission from Wikipedia/U. Kils). e, Thoracopod from the cirripede Darwiniella angularis (image reproduced with permission from ref. 42).
Extended Data Figure 6 Schematic drawings of different anomalocarid frontal appendages.
a, T. borealis. b, A. briggsi. c, A. canadensis. d, A. cf. saron, NIGP 154565. e, Amplectobelua symbrachiata. f, A. stephenensis. g, Hurdia victoria. h, Stanleycaris hirpex.
Extended Data Figure 7 A schematic overview of some of the known components the Early Cambrian pelagic food web.
At the base of the food chain were phytoplankton in the form of acritarchs and probably other forms with no apparent fossil record. Diverse mesozooplankton were present as copepod and branchiopod-like crustaceans feeding on phytoplankton, along with vetulicolians, which exhibit a morphology suggesting suspension feeding similar to basal chordates. Larger pelagic predators such as chaetognaths, larger arthropods and potentially also ctenophores preyed upon the mesozooplankton. Tamisiocaris would similarly have fed on the mesozooplankton. The presence of a large nektonic suspension feeder suggests a high abundance of primary producers and mesozooplankton. Other anomalocarids, such as Anomalocaris and Amplectobelua were present as some of the macrophagous apex predators at this time.
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Supplementary Information
Taxonomy and Systematics: This file contains information on the cladistics analysis, the characters and taxa used, as well as our suggested revised higher taxonomy. (PDF 573 kb)
Supplementary Information
This Nexus file contains the character codings by taxon used in the cladistics analyses. (TXT 14 kb)
An animation of the frontal appendages of Tamisiocaris borealis based on the single articulated assemblage and in relation to a radiodont mouth apparatus.
This video shows the ventral view and serves to illustrate schematically the range of motions inferred by T. borealis. (MOV 10846 kb)
An animation of the frontal appendages of Tamisiocaris borealis based on the single articulated assemblage and in relation to a radiodont mouth apparatus.
This video shows the lateral view and serves to illustrate schematically the range of motions inferred performed by T. borealis. (MOV 14305 kb)
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Vinther, J., Stein, M., Longrich, N. et al. A suspension-feeding anomalocarid from the Early Cambrian. Nature 507, 496–499 (2014). https://doi.org/10.1038/nature13010
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DOI: https://doi.org/10.1038/nature13010
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