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Fruitbat's Pdf Library - General Arthropoda

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These are a few of the pdf files (and a few Microsoft Word documents) that I've accumulated in my web browsing.

MOST of these are hyperlinked to their source.

If you want one that is not hyperlinked or if the link isn't working, e-mail me at joegallo1954@gmail.com and I'll be happy to send it to you


Please note that this list will be updated continuously as I find more available resources. All of these files are freely available on the Internet so there should be no copyright issues.


Articles with author names in RED are new additions since June 27, 2017.


Phylum Arthropoda - General

Precambrian (Ediacaran) 'Arthropods'


Glaessner, M.F. (1980). Parvancorina - an arthropod from the Late Precambrian (Ediacarian) of South Australia. Ann.Naturhist.Mus. Wien, 83.


Cambrian 'Arthropods'


Cambrian 'Arthropods' - Asia/Malaysia/Pacific Islands


Fu, D.-J., X.-L. Jiang and D.-G. Shu (2011). Soft anatomy of the Early Cambrian arthropod Isoxys curvirostratus from the Chengjiang biota of South China with a discussion on the origination of great appendages. Acta Palaeontologica Polonica, 56(4).

Fu, D.-J., X. Zhang and G.E. Budd (2014). The first dorsal-eyed bivalved arthropod and its significance for early arthropod evolution. GFF, Vol.136, Number 1.

Fu, D.-J., et al. (2014). Ontogeny and dimorphism of Isoxys auritus (Arthropoda) from the Early Cambrian Chengjiang biota, South China. Gondwana Research, 25.

Hou, X. and J. Bergstrom (1997). Arthropods of the Lower Cambrian Chengjiang fauna , southwest China. Fossils & Strata, Number 45. (Thanks to oilshale for locating this one!)

Lin, J.-P., et al. (2006). A Parvancorina-like arthropod from the Cambrian of South China. Historical Biology, 18(1).

Liu, Y., et al. (2016). Three-dimensionally preserved minute larva of a great-appendage arthropod from the early Cambrian Chengjiang biota. PNAS, Early Edition.

Luo, H., et al. (2007). A New Arthropod, Guangweicaris Luo, Fu et Hu gen.nov. from the Early Cambrian Guanshan Fauna, Kunming, China. Acta Geologica Sinica, Vol.81, Number 1.

Ma, X., et al. (2015). Preservational Pathways of Corresponding Brains of a Cambrian Euarthropod. Current Biology, 25.

Ma, X., et al. (2014). An exceptionally preserved arthropod cardiovascular system from the early Cambrian. Nature Communications, 5:3560.

Peng, J., Y.-L. Zhao and H.-J. Sun (2012). Discovery and significance of Naraoia from the Quiandongian (lower Cambrian) Balang Formation, Eastern Guizhou, South China. Bulletin of Geosciences, 87(1). (Thanks to piranha for finding this one!)

Tanaka, G., et al. (2013), Chelicerate neural ground pattern in a Cambrian great appendage arthropod. Nature, Vol.502.

Wu, Y., et al. (2016). Dimorphism of Bivalved Arthropod ?Branchiocaris yunnanensis from the Early Cambrian Chengjiang Biota, South China. Acta Geologica Sinica, Vol.90, Number 3.

Zhang, X. and D. Shu (2007). Soft Anatomy of Sunellid Arthropods from the Chengjiang Lagerstätte, Lower Cambrian of Southwest China. J.Paleont., 81(6).

Zhang, X., J. Han and D. Shu (2000). A New Arthropod Pygmaclypeatus daziensis from the Early Cambrian Chengjiang Lagerstätte, South China. J.Paleont., 74(5).


Cambrian 'Arthropods' - Australia/New Zealand


Edgecombe, G.D., D.C. Garcia-Bellido and J.R. Paterson (201X). A new leanchoiliid megacheiran arthropod from the lower Cambrian Emu Bay Shale, South Australia. Acta Palaeontologica Polonica, 5X(X).

Lee, M.S.Y., et al. (2011). Modern optics in exceptionally preserved eyes of Early Cambrian arthropods from Australia. Nature, Vol.474(7353).

Paterson, J.R., et al. (2008). Early Cambrian Arthropods from the Emu Bay Shale Lagerstätte, South Australia. In: Advances in trilobite research. Rabano, I., R. Gozalo and D. Garcia-Bellido (eds.), Cuademos del Museo Geominero, Number 9. Instituto Geologico y Minero de Espana, Madrid.


Cambrian 'Arthropods' - Europe (including Greenland and Siberia)


Chulpáč, I. and V. Kordule (2002). Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic). Bulletin of the Czech Geological Survey, Vol.77, Number 3.

Dzik, J. and K. Lendzion (1988). The oldest arthropods of the East European Platform. Lethaia, Vol.21.

Eriksson, M.E., et al. (2012). Internal Soft-Tissue Anatomy of Cambrian 'Orsten' Arthropods as Revealed by Synchrotron X-Ray Tomographic Microscopy. PLoS ONE, 7(8).

Lamsdell, J.C., M. Stein and P.A. Selden (2013). Kodymirus and the case for convergence of raptorial appendages in Cambrian arthropods. Naturwissenschaften, 800.

Stein, M. (2010). A new arthropod from the Early Cambrian of North Greenland, with a 'great appendage'-like antennula. Zoological Journal of the Linnean Society, 158.

Stein, M. (2008). Evolution and taxonomy of Cambrian arthropods from Greenland and Sweden. Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology, 558.

Stein, M., et al. (2013). Arthroaspis n.gen., a common element of the Sirius Passet Lagerstătte (Cambrian, North Greenland), sheds light on trilobite ancestry. BMC Evolutionary Biology, 13:99.


Cambrian 'Arthropods' - North America


Aria, C. and J-B. Caron (2015). Cephalic and Limb Anatomy of a New Isoxyid from the Burgess Shale and the Role of "Stem Bivalved Arthropods" in the Disparity of the Frontalmost Appendage. PLoS ONE, 10(6).

Briggs, D.E.G. (1977). Bivalved Arthropods from the Cambrian Burgess Shale of British Columbia. Palaeontology, Vol.20, Part 3.

Briggs, D.E.G. and R.A. Robison (1984). Exceptionally Preserved Nontrilobite Arthropods and Anomalocaris from the Middle Cambrian of Utah. The University of Kansas Paleontological Contributions, Paper 111.

Briggs, D.E.G., et al. (2008). Middle Cambrian Arthropods from Utah. J.Paleont., 82(2).

Collette, J.H., J.W. Hagadorn and M.A. Lacelle (2010). Dead in Their Tracks - Cambrian Arthropods and Their Traces from Intertidal Sandstones of Quebec and Wisconsin. Palaios, Vol.25.

Garcia-Bellido, D.C., J. Vannier, and D. Collins (2009). Soft-part preservation in two species of the arthropod Isoxys from the middle Cambrian Burgess Shale of British Columbia, Canada. Acta Palaeontologica Polonica, 54(4).

Gould, S.J. (1991). The disparity of the Burgess Shale arthropod fauna and the limits of cladistic analysis: why we must strive to quantify morphospace. Paleobiology, 17(4).

Lin, J.-P. (2009). Function and hydrostatics in the telson of the Burgess Shale arthropod Burgessia. Biology Letters, 5.

Ortega-Hernandez, J. (2015). Homology of Head Sclerites in Burgess Shale Euarthropods. Current Biology, 25.

Stein, M. and P.A. Selden (2011). A restudy of the Burgess Shale (Cambrian) arthropod Emeraldella brocki and reassessment of its affinities. Journal of Systematic Paleontology, iFirst 2011.


General Cambrian 'Arthropods'


Hendricks, J.R. and B.S. Lieberman (2008). New Phylogenetic Insights into the Cambrian Radiation of Arachnomorph Arthropods. J. Paleont., 82(3).

Hendricks, J.R. and B.S. Liebermann (2007). Biogeography and the Cambrian radiation of arachnomorph arthropods. Memoirs of the Association of Australasian Palaeontologists, 34.

Hendricks, J.R., B.S. Lieberman and A.L. Stigall (2008). Using GIS to study palaeobiogeographic and macroevolutionary patterns in soft-bodied Cambrian arthropods. Palaeogeography, Palaeclimatology, Palaeoecology, 264.

Legg, D.A. and J. Vannier (2013). The affinities of the cosmopolitan arthropod Isoxys and its implications for the origin of arthropods. Lethaia, Vol.46.

Lin, J.-P. and D.E.G. Briggs (2010). Burgess shale-type preservation: a comparison of naraoiids (Arthropoda) from three Cambrian localities. PALAIOS, Vol.25.

Ma, X., et al. (2012). Complex brain and optic lobes in an early Cambrian arthropod. Nature, Vol.490.

Strausfeld, N.J. (2015). Palaeontology: Clearing the Heads of Cambrian Arthropods. Current Biology, 25.

Strausfeld, N.J., et al. (2016). Arthropod eyes: The early Cambrian fossil record and divergent evolution of visual systems. Arthropod Structure & Development, 45.

Vannier, J., et al. (2009). Arthropod visual predators in the early pelagic ecosystem: evidence from the Burgess Shale and Chengjiang biotas. Proc.R.Soc. B, 276.

Waloszek, D., et al. (2005). Early Cambrian arthropods-new insights into arthropod head and structural evolution. Arthropod Structure & Evolution, 34.

Zhang, X.-g. and B.R. Pratt (1994). Middle Cambrian Arthropod Embryos with Blastomeres. Science, Vol.266.

Zhu, M.-Y., et al. (2004). Direct evidence for predation on trilobites in the Cambrian. Proc.R.Soc.Lond. B (Suppl.), 271.


Ordovician 'Arthropods'


Chulpáč, I. (1999). Some problematical arthropods from the Upper Ordovician Letna Formation of Bohemia. Journal of the Czech Geological Society, 44/1-2.

Chulpáč, I. (1988). The Enigmatic Arthropod Duslia from the Ordovician of Czechoslovakia. Palaeontology, Vol.31, Part 3.

Ortega-Hernandez, J., S.J. Braddy and S. Rak (2010). Trilobite and xiphosuran affinities for putative aglaspidid arthropods Caryon and Drabovaspis, Upper Ordovician, Czech Republic. Lethaia Seminar.

Rak, S., J. Ortega-Hernandez and D.A.Legg (2013). A revision of the Late Ordovician arthropod Furca bohemica from Czech Republic. Acta Palaeontologica Polonica, 58(3). (Thanks to piranha for finding this one!)

Vannier, J., et al. (2003). Cosmopolitan arthropod zooplankton in the Ordovician seas. Palaeogeography, Palaeoclimatology, Palaeoecology, 195.


Silurian 'Arthropods'


Briggs, D.E.G., et al. (2016). Tiny individuals attached to a new Silurian arthropod suggest a unique mode of brood care. PNAS, Vol.113, Number 16.


Devonian 'Arthropods'


Dunlop, J.A. (2002). Arthropods from the Lower Devonian Severnaya Zemlya Formation of October Revolution Island (Russia). Geodiversitas, 24(2).

Lagebro, L. (2015). The Arthropod Assemblage of the Upper Devonian Strud locality and its Ecology. Ph.D. Dissertation - Uppsala Universitet. (Summary only)

Tanaka, G., D.J. Siveter and M. Williams (2013). A new bivalved arthropod from the Devonian of Japan. Paleontological Research, Vol.17, Number 3.


Carboniferous 'Arthropods'


Atkinson, T.P. (2005). Arthropod Body Fossils from the Union Chapel Mine. In: Pennsylvanian Footprints in the Black Warrior Basin of Alabama. Buta, R.J., A.K. Rindsberg, and D.C. Kopaska-Merkel (eds.), Alabama Paleontological Society Monograph Number 1.

Haug, J.T., et al. (2014). Arthropod cuticles from the upper Visean (Mississippian) of eastern Germany. Bulletin of Geosciences, 89(3).


Triassic 'Arthropods'


Schmidt, A.R., et al. (2012). Arthropods in amber from the Triassic Period. PNAS, Vol.109, Number 37.


Jurassic 'Arthropods'


Charbonnier, S., et al. (2010). Ecological Significance of the arthropod fauna from the Jurassic (Callovian) La Voulte Lagerstätte. Acta Palaeontologica Polonica, 55(1).


Miocene 'Arthropods'


Pierce, W.D. and J. Gibron (1962). Fossil Arthropods of California, 24. Some Unusual Fossil Arthropods from the Calico Mountain Nodules. Bulletin. So. Calif. Academy Sciences, Vol.61, Part 3.


Arthropod Origins and Evolution


Bergstrom, J. and X.-G. Hou (2003). Arthropod origins. Bulletin of Geosciences, Vol.78, Number 4.

Budd, G.E. (2001). Tardigrades as 'Stem-Group Arthropods': The Evidence from the Cambrian Fauna. Zool.Anz., 240(2001).
Budd, G.E. and M.J. Telford (2009). The origin and evolution of arthropods. Nature, Vol.457.

Edgecombe, G.D. and D.A. Legg (2014). Origins and Early Evolution of Arthropods. Palaeontology, 57(3).

Edgecombe, G.D. and D.A. Legg (2013). 15. The Arthropod Fossil Record. In: Arthropod Biology and Evolution - Molecules, Development, Morphology. Minelli, A. G. Boxshall and G. Fusco (eds.), Springer-Verlag.

Grosberg, R.K. (1990). Out on a limb: Arthropod Origins. Science, New Series, Vol.250, Number 4981.

Haug, J.T., et al. (2011). Sarotrocercus oblitus - Small arthropod with great impact on the understanding of arthropod evolution? Bulletin of Geosciences, 86(4).

Lozano-Fernandez, et al. (2016). A molecular palaeobiological exploration of arthropod terrestrialization. Phil.Trans.R.Soc.B, 371.

Perrier, V., M. Williams and D.J. Siveter (2015). The fossil record and palaeoenvironmental significance of marine arthropod zooplankton. Earth-Science Reviews, 146.

Pisani, D., et al. (2004). The colonization of land by animals: molecular phylogeny and divergence times among arthropods. BMC Biology, 2:1.

Schwermann, A.H., et al. (2016). Preservation of three-dimensional anatomy in phosphatized fossil arthropods enriches evolutionary inference. eLife Sciences, 5(e12129).

Williams, M., et al. (2011). Oxygen as a Driver of Early Arthropod Micro-Benthos Evolution. PLoS ONE, 6(12).

Wolfe, J.M., et al. (2016). Fossil calibrations for the arthropod Tree of Life. Earth-Science Reviews, 160.


Arthropod Anatomy/Morphology


Bartram, K.M., A.J. Jeram and P.A. Selden (1987). Arthropod cuticles in coal. Journal of the Geological Society, London, Vol.144.

Edgecombe, G.D., X. Ma and N.J. Strausfeld (2015). Unlocking the early fossil record of the arthropod central nervous system. Phil.Trans.Roy.Soc. B, 370.

Ortega-Hernandez, J. and G. Budd (2016). The nature of non-appendicular anterior paired projections in Palaeozoic total-group Euarthropoda. Arthropod Structure & Development, xxx. (Article in Press)

Ramskold, L., et al. (1996). Preservational folds simulating tergite junctions in tegopeltid and naraoiid arthropods. Lethaia, Vol.29.

Strausfeld, N.J., X. Ma and G.D. Edgecombe (2016). Fossils and the Evolution of the Arthropod Brain. Current Biology, 26.

Vannier, J., et al. (2016). Exceptional preservation of eye structure in arthropod visual hunters from the Middle Jurassic. Nature Communications, 7:10320.

Waloszek, D., et al. (2007). Evolution of cephalic feeding structures and the phylogeny of Arthropoda. Palaeogeography, Palaeoclimatology, Palaeoecology, 254.


General Arthropods


Chipman, A.D. (2015). An embryological perspective on the early arthropod fossil record. BMC Evolutionary Biology, 15:285.
Legg, D.A. (2013). The Impact of Fossils on Arthropod Phylogeny. Ph.D. Thesis - Imperial College, London.Ortega-Hernandez, J. (2014). Making sense of 'lower' and 'upper' stem-group Euarthropoda, with comments on the strict use of the name Arthropoda von Siebold, 1848. Biological Reviews, 91(1).

Palmer, D. (2010). Chapter 1. An introduction to the fossil arthropods of Great Britain. In: Fossil Arthropods of Great Britain. Jarzembowski, E.A., et al., Geological Conservation Review Series, Number 35, Joint Nature Conservation Committee, Peterborough.
Perkovsky, E.E., et al. (2007). A comparative analysis of the Baltic and Rovno amber arthropod faunas: representative samples. African Invertebrates, 48(1).
Racheboeuf, P.R., et al. (2008). The euthycarcinoid arthropods from Montceau-les-Mines, France: functional morphology and affinities. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 99.

Shear, W.A. and J. Kukalova-Peck (1990). The ecology of Paleozoic terrestrial arthropods: the fossil evidence. Can.J.Zool., Vol.68.

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Updated December 23, 2011.


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Updated March 1, 2012.


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Updated March 26, 2012.


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Updated October 23, 2012.


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Updated August 5, 2014.


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Updated August 18, 2016.


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Updated October 9, 2016.



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Updated April 20, 2017.



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