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Surprisingly complex community discovered in the mid-Devonian fossil forest at Gilboa


The origin of trees by the mid-Devonian epoch (398–385 million years ago) signals a major change in terrestrial ecosystems with potential long-term consequences including increased weathering, drop in atmospheric CO2, modified climate, changes in sedimentation patterns and mass extinction1,2,3. However, little is known about the ecology of early forests or how changes in early terrestrial ecosystems influenced global processes. One of the most famous palaeontological records for this time is the ‘oldest fossil forest’ at Riverside Quarry, Gilboa, New York, USA, discovered in the 1920s4,5. Hundreds of large Eospermatopteris sandstone casts, now thought to represent the bases of standing cladoxylopsid trees6, were recovered from a horizon that was originally interpreted as a muddy swamp. After quarry operations ceased, relatively minor outcrops of similar fossils at nearby localities have provided limited opportunities to evaluate this pervasive view using modern methods7,8. In 2010, removal of the quarry backfill enabled reappraisal of the palaeoecology of this important site. Here we describe a 1,200 m2 map showing numerous Eospermatopteris root systems in life position within a mixed-age stand of trees. Unexpectedly, large woody rhizomes with adventitious roots and aerial branch systems identified as aneurophytalean progymnosperms run between, and probably climb into, Eospermatopteris trees. We describe the overall habit for these surprisingly large aneurophytaleans, the earliest fossil group having wood produced by a bifacial vascular cambium. The site also provides evidence for arborescence within lycopsids, extending the North American range for trees in this ecologically critical group. The rooting horizon is a dark grey sandy mudstone showing limited root penetration. Although clearly belonging to a wetland coastal plain environment9, the forest was probably limited in duration and subject to periodic disturbance. These observations provide fundamental clarification of the palaeoecology of this mixed-group early forest, with important implications for interpreting coeval assemblage data worldwide.

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Figure 1: Eospermatopteris cast, moulds and root system, Riverside Quarry, New York, USA.
Figure 2: Plan map of part of the quarry floor showing original rooting horizon.
Figure 3: Aneurophytalean rhizomes.
Figure 4: Arborescent lycopsid.

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  1. Algeo, T. J., Scheckler, S. E. & Maynard, J. B. in Plants Invade the Land: Evolutionary and Environmental Approaches (eds Gensel, P. G. & Edwards, D. ) 213–236 (Columbia Univ. Press, 2001)

  2. Davies, N. S. & Gibling, M. R. Paleozoic vegetation and the Siluro–Devonian rise of fluvial lateral accretion sets. Geology 38, 51–54 (2010)

    Article  ADS  Google Scholar 

  3. Driese, S. G. & Mora, C. I. in Plants Invade the Land: Evolutionary and Environmental Approaches (eds Gensel, P. G. & Edwards, D. ) 237–253 (Columbia Univ. Press, 2001)

  4. Goldring, W. The Upper Devonian forest of seed ferns in eastern New York. NY State Mus. Bull. 251, 50–72 (1924)

    Google Scholar 

  5. Goldring, W. The oldest known petrified forest. Sci. Mon. 24, 514–529 (1927)

    ADS  Google Scholar 

  6. Stein, W. E., Mannolini, F., Hernick, L. V., Landing, E. & Berry, C. M. Giant cladoxylopsid trees resolve the enigma of the Earth’s earliest forest stumps at Gilboa. Nature 446, 904–907 (2007)

    Article  ADS  CAS  Google Scholar 

  7. Driese, S., Mora, C. I. & Elick, J. M. Morphology and taphonomy of root and stump casts of the earliest trees (Middle to Late Devonian), Pennsylvania and New York, U.S.A. Palaios 12, 524–537 (1997)

    Article  ADS  Google Scholar 

  8. Mintz, J. S., Driese, S. G. & White, J. D. Environmental and ecological variability of Middle Devonian (Givetian) forests in Appalachian Basin paleosols, New York, United States. Palaios 25, 85–96 (2010)

    Article  ADS  Google Scholar 

  9. Greb, S. F., DiMichele, W. A. & Gastaldo, R. A. Evolution and importance of wetlands in earth history. Geol. Soc. Am. Spec. Pap. 399, 1–40 (2006)

    Google Scholar 

  10. Berry, C. M. & Fairon-Demaret, M. The architecture of Pseudosporochnus nodosus Leclercq et Banks: a Middle Devonian Cladoxylopsid from Belgium. Int. J. Plant Sci. 163, 699–713 (2002)

    Article  Google Scholar 

  11. Beck, C. B. Current status of the Progymnospermopsida. Rev. Palaeobot. Palynol. 21, 5–23 (1976)

    Article  Google Scholar 

  12. Meyer-Berthaud, B., Scheckler, S. E. & Wendt, J. Archaeopteris is the earliest known modern tree. Nature 398, 700–701 (1999)

    Article  ADS  CAS  Google Scholar 

  13. Retallack, G. J. & Huang, C. Ecology and evolution of Devonian trees in New York, USA. Palaeogeogr. Palaeoclimatol. Palaeoecol. 299, 110–128 (2011)

    Article  Google Scholar 

  14. Banks, H. P., Grierson, J. D. & Bonamo, P. M. The flora of the Catskill clastic wedge. Geol. Soc. Am. Spec. Pap. 201, 125–141 (1985)

    Google Scholar 

  15. Beck, C. B. & Wight, D. C. in Origin and Evolution of Gymnosperms (ed. Beck, C.B. ) 1–84. Columbia Univ. Press, New York, 1988)

  16. Dannenhoffer, J. M., Stein, W. & Bonamo, P. M. The primary body of Rellimia thomsonii: integrated perspective based on organically connected specimens. Int. J. Plant Sci. 168, 491–506 (2007)

    Article  Google Scholar 

  17. Hammond, S. E. & Berry, C. M. A new species of Tetraxylopteris (Aneurophytales) from the Devonian of Venezuela. Bot. J. Linn. Soc. 148, 275–303 (2005)

    Article  Google Scholar 

  18. Pigg, K. B. Evolution of Isoetalean lycopsids. Ann. Mo. Bot. Gard. 79, 589–612 (1992)

    Article  Google Scholar 

  19. Gensel, P. G. & Berry, C. M. Early lycophyte evolution. Am. Fern J. 91, 74–98 (2001)

    Article  Google Scholar 

  20. Grierson, J. D. & Banks, H. P. Lycopods of the Devonian of New York State. Palaeontogr. Am. 4, 217–295 (1963)

    Google Scholar 

  21. Berry, C. M. & Edwards, D. A new species of the lycopsid Gilboaphyton Arnold from the Devonian of Venezuela and New York State with a revision of the closely related genus Archaeosigillaria Kidston. Rev. Palaeobot. Palynol. 96, 47–70 (1997)

    Article  Google Scholar 

  22. White, D. A remarkable fossil tree trunk from the middle Devonic of New York. NY State Mus. Bull. 107, 328–340 (1907)

    Google Scholar 

  23. Miller, M. F. & Woodrow, D. L. in Dynamic Stratigraphy and Depositional Environments of the Hamilton Group (Middle Devonian) in New York State, Part II (eds Landing, E. & Brett, C.E. ) 153–177 (State Education Department, 1991)

  24. Bridge, J. S. The geometry, flow patterns and sedimentary processes of Devonian rivers and coasts, New York and Pennsylvania, USA. Geol. Soc. Lond. Spec. Publ. 180, 85–108 (2000)

    Article  ADS  Google Scholar 

  25. Berry, C. M. & Fairon-Demaret, M. in Plants Invade the Land: Evolutionary and Environmental Approaches (eds Gensel, P. G. & Edwards, D.) 120–139 (Columbia University Press, 2001)

  26. Bridge, J. S. & Willis, B. J. Marine transgressions and regressions recorded in Middle Devonian shore-zone deposits of the Catskill clastic wedge. Geol. Soc. Am. Bull. 106, 1440–1458 (1994)

    Article  ADS  Google Scholar 

  27. Baird, G. C. & Brett, C. E. Late Givetian (Taghanic) bioevents in New York State: new discoveries and questions. Bull. Geosci. 83, 357–375 (2008)

    Article  Google Scholar 

  28. Wright, V. P., Taylor, K. G. & Beck, V. H. The paleohydrology of lower Cretaceous seasonal wetlands, Isle of Wight, southern England. J. Sediment. Res. 70, 619–632 (2000)

    Article  ADS  Google Scholar 

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Financial support was provided by the New York State Museum. C.M.B. acknowledges support from the UK Natural Environment Research Council (NERC) grant NE/F010699/1. Permission to work at the site by New York City Department of Environmental Protection (DEP) is acknowledged. On-site and other assistance was provided by E. Chase, P. Costa and G. Heath (DEP), as well as by P. Kollak and C. Tompkins of the Thalle Construction Company. We also thank P. Wright for advice and guidance on the major palaeosol horizon.

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Authors and Affiliations



L.V.H. and F. M. were responsible for collections and analysis, W.E.S. for mapping, C.M.B. for geological section, and W.E.S. and C.M.B. for palaeoecological interpretation. W.E.S. led the writing of the paper with substantial contributions from C.M.B.

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Correspondence to William E. Stein.

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The authors declare no competing financial interests.

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Supplementary Information

This file contains a Supplementary Discussion comprising: Survey data and site comparison; Geology, palaeoenvironmental setting and age; Use of names for the fossil remains; Criteria for recognizing Eospermatopteris root systems; Difficulties in estimating height for Eospermatopteris trees; Identity of aneurophytaleans at Gilboa; Aneurophytalean rhizome compression collected by Goldring; The arborescent lycopsid at Gilboa and Supplementary References. Supplementary Figures 1-27 with legends are also included. (PDF 10436 kb)

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Stein, W., Berry, C., Hernick, L. et al. Surprisingly complex community discovered in the mid-Devonian fossil forest at Gilboa. Nature 483, 78–81 (2012).

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