Skip to main content

Thank you for visiting You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

Giant cladoxylopsid trees resolve the enigma of the Earth’s earliest forest stumps at Gilboa


The evolution of trees of modern size growing together in forests fundamentally changed terrestrial ecosystems1,2,3. The oldest trees are often thought to be of latest Devonian age (about 380–360 Myr old) as indicated by the widespread occurrence of Archaeopteris (Progymnospermopsida)4. Late Middle Devonian fossil tree stumps, rooted and still in life position, discovered in the 1870s from Gilboa, New York5, and later named Eospermatopteris, are widely cited as evidence of the Earth’s ‘oldest forest’6,7. However, their affinities and significance have proved to be elusive because the aerial portion of the plant has been unknown until now. Here we report spectacular specimens from Schoharie County, New York, showing an intact crown belonging to the cladoxylopsid Wattieza (Pseudosporochnales)8 and its attachment to Eospermatopteris trunk and base. This evidence allows the reconstruction of a tall (at least 8 m), tree-fern-like plant with a trunk bearing large branches in longitudinal ranks. The branches were probably abscised as frond-like modules. Lower portions of the trunk show longitudinal carbonaceous strands typical of Eospermatopteris, and a flat bottom with many small anchoring roots. These specimens provide new insight into Earth’s earliest trees and forest ecosystems. The tree-fern-like morphology described here is the oldest example so far of an evolutionarily recurrent arborescent body plan within vascular plants. Given their modular construction, these plants probably produced abundant litter, indicating the potential for significant terrestrial carbon accumulation and a detritus-based arthropod fauna by the Middle Devonian period.

This is a preview of subscription content, access via your institution

Relevant articles

Open Access articles citing this article.

Access options

Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.

Figure 1: Wattieza Stockmans from South Mountain, New York, NYSM 17039.
Figure 2: Large trunk and reconstruction of plant from South Mountain, New York.
Figure 3: Trunk top and base of Wattieza Stockmans from South Mountain, New York; details of the large trunk specimen, NYSM 17040.


  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, New York, 2001)

    Google Scholar 

  2. 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, New York, 2001)

    Google Scholar 

  3. Greb, S. F., DiMichele, W. A. & Gastaldo, R. A. in Wetlands Through Time (eds DiMichele, W.A., & Greb, S.) 1–40 (Geol. Soc. Am. Special Paper no. 399, Boulder, Colorado, 2006)

    Book  Google Scholar 

  4. Meyer-Berthaud, B., Scheckler, S. E. & Wendt, J. Archaeopteris is the earliest known modern tree. Nature 938, 700–701 (1999)

    ADS  Article  Google Scholar 

  5. Dawson, J. W. On new tree ferns and other fossils from the Devonian. Q. J. Geol. Soc. Lond. 27, 269–275 (1871)

    Article  Google Scholar 

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

    ADS  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, USA. Palaios 12, 524–537 (1997)

    ADS  Article  Google Scholar 

  8. Berry, C. M. A reconsideration of Wattieza Stockmans (here attributed to Cladoxylopsida) based on a new species from the Devonian of Venezuela. Rev. Palaeobot. Palynol. 112, 125–146 (2000)

    CAS  Article  Google Scholar 

  9. Goldring, W. The Upper Devonian forest of seed ferns. NY State Mus. Bull. 521, 50–72 (1924)

    Google Scholar 

  10. Kräusel, R. & Weyland, H. Pflanzenreste aus dem Devon IX. Ein Stamm von Eospermatopteris Bau aus dem Mittledevon des Kirberges, Elberfeld. Senckenbergiana 17, 1–20 (1935)

    Google Scholar 

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

    Article  Google Scholar 

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

    Article  Google Scholar 

  13. Boyer, J. S. & Matten, L. C. Anatomy of Eospermatopteris eriana from the Upper Middle Devonian ( = Givetian) of New York. Int. Org. Palaeobot. 5, 13 (1996)

    Google Scholar 

  14. Berry, C. M. & Fairon-Demaret, M. in Plants Invade the Land: Evolutionary and Environmental Approaches (eds Gensel, P. G. & Edwards, D.) 120–158 (Columbia Univ. Press, New York, 2001)

    Google Scholar 

  15. Lemoigne, Y. & Iurina, Y. A. Xenocladia medullosina Ch. A. Arnold (1940) 1952 du Dévonien moyen du Kazakhstan (URSS). Geobios 16, 513–547 (1983)

    Article  Google Scholar 

  16. Stein, W. E. & Hueber, F. M. The anatomy of Pseudosporochnus: P. hueberi from the Devonian of New York. Rev. Palaeobot. Palynol. 60, 311–359 (1989)

    Article  Google Scholar 

  17. Berry, C. M. & Yi, W. A new plant attributed to Cladoxylopsida from the Middle Devonian of Yunnan province, China. Rev. Palaeobot. Palynol. 142, 63–78 (2006)

    Article  Google Scholar 

  18. Soria, A., Meyer-Berthaud, B. & Scheckler, S. E. Reconstructing the architecture and growth habit of Pietschia levis sp. nov. (Cladoxylopsida) from the Late Devonian of southeastern Morocco. Int. J. Plant Sci. 162, 911–926 (2001)

    Article  Google Scholar 

  19. Hilton, J., Geng, B. & Kenrick, P. A novel late Devonian (Frasnian) woody cladoxylopsid from China. Int. J. Plant Sci. 164, 793–805 (2003)

    Article  Google Scholar 

  20. Leclercq, S. & Banks, H. P. Pseudosporochnus nodosus sp. nov., a Middle Devonian plant with cladoxylalean affinities. Palaeontographica B 110, 1–34 (1962)

    Google Scholar 

  21. Berry, C. M. & Fairon-Demaret, M. A reinvestigation of the cladoxylopsid Pseudosporochnus nodosus Leclercq et Banks from the Middle Devonian of Goé, Belgium. Int. J. Plant Sci. 158, 350–372 (1997)

    Article  Google Scholar 

  22. 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 

  23. 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)

    ADS  Article  Google Scholar 

  24. Banks, H. P., Grierson, J. D. & Bonamo, P. M. in The Catskill Delta (eds Woodrow, D. L. & Sevon, W. D.) 125–141 (Geol. Soc. Am. Special Paper no. 201, Boulder, Colorado, 1985)

    Book  Google Scholar 

  25. Traverse, A. & Schuyler, A. Palynostratigraphy of the Catskill and part of the Chemung magnifacies, southern New York State, USA. Courier Forschungsinstitut Senckenberg 169, 261–274 (1994)

    Google Scholar 

  26. Soria, A. & Meyer-Berthaud, B. Tree fern growth strategy in the late Devonian cladoxylopsid species Pietzschia levis from the study of its stem and root system. Am. J. Bot. 91, 10–23 (2004)

    Article  Google Scholar 

  27. 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 

Download references


Financial support was provided by the New York State Museum.

Author Contributions L.V.H. and F.M. were responsible for initial discovery, field work and museum curation, E.L. for geological interpretation, W.E.S. and C.M.B. for palaeobotanical interpretation, F.M. for drawing the reconstruction in Fig. 2, and W.E.S. for writing the paper.

Author information

Authors and Affiliations


Corresponding authors

Correspondence to William E. Stein or Christopher M. Berry.

Ethics declarations

Competing interests

Reprints and permissions information is available at The authors declare no competing financial interests.

Supplementary information

Supplementary Information

This file contains Supplementary Discussion and Supplementary Figures S1-S6 with Legends. (PDF 12623 kb)

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Stein, W., Mannolini, F., Hernick, L. et al. Giant cladoxylopsid trees resolve the enigma of the Earth’s earliest forest stumps at Gilboa. Nature 446, 904–907 (2007).

Download citation

  • Received:

  • Accepted:

  • Issue Date:

  • DOI:

Further reading


By submitting a comment you agree to abide by our Terms and Community Guidelines. If you find something abusive or that does not comply with our terms or guidelines please flag it as inappropriate.


Quick links

Nature Briefing

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.

Get the most important science stories of the day, free in your inbox. Sign up for Nature Briefing