Fragments of the earliest land plants


The earliest fossil evidence for land plants comes from microscopic dispersed spores1,2,3. These microfossils are abundant and widely distributed in sediments, and the earliest generally accepted reports are from rocks of mid-Ordovician age (Llanvirn, 475 million years ago)4. Although distribution, morphology and ultrastructure of the spores indicate that they are derived from terrestrial plants, possibly early relatives of the bryophytes, this interpretation remains controversial5 as there is little in the way of direct evidence for the parent plants. An additional complicating factor is that there is a significant hiatus between the appearance of the first dispersed spores and fossils of relatively complete land plants (megafossils)6: spores predate the earliest megafossils (Late Silurian, 425 million year ago) by some 50 million years7. Here we report the description of spore-containing plant fragments from Ordovician rocks of Oman. These fossils provide direct evidence for the nature of the spore-producing plants. They confirm that the earliest spores developed in large numbers within sporangia, providing strong evidence that they are the fossilized remains of bona fide land plants. Furthermore, analysis of spore wall ultrastructure supports liverwort affinities.

Access options

Rent or Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.

Figure 1: Fossil plant fragments from borehole Ghaba-1, core 21.
Figure 2: Specimen CW47c.


  1. 1

    Gray, J. & Boucot, A. J. Early Silurian spore tetrads from New York: earliest New World evidence for vascular plants? Science 173, 918–921 (1971)

    ADS  CAS  Article  Google Scholar 

  2. 2

    Gray, J. The microfossil record of early land plants: advances in understanding of early terrestrialization, 1970–1984. Phil. Trans. R. Soc. Lond. B 309, 167–195 (1985)

    ADS  Article  Google Scholar 

  3. 3

    Wellman, C. H. & Gray, J. The microfossil record of early land plants. Phil. Trans. R. Soc. Lond. B 355, 717–732 (2000)

    CAS  Article  Google Scholar 

  4. 4

    Strother, P. K., Al-Hajri, S. & Traverse, A. New evidence for land plants from the lower Middle Ordovician of Saudi Arabia. Geology 24, 55–58 (1996)

    ADS  Article  Google Scholar 

  5. 5

    Banks, H. P. The oldest vascular land plants: a note of caution. Rev. Palaeobot. Palynol. 20, 13–25 (1975)

    Article  Google Scholar 

  6. 6

    Edwards, D. & Feehan, J. Records of Cooksonia-type sporangia from late Wenlock strata in Ireland. Nature 287, 41–42 (1980)

    ADS  Article  Google Scholar 

  7. 7

    Edwards, D. & Wellman, C. H. in Plants Invade the Land (eds Gensel, P. G. & Edwards, D.) 3–28 (Columbia Univ. Press, New York, 2001)

    Google Scholar 

  8. 8

    Droste, H. H. J. Stratigraphy of the Lower Palaeozoic Haima Supergroup of Oman. GeoArabia 2, 419–468 (1997)

    Google Scholar 

  9. 9

    Mohiuddin, U. . Quantitative Analysis of Ordovician Palynomorphs of Oman Thesis (Univ Sheffield)

  10. 10

    Edwards, D., Selden, P. A., Richardson, J. B. & Axe, L. Coprolites as evidence for plant–animal interaction in Siluro-Devonian terrestrial ecosystems. Nature 377, 329–331 (1995)

    ADS  CAS  Article  Google Scholar 

  11. 11

    Taylor, W. A. Studies in cryptospore ultrastructure: variability in the tetrad genus Tetrahedraletes and type material of the dyad Dyadospora murusattenuata. Rev. Palaeobot. Palynol. 119, 325–334 (2002)

    Article  Google Scholar 

  12. 12

    Pacini, E. in Microspores: Evolution and Ontogeny (eds Blackmore, S. & Knox, R. B.) 213–237 (Academic, London, 1990)

    Google Scholar 

  13. 13

    Edwards, D. New insights into early land ecosystems: a glimpse of a Lilliputian world. Rev. Palaeobot. Palynol. 90, 159–174 (1996)

    Article  Google Scholar 

  14. 14

    Richardson, J. B. in Subsurface Palynostratigraphy of Northeast Libya (eds El-Arnauti, B., Owens, B. & Thusu, B.) 89–109 (Garyounis Univ., Benghazi, Libya, 1988)

    Google Scholar 

  15. 15

    Steemans, P. Paléodiversification des spores et des cryptospores de L'Ordovician au Dévonien inferieur. Geobios 32, 341–352 (1999)

    Article  Google Scholar 

  16. 16

    Taylor, W. A. Spores in earliest land plants. Nature 373, 391–392 (1995)

    ADS  CAS  Article  Google Scholar 

  17. 17

    Edwards, D., Duckett, J. G. & Richardson, J. B. Hepatic characters in the earliest land plants. Nature 374, 635–636 (1995)

    ADS  CAS  Article  Google Scholar 

  18. 18

    Wellman, C. H., Edwards, D. & Axe, L. Ultrastructure in laevigate hilate spores from sporangia and spore masses from the Upper Silurian and Lower Devonian of the Welsh Borderland. Phil. Trans. R. Soc. Lond. B 153, 1983–2004 (1998)

    Article  Google Scholar 

  19. 19

    Wellman, C. H., Edwards, D. & Axe, L. Permanent dyads in sporangia and spore masses from the Lower Devonian of the Welsh Borderland. Bot. J. Linn. Soc. 127, 117–147 (1998)

    Article  Google Scholar 

  20. 20

    Edwards, D., Wellman, C. H. & Axe, L. Tetrads in sporangia and spore masses form the Upper Silurian and Lower Devonian of the Welsh Borderland. Bot. J. Linn. Soc. 130, 111–156 (1999)

    Article  Google Scholar 

  21. 21

    Graham, L. E. Origin of Land Plants (Wiley, New York, 1993)

    Google Scholar 

  22. 22

    Brown, R. C. & Lemmon, B. E. in Microspores: Evolution and Ontogeny (eds Blackmore, S. & Knox, R. B.) 55–94 (Academic, London, 1990)

    Google Scholar 

  23. 23

    Kenrick, P. The relationships of vascular plants. Phil. Trans. R. Soc. Lond. B 355, 847–855 (2000)

    CAS  Article  Google Scholar 

  24. 24

    Kenrick, P. & Crane, P. R. The Origin and Early Diversification of Land Plants (Smithsonian Institution, Washington, 1997)

    Google Scholar 

  25. 25

    Nickrent, D. L. et al. Multigene phylogeny of land plants with special reference to the bryophytes and the earliest land plants. Mol. Biol. Evol. 17, 1885–1895 (2000)

    CAS  Article  Google Scholar 

  26. 26

    Qiu, Y. L. & Lee, J. Transition to a land flora: a molecular phylogenetic perspective. J. Phycol. 36, 799–802 (2000)

    CAS  Article  Google Scholar 

  27. 27

    Kroken, S. B., Graham, L. E. & Cook, M. E. Occurrence and evolutionary significance of resistant cell walls in charophytes and bryophytes. Am. J. Bot. 83, 1241–1254 (1996)

    Article  Google Scholar 

Download references


This research was supported by a NERC grant. We would like to thank Petroleum Development Oman and Oman Ministry of Oil and Gas for making the material available for study and for permission to publish.

Author information



Corresponding author

Correspondence to Charles H. Wellman.

Ethics declarations

Competing interests

The authors declare that they have no competing financial interests.

Rights and permissions

Reprints and Permissions

About this article

Cite this article

Wellman, C., Osterloff, P. & Mohiuddin, U. Fragments of the earliest land plants. Nature 425, 282–285 (2003).

Download citation

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