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Research Article

Low atmospheric CO2 levels during the Permo- Carboniferous glaciation inferred from fossil lycopsids

D. J. Beerling
  1. Department of Animal and Plant Sciences, University of Sheffield, Sheffield S10 2TN, United Kingdom

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PNAS October 1, 2002 99 (20) 12567-12571; https://doi.org/10.1073/pnas.202304999
D. J. Beerling
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  1. Edited by Robert A. Berner, Yale University, New Haven, CT, and approved August 6, 2002 (received for review May 21, 2002)

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    Figure 1

    Relationship between SI (fraction of epidermal cells that are stomata) and atmospheric CO2 partial pressure of different tropical populations of L. cernuum. Solid line denotes the linear regression, dotted lines indicate the ±95% confidence limits. Central America (●), South America (□), and Asia (●) (n = 21, r2 = 0.73, P < 0.01). All leaves were obtained from the herbarium of the Natural History Museum, London. SI measurements were made on cleared leaves (n = 5–8 per date) of each species collected over the past 150 years of CO2 increase. Corresponding mean global atmospheric CO2 concentrations were determined from ice core studies (13) or instrumental records (available at http://cdiac.esd.ornl.gov), after appropriate correction for altitude.

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    Figure 2

    Comparison of the carbon isotope discrimination (▵) of modern (□) and extinct (■) lycopsids. Each box delineates the 25th and 75th percentiles, errors bars show the fifth and 95th percentiles, the horizontal line represents the median value of the sample set. Δ calculated for herbarium leaf sequences of L. cernuum (Central America populations, n = 9, 1868–1995; South American populations, n = 8, 1853 – 1987; Asian populations, n = 12, 1822–1981). Changes in Δ were not correlated with historical atmospheric CO2 increase in the extant lycopsid species. Therefore, intraspecific variations reflect differences in microhabitats and interannual climate fluctuations.

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    Figure 3

    Late Palaeozoic evidence for ice (summarized from refs. 23, 25, and 27) and atmospheric CO2 concentrations reconstructed from paleosols and predicted by a geochemical model of the long-term global carbon cycle (solid line). Red boxes indicate CO2 estimates from the current study. The dashed horizontal line indicates the threshold atmospheric CO2 level above which theoretical studies predict deglaciation on Pangea and the maintenance of an ice-free state (24).

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    Table 1

    Stomatal characteristics of fossil Carboniferous (C) and Permian (P) lycopsid microphyll leaves and leaf bases

    Fossil specimenPeriodStageAge, Myr agoOrganSI*, %Atmospheric CO2 concentration (ppm) ± 95% cls
    Lepidodendron rhodianum CNamurian331–318Leaf scars4.9385  ± Embedded Image
    L. veltheimii CNamurian331–318Leaf cushions4.0392  ± Embedded Image
    Lepidophloios grangeri CNamurian331–318Leaf cushions8.1359  ± Embedded Image
    Bothrodendron minutifolium CWestphalian318–303Leaves9.4348  ± Embedded Image
    Lepidodendron aculeatum CWestphalian318–303Leaf cushions9.0351  ± Embedded Image
    L. arberi CWestphalian318–303Leaf cushions10.9336  ± Embedded Image
    L. dichotomum CWestphalian318–303Leaf cushions14.4308  ± Embedded Image
    L. feistmantelii CWestphalian318–303Leaf cushions9.2350  ± Embedded Image
    L. mannabachense CWestphalian318–303Leaf scars7.3366  ± Embedded Image
    L. peachii CWestphalian318–303Leaf scars6.9368  ± Embedded Image
    L. subdichotomum CWestphalian318–303Leaf cushions8.2358  ± Embedded Image
    Lepidophloios acadianus CWestphalian318–303Leaf cushions15.0303  ± Embedded Image
    L. acerosus CWestphalian318–303Leaf cushions7.1367  ± Embedded Image
    L. laricinus CWestphalian318–303Leaf cushions9.2350  ± Embedded Image
    L. macrolepidotus CWestphalian318–303Leaf cushions2.9401  ± Embedded Image
    Ulodendron landsbergii CWestphalian318–303Leaf cushions17.0287  ± Embedded Image
    U. landsbergii CWestphalian318–303Leaves14.0311  ± Embedded Image
    U. majus CWestphalian318–303Leaf cushions17.3284  ± Embedded Image
    U. majus CWestphalian318–303Leaves12.3325  ± Embedded Image
    Synchysidendron sp.PKazanian268Leaf cushions10.1342  ± Embedded Image
    Paralycopodites sp.PKazanian266Leaf cushions12.1326  ± Embedded Image
    Dispersed cuticlesPKazanian263Leaf cushions (?)16.6290  ± Embedded Image
    Synchysidendron baodeense PKazanian260Leaf cushions10.1343  ± Embedded Image
    Dispersed cuticlesPKazanian260Leaf cushions (?)19.6266  ± Embedded Image
    • ↵* Calculated as (SD/(SD + ED) × 100), where SD is stomatal density (mm−2), measured directly on fossil cuticles, and ED is epidermal cell density, calculated from cell dimensions for the Carboniferous plant cuticles. 

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    Table 2

    Carbon isotope characteristics of Carboniferous compression fossil lycopsid shoots

    Fossil specimenSpecimen numberOrganδpcorr, ‰Δ, ‰ci/ca, unitless
    Bothrodendraceae
     Bothrodendron minutifolium RC2872Leafy shoot−23.522.00.78
     B. minutifolium 51813Leafy shoot−24.022.50.80
     B. minutifolium 1292Leafy shoot−23.421.90.77
     B. minutifolium 1300Leafy shoot−22.921.40.75
     B. minutifolium 1201Leafy shoot−23.622.10.78
    Lepidodendraceae
     Lepidostrobophyllum alatum RC4838Reproductive shoot−23.121.60.76
     L. lanceolatum RC3134Reproductive shoot−22.921.40.75
     Lepidophyllum sp.6127Reproductive shoot−23.021.50.76
     Lepidodendron ophiurus 5284Leafy shoot−23.321.80.77
     L. lanceolatum JP218Leafy shoot−23.922.40.80
     L. lycopodioides 18239Leafy shoot−23.221.70.77
     Lepidostrobus sp.6276Reproductive shoot−24.122.60.81
    Sigillariaceae
     Sigillariostrobus rhombibracteatus 1177Cone−23.021.50.76
     Sigillariostrobus sp.1656Cone−23.722.20.79
     Sigillaria mamillaris RC2875Leafy shoot−22.721.20.74
    Other lycopsids
     Cyperites bicarinatus 4907Leafy shoot−24.723.30.84
     C. bicarinatus RG4477Leafy shoot−24.322.90.80
     C. ciliatum 76313Leafy shoot−24.523.10.83
    • All specimens are of Westphalian age (303–318 Myr BP) and held at the British Geological Survey, Keyworth, United Kingdom. 

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Low atmospheric CO2 levels during the Permo- Carboniferous glaciation inferred from fossil lycopsids
D. J. Beerling
Proceedings of the National Academy of Sciences Oct 2002, 99 (20) 12567-12571; DOI: 10.1073/pnas.202304999

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Low atmospheric CO2 levels during the Permo- Carboniferous glaciation inferred from fossil lycopsids
D. J. Beerling
Proceedings of the National Academy of Sciences Oct 2002, 99 (20) 12567-12571; DOI: 10.1073/pnas.202304999
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