Paper · 2026 · White Rose Research Online (University of Leeds, The University of Sheffield, University of York)

N2 fixation is linked to the ability to encroach in African savanna trees

Telford, E.M.; Simpson, K.; Street, L.; et al.

8
claims
19
verbatim quotes
0
citations
3
research areas
—
Not open access

Abstract

1. Encroachment is a globally ubiquitous phenomenon, characterised by increasing indigenous tree densities in savanna and grassland. Encroachment has been attributed to rising atmospheric CO2 concentrations fertilising tree growth and shifting the competitive balance between trees and grasses. However, only a subset of savanna tree species are currently described as encroachers, raising the hypothesis that CO2 responsiveness differs among species. Within southern African savannas, encroachment is driven primarily by nitrogen (N2)-fixing species, implying the CO2 response may be mediated via traits that enhance plant-available N. 2. Using an open-topped chamber system, we experimentally manipulated atmospheric CO2 concentrations and soil moisture for 12 savanna tree species (six encroachers and six non-encroachers) under ambient (a)CO2 (~397.9 ppm) or elevated (e)CO2 (~545.1 ppm) treatments and water limited or well-watered soil moisture treatments. We measured N-dynamics traits including nodule mass fraction (NMF), leaf δ15N, stem δ15N, and percentage of N derived from fixation (%Ndfa). 3. We found that encroachers and non-encroachers differ in short-term N-dynamics but share similar long-term N allocation strategies. Encroachers exhibited lower leaf δ15N, indicating greater utilisation of N2 fixation products to meet immediate short-term protein synthesis. Long-term N allocation strategies (NMF, stem δ15N, and %Ndfa) were similar between encroachers and non-encroachers, with plants fixing more N2 under the eCO2 and well-watered treatment. In encroachers, leaf and stem δ¹⁵N were unrelated, in contrast to the positive relationship in non-encroachers, pointing to distinct tissue-level N allocation, possibly reflecting differential N utilisation and retention. 4. We demonstrate the significance of N2 fixation in mediating the CO2 responsiveness of encroaching savanna trees. N2 fixation increases plant-available N, likely enabling encroachers to meet immediate N demands even under water limitation and increasing CO2. The potential feedback loop, where eCO2 enhances photosynthesis, facilitating greater C allocation for N2 fixation, helps to explain the ecological success of the subset of species driving encroachment under increased atmospheric CO2.

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Claims 8

Observationobservedoriginal resultmeasured

Claim · model-written summary

Encroachers had lower leaf δ15N than non-encroachers.

Taxon
African savanna treest0
Material
Encroacherse0
Outcome
leaf δ15Ne0
Direction
lowere0

Verbatim

e0

“Encroachers exhibited lower leaf δ15N”

✓ located verbatim in the source · abstract, characters 1238–1275 · show in text ↑
t0

“African savanna trees”

✓ located verbatim in the source · title, characters 52–73 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Nodule mass fraction, stem δ15N and percentage of N from fixation were similar across encroachers and non-encroachers.

Taxon
African savanna treest0
Outcome
NMF, stem δ15N, and %Ndfae0
Direction
similare0
Comparator
encroachers and non-encroacherse0

Verbatim

e0

“Long-term N allocation strategies (NMF, stem δ15N, and %Ndfa) were similar between encroachers and non-encroachers”

✓ located verbatim in the source · abstract, characters 1380–1494 · show in text ↑
t0

“African savanna trees”

✓ located verbatim in the source · title, characters 52–73 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Plants fixed more N2 under elevated CO2.

Taxon
African savanna treest0
Treatment
eCO2e0
Outcome
N2e0
Direction
moree0
Comparator
ambient (a)CO2e1

Verbatim

e0

“plants fixing more N2 under the eCO2 and well-watered treatment”

✓ located verbatim in the source · abstract, characters 1501–1564 · show in text ↑
e1

“ambient (a)CO2 (~397.9 ppm) or elevated (e)CO2 (~545.1 ppm) treatments”

✓ located verbatim in the source · abstract, characters 832–902 · show in text ↑
t0

“African savanna trees”

✓ located verbatim in the source · title, characters 52–73 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Plants fixed more N2 under well-watered conditions.

Taxon
African savanna treest0
Treatment
well-waterede0
Outcome
N2e0
Direction
moree0
Comparator
water limitede1

Verbatim

e0

“plants fixing more N2 under the eCO2 and well-watered treatment”

✓ located verbatim in the source · abstract, characters 1501–1564 · show in text ↑
e1

“water limited or well-watered soil moisture treatments”

✓ located verbatim in the source · abstract, characters 907–961 · show in text ↑
t0

“African savanna trees”

✓ located verbatim in the source · title, characters 52–73 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Leaf and stem δ15N were unrelated in encroachers.

Taxon
African savanna treest0
Material
encroacherse0
Outcome
leaf and stem δ¹⁵Ne0

Not reportedDirection

Verbatim

e0

“In encroachers, leaf and stem δ¹⁵N were unrelated”

✓ located verbatim in the source · abstract, characters 1566–1615 · show in text ↑
t0

“African savanna trees”

✓ located verbatim in the source · title, characters 52–73 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Leaf and stem δ15N were positively related in non-encroachers.

Taxon
African savanna treest0
Material
non-encroacherse0
Outcome
leaf and stem δ¹⁵Ne1
Direction
positivee0

Verbatim

e0

“positive relationship in non-encroachers”

✓ located verbatim in the source · abstract, characters 1636–1676 · show in text ↑
e1

“leaf and stem δ¹⁵N were unrelated, in contrast to the positive relationship in non-encroachers”

✓ located verbatim in the source · abstract, characters 1582–1676 · show in text ↑
t0

“African savanna trees”

✓ located verbatim in the source · title, characters 52–73 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivebasis n/a

Claim · model-written summary

Authors interpret N2 fixation as enabling encroachers to meet immediate nitrogen demands under water limitation and increasing CO2.

Taxon
African savanna treest0
Outcome
meet immediate N demandse0
Direction
increasese0
Qualifier
N2 fixation increases plant-available Ne0

Verbatim

e0

“N2 fixation increases plant-available N, likely enabling encroachers to meet immediate N demands even under water limitation and increasing CO2”

✓ located verbatim in the source · abstract, characters 1904–2047 · show in text ↑
t0

“African savanna trees”

✓ located verbatim in the source · title, characters 52–73 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivebasis n/a

Claim · model-written summary

Authors propose that elevated CO2 enhances photosynthesis and carbon allocation for N2 fixation, helping explain encroachment.

Taxon
African savanna treest0
Outcome
ecological successe0
Qualifier
eCO2 enhances photosynthesis, facilitating greater C allocation for N2 fixatione0

Verbatim

e0

“eCO2 enhances photosynthesis, facilitating greater C allocation for N2 fixation, helps to explain the ecological success of the subset of species driving encroachment under increased atmospheric CO2”

✓ located verbatim in the source · abstract, characters 2084–2282 · show in text ↑
t0

“African savanna trees”

✓ located verbatim in the source · title, characters 52–73 · show in text ↑