Paper · 2026 · Forest Ecology and Management

Shaping gas exchange under drought: Seasonal and interspecific variation in photosynthetic traits of Pinus halepensis and Pinus brutia in Mediterranean forests

Natasa Kiorapostolou; Nikoleta Eleftheriadou; Christodoulos Sazeides; et al.

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Open access · cc-by

Abstract

As Mediterranean pine forests are increasingly exposed to intense drought events that may threaten their survival, it is important to improve our understanding of their ecophysiological responses to climatic drivers under different temporal and spatial scales. We studied key photosynthetic traits in different ecosystems of Pinus halepensis and the less studied Pinus brutia by applying an integrated, multi-site and multi-season comparative approach. For this purpose, we performed seasonal gas exchange measurements across three sites (Lesvos, Sani, and Xanthi) and ten plots in Greece to assess how species-specific stomatal strategies, microclimatic drivers, and stand structure shape carbon assimilation. We assessed CO 2 - and light-saturated photosynthetic capacity (A max ), stomatal conductance (g s ), and intrinsic water use efficiency (iWUE ) , and their responses to vapor pressure deficit (VPD) and the land surface water index (LSWI). No effect of stand structure, but significant variation among sites and seasons in A max , g s , and iWUE was detected. P. halepensis in Sani and P. brutia in Xanthi exhibited a bimodal photosynthetic activity, showing the highest A max in spring and its recovery in autumn or winter under favorable microclimate, reflecting an adaptive strategy to exploit favorable microclimatic windows. P. brutia in Lesvos, characterized by the highest VPD, exhibited a more conservative seasonal pattern with a peak of A max only in spring, reflecting prolonged atmospheric constraints on stomatal opening. Overall, atmospheric drought vs. land surface water availability had a stronger control on the A max of both species, reflecting a prioritization of hydraulic safety, in which both species employ isohydric stomatal regulation to increase their intrinsic water use efficiency at the cost of carbon uptake. However, P. brutia showed a less steep decline in A max under high atmospheric water demand, indicating a less isohydric response than P. halepensis that allows the maintenance of photosynthetic activity over a wider range of VPD and offers an advantage under increasingly intensified drought regimes. • Microclimate rather than stand structure drives gas exchange at the studied pines. • A bimodal photosynthesis pattern occurs if drought is alleviated in winter or autumn. • Atmospheric drought controlled gas exchange more than land moisture across sites. • P. brutia showed less isohydric responses than P. halepensis . • P. brutia exhibited a slower photosynthetic decline over wider atmospheric drought.

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Claims 7

Associationobservedoriginal resultno effectmeasured

Claim · model-written summary

Stand structure had no detected effect on measured gas exchange traits.

Outcome
A max , g s , and iWUEe0
Direction
No effecte0
Setting
sites and seasonse0

Not reportedTaxon

Verbatim

e0

“No effect of stand structure, but significant variation among sites and seasons in A max , g s , and iWUE was detected”

✓ located verbatim in the source · abstract, characters 951–1069 · show in text ↑
Observationobservedoriginal resultmixedsignificant (as stated)measured

Claim · model-written summary

A max, stomatal conductance and iWUE varied significantly among sites and seasons.

Outcome
A max , g s , and iWUEe0
Setting
sites and seasonse0

Not reportedTaxon

Verbatim

e0

“significant variation among sites and seasons in A max , g s , and iWUE was detected”

✓ located verbatim in the source · abstract, characters 985–1069 · show in text ↑
Observationobservedoriginal resultmixedmeasured

Claim · model-written summary

P. halepensis at Sani showed spring maxima and autumn or winter recovery in A max.

Taxon
P. halepensisPinus halepensise0
Outcome
A maxe0
Direction
higheste0
Qualifier
springe0
Setting
Sanie0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Pinus halepensis
Pinus halepensis · Taxonspacebirdy (also known as geimfyglið (:> )=| made with Sternenlaus -spirit) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“P. halepensis in Sani and P. brutia in Xanthi exhibited a bimodal photosynthetic activity, showing the highest A max in spring and its recovery in autumn or winter”

✓ located verbatim in the source · abstract, characters 1071–1234 · show in text ↑
Observationobservedoriginal resultmixedmeasured

Claim · model-written summary

P. brutia at Xanthi showed spring maxima and autumn or winter recovery in A max.

Taxon
P. brutiaPinus brutiae0
Outcome
A maxe0
Direction
higheste0
Qualifier
springe0
Setting
Xanthie0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Pinus brutia
Pinus brutia · TaxonZeynel Cebeci · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“P. halepensis in Sani and P. brutia in Xanthi exhibited a bimodal photosynthetic activity, showing the highest A max in spring and its recovery in autumn or winter”

✓ located verbatim in the source · abstract, characters 1071–1234 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

P. brutia at Lesvos showed a single spring A max peak.

Taxon
P. brutiaPinus brutiae0
Outcome
A maxe0
Direction
peake0
Qualifier
only in springe0
Setting
Lesvose0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Pinus brutia
Pinus brutia · TaxonZeynel Cebeci · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“P. brutia in Lesvos, characterized by the highest VPD, exhibited a more conservative seasonal pattern with a peak of A max only in spring”

✓ located verbatim in the source · abstract, characters 1341–1478 · show in text ↑
Associationobservedoriginal resultnegativemeasured

Claim · model-written summary

Atmospheric drought exerted stronger control on A max than land surface water availability in both pines.

Taxon
both speciese0
Outcome
A maxe0
Direction
strongere0
Comparator
land surface water availabilitye0

Verbatim

e0

“atmospheric drought vs. land surface water availability had a stronger control on the A max of both species”

✓ located verbatim in the source · abstract, characters 1555–1662 · show in text ↑
Associationobservedoriginal resultnegativemeasured

Claim · model-written summary

P. brutia A max declined less steeply under high atmospheric water demand than P. halepensis.

Taxon
P. brutiaPinus brutiae0
Outcome
A maxe0
Direction
less steep declinee0
Comparator
P. halepensise0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Pinus brutia
Pinus brutia · TaxonZeynel Cebeci · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“P. brutia showed a less steep decline in A max under high atmospheric water demand, indicating a less isohydric response than P. halepensis”

✓ located verbatim in the source · abstract, characters 1860–1999 · show in text ↑