Thermal acclimation fails to confer a carbon budget advantage to invasive species over natives

Thermal acclimation fails to confer a carbon budget advantage to invasive species over natives | bioRxiv /* */ /* */ <!-- <!-- /*! * yepnope1.5.4 * (c) WTFPL, GPLv2 */ (function(a,b,c){function d(a){return"[object Function]"==o.call(a)}function e(a){return"string"==typeof a}function f(){}function g(a){return!a||"loaded"==a||"complete"==a||"uninitialized"==a}function h(){var a=p.shift();q=1,a?a.t?m(function(){("c"==a.t?B.injectCss:B.injectJs)(a.s,0,a.a,a.x,a.e,1)},0):(a(),h()):q=0}function i(a,c,d,e,f,i,j){function k(b){if(!o&&g(l.readyState)&&(u.r=o=1,!q&&h(),l.onload=l.onreadystatechange=null,b)){"img"!=a&&m(function(){t.removeChild(l)},50);for(var d in y[c])y[c].hasOwnProperty(d)&&y[c][d].onload()}}var j=j||B.errorTimeout,l=b.createElement(a),o=0,r=0,u={t:d,s:c,e:f,a:i,x:j};1===y[c]&&(r=1,y[c]=[]),"object"==a?l.data=c:(l.src=c,l.type=a),l.width=l.height="0",l.onerror=l.onload=l.onreadystatechange=function(){k.call(this,r)},p.splice(e,0,u),"img"!=a&&(r||2===y[c]?(t.insertBefore(l,s?null:n),m(k,j)):y[c].push(l))}function j(a,b,c,d,f){return q=0,b=b||"j",e(a)?i("c"==b?v:u,a,b,this.i++,c,d,f):(p.splice(this.i++,0,a),1==p.length&&h()),this}function k(){var a=B;return a.loader={load:j,i:0},a}var l=b.documentElement,m=a.setTimeout,n=b.getElementsByTagName("script")[0],o={}.toString,p=[],q=0,r="MozAppearance"in l.style,s=r&&!!b.createRange().compareNode,t=s?l:n.parentNode,l=a.opera&&"[object Opera]"==o.call(a.opera),l=!!b.attachEvent&&!l,u=r?"object":l?"script":"img",v=l?"script":u,w=Array.isArray||function(a){return"[object Array]"==o.call(a)},x=[],y={},z={timeout:function(a,b){return b.length&&(a.timeout=b[0]),a}},A,B;B=function(a){function b(a){var a=a.split("!"),b=x.length,c=a.pop(),d=a.length,c={url:c,origUrl:c,prefixes:a},e,f,g;for(f=0;f<d;f++)g=a[f].split("="),(e=z[g.shift()])&&(c=e(c,g));for(f=0;f<b;f++)c=x[f](c);return c}function g(a,e,f,g,h){var i=b(a),j=i.autoCallback;i.url.split(".").pop().split("?").shift(),i.bypass||(e&&(e=d(e)?e:e[a]||e[g]||e[a.split("/").pop().split("?")[0]]),i.instead?i.instead(a,e,f,g,h):(y[i.url]?i.noexec=!0:y[i.url]=1,f.load(i.url,i.forceCSS||!i.forceJS&&"css"==i.url.split(".").pop().split("?").shift()?"c":c,i.noexec,i.attrs,i.timeout),(d(e)||d(j))&&f.load(function(){k(),e&&e(i.origUrl,h,g),j&&j(i.origUrl,h,g),y[i.url]=2})))}function h(a,b){function c(a,c){if(a){if(e(a))c||(j=function(){var a=[].slice.call(arguments);k.apply(this,a),l()}),g(a,j,b,0,h);else if(Object(a)===a)for(n in m=function(){var b=0,c;for(c in a)a.hasOwnProperty(c)&&b++;return b}(),a)a.hasOwnProperty(n)&&(!c&&!--m&&(d(j)?j=function(){var a=[].slice.call(arguments);k.apply(this,a),l()}:j[n]=function(a){return function(){var b=[].slice.call(arguments);a&&a.apply(this,b),l()}}(k[n])),g(a[n],j,b,n,h))}else!c&&l()}var h=!!a.test,i=a.load||a.both,j=a.callback||f,k=j,l=a.complete||f,m,n;c(h?a.yep:a.nope,!!i),i&&c(i)}var i,j,l=this.yepnope.loader;if(e(a))g(a,0,l,0);else if(w(a))for(i=0;i (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0];var j=d.createElement(s);var dl=l!='dataLayer'?'&l='+l:'';j.src='//www.googletagmanager.com/gtm.js?id='+i+dl;j.type='text/javascript';j.async=true;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-M677548'); Skip to main content Home About Submit ALERTS / RSS Search for this keyword Advanced Search New Results Thermal acclimation fails to confer a carbon budget advantage to invasive species over natives View ORCID Profile Thibaut Juillard , Christoph Bachofen , Marco Conedera , Mattéo Dumont , Jean-Marc Limousin , Arianna Milano , Gianni Boris Pezzatti , Alberto Vilagrosa , Charlotte Grossiord doi: https://doi.org/10.1101/2025.07.21.665833 Thibaut Juillard 1 Plant Ecology Research Laboratory PERL, School of Architecture, Civil and Environmental Engineering ENAC , EPFL, CH-1015 Lausanne, Switzerland 2 Community Ecology Unit, Swiss Federal Institute for Forest, Snow and Landscape Research WSL , CH-8903, Birmensdorf, Switzerland Find this author on Google Scholar Find this author on PubMed Search for this author on this site ORCID record for Thibaut Juillard For correspondence: thibaut.juillard{at}epfl.ch Christoph Bachofen 1 Plant Ecology Research Laboratory PERL, School of Architecture, Civil and Environmental Engineering ENAC , EPFL, CH-1015 Lausanne, Switzerland 2 Community Ecology Unit, Swiss Federal Institute for Forest, Snow and Landscape Research WSL , CH-8903, Birmensdorf, Switzerland Find this author on Google Scholar Find this author on PubMed Search for this author on this site Marco Conedera 2 Community Ecology Unit, Swiss Federal Institute for Forest, Snow and Landscape Research WSL , CH-8903, Birmensdorf, Switzerland Find this author on Google Scholar Find this author on PubMed Search for this author on this site Mattéo Dumont 3 Institut National des Sciences Appliquées de Lyon , INSA, 69100 Villeurbanne, France Find this author on Google Scholar Find this author on PubMed Search for this author on this site Jean-Marc Limousin 4 CEFE, Univ Montpellier , CNRS, EPHE, IRD, Montpellier, France Find this author on Google Scholar Find this author on PubMed Search for this author on this site Arianna Milano 1 Plant Ecology Research Laboratory PERL, School of Architecture, Civil and Environmental Engineering ENAC , EPFL, CH-1015 Lausanne, Switzerland 2 Community Ecology Unit, Swiss Federal Institute for Forest, Snow and Landscape Research WSL , CH-8903, Birmensdorf, Switzerland Find this author on Google Scholar Find this author on PubMed Search for this author on this site Gianni Boris Pezzatti 2 Community Ecology Unit, Swiss Federal Institute for Forest, Snow and Landscape Research WSL , CH-8903, Birmensdorf, Switzerland Find this author on Google Scholar Find this author on PubMed Search for this author on this site Alberto Vilagrosa 5 Mediterranean Center for Environmental Studies (CEAM Foundation). Joint Research Unit University of Alicante-CEAM, University of Alicante , 03690, Sant Vicent del Raspeig, Alicante, Spain 6 CEAM-Department de Ecologia, Universitat d’Alacant , POB 99, E-03080 Alacant, Spain Find this author on Google Scholar Find this author on PubMed Search for this author on this site Charlotte Grossiord 1 Plant Ecology Research Laboratory PERL, School of Architecture, Civil and Environmental Engineering ENAC , EPFL, CH-1015 Lausanne, Switzerland 2 Community Ecology Unit, Swiss Federal Institute for Forest, Snow and Landscape Research WSL , CH-8903, Birmensdorf, Switzerland Find this author on Google Scholar Find this author on PubMed Search for this author on this site Abstract Full Text Info/History Metrics Supplementary material Data/Code Preview PDF Abstract Both native and invasive plants can adjust photosynthesis and respiration when exposed to warmer temperatures. However, it is uncertain if invasive plants are more plastic and exhibit higher acclimation to rising temperatures than native ones, a trait that could contribute to their invasive behavior in novel environments. We compared the capacity of a highly invasive palm in central Europe ( Trachycarpus fortunei ) and two native co-occurring species ( Ilex aquifolium and Tilia cordata ) to acclimate photosynthesis and respiration to air temperature changes using a two-year-long transplant experiment across Europe (mean temperatures ranging from 8.4 to 21.8°C). We measured the optimal temperature of photosynthesis (T opt ), the assimilation at optimal temperature (A opt ), the thermal breath of photosynthesis (T 80 ), the respiration at 25°C (R 25 ), the temperature sensitivity of respiration (Q 10 ), and simulated the whole-plant carbon budget. For all species, T opt , A opt, and T 80 increased with warming, while R 25 decreased in the native species and Q 10 decreased in the invasive species only. Consequently, acclimation enhanced the carbon budget of the invasive and native plants in the warm and hot sites. The invasive palm had a similar or lower acclimation capacity than other species and a lower but constant carbon budget across the European temperature gradient. Our work reveals that not all invasive plants exhibit greater photosynthetic plasticity than native ones, suggesting that temperature-driven enhancement of their carbon budget may play a limited role in future invasion processes. 1. Introduction Global warming promotes the spread of invasive plant species worldwide, representing one of the most significant threats to plant biodiversity ( Walther et al ., 2009 ; Liu et al ., 2017 ; Pyšek et al ., 2020 ). Among possible factors, long-lived invasive plants can outperform native species under a warmer climate as they tend to be adapted to broader air temperature (T air ) ranges ( e . g ., Higgins and Richardson, 2014 ; Finch et al ., 2021 ; Turbelin and Catford, 2021 ), can extend their growing seasons more significantly ( e . g ., Fridley, 2012 ; Juillard et al ., 2024 ), and acclimate various functional traits more extensively ( e . g . Davidson, Jennions and Nicotra, 2011 ; Gioria et al ., 2023 ). Higher carbon (C) uptake and plant productivity following a better thermal acclimation of photosynthesis and respiration may, in turn, enhance species competitiveness ( Duan et al ., 2013 ; Yu et al ., 2019 ; Grossiord et al ., 2022 ). Still, few studies investigated photosynthetic and respiratory thermal acclimation in the context of species invasiveness ( Verlinden and Nijs, 2010 ; Godoy, Valladares and Castro-Díez, 2011 ; Ripley et al ., 2020 ), with none addressing if invasive plants are more plastic than native ones with increasing temperature. Yet, understanding and predicting these processes would help reduce uncertainties in climate-vegetation models, where acclimation is often simplified or fully neglected ( Crous, Uddling and De Kauwe, 2022 ) and would allow finding appropriate conservation strategies for forests highly susceptible to invasion as a consequence of global warming. Plants generally acclimate rapidly to higher T air (within one month; e . g ., Kattge and Knorr, 2007 ; Kumarathunge et al ., 2019 ) by increasing their optimal temperature (T opt , i . e ., the temperature at which the net assimilation (A net ) reaches its maximum), its corresponding optimal net assimilation value (A opt ), and their thermal breath (T 80 ), which represents the temperature range where photosynthesis reaches >80 % of its maximum ( Yamori, Hikosaka and Way, 2014 ), thereby maintaining C gain despite warmer air ( Kruse et al ., 2019 ; Vico et al ., 2019 ; Choury et al ., 2022 ). Thermal acclimation of photosynthesis also depends on stomatal conductance ( Kruse et al ., 2019 ; Kullberg, Slot and Feeley, 2023 ) and phenology ( Grossiord et al ., 2022 ), which vary interannually ( Petrik et al ., 2022 ; Didion-Gency et al ., 2024 ), leading to significant uncertainties in plant’s thermal acclimation capacity. Shifts in A opt , T opt , and T 80 are mainly driven by an increase in the maximum catalytic activity of Rubisco (V Cmax ) and the maximum ratio of electron transport (J max ). Further, V Cmax and J max are both limited by nitrogen availability, therefore, thermal acclimation of photosynthesis also depends on nitrogen allocation, for which invasive plants tend to be more plastic than native ones ( Feng and Fu, 2008 ; Funk, Glenwinkel and Sack, 2013 ). In parallel, acclimation to higher T air can also involve lowering the rate of respiration at 25°C (R 25 ) and the respiration yield every 10°C (Q 10 ) ( Atkin and Tjoelker, 2003 ; Atkin, Bruhn and Tjoelker, 2005 ), thereby limiting C loss at high temperatures as respiration increases exponentially ( Atkin and Tjoelker, 2003 ; Way and Yamori, 2014 ; Crous, Uddling and De Kauwe, 2022 ). Plants with higher photosynthesis acclimation can show higher respiration acclimation ( Dusenge, Duarte and Way, 2019 ), although respiration can also acclimate more extensively to T air than assimilation ( Campbell et al ., 2007 ; Way and Oren, 2010 ; Crous, Uddling and De Kauwe, 2022 ). Just as for photosynthesis, the duration of exposure to changed temperatures can influence respiration acclimation. Typically, acclimation of R 25 to a particular T air occurs during tissue development, while Q 10 varies more rapidly with seasonal changes in ambient T air ( Atkin, Bruhn and Tjoelker, 2005 ). Some studies found that respiration tends to acclimate universally among plant species ( Crous, Uddling and De Kauwe, 2022 ), and others reported acclimation to be species-specific independently of biomes or functional groups ( e.g ., evergreen vs . deciduous) ( Slot and Kitajima (2015) ). However, respiratory rates are inversely proportional to the leaf nitrogen content and the specific leaf area ( Lee, Reich and Bolstad, 2005 ; Xu, Schuster and Griffin, 2007 ), frequently found to be higher in invasive species than in native ones ( Baruch and Goldstein, 1999 ; Leishman et al ., 2007 ). Therefore, temperature-induced reduction of respiratory rates could favor invasive species at high temperatures, but this remains largely unknown. In this study, we compared the acclimation of leaf photosynthesis and respiration of the invasive Trachycarpus fortunei , a palm native from Southeastern China, with two co-occurring natives growing in the southern Alps in a sub-mediterranean climate ( i . e ., the evergreen Ilex aquifolium and deciduous Tilia cordata ). Since the 2000s, T. fortunei has been colonizing natural forests worldwide, impacting the regeneration of native species ( Fehr et al ., 2023 ). In the southern Alps, the spread of T. fortunei has been correlated with the rise in T air since the 1970s (ΔMAT: +1.7°C) ( Walther et al ., 2007 ; Fehr et al ., 2023 ). As such, T. fortunei is suspected to benefit from warmer temperatures to outcompete its native competitors, but whether this includes enhanced photosynthetic and respiratory thermal acclimation is unknown. Using a transplant experiment in five sites covering a large range of mean T air across Europe, we measured the responses of photosynthesis and respiration to temperature over two years. A soil-plant-atmosphere continuum (SPAC) model was used to estimate the impact of temperature acclimation on the leaf and whole-plant C budget over an entire growing season. We hypothesized that (1) the invasive T . fortunei acclimates more extensively than the native species to increased T air , leading to higher A opt , T opt , and T 80 . Similarly, (2) the invasive T . fortunei displays lower R 25 and Q 10 than native species with higher T air and that, as such, (3) T. fortunei assimilates more C than native species in warmer sites. 2. Materials and methods 2.1 Experimental design We selected five sites across a temperature gradient in Europe for a transplant experiment: one reference site located in the sub-Mediterranean climate where T. fortunei is the most invasive in Europe ( i . e ., the lowlands of the southern slope of the Swiss Alps), and four sites with differences of respectively −6°, −3°, +3°, and +6°C mean summer T air (from April to October) compared to the reference site in the last ten years (2010-2020). This allowed us to assess the acclimation of the photosynthetic and respiration properties along a large T air gradient, thereby covering regions where T. fortunei already has or could become invasive in future years ( Fehr et al ., 2023 ). In addition to the reference site in southern Switzerland (sub-Mediterranean climate), sites spanned from south-eastern Spain (thermic semi-arid), southern France (Mediterranean) to the Swiss Plateau (temperate), and the Swiss Alps (cold) ( Fig. 1 ). The elevation of the sites ranged between 1 and 965 m a.s.l. (see Table 1 for more details). During the measurement period (2022 and 2023), mean annual T air were 8.4, 13.3, 15.2, 17.7, and 21.8°C from the coldest to the warmest site, respectively, with slightly warmer conditions in 2022 than in 2023 (Fig. S1). Download figure Open in new tab Fig. 1: Map of the five experimental sites in Europe (from the coldest to the warmest: Filisur (cold, dark blue), Birmensdorf (temperate, light blue), Cadenazzo (sub-Mediterranean, yellow), Montpellier (Mediterranean, orange), and Guardamar del Segura (semi-arid, red). On the left are pictures of the three focal species included in the study. The right panels show the mean air temperature and VPD within each shading infrastructure from April to October 2022 and 2023. At the bottom right is a picture showing the shading infrastructure. View this table: View inline View popup Download powerpoint Table 1: Location of the experimental sites and associated climatic conditions. T. fortunei typically co-occurs with other species in the forest understory ( e . g ., the native evergreen Ilex aquifolium and deciduous Tilia cordata ), which it occasionally excludes when dominating the canopy ( Fehr et al ., 2023 ). To simulate the natural half-shady understory environment, we constructed a wooden shading infrastructure (6 x 4 x 1.8 m) in an unobstructed area on all sites. The top and sides were covered with a permeable shading tissue obstructing 70% of sunlight. Mean yearly incoming solar radiation for 2022-2023 was similar among sites (Fig. S1). In November 2021, we dug out 200 saplings of T. fortunei of 50-70 cm in height in a freshly invaded sub-Mediterranean forest (Cugnasco, Switzerland; 46°10’15’’ N, 8°55’49’’ E, 205 m a.s.l., MAT 12.2 °C, MAP: 1757 mm; 1.1 km from our reference site). We further bought 200 saplings of T. cordata (Morbio Superiore, Switzerland) and 150 saplings of I. aquifolium (Wiler bei Utzenstorf, Switzerland) from commercial producers. All individuals were immediately potted in 7 L pots with generic sandy forest soil made of 20% peat and mineral substrate (pH = 6.3; Ökohum; DE) and overwintered at the reference site. In March 2022, 100 healthy individuals of each species ( i . e ., 300 saplings) were randomly separated into five groups of 20 individuals per species and transported to the five sites (n = 60 saplings per site). The plants for the coldest site were kept at the second coldest one (temperate climate) until April 2022 and from December 2022 to March 2023 to avoid exposure to temperatures below −15°C as it could have killed T. fortunei ( Fehr and Burga, 2016 ). Despite this, 75% of the individuals of T. fortunei at the two coldest sites suffered frost damage during the winter of 2022-2023, limiting the number of replicates to four in 2023. At each site, pots were placed inside the shading infrastructure in four rows of 15 individuals each, alternating species to randomize potential differences in light and wind conditions. Each pot was individually connected to a drip irrigation system equipped with individual pressure resistances (Allenspach GreenTech AG, CH) to guarantee the same irrigation for each pot. Plants were watered automatically every two days at dawn to field capacity to ensure water availability throughout the experiment. We recorded T air , RH, and solar radiation (data logger: EM-100; air temperature and humidity: Atmos-14, Meter Group Inc.; USA; light quantum sensor: SQ-100X-SS, Apogee Instruments; USA) inside each shading infrastructure every 30 minutes at 1,5 m above the ground. We also monitored the soil moisture of each plant’s pot at 10 cm depth monthly with a soil moisture meter (TDR-100, Spectrum Technologies; USA). For two years (2022-2023), the measurement campaigns were conducted in May, July, and September (six in total) during two consecutive growing seasons ( i . e ., 2022 and 2023). We tracked leaf flush and senescence at each site with phenocams (IPX5, VisorTech; DE). To guarantee that we measured fully expanded leaves, the first campaign of each year in May was conducted one month after all individuals of T. cordata had flushed. 2.2 CO 2 assimilation and respiration responses to air temperature We measured the optimal temperature (T opt ), net assimilation at the optimal temperature (A opt ), thermal breath (T 80 ), dark respiration at 25°C (R 25 ), and respiration yield (Q 10 ) of one-year-old (or fully developed of the current year in deciduous T. cordata ), undamaged leaves of 10 individuals per species (4 individuals of T. fortunei in 2023). Hence, every year in May, one leaf per individual was selected for T opt , A opt , and T 80 measurements, while an adjacent leaf was chosen for R 25, and Q 10. T opt , A opt , and T 80 were obtained through CO 2 assimilation response to T air curves using portable photosynthesis systems (Li-6800, Licor Biosciences; USA) similar to Gauthey et al ., (2023) and Deluigi et al., (2025). Measurements were conducted approx. every 1-2 hours at five or six time points during the day, reflecting different ambient T air from the sunrise ( i.e. , the coldest T air of the day) to the middle of the afternoon ( i.e. , the warmest T air of the day) to obtain the possible largest T air range (from 12 to 35°C on average across sites and campaigns). Air temperature within the Li-6800 chamber was set according to the ambient T air , which was measured continuously (RS-91, RS Instruments; DE). By doing so, we ensured to track the temperature response curve of the ambient T air and to avoid possible artifacts associated with differences between the ambient temperature experienced by the plants and the conditions in the cuvette, as well as bias associated with the calculations of T L within gas exchange systems ( Still et al ., 2019 ). To extend the temperature range, we additionally decreased and increased the temperature of the cuvette by 5°C during the coldest and warmest time of the day, respectively. Relative humidity within the cuvette was increased at high temperatures to avoid stomatal closure due to high VPD. Hence, VPD ranged between 1 (fixed minimum) and maximum 3,5 kPa (when T air reached > 38°C). T opt , A opt , and T 80 were measured at saturating light (PPFD of 1500 μmol m −2 s −1 ) and ambient CO 2 (400 ppm). Measurements of R 25 and Q 10 were conducted as described above, except that the leaf of each individual was wrapped in aluminum foil for at least 30 minutes before each measurement to dark-acclimate the leaves and that light was reduced to 0 μmol m −2 s −1 inside the cuvette during the measurements. All measurements were taken after the gas exchange rates had stabilized for at least 5 min. We extracted T opt , A opt , and T 80 of each individual by fitting our measurements of assimilation at different T air with a parabolic curve in R (4.1.1, R Core Team, 2021), using the following equation as in Choury et al ., (2022) : where A i is the net assimilation at temperature T i and b is the width of the curve. T 80 was then calculated after resolving equation (1) for A i = 0.8 A opt and isolating T i . Similarly, R 25 and Q 10 were obtained after fitting the respiration measurements at different T air with an exponential curve in R, using the following equation ( Choury et al ., 2022 ): 2.3 SPAC modeling of leaf-level photosynthesis and respiration To assess the temperature effect on the annual leaf-level C uptake (C leaf ), we modeled the instantaneous leaf-level net photosynthesis at 30-minute intervals from leaf flushing to senescence with a mechanistic soil-plant-atmosphere continuum (SPAC) model proposed by Garcia-Tejera et al . (2017). The model was modified to account for the acclimation of temperature responses of photosynthesis and respiration ( Fig. 2 ). The original model calculates iteratively net assimilation (A net ) and stomatal conductance (g s ) based on environmental drivers and physiological parameters. A net is calculated as: where C a is the ambient CO 2 concentration, C i is the intracellular CO 2 concentration, and R d is the dark respiration rate. g s is derived from its theoretical value at unlimited water availability (g s,max ): where B is the CO 2 uptake limiting rate of either J max or V Cmax , Γ is the CO 2 compensation point of photosynthesis, E is a metric of carboxylation and oxygenation rates, and D and F are constants. g s is then derived from g s,max based on hydraulic parameters, water transport from the soil to the leaf, and transpiration (Fig. S2, equations in Tables S1 & S2). Download figure Open in new tab Fig. 2: Assimilation-temperature response curves for the summer months (mean of May-October) of both years (2022-2023) for T. fortunei , I. aquifolium , and T. cordata at the five experimental sites (indicated with colors from blue to red going from the coldest to the warmest). On the right panels, the mean and standard errors of A opt , T opt , and T 80 at each site are shown (n = 4-10 individuals per species). Different letters indicate significative differences ( p < 0.05) between sites for each species. To model leaf temperature (T L ), we used a leaf energy balance model by Kevin Tu ( http://landflux.org/Tools.php ) as in Marias, Meinzer and Still (2017) . As such, T L varied with T air , light, RH, and g s following the equations from Jones (1992) , Sridhar and Elliott (2002) , and Monteith and Reifsnyder (2007) (Table S1). As T L impacts g s , and vice versa, we nested the SPAC optimization process for g s calculations into a second optimization process to calculate T L ( Fig. 2 ). R d was calculated with T L following equation (2) . V Cmax and J max were adjusted for T L following the peaked Arrhenius function equation as in Kumarathunge et al . (2019) . To incorporate T L acclimation in our model, we adjusted the entropy factor (ΔS, J mol -1 K -1 ) and the activation energy (Ha, kJ mol -1 ) in the peaked Arrhenius function with the general coefficients proposed by Kumarathunge et al . (2019) (Tables S2 & S3). For respiration, we adjusted R 25 with T air with linear species-specific equations that we derived from our results (Tables S1): To parametrize the model, we measured J max,25 and V c,max,25 on five individuals of the three species in July 2023 at our reference site with the Li-6800 following the methodology used in Didion-Gency et al ., (2022) . In May 2022, we measured the minimum stomatal conductance (g min ) on 4-5 saplings per species at EPFL (46°31’15.3"N, 6°34’04.0"E, Lausanne, CH). In September each year (2022 and 2023), we measured the leaf area of 4-10 individuals of each species at every site. For this, we photographed ten representative leaves of I. aquifolium and T. cordata, and all the leaves of T. fortunei . We extracted the leaf area using the software Fiji ( Schindelin et al ., 2012 ). The mean leaf area of I. aquifolium and T. cordata was multiplied by the total number of leaves of each individual to compute the total leaf area. Finally, the leaf area index (LAI) was obtained by dividing the total leaf area by the pot’s upper surface. We calibrated and validated the model with A net measurements that we took diurnally at the five experimental sites during every campaign. A net was measured approximately every hour with the Li-6800 under ambient light, T air , and RH. We randomly selected 70 % of the measurements to calibrate influential model parameters that we could not measure similarly as in Grossiord et al . (2022) and Deluigi et al ., (2025). The model was calibrated using a differential evolution (DEzs) Markov chain Monte Carlo (MCMC) sampler ( Ter Braak and Vrugt, 2008 ) using the R package BayesianTools ( Hartig et al ., 2023 ). For each species, we run 10,000 iterations of three independent chains. We confirmed the convergence of the calibration with the Gelman–Rubin diagnostic and a threshold of 1.1 ( Gelman and Rubin, 1992 ). We used the 30 % remaining measurements to assess the goodness-of-fit (RMSE, percent bias, and Nash-Sutcliff efficiency) between the measured and modeled A net at each site. To estimate the effect of photosynthetic temperature acclimation on the plant C budget, we ran the model without acclimation by keeping V C,max , J max , R 25 , and Q 10 constant (mean of the reference site). 2.4 Statistical analyses Species and site differences in T opt , T 80 , A opt , R 25 , and Q 10 were tested through analysis of variance by using species ( i . e ., T. fortunei , I. aquifolium , and T. cordata ) and climate (semi-arid, Mediterranean, sub-Mediterranean, temperate, and cold) and their interaction as fixed effects. No variable transformation was required to ensure homoscedasticity. Tukey’s HSD post hoc tests were used to separately estimate differences between species or climates. Linear regressions were used to test the relationships between T opt , A opt , R 25 , Q 10, and T air (mean of the two weeks preceding the measurements) and between A opt and T opt . All statistical tests were done using the software R (4.1.1, R Core Team, 2021). 3. Results 3.1 Photosynthetic and respiration responses to air temperature In both years, A opt , T opt , and T 80 differed between species and climates ( p < 0.001 for A opt and T opt and p < 0.05 for T 80 ; Fig. 2 & Table S4), with generally higher values in the warmest site and lower values in the coldest one (Fig. S3 & Table S5). In all sites, A opt and T opt were lower in T. fortunei (2.86 µmol m 2 s -1 and 21.3°C on average, respectively) than in I. aquifolium (5.5 µmol m 2 s -1 and 22.7°C on average, respectively) and T. cordata (6.96 µmol m 2 s -1 and 23.9°C on average, respectively), even in the reference sub-Mediterranean climate where T. fortunei is highly invasive ( p < 0.001; Fig 2 ). Similarly, T. fortunei had a lower T 80 (11.1°C on average) than I. aquifolium (13.9°C on average) and T. cordata (15.2°C on average) in all sites ( Fig. 2 ). A opt and T opt were strongly correlated with the mean T air of the two preceding weeks, suggesting rapid photosynthetic acclimation ( Fig. 3 ). The relationship between mean T air and T opt was significant for all species ( p < 0.01), but steeper for the two evergreen species than for the deciduous one (+0.60, +0.55, and + 0.33°C T opt per °C mean T air for T . fortunei , I . aquifolium , and T . cordata , respectively). The same relationship with A opt was significant in T. fortunei (+0.16 µmol m 2 s -1 per °C; p < 0.001) and I. aquifolium (+0.19 µmol m 2 s -1 per °C; p < 0.001), but not for T. cordata ( Fig. 3 ). Air temperature and T 80 were only correlated in T. fortunei (+ 0.19°C per °C air temperature, R 2 = 0.25, p < 0.01; Fig. S4). A opt , T opt , and T 80 were positively correlated in all species ( p < 0.05; Fig. 4 & Fig. S5), except for T opt and T 80 in T. fortunei . In T. fortunei , A opt increased by 0.12 µmol m 2 s -1 per °C positive shift in T opt . The same correlation was steeper in I. aquifolium and T. cordata (+0.23 and +0.27 µmol m 2 s -1 per °C shift in T opt , respectively; Fig. 4 ). Download figure Open in new tab Fig. 3: A opt and T opt (n = 4-10 individuals per species) averaged by campaigns in relation to air temperature of the two weeks preceding the measurements for T . fortunei , I . aquifolium , and T . cordata . Colors represent climates from blue to red, from the coldest to the warmest. Linear regression lines were added when significant. Download figure Open in new tab Fig. 4: Assimilation at the optimal temperature (A opt ) in function of the optimal temperature of photosynthesis (T opt ) (n = 4-10 individuals per species) in T. fortunei , I. aquifolium , and T. cordata during all campaigns in 2022 and 2023. Colors represent sites from blue to red, from the coldest to the warmest. Linear regression lines were added when significant. R 25 varied across sites and species ( p < 0.001; Fig. 5 & Table S4) with higher values in the colder sites, except in T . fortunei . On average, R 25 was almost half as high in T. fortunei (0.40 µmol m 2 s -1 ) than in I. aquifolium and T. cordata (0.85 and 0.75 µmol m 2 s -1 , respectively) ( Fig. 5 ). R 25 decreased with increasing mean T air of the two preceding weeks in I. aquifolium and T. cordata (−0.03 µmol m 2 s -1 / °C in both species, P < 0.01; Fig. 6 ) but not in T. fortunei, suggesting limited temperature acclimation in that species. On the other hand, while Q 10 was different between species and sites ( p < 0.001; Fig. 5 & Table S4), Q 10 decreased with increasing mean T air only in T. fortunei ( Fig. 6 ). Download figure Open in new tab Fig. 5: Respiration-temperature response curves for the summer months (mean of May-October) of both years (2022-2023) for T. fortunei , I. aquifolium , and T. cordata at the five experimental sites (indicated with colors from blue to red, from the coldest to the warmest). The right panels show the mean and standard errors of R 25 and Q 10 at each site (n = 4-10 individuals per species). Different letters indicate significant differences ( p < 0.05) between the sites. Download figure Open in new tab Fig. 6: R 25 and Q 10 (n = 4-10 individuals per species) averaged by campaigns in relation to the air temperature of the two weeks preceding the measurements for T. fortunei , I. aquifolium , and T. cordata . Colors represent sites from blue to red, from the coldest to the warmest. Linear regression lines were added when significant. 3.2 Modelled carbon uptake The SPAC model produced more precise predictions for I. aquifolium and T. cordata than for T. fortunei (R 2 = 0.61, 0.66, and 0.17; NSE = 0.47, 0.62, and 0.11, respectively). Still, bias was low in all species (−0.61, 1.41, and 1.72%, respectively, Table S6), indicating a high accuracy of the model. Overall, the model revealed significant differences in the annual leaf-level C uptake (C leaf ) between species ( i . e ., 360, 345, and 587 gC m -2 on average for T . fortunei , I . aquifolium , and T . cordata , respectively; Fig. 7 ) and climates ( p < 0.001), as well as an interaction between species and climate ( P = 0.016, Table S7). At the reference site, T. fortunei had the lowest C leaf (321 gC m -2 ) compared to I. aquifolium and T . cordata (418 and 615 gC m -2 , respectively; Fig. 7 ). However, T . fortunei maintained a relatively constant C leaf between the sites except at the warmest site, where a high R resulted in a lower C leaf ( Fig. 7 & Table S8). In contrast, both native species show the lowest C leaf at the coldest site, a progressively higher C leaf until a peak at the Mediterranean site, and a subsequent decrease at the warmest site ( i . e ., semi-arid) ( Fig. 7 ). Notably, T . cordata was the only species that kept a similarly high C leaf at the warmest site as at the reference sites despite higher R ( Fig. 7 ). Download figure Open in new tab Fig. 7: Daily mean leaf-level C budget in T. fortunei , I. aquifolium , and T. cordata at the five experimental sites from 15 th October 2022 to 15 th October 2023. The right panels show the annual leaf-level C uptake for each site over the same period. Net assimilation corresponds to the plain bar, whereas respiration is dashed. Error bars indicate the uncertainty of J max,25 , V Cmax,25 , R 25 , and Q 10 (n = 37–57). Different letters indicate significant differences ( p < 0.05) between the sites. Total leaf area did not vary largely between sites, apart for T . fortunei , which had a larger leaf area in the three warmest sites compared to the two colder ones ( i . e ., because of frost damage in winter 2022-2023; Fig. S6). As a consequence, total C uptake (C tot ) showed similar patterns as C leaf across sites for all species ( i . e ., peaking at the Mediterranean site; Fig. S7) with higher C tot in T. cordata ( i . e ., because of its higher leaf area; Fig. S6) compared to the other species. In contrast to C leaf , T . fortunei had lower C tot in the two coldest sites because of reduced leaf area. Acclimation of photosynthesis and respiration contributed strongly to the variation in C leaf . Without considering acclimation, C leaf of T . fortunei would have decreased at the temperate and Mediterranean sites ( p < 0.05; Fig. 8 and Table S9) and remained similar than the C leaf with acclimation at more extreme climates (cold and semi-arid conditions). Similarly, T . cordata would have had a significantly lower C leaf at the Mediterranean and semi-arid sites. Contrastingly, acclimation did not modify the C leaf in I. aquifolium ( Fig. 8 ). Download figure Open in new tab Fig. 8: Differences between the modeled annual leaf-level C uptake (ΔA net ) at every site and the reference site (sub-Mediterranean). Positive values indicate higher annual leaf-level C uptake in the respective sites compared to the reference one. For each site, the lighter bars show results obtained with non-acclimated physiological traits ( i . e ., the traits from the reference site), while the darker bars correspond to the outputs of the model with acclimated physiology. Error bars indicate the uncertainty of modeled J max,25 and V Cmax,25 , R 25 , and Q 10 (n = 37–57). Significant differences between acclimated and non-acclimated simulations are represented with stars ( P < 0.05 = *; P < 0.01 = **; P < 0.001 = ***). 4. Discussion 4.1 Photosynthetic and respiratory acclimation is not higher in invasive palms than in native species Against our initial hypothesis, the invasive T . fortunei did not show a higher photosynthetic acclimation than the native species, especially compared to the evergreen I . aquifolium that showed a similar thermal plasticity ( Figs 3 & 4 ). All species had higher T opt in the warmer sites than the colder ones ( Fig. 3 ), but the two evergreen species showed similar T opt acclimation rates at almost double that of the deciduous one (about +0.6°C vs . +0.3°C per degree T air ; Fig 4 .). Such differences between functional groups can be explained by the higher need for species with long-lasting tissues to tolerate a broader range of thermal conditions ( Yamori, Hikosaka and Way, 2014 ). Despite those differences between functional groups, all our shifts in T opt were within the range of +0.3 to 0.8 °C per degree T air , as found in most studies on leaf thermal acclimation ( e . g ., Kumarathunge et al ., 2019 ; Choury et al ., 2022 ; Crous, Uddling and De Kauwe, 2022 ). The T opt shifts were probably driven by changes in the ratio between V C,max and J max , as commonly found in other C3 species ( Kattge and Knorr, 2007 ; Smith and Dukes, 2017 ), even if more work would be needed to confirm this. In contrast to previous studies ( Gunderson et al ., 2010 ; Sendall, Lusk and Reich, 2016 ; Kruse et al ., 2019 ), we also observed an increase of A opt with rising T air ( Fig. 4 ) and generally higher values at warmer sites ( Fig. 3 ). Several studies have underpinned the important role of stomatal aperture on A opt shifts with temperature, especially compared to T opt, which is generally less impacted by stomatal closure ( Kruse et al ., 2019 ; Akaji, Torimaru and Akada, 2024 ; Slot et al ., 2024 ). In our study, plants were well-watered, and VPD was as low as technically possible during the measurements, allowing us to measure the thermal acclimation of A opt mostly without the bias of increasing stomatal closure. These results are far-reaching as they suggest that VPD effects on stomata may hide the actual temperature acclimation potential of A opt driven by higher enzymatic activity in warmer environments ( Berry and Bjorkman, 1980 ). Still, similarly to T opt , T . fortunei did not show larger A opt shifts than the other species ( Fig. 3 ). Higher T opt also led to higher A opt , and for every °C change in T opt , A opt only increased by 0.11 µmol m 2 s -1 in the invasive species, while it increased up to three times more for T. cordata ( Fig. 4 ). However, T . fortunei was the only species increasing its T 80 with T air (+ 0.19°C per °C air temperature; Fig. S3). Slot and Winter (2017) found an increase in T 80 that was accompanied by the rise in the J max :V C,max ratio. Hence, a higher T 80 could be either related to a higher J max , or a lower V C,max . The higher T 80 in the invasive palm could thus indicate a decrease in V C,max or increase in J max with higher T air . Contrary to our initial hypothesis, R 25 in T . fortunei did not decrease with higher T air , showing no evidence of acclimation ( Figs 5 & 6 ). Wang et al . (2020) observed that V C,max and R 25 were decreasing with increasing T air as less active Rubisco is needed for a given value of V c,max at higher temperatures, decreasing the respiratory costs for Rubisco turnover and therefore R 25 . As R 25 of T . fortunei remained relatively constant across temperatures, it supports the notion that V C,max remained stable as well in that species. The constant R 25 may also indicate a lack of acclimation pressure, as at temperatures below 30°C, respiratory rates were low in this species ( Fig. 5 ). This corroborates that palms can display lower foliar respiration rates than dicot species below 35 °C ( Cavaleri, Oberbauer and Ryan, 2008 ). In contrast to T . fortunei , in the two native species, R 25 decreased with higher T air (−0.03 μmol CO 2 m −2 s −1 per degree T air on average; Fig. 6 ), with similar rates as in other studies ( e . g ., Reich et al ., 2016 ; Zhu et al ., 2020 ) following the consistent shifts of R 25 with T air found in a wide range of species and diverse experimental designs ( Slot and Kitajima, 2015 ; Crous, Uddling and De Kauwe, 2022 ). Contrastingly, T . fortunei shifted Q 10 while the native species did not ( Figs 5 & 6 ). Similar acclimation of Q 10 was found in other works focusing on evergreen species ( e . g ., Quan, Wang and Wang, 2020 ; Crous, Uddling and De Kauwe, 2022 ) and typically occurs when respiration is limited by low substrate availability ( Atkin and Tjoelker, 2003 ; van de Weg, Fetcher and Shaver, 2013 ). The decrease in Q 10 at high temperatures allowed the invasive palm to increase its A net at warmer sites and could partially explain the broader T 80 we observed at high temperatures (Fig. S3). Q 10 shifts have been associated with short-term changes of respiration with T air ( Atkin and Tjoelker, 2003 ) and our findings contrast with studies measuring evergreen species in climatic chambers that observed no acclimation with temperature ( e . g ., Dusenge, Madhavji and Way, 2020 ; Choury et al ., 2022 ), potentially because Q 10 also varies seasonally and its acclimation depends on the warming duration and intensity ( Atkin, Bruhn and Tjoelker, 2005 ; O’Sullivan et al ., 2013 ). In fact, all our measured parameters (A opt , T opt , T 80 , R 25 , and Q 10 ) shifted systematically with T air within two weeks in the three species. These results align with other studies ( e.g ., Kattge and Knorr, 2007 ; Kumarathunge et al ., 2019 ) stating that the commonly used 30-day acclimation period may be generally overestimated and that future works should consider using shorter acclimation periods ( Smith and Dukes, 2013 ; Reich et al ., 2016 ; Vico et al ., 2019 ). Second, all species ( i . e ., invasive and native ones) demonstrated a short acclimation duration and the invasive palm did not benefit from a more extensive photosynthetic and respiratory acclimation than native tree species. 4.2 Impact of photosynthetic and respiratory acclimation on the leaf-level and plant C budget Contrary to our expectations, the invasive palm generally showed lower C leaf and C tot than native species across all sites, even at the reference site where the palm invades the natural ecosystem ( Figs. 7 and S6). While higher C gains increase the competitivity of plant species ( Pearcy et al ., 1987 ), lower C costs in invasive plants also play a significant role in invasion processes, and T . fortunei may have benefitted from other characteristics ( e . g ., longer photosynthetic period in autumn, lower whole-plant respiration, herbivory, and tissue turnover ( Fridley et al ., 2022 ; Juillard et al ., 2024 )) to be invasive under current climatic conditions. At the reference site, T . fortunei also shares these characteristics with several other natives ( e . g ., Ilex aquifolium , Hedera helix ) and non-native evergreen species ( e . g ., Prunus laurocerasus ) currently spreading in the understory of sub-Mediterranean deciduous forests, which could confirm the important role of these characteristics in the forest composition changes ( Conedera et al ., 2018 ). The similar C leaf of T . fortunei between the sites ( Fig. 7 ) can be explained by the relatively low increase in A opt per °C T opt (+0.11 µmol m 2 s -1 per °C T opt ; Fig. 4 ), while T opt showed the highest increase with T air among the compared species (+0.60°C per °C T air ). Hence, accurate A opt measurements not biased by stomatal closure at high VPD are crucial to predict whole tree carbon exchange acclimation realistically. At the leaf level, the invasive palm performed similarly well at the reference site than in the two colder sites, while the native species had a lower C leaf ( Fig. 7 ), which shows that if T . fortunei is not damaged by frost in colder climates (Figs S6 & S7), its photosynthetic capacity allows to perform equally well as the native species. In contrast, native species acclimated more to the higher T air at the Mediterranean site than the invasive palm ( Fig 7 & S1), potentially limiting the capacity of the invasive palm to invade those environments. At the warmest site, C leaf was lower in all species. Still, high temperatures especially limited the performance of I . aquifolium due to high respiratory rates ( Figs 7 & 8 ), highlighting the critical role of respiration on the C budget at temperatures above 30°C. Acclimation of photosynthesis and respiration allowed the invasive palm to increase C uptake and reduce C losses (leading to a higher C budget) in intermediate sites with milder conditions ( Fig. 8 ). Nevertheless, the importance of acclimation was limited in extreme climates (both cold and hot ones), suggesting that shifts in photosynthetic and respiratory patterns are insufficient to compensate for the harsher climatic conditions. In contrast, acclimation allowed the native T . cordata to increase the C budget in the hottest environments ( Fig. 8 ). Still, the highest C budget was observed in the Mediterranean climate (+2.5°C), suggesting a limited enhancement of the C budget under global warming. Previous work also reported increased C gains in some temperate trees with a rise between +2 and +5 °C T air , especially because warming also leads to a certain extent to a longer growing season in deciduous trees ( Duan et al ., 2013 ; Grossiord et al ., 2022 ). On the other hand, the absence of photosynthetic and respiratory acclimation benefits for I. aquifolium ( Fig. 8 ) could be explained by the high respiratory rates that drastically reduced the C leaf in all sites, even the cold ones ( Fig 7 ). As a shade-tolerant species, I . aquifolium tends to have high leaf N content, which has been associated with high respiration ( Reich et al ., 1998 ; Niinemets, Valladares and Ceulemans, 2003 ), potentially constraining the positive effect of T air on its C budget. Overall, our results suggest that photosynthetic and respiration thermal acclimation may not be the primary underlying mechanisms driving the fast propagation of the invasive palm in recent decades. Those findings are consistent with reports of invasive species conserving their initial niche ( Verlinden and Nijs, 2010 ; Liu et al ., 2020 ; Ripley et al ., 2020 ). In addition, our results confirm that some long-lived invasive species have not reached their potential range yet, as observed in Bradley, Early and Sorte (2015) . Their range expansion could, therefore, be predicted from physiological measurements directly but not from the extrapolation of their native or current range ( Walther et al ., 2009 ). Invasive plants often depend on many traits to colonize a new environment, and future invasion patterns in a warmer environment are difficult to predict. Our results suggest that low foliar respiratory rates may provide an advantage to the invasive palm over the native plants. Given that respiration plays a vital role at high temperatures, high respiratory rates often observed in fast-growing invasive species ( Montesinos, 2022 ), could, therefore, reduce the C budget of fast-growing invasive species under higher temperatures. 5. Conclusion Using a natural temperature gradient across European study sites, we showed that acclimation of photosynthesis and respiration rapidly occurs in young plants exposed to understory conditions and can play a significant role in the C budget of both invasive and native plant species. Contrary to our expectations, the invasive palm T. fortunei did not demonstrate higher acclimation capacity than its native competitors and had a lower (but more constant) C budget overall. In central and southern Europe, T air during summer could increase by 5–7°C by the end of the century compared to 1995–2014 (SSP5-8.5) ( Carvalho, Cardoso Pereira and Rocha, 2021 ), which would decrease the C budget of all species measured in this study. It is, therefore, likely that native sub-Mediterranean species will be outcompeted by more thermophilic invasive species in the future ( Walther et al ., 2009 ; Liu et al ., 2017 ), which will likely lead to species composition changes in these forests. Data availability statement Data will be made available on the Dryad Digital Repository. Author Contributions TJ, CG, and CB planned and designed the research. All authors carried out the fieldwork. TJ, CB, and MD performed data analyses. TJ wrote the manuscript draft. All authors contributed to the review and editing of the manuscript. Acknowledgments This study was supported by the Sandoz Family Foundation. CG and CB were supported by the Swiss National Science Foundation (310030_204697). AV was supported by EVER project (CIPROM/2022/37-Prometeu program, GVA). CEAM is funded by Generalitat Valenciana. We thank Janisse Deluigi, Stéphane Jenni, Samuel Reyes, Alex Tunas Corzon, Gabor Reiss, Luis Pizarro Pietro, Eugénie Mas, Kevin Knecht, Elias Ackermann, Omar Basquet, Alice Gauthey, Krzysztof Wroński, Jolan Wicht, Margaux Didion-Gency, Valentin Meister, Patrick Favre, and Maxwell Bergström for their precious help during the management of the experimental sites and the fieldwork. We also thank Mercedes Carmona and Diana Fernández in the Forestry nursery of Guardamar del Segura (Conselleria de Medio Ambiente, Infraestructuras y Territorio, Generalitat Valenciana, ES;), Schutz Filisur Alpin Gartencenter at Filisur (CH), and the CEFE CNRS at Montpellier (FR) for hosting the experimental sites and providing water access all along the experiment. References ↵ Akaji , Y. , Torimaru , T. and Akada , S . 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