Ecophysiological responses of <i>Stipa tenacissima</i> L. under grassland degradation in a Mediterranean arid bioclimate

doi:10.1016/j.jaridl.2026.04.006

Journal of Arid Land

2026, Vol. 18

Issue (4): 657-675 DOI: 10.1016/j.jaridl.2026.04.006 CSTR: 32276.14.JAL.20250447

Research article

Ecophysiological responses of Stipa tenacissima L. under grassland degradation in a Mediterranean arid bioclimate

Khouloud KRICHEN^*(

), Mohamed CHAIEB

Laboratory of Biodiversity and Ecosystems in Arid Environments (LEBIOMAT), Faculty of Sciences, University of Sfax, Sfax 3000, Tunisia

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Abstract

Under arid and semi-arid bioclimates, steppes are increasingly threatened by anthropogenic disturbance and climatic variability, which strongly affects ecosystem functioning and subsequently leads to desertification. We investigated the morphological and physiological responses of Stipa tenacissima L. across three disturbance levels (undisturbed, slightly disturbed, and highly disturbed) in three Tunisian steppe areas (Kasserine, Sidi Bouzid, and Sfax). Morphological and physiological traits were monitored over one year, together with microclimatic variables. Result showed that disturbance was a strong driver of plant functional dynamics, with significant effects on all traits and strong interactions with site and season. Disturbance reduced photosynthetic activity and water use efficiency, particularly in Sfax, where plants adopted conservative strategies (i.e., higher leaf dry matter content and reduced leaf area). In contrast, undisturbed populations maintained a stronger coordination between physiological and morphological traits. Seasonal analyses revealed that disturbance amplified physiological stress with limited recovery. Heatmap analyses further showed that disturbance weakened trait coordination and reshaped trade-offs between acquisitive and conservative traits. Partial least squares-path modeling showed that morphology strongly drove physiological performance (path coefficient=0.48). Disturbance (path coefficient=0.41) and tussock cover (path coefficient=0.47) influenced morphology both directly and indirectly through their effects on physiology. In conclusion, S. tenacissima adjusts physiological and morphological traits under disturbance, favoring stress tolerance, while undisturbed sites maintain high physiological efficiency and coordinated trait integration, reflecting a trade-off between survival and performance while overriding local site differences. Disturbance strongly restructures trait networks, drives site-specific adjustments, and modulates the seasonal balance between morphological stability and physiological flexibility.

Key words： disturbance physiological traits morphological traits arid and semi-arid steppes plant adaptation

Received: 09 September 2025 Published: 30 April 2026

Corresponding Authors: ^*Khouloud KRICHEN (E-mail: kh.krichen@gmail.com)

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Cite this article:

Khouloud KRICHEN, Mohamed CHAIEB. Ecophysiological responses of Stipa tenacissima L. under grassland degradation in a Mediterranean arid bioclimate. Journal of Arid Land, 2026, 18(4): 657-675.

URL:

http://jal.xjegi.com/10.1016/j.jaridl.2026.04.006 OR http://jal.xjegi.com/Y2026/V18/I4/657

Fig. 1 Study area including Kasserine, Sidi Bouzid, and Sfax sites in Tunisia, and classification of disturbance level

Fig. 2 Daily estimated photosynthetically active radiation (PAR; a), precipitation (b), relative humidity (RH; c), and air temperature (d) from January to December 2024 across Kasserine, Sidi Bouzid, and Sfax sites in Tunisia

Table 1 Classification of disturbance levels based on normalized difference vegetation index (NDVI), difference in NDVI (DNDVI), and field-measured tussock cover (TC) of Stipa tenacissima L. across Kasserine, Sidi Bouzid, and Sfax sites in Tunisia

Trait	Site	Parameter	Disturbance level
Trait	Site	Parameter	Disturbed	Slightly disturbed	Undisturbed
Physiological trait	Sidi Bouzid	A (µmol CO₂/(m²•s))	1.199±0.081^a	1.446±0.066^a	2.720±0.082^b
		g_s (mol H₂O/ (m²•s))	0.075±0.002^ab	0.072±0.003^a	0.082±0.002^b
		E (mmol H₂O/ (m²•s))	0.264±0.019^a	0.292±0.014^a	0.255±0.009^a
		C_i(µmol/mmol)	115.919±8.290^a	160.749±6.852^b	134.466±3.814^ab
		WUE (µmol CO₂/mol H₂O)	20.280±1.636^a	24.201±1.322^a	33.391±1.280^b
	Kasserine	A (µmol CO₂/(m²•s))	1.459±0.073^a	2.242±0.078^b	2.722±0.091^c
		g_s (mol H₂O/(m²•s))	0.056±0.002^a	0.055±0.001^a	0.057±0.002^a
		E (mmol H₂O/(m²•s))	0.306±0.024^a	0.334±0.022^ab	0.396±0.022^b
		C_i(µmol/mmol)	166.730±10.296^a	208.422±8.218^b	234.178±8.999^b
		WUE (µmol CO₂/mol H₂O)	26.036±0.959^a	42.755±1.819^b	49.342±1.703^c
	Sfax	A (µmol CO₂/(m²•s))	1.606±0.117^a	1.452±0.063^a	2.314±0.101^b
		g_s (mol H₂O/(m²•s))	0.072±0.005^a	0.067±0.004^a	0.099±0.003^b
		E (mmol H₂O/(m²•s))	0.241±0.022^a	0.315±0.018^b	0.198±0.016^a
		C_i(µmol/mmol)	129.881±7.321^a	160.660±7.097^b	151.702±6.857^ab
		WUE (µmol CO₂/mol H₂O)	22.696±1.116^a	28.981±1.551^b	23.977±0.997^a
Morphological trait	Sidi Bouzid	BL (cm)	3.111±0.119^a	2.219±0.108^b	1.755±0.089^c
		GL (cm)	1.435±0.077^a	2.714±0.086^b	2.794±0.101^b
		LL (cm)	22.186±0.647^a	25.45±0.585^b	30.606±0.727^c
		SLA (cm²/g)	16.818±0.228^a	14.710±0.175^b	14.407±0.185^b
		LDMC (g)	604.589±5.398^a	561.642±5.858^b	592.480±6.054^b
		LTh (cm)	0.010±0.0002^a	0.012±0.0002^b	0.012±0.0002^b
		D (m)	1.604±0.034^a	2.929±0.026^b	3.648±0.081^c
		H (m)	0.623±0.017^a	0.891±0.013^b	1.017±0.014^c
	Kasserine	BL (cm)	2.722±0.098^a	1.253±0.088^b	1.111±0.074^b
		GL (cm)	1.870±0.092^a	3.083±0.109^b	4.354±0.130^c
		LL (cm)	19.318±0.266^a	34.277±0.915^b	34.394±0.969^b
		SLA (cm²/g)	17.101±0.233^a	15.727±0.200^b	13.564±0.121^c
		LDMC(g)	543.438±3.480^a	559.023±4.311^ab	566.750±5.716^b
		LTh (cm)	0.011±0.0002^a	0.012±0.0002^a	0.013±0.0002^b
		D (m)	1.521±0.020^a	1.919±0.050^b	1.983±0.054^b
		H (m)	0.389±0.004^a	0.845±0.020^b	1.020±0.021^c
	Sfax	BL (cm)	2.769±0.103^a	2.000±0.100^b	2.287±0.074^b
		GL (cm)	1.935±0.078^a	2.852±0.063^b	2.148±0.081^b
		LL (cm)	20.829±0.309^a	26.621±0.343^b	30.177±0.367^c
		SLA (cm²/g)	18.925±0.139^a	16.221±0.121^b	16.332±0.110^b
		LDMC (g)	480.598±4.142^a	556.711±4.915^b	566.194±4.806^b
		LTh (cm)	0.011±0.0001	0.011±0.0001	0.011±0.0001
		D (m)	0.779±0.040^a	1.379±0.066^b	2.276±0.061^c
		H (m)	0.597±0.014^a	0.799±0.011^b	0.832±0.015^b

Table 2 Morphological and physiological traits of S. tenacissima across Sidi Bouzid, Kasserine, and Sfax sites under different disturbance levels

Variable	Effect	Chi-square	df	P
Physiological trait
A	Site	125.86	2	<0.001
	Disturbance	1009.84	2	<0.001
	Month	1510.82	11	<0.001
	Site×Disturbance	92.14	4	<0.001
	Site×Month	257.97	22	<0.001
	Disturbance×Month	612.02	22	<0.001
	Site×Disturbance×Month	383.76	44	<0.001
g_s	Site	174.01	2	<0.001
	Disturbance	90.60	2	<0.001
	Month	267.60	11	<0.001
	Site×Disturbance	84.83	4	<0.001
	Site×Month	264.59	22	<0.001
	Disturbance×Month	136.62	22	<0.001
	Site×Disturbance×Month	568.58	44	<0.001
E	Site	75.96	2	<0.001
	Disturbance	52.13	2	<0.001
	Month	427.12	11	<0.001
	Site×Disturbance	59.98	4	<0.001
	Sites×Month	194.82	22	<0.001
	Disturbance×Month	169.61	22	<0.001
	Site×Disturbance×Month	219.32	44	<0.001
C_i	Site	161.03	2	<0.001
	Disturbance	130.11	2	<0.001
	Month	162.62	11	<0.001
	Site×Disturbance	30.69	4	<0.001
	Site×Month	434.12	22	<0.001
	Disturbance×Month	446.95	22	<0.001
	Site×Disturbance×Month	1333.33	44	<0.001
WUE	Site	297.81	2	<0.001
	Disturbance	232.47	2	<0.001
	Month	425.16	11	<0.001
	Site×Disturbance	102.05	4	<0.001
	Site×Month	129.13	22	<0.001
	Disturbance×Month	322.01	22	<0.001
	Site×Disturbance×Month	315.66	44	<0.001
Morphological trait
BL	Site	138.82	2	<0.001
	Disturbance	319.66	2	<0.001
	Month	74.87	11	<0.001
	Site×Disturbance	77.25	4	<0.001
	Site×Month	110.64	22	<0.001
	Disturbance×Month	103.63	22	<0.001
	Site×Disturbance×Month	112.43	44	<0.001
GL	Site	155.71	2	<0.001
	Disturbance	639.27	2	<0.001
	Month	49.45	11	<0.001
	Site×Disturbance	145.51	4	<0.001
	Site×Month	79.87	22	<0.001
	Disturbance×Month	266.97	22	<0.001
	Site×Disturbance×Month	179.39	44	<0.001
LL	Site	47.58	2	<0.001
	Disturbance	773.08	2	<0.001
	Month	19.36	11	<0.001
	Site×Disturbance	130.50	4	<0.001
	Site×Month	58.16	22	<0.001
	Disturbance×Month	109.21	22	<0.001
	Site×Disturbance×Month	210.34	44	<0.001
SLA	Site	329.43	2	<0.001
	Disturbance	637.76	2	<0.001
	Month	98.51	11	<0.001
	Site×Disturbance	82.86	4	<0.001
	Site×Month	193.06	22	<0.001
	Disturbance×Month	69.21	22	<0.001
	Site×Disturbance×Month	268.07	44	<0.001
LDMC	Site	261.79	2	<0.001
	Disturbance	103.55	2	<0.001
	Month	74.50	11	<0.001
	Site×Disturbance	283.73	4	<0.001
	Site×Month	234.33	22	<0.001
	Disturbance×Month	115.75	22	<0.001
	Site×Disturbance×Month	165.22	44	<0.001
LTh	Site	60.74	2	<0.001
	Disturbance	168.70	2	<0.001
	Month	119.36	11	<0.001
	Site×Disturbance	166.21	4	<0.001
	Site×Month	191.73	22	<0.001
	Disturbance×Month	77.15	22	<0.001
	Site×Disturbance×Month	242.78	44	<0.001
D	Site	1596.62	2	<0.001
	Disturbance	1858.59	2	<0.001
	Month	197.36	11	<0.001
	Site×Disturbance	539.10	4	<0.001
	Site×Month	120.28	22	<0.001
	Disturbance×Month	168.93	22	<0.001
	Site×Disturbance×Month	636.17	44	<0.001
H	Site	203.04	2	<0.001
	Disturbance	2285.14	2	<0.001
	Month	101.31	11	<0.001
	Site×Disturbance	580.67	4	<0.001
	Site×Month	60.43	22	<0.001
	Disturbance×Month	74.09	22	<0.001
	Site×Disturbance×Month	498.82	44	<0.001

Table 3 Results of analysis of variation (ANOVA) for physiological and morphological variables among different sites, disturbance, and months

Fig. 3 Correlation heatmaps showing pairwise Spearman's correlation coefficients among physiological, morphological, and environmental variables in disturbed (a), undisturbed (b), and slightly disturbed (c) sites. T, temperature; PAR, photosynthetically active radiation; P, precipitation; RH, relative humidity. *, P<0.050 level; **, P<0.010 level; ***, P<0.001 level.

Fig. 4 Seasonal variation of morphological (a) and physiological (b) indices across disturbed, slightly disturbed, and undisturbed sites, as predicted by generalized additive models (GAMs). Shaded areas represent 95% confidence intervals.

Fig. 5 Partial least squares-path modeling (PLS-PM) linking disturbance indicators (DNDVI and TC (tussock cover)) to Physio (physiological) and Morpho (morphological) traits. Standardized path coefficients are shown on the arrows. Disturbance indicators affected morphology through two pathways: a moderate effect of DNDVI and a stronger effect of TC.


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