# Defense and Adaptive Strategies of Crithmum maritimum L. Against Insect Herbivory: Evidence of Phenotypic Plasticity

## TL;DR

This study shows how sea fennel adapts to insect attacks by changing its traits depending on environmental conditions and herbivory levels.

## Contribution

The paper provides the first evidence of insect herbivory in Crithmum maritimum and reveals its adaptive defense strategies.

## Key findings

- Low-herbivory populations show tolerance strategies with higher phenolics and antioxidants.
- High-herbivory populations exhibit resistance strategies with elevated apiol and terpenes.
- Phenotypic plasticity is evident in populations exposed to varying herbivory and climate conditions.

## Abstract

Insect herbivory exerts strong selective pressure on plants, yet no study has documented its effects on the halophytic Apiaceae Crithmum maritimum L. (sea fennel). Here, we present the first evidence of natural insect attack on this species, based on five Tunisian coastal populations distributed along a transparent bioclimatic gradient—from sub-humid to semi-arid—and exposed to different levels of herbivory. We implemented an integrative, multi-trait analytical design encompassing morphological, biochemical, mineral, and lipophilic datasets. Each dataset was explored through a suite of complementary multivariate analyses, including ANOVA coupled with Tukey’s HSD, principal component analysis (PCA), partial least squares discriminant analysis (PLS-DA) with variable-importance-in-projection (VIP) scores, correlation matrices, hierarchical clustering, and distance-based redundancy analysis (dbRDA). This integrative strategy provided a robust framework for disentangling the complex trait associations underlying two distinct defense syndromes. Populations from low-herbivory, sub-humid sites (Tabarka, Bizerte, Tunisia) showed higher levels of phenolics, tannins, antioxidants, sterols, PUFA, and structural robustness, indicating a tolerance strategy. Conversely, high-herbivory, semi-arid sites (Haouaria, Monastir, Tunisia) were marked by elevated apiol and terpene levels, sodium and phosphorus accumulation, and reproductive adjustments, reflecting a resistance strategy. The site Cap Negro exhibited a transitional expression, revealing intermediate phenotypic plasticity. These findings show that herbivory intensity and bioclimatic conditions jointly influence the defense syndromes of C. maritimum, emphasizing its remarkable phenotypic plasticity and providing the first ecological evidence of insect herbivory in sea fennel.