Modelling prey-predator interactions in Messina beachrock pools
Introduction
Interaction among species, populations and communities are at the base of ecosystem equilibria (Deangelis, Waterhouse, 1987, Loreau, 2000). Among the different interactions, the prominent role played by prey-predator relationships is basic to gain an understanding on ecosystem dynamics (Ortiz et al., 2013).
Prey-predator interaction is a well-known topic, both in wild and experimental conditions. In wild environments it involves several behavioral strategies, such as selection of prey according to suitable morphological and behavioral features, detection and capture, camouflage, cooperation and defense mechanisms (e.g. release of repulsive chemical compounds, evasive movements, etc.).
In a natural environment, the interaction between two species can be hard to determine, due to the numerous factors affecting the whole ecosystem balance (Gabel et al., 2011). This is especially true in high biodiversity areas characterized by a rich variety of mutually-interacting plant and animal communities.
The Strait of Messina (Sicily, Italy), known for the peculiar and diversified marine habitats (Spanó, 1998, Spanó, De Domenico, 2017), includes a long-shore rocky structure known as “Messina beachrock”that represents, in this respect, an appropriate case-study (Cosentino and Giacobbe, 2015). Such worldwide distributed structure may occur from the intertidal to the upper subtidal zone (Al-Zamel, Al Sarkawi, Khader, 2007, Spurgeon, Davis Jr., Shinnu, 2003) and are characterized by large fluctuations of both biotic and abiotic conditions due to, for example, tidal cycles and storms (Kelletat, 2006, Russell, McIntire, 1965, Wainwright, Koehl, 1976). Messina beachrock, preliminary described in Bottari et al. (2005), is characterized by a microtidal regime, similar to that encountered in other known formations, e.g., in eastern Mediterranean (Vousdoukas et al., 2009). The interest in studying such zones also arises from the fact that they represent sensitive indicators of the effects of climate change in coastal ecosystems (Mauersberger, 1993, Menge, Chan, Lubchenco, 2008, Wethey, Brin, Helmuth, Mislan, 2011, Xu, Chen, Li, Huang, Li, 2011).
The species inhabiting these extreme environments need morphological, physiological and behavioral adaptations (Gomes et al., 1998). Consequently, settled communities are markedly oligotypic. In this ultra-simplified context, some evidences of prey-predator interactions between benthic species have been preliminary reported (Capillo et al., 2018a). The case-study concerns the top-snail, Phorcus turbinatus (Born, 1778), and the hermit crab, Clibanarius erythropus (Latreille, 1818). Indeed, it is known that this latter feeds upon the gastropod and steals the molluscan shell as an additional protection against predators (Abrams, 2019).
The present study aims at elucidating some ecosystem dynamics taking place in the peculiar beachrock formation. In particular, the main goals are: (i) to describe the local megabenthic communities; (ii) to individuate possible prey-predator dynamics on the basis of the analysis of the factors influencing and determining organisms distribution and abundance; (iii) to theoretically describe such prey-predator interactions through a mathematical model able to qualitatively capture the most relevant behavioral aspects.
The paper is organized as follows. In Section 2, details on the investigated area, sampling methodology, variables, statistical tools used in data analysis, mathematical model and numerical simulations tools are provided. In particular, we discuss a simple prey-predator model and address a linear stability analysis to determine the local stability character of the equilibria against small perturbations as well as to explore the occurrence of Hopf bifurcation responsible for oscillatory behaviors. In Section 3, results of statistical analysis on ecological parameters are exposed, the outcome of visual census procedure is discussed, a multivariate analysis is performed and the occurrence of possible prey-predator interactions is assessed. In the same section, the different dynamical regimes observed in situ are mathematically described, all the model parameters are identified and numerical simulations are carried out to qualitatively reproduce the various population dynamics that followed one another throughout the sampling period. Discussions on ecological implications, mathematical assumptions and future developments of this work are reported in the last section.
Section snippets
Materials and methods
In this section, we first briefly describe the investigated habitat. Then, the experimental and statistical tools adopted in the post-processing analyses are illustrated.
Results
In this section we report the outcomes of in situ measurements and theoretical investigations. In particular, we inspect temporal and spatial variations of abiotic parameters and of the main biological indexes characterizing the macrofaunal community. Then we address a multivariate analysis aimed at evaluating the most statistically significant species inhabiting Messina beachrock pools. Finally, we provide a mathematical description of prey-predator interactions, carry out the identification
Discussion
Although beachrock formations are known from several Mediterranean coasts, the related benthic assemblages have been poorly investigated. Such environments, similarly to other shallower rocky formations (Milazzo et al., 2004), is subject to strong environmental (e.g. waves, drying, rain) and anthropogenic stressors (e.g. trampling), allowing the settlement of strictly oligotypic communities.
The Messina beachrock, scarcely affected by the local microtidal regime but exposed to seasonal
CRediT authorship contribution statement
S. Savoca: Conceptualization, Software, Formal analysis, Writing - original draft. G. Grifó: Software, Formal analysis, Writing - original draft. G. Panarello: Investigation. M. Albano: Investigation, Visualization. S. Giacobbe: Supervision, Writing - review & editing. G. Capillo: Conceptualization, Validation, Supervision, Writing - review & editing. N. Spanó: Supervision, Writing - review & editing. G. Consolo: Conceptualization, Methodology, Supervision, Writing - review & editing.
Declaration of Competing Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Acknowledgments
The authors gratefully thank the anonymous reviewers for their insightful comments. G. Grifó and G. Consolo acknowledge support from INdAM-GNFM and from Italian MIUR through project PRIN2017 “Multiscale phenomena in Continuum Mechanics: singular limits, off-equilibrium and transitions” (project number:2017YBKNCE).
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