Characterization, dynamics and trophic ecology of macrofauna associated to seagrass macrophytodetritus accumulations (Calvi Bay, Mediterranean Sea)

Posidonia oceanica meadows are a major coastal Mediterranean ecosystem. Although highly productive this Mediterranean marine flower plant is not much consumed by herbivore organisms. During autumnal senescence, most (up to 90%) of the foliar primary production of P. oceanica ends in the “detrital compartment”. These dead leaves, also called “macrophytodetritus”, begin to degrade immediately inside the meadow, but a large amount will be rapidly exported to adjacent unvegetated accumulation zones, such as bare sand patches. Associated to drift macroalgae, living detached P. oceanica shoots, micro-organisms and fine sediment, these macrophytoderitus form what we call “exported P. oceanica litter”.
This exported litter is a highly dynamic habitat for a whole community of invertebrates: meiofauna (38µm < size < 500µm) and macrofauna (size ≥ 500µm) on which we focused on. This dynamic nature of exported litter could play a major structuring role in terms of abundance, diversity and trophic ecology of this vagile macrofauna community at a seasonal, annual or spatial scale, but also during stochastic, brief and very strong perturbations: resource pulses.
In this context, this PhD Thesis had 7 main objectives:

i. Characterize for the first time exhaustively the macrofauna community.
ii. Evaluate the spatiotemporal changes occurring at two different time scales in the detritus themselves and in the macrofauna community.
iii. Relating these variations with measured environmental parameters.
iv. Experimentally demonstrate the stratification occurring in a stable P. oceanica litter accumulation and the impact of this stratification on environmental conditions and on the macrofauna.
v. Experimentally demonstrate the impact of resource pulses on the exported P. oceanica litter macrofauna community.
vi. Unravel for the first time the global P. oceanica litter macrofauna food web using gut contents examinations and stable isotopes (C and N).
vii. Evaluate the spatiotemporal changes of diet preferences of this community and determine if the observed changes are really synonym of true diet changes.

This PhD Thesis demonstrated that exported P. oceanica litter was mainly composed of dead P. oceanica leaves (70-80%). It followed the natural annual cycle of P. oceanica and presented a maximum abundance in autumn just after leaves senescence. Measured environmental parameters also showed important variations linked to different factors such as force and direction of the wind, litter abundance and probably temperature. The continuous presence of the vagile macrofauna community throughout the year was demonstrated as well. This community was composed of 115 species and largely dominated by arthropods (77%), followed by annelids (12%) and mollusks (7%), while other taxa were much more anecdotal. Even if diversity is quite important, only a few species dominate largely the community. Indeed, 19 species represent more than 90% of the total abundance. One species to keep in mind: Gammarella fucicola, the most typical dominant and abundant amphipod species, representing 40-50% of the total abundance.
In addition to this general pattern, litter vagile macrofauna presented important seasonal and annual variations. In the case of several species, these variations could be linked to some measured environmental parameters, but we had to recognize that most species did not seem to be influenced by environmental parameters measured during this PhD. However, oxygen concentration was the most important environmental parameter, potentially influencing 7 of the 19 most dominant and abundant species. The experimentally demonstrated physico-chemical stratification occurring inside litter accumulations was strongly related to this oxygen parameter. Indeed we demonstrated that several species were distributed in the different layers of a litter accumulation according to oxygen concentration and to a lesser extent, to nutrients concentration (mostly NH4).
Besides, smaller time scale sampling allowed the identification of several stormy events corresponding to the definition of resource pulses. These pulses were demonstrated to play a potentially important role on the structure of the macrofauna community, favoring importantly the detritivore species and hypoxia tolerant species. It was also demonstrated that resource pulses could induce diet switching increasing the consumption of dead P. oceanica leaves just after the events, potentially increasing the litter decomposition by the macrofauna.

The trophic web described in this PhD Thesis was composed of several trophic levels, from the primary herbivore/detritivore consumer, to second order carnivore predators. Different dietary preferences were highlighted, but major information was that dead P. oceanica leaves were ingested by a majority (85%) of the sampled species. Moreover, stable isotope analysis confirmed that P. oceanica litter was assimilated by most primary consumers and this “detrital signal” could be identified to the upper trophic levels, which is an argument in favor of the importance of macrofauna as major dead P. oceanica leaves decomposers. This also highlighted their potential role in terms of organic matter transfer from the P. oceanica meadow itself to the Mediterranean coastal food webs. Seasonal variations were observed in terms of trophic niches, and SIAR mixing model confirmed that this variability was sometimes caused by real diet modifications, potentially linked to the variable availability of food sources.
This PhD Thesis, combining standardized sampling at two different time scales, trophic web analysis (gut contents and stable isotopes) and original experimentation allowed us to describe a diverse and abundant macrofauna community associated to P. oceanica exported litter, its temporal variations, potential responses to resource pulses as well as the link existing between some species and measured environmental parameters. This PhD also described the food web of this community and demonstrated the importance of dead P. oceanica leaves as food source for many invertebrates composing this community. These invertebrates thus seemed to play an important role in both litter decomposition and organic matter flux from the P. oceanica meadow to the Mediterranean coastal food webs.