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Ecol Evol
2017 Jun 15;715:5542-5559. doi: 10.1002/ece3.2857.
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Functional roles of an engineer species for coastal benthic invertebrates and demersal fish.
Chaalali A
,
Brind'Amour A
,
Dubois SF
,
Le Bris H
.
Abstract
Through their tissues or activities, engineer species create, modify, or maintain habitats and alter the distribution and abundance of many plants and animals. This study investigates key ecological functions performed by an engineer species that colonizes coastal ecosystems. The gregarious tubiculous amphipod Haploops nirae is used as a biological model. According to previous studies, the habitat engineered by H. nirae (i.e., Haploops habitat) could provide food and natural shelter for several benthic species such as benthic diatoms belonging to the gender Navicula, the micrograzer Geitodoris planata, or the bivalve Polititapes virgineus. Using data from scientific surveys conducted in two bays, this study explored whether (1) the Haploops sandy-mud community modifies invertebrate and ichthyologic community structure (diversity and biomass); (2) H. nirae creates a preferential feeding ground; and (3) this habitat serves as a refuge for juvenile fish. Available Benthic Energy Coefficients, coupled with more traditional diversity indices, indicated higher energy available in Haploops habitat than in two nearby habitats (i.e., Sternaspis scutata and Amphiura filiformis/Owenia fusiformis habitats). The use of isotopic functional indices (IFIs) indicated (1) a higher functional richness in the Haploops habitat, related to greater diversity in food sources and longer food chains; and (2) a higher functional divergence, associated with greater consumption of a secondary food source. At the invertebrate-prey level, IFIs indicated little specialization and little trophic redundancy in the engineered habitat, as expected for homogenous habitats. Our results partly support empirical knowledge about engineered versus nonengineered habitats and also add new perspectives on habitat use by fish and invertebrate species. Our analyses validated the refuge-area hypothesis for a few fish species. Although unique benthic prey assemblages are associated with Haploops habitat, the hypothesis that it is a preferential feeding area was not verified. However, specialist feeding behavior was observed for predators, which calls for further investigation.
Figure 1. Maps of the two studied bays with local bathymetry with an additional map of France showing their respective locations. (a) Bay of Vilaine. (b) Bay of Concarneau. The three study habitats, that is, Haploops habitat, Amphiura/Owenia habitat, and Sternaspis habitat, are distinguished using a gradient of gray colors. The mean locations of hauls are presented in red circles numbered from 1 to 25
Figure 2. (a) Factorial correspondence analysis of fish community abundances for the Bay of Concarneau. (b) Factorial correspondence analysis of fish community abundances for the Bay of Vilaine. Both factorial maps display the sampling sites and the fish species (with a cosine2 > 0.4) with the addition of the ellipses of dispersion of each habitat. Megafaunal invertebrate species were considered as supplementary variables and superimposed to the factorial map color‐coded in violet and numbered according to Appendix Table S1
Figure 3. Two‐dimensional isotopic δ‐spaces (δ15N–δ13C) including the stable isotopic compositions of producers (organic matter sources), and first and second consumers (benthic mega‐invertebrates and fish). Species (labeled with numbers, see Appendix Table 1) were color‐coded according to the HAC grouping. All habitats for each bay were represented as specified in the top‐right corner of each biplot. It is worth noting that each bay has its own δ13C and δ15N scales
Figure 4. Contribution of the Available Benthic Energy Coefficient (ABEC) groups to the diet of predators (fish and invertebrates) in the Haploops habitat of the Bay of Vilaine using SIAR mixing model. Box plots illustrate the 25th, 50th, and 75th percentiles; the whiskers indicate the 10th and 90th percentiles. The details of ABEC‐based sources’ compositions (i.e., ABEC groups 1–3) are given in the Appendix Table S1. Top left to bottom right panels display the results for mobility group 1 (up to 100 m), mobility group 2 (between 100 m and 1 km), mobility group 3 (between 1 and 10 km), and mobility group 4 (above 10 km). The black circle indicates the ABEC source associated with the highest accessible biomass, and the black square indicates the source with the highest energetic supply (see Section 2 for complementary information)
Figure 5. Contribution of the Available Benthic Energy Coefficient (ABEC) groups to the diets of predators in the Haploops habitat of the Bay of Concarneau, using SIAR mixing model. Box plots illustrate the 25th, 50th, and 75th percentiles; the whiskers indicate the 10th and 90th percentiles. The details of ABEC‐based sources’ compositions (i.e., ABEC groups 1–3) are given in the Appendix Table S1. From top left to bottom are displayed the results for mobility group 1 (up to 100 m), mobility group 2 (between 100 m and 1 km), and mobility group 4 (above 10 km). Due to a limited number of data to perform the analysis of mobility group 3, result was not presented here. The black circle indicates the ABEC source associated with the highest accessible biomass, and the black square indicates the source with the highest energetic supply (E
i × πi × R
i)
Figure 6. Size spectra of the whole fish community in Haploops habitat (in white), and outside Haploops habitats together (in black) (a) For the Bay of Concarneau. Description of species composition for the Bay of Concarneau: (b) Outside Haploops. (c) For Haploops habitat
Figure 7. Diagram synthesizing the functional roles of the ecosystem engineer Haploops nirae based on the results found in this study (in italics) and from Rigolet (2013). The scheme is adapted from Jones et al. (1994). (1) indicates the direct alteration of the physical environment via tubes building. (2) indicates the modification of organic matter sources (filtering activity and fecal pellets production). The two effects have consequences in terms of diversity of invertebrates and fish species inhabiting (temporarily or not) Haploops habitats. They also result in functional changes (in green). H1: the hypothesis on the preferential feeding ground. H2: hypothesis on the refuge area
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