Despite the great interest in characterizing the functional structure and resilience of functional groups in natural communities, few studies have examined in which way the roles and relationships of coexisting species change during community succession, a fundamental and natural process that follows the release of new resources in terrestrial and aquatic ecosystems. Variation in algal traits that characterize different phases and stages of community succession on rocky shores are likely to influence the magnitude, direction of effects, and the level of redundancy and complementarity in the diverse assemblage of herbivores. Two separate field experiments were conducted to quantify per capita and population effects and the functional relationship (i.e., redundancy or complementarity) of four herbivore species found in central Chile during early and late algal succession. The first experiment examined grazer effects on the colonization and establishment of early-succession algal species. The second experiment examined effects on the late-successional, dominant corticated alga Mazzaella laminarioides. Complementary laboratory experiments with all species and under natural environmental conditions allowed us to further characterize the collective effects of these species. We found that, during early community succession, all herbivore species had similar effects on the ephemeral algae, ulvoids, but only during the phase of colonization. Once these algae were established, only a subset of the species was able to control their abundance. During late succession, only the keyhole limpet Fissurella crassa could control corticated Mazzaella. The functional relationships among these species changed dramatically from redundant effects on ephemeral algae during early colonization, to a more complementary role on established early-successional algae, to a dominant (i.e., keystone) effect on late succession. This study highlights that functional relationship within consumer assemblages can vary at different phases and times of community succession. Differentiation in herbivore roles emphasizes the need to evaluate consumer's impacts through different times of community succession, and through experimental manipulations to make even broad predictions about the resilience or vulnerability of diverse intertidal assemblages to human disturbances.
In consumer assemblages, the organization of individual foraging behavior, as well as spatial distribution, can largely determine environmental risks, and the intensity of intra-and interspecific interactions. We characterized distributional and behavioral patterns of the most common benthic grazers coexisting in the rocky shores of central Chile: the chiton Chiton granosus, the pulmonate limpet Siphonaria lessoni, the scurrinid limpet Scurria araucana and the keyhole limpet Fissurella crassa. C. granosus and F. crassa were strictly nocturnal foragers whereas S. lessoni foraged during daytime. Most S. araucana individuals foraged at night, but daytime foraging was also common. The spatial distribution at resting varied from aggregated for C. granosus and S. lessoni to a more dispersive pattern for F. crassa and S. araucana. C. granosus dispersed slightly from aggregation when foraging whereas S. lessoni foraged in tight conspecific aggregations. Foraging excursions varied from over 60 cm in F. crassa to less than 7 cm in S. araucana. Homing behavior ranged from extreme fidelity in F. crassa to low fidelity in S. lessoni. Positive associations were observed between C. granosus and F. crassa during resting and foraging whereas negative associations were observed between these species and S. lessoni. These general patterns varied little between 2 sites separated by a few kilometers. Interspecific competition might be important in structuring this guild, but it may affect only some species pairs. Direct interference in the use of shelter or while foraging, rather than food exploitation, seems the most likely mechanism. The marked differences in individual behavior among species, despite ample diet overlap, might translate into different functional effects, which should be explored in future experiments.
Aim Topographic complexity is widely accepted as a key driver of biodiversity, but at the patch‐scale, complexity–biodiversity relationships may vary spatially and temporally according to the environmental stressors complexity mitigates, and the species richness and identity of potential colonists. Using a manipulative experiment, we assessed spatial variation in patch‐scale effects of complexity on intertidal biodiversity. Location 27 sites within 14 estuaries/bays distributed globally. Time period 2015–2017. Major taxa studied Functional groups of algae, sessile and mobile invertebrates. Methods Concrete tiles of differing complexity (flat; 2.5‐cm or 5‐cm complex) were affixed at low–high intertidal elevation on coastal defence structures, and the richness and abundance of the colonizing taxa were quantified after 12 months. Results The patch‐scale effects of complexity varied spatially and among functional groups. Complexity had neutral to positive effects on total, invertebrate and algal taxa richness, and invertebrate abundances. However, effects on the abundance of algae ranged from positive to negative, depending on location and functional group. The tidal elevation at which tiles were placed accounted for some variation. The total and invertebrate richness were greater at low or mid than at high intertidal elevations. Latitude was also an important source of spatial variation, with the effects of complexity on total richness and mobile mollusc abundance greatest at lower latitudes, whilst the cover of sessile invertebrates and sessile molluscs responded most strongly to complexity at higher latitudes. Conclusions After 12 months, patch‐scale relationships between biodiversity and habitat complexity were not universally positive. Instead, the relationship varied among functional groups and according to local abiotic and biotic conditions. This result challenges the assumption that effects of complexity on biodiversity are universally positive. The variable effect of complexity has ramifications for community and applied ecology, including eco‐engineering and restoration that seek to bolster biodiversity through the addition of complexity.
Aim To examine the role of ocean temperature and chemistry as drivers of interpopulation differences in multiple phenotypic traits between rear and leading edge populations of two species of limpet. Location The coast of north‐central Chile, western South America. Taxon Mollusca, Gastropoda (Lottidae). Methods We used field and laboratory experiments to study the ecology and physiology of individuals from populations located at the overlap of the rear and leading edges of their respective geographical distributions. At the same time, we characterized local environmental regimes, measuring seawater physical and chemical properties. Results Towards the edge of their range, individuals from the leading edge species gradually reduced their shell length, metabolic rate and thermal response capacity, and increased carbonate content in their shells. Individuals of the rear edge species showed dissimilar responses between sites. Contrasting behavioural responses to experimental heating reconciled observations of an unintuitive higher maximal critical temperature and a smaller thermal safety margin for individuals of the rear edge populations. Physical–chemical characterization of seawater properties at the site located on the core of the upwelling centre showed extreme environmental conditions, with low oxygen concentration, high pCO2 and the episodic presence of corrosive seawater. These challenging environmental conditions were reflected in reduced growth for both species. Main conclusions We found different spatial patterns of phenotypic plasticity in two sister species around the leading and trailing edges of their distributions. Our results provide evidence that environmental conditions around large upwelling centres can maintain biogeographical breaks through metabolic constraints on the performance of calcifying organisms. Thus, local changes in seawater chemistry associated with coastal upwelling circulation emerge as a previously overlooked driver of marine range edges.
Examining the co-occurrence of taxonomically similar species can provide important information about their niches and coexistence. Segregation at smaller scales can be especially relevant for grazers living at the edge of their geographic distribution, because environmental factors can lead to similar distribution. Related grazer species may show dispersive, i.e. uniform, distribution at small scales (few centimetres) to reduce interference among individuals. We examined intra-and interspecific spatial distribution and habitat use in 2 phylogenetically related intertidal limpets, Scurria viridula and S. zebrina, at the polar and equatorial edge of their geographic distribution, respectively, and in S. araucana, a widely distributed species that overlaps the range of the other 2 species across the southeastern Pacific. S. viridula and S. zebrina overlapped in a narrow geographic zone (ca. 250 km) and reached relatively similar densities and sizes. Intraspecific spatial structure estimated through autocorrelation and individual-to-indi vidual distances was random for S. viridula and variable for S. zebrina and S. araucana, depending on the scale considered; S. zebrina was aggregated at the individual-to-individual distances, while S. araucana was mostly random at this scale. Segregated distribution between S. viridula and S. zebrina was observed at the finer scale, whereas the association with S. araucana was random. Abundance of limpets loosely followed major habitats, namely bare rock and the alga Mazzaella laminarioides, which showed patchy distributions. We suggests that similarity in population traits between S. viridula and S. zebrina found in the overlap zone may be compensated by small-scale spatial segregation. Thus, at the edge of their geographic distribution, co existence between related species may be influenced by spatial niche differentiation driven by habitat suitability or competition.
Understanding the ecological factors regulating exploited natural communities is important in establishing conservation and management strategies. Since the 1960s, artisanal fishermen have harvested up to 300 000 dry tons yr −1 of wild populations of Lessonia spp. kelps. Adult kelps form a key habitat on the rocky shores of Chile, and benthic grazers regulate kelp populations by grazing or bulldozing microscopic and juvenile stages. To establish the role of the grazer assemblage in the recovery of kelp stands following harvesting, we conducted a manipulative experiment simulating artisanal kelp extraction and then manipulating the presence of grazers. We followed community succession and spatial distribution of L. berteroana recruits in manipulated and control rocky shore platforms for ca. 12 mo. Inter-individual distances of recruits and spatial autocorrelation analyses showed that the presence of grazers determined a patchy distributional pattern of L. berteroana. The aggregated spatial pattern of kelp recruits in the presence of grazers was followed by numerous coalescence events between small holdfasts, but no coalescence events were observed between recruits in the grazer removal areas. Our results suggest that grazing and recruit coalescence play an important role in the recovery of kelp stands following artisanal harvesting. Incorporating these processes into conservation and management strategies may bolster current strategies, which are based solely on the spatial structure of kelp stands.
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