Intraspecific physiological variability of the gastropod Littorina saxatilis related to the vertical shore gradient in the White and North Seas

 Physiological responses to desiccation and temperature stress as well as behavioural responses to fast and abrupt environmental changes were investigated in high- and low-shore Littorina saxatilis (Olivi) from several populations from the White and North Seas. Variations in evaporation rates, resistance to air exposure and to acute and chronic temperature stress between animals from different shore levels were similar in White and North Sea periwinkles, consistent with the adaptive nature of these variations. High-shore snails were found to be able to conserve body water reserves better, to resist higher temperatures and to survive longer under conditions of combined temperature and desiccation stress than their low-shore counterparts. In a temperature range of 25 to 35 °C, the rate of evaporative water loss was positively correlated with temperature in low-shore snails while being largely temperature-independent in high-shore snails. Median lethal time during air exposure in L. saxatilis was negatively but not linearly related to the temperature of exposure. In a temperature range of 30 to 38 °C, the resistance to heat exposure in air was only slightly dependent on the temperature, with Q ₁₀ = 1.4 for the median lethal time; the heat resistance dropped drastically at temperatures above 38 °C, with Q ₁₀ = 593.8. This suggests different mechanisms of temperature resistance in different parts of the studied temperature range. In contrast, behavioural response to extreme salinity fluctuations was not uniform in the high- and low-shore periwinkles from the White and North Seas, which may reflect specific environmental conditions at different shore levels in the two areas studied. Observed physiological and behavioural variations are discussed from the viewpoint of different adaptive strategies employed by eulittoral and eulittoral-fringe animals within populations of a single species. Received: 13 December 1999 / Accepted: 11 April 2000     

Importance of grazing on the salt-marsh grassSpartina alterniflora to nitrogen turnover in a macrofaunal consumer,Littorina irrorata,and to decomposition of standing-deadSpartina

Dying leaves ofSpartina alterniflora Loisel (hereafterSpartina) do not undergo abscission and consequently are at least partially degraded while remaining attached to the shoot, i.e., under conditions which may be very different from those occurring in litterbags used to measure decomposition ofSpartina at the sediment surface. Attached living and dead leaves in high-marsh areas are subject to grazing by the abundant gastropodLittorina irrorata Say (hereafterLittorina), a salt marsh periwinkle. In 1986, nitrogen assimilation from living and standing-deadSpartina byLittorina was examined in Sapelo Island (Georgia, USA) salt marshes by labelling plants with the stable nitrogen isotope¹⁵N and measuring the transfer into grazing snails in the field. The initial label of ca 8% total plant nitrogen declined to ca 1% over 5 mo, perhaps due to label dilution by less enriched nitrogen taken up and translocated from below- to above-groundSpartina biomass. Snails incorporatedSpartina-derived nitrogen into tissues at rates equal to 10 to 20% of total snail nitrogen 30-d^–1 in summer and fall, and 2 to 5% 30-d^–1 in winter. In the absence of measurable growth, these high nitrogen incorporation rates may indicate a large reproductive effort, or substantial turnover of somatic tissue nitrogen. The annual total assimilation ofSpartina-derived nitrogen was equal to theLittorina-nitrogen biomass. Assimilation of nitrogen in the presence of livingSpartina material (dead material removed) was reduced substantially below that in the presence of intact plants (living and dead material present).Littorina populations at abundances found in Georgia would assimilate ca 3.4% of above-groundSpartina-nitrogen production annually in high-marsh, short-Spartina areas. Based on preliminary estimates of nitrogen assimilation efficiency, 13.2 to 27.2% of short-Spartina production could be ingested annually by Georgia populations ofLittorina. Most of this ingestion would be concentrated in the summer and early fall, when monthly ingestion could equal 100% of deadSpartina biomass. The impact of grazing byLittorina onSpartina decomposition may be greatest on these early-senescing leaves. Grazing may have little impact on the early stages of decomposition of the bulk of the shoots that senesce later in fall, but may be important in the later stages of decomposition of dead shoots that persist through winter until the following spring and summer.     

Biodiversity in Littorina species (Mollusca: Gastropoda): a physiological approach using heat-coma

 The upper critical thermal limits were measured as heat-coma temperatures in Littorina species as a tool for examining physiological diversity. Thermal tolerance was found to be stable within a population, but extensive variation occurred between populations and species. All species examined displayed heat-coma values of ∼30 °C and did not show a positive correlation with shore height. Eulittoral-fringe species tended to have higher coma temperatures than eulittoral species. Coma temperatures varied both seasonally and geographically. Lethal thermal limits were also investigated; these were ∼10 C° higher than coma temperatures. Received: 16 August 1999 / Accepted: 13 June 2000     

Microscale variation in<Emphasis Type="Italic"> Aat</Emphasis> (aspartate aminotransferase) is supported by activity differences between upper and lower shore allozymes of<Emphasis Type="Italic"> Littorina saxatilis</Emphasis>

The role of natural selection in maintaining inherited variation in morphological characters is indisputable. In contrast, genetic variation in allozyme loci is mostly assumed neutral or under weak selection, although enzyme functions are critical to central metabolic processes. A locus coding for aspartate aminotransferase (AAT) is polymorphic with two alleles in the rocky shore gastropod Littorina saxatilis. One allele (Aat ¹²⁰ ) is most frequent (0.8–0.9) in upper shore subpopulations, while the other (Aat ¹⁰⁰ ) dominates (0.7–0.8) lower shore subpopulations only meters below. Earlier studies show this variation to be maintained by divergent selection, but the mechanism is unknown. We measured enzyme activities in AAT extracts from individuals of known Aat genotype in populations from two islands and at three different temperatures. In all treatments lower shore homozygotes (Aat ^100/100 ) had higher enzyme activity than upper shore homozygotes (Aat ^120/120 ), with heterozygotes having intermediate levels of activity. AAT has a central role in anaerobic energy production in marine mollusks during low-tide periods of emergence, and it might appear surprising that upper shore snails, which are most likely to be emerged, have lower enzyme activity. However, upper shore snails are forced to survive long periods of desiccation, and a low metabolic rate will help to conserve limited resources of energy and nutrients and reduce accumulation of waste products.Communicated by L. Hagerman, Helsingør     

Are direct developers more locally adapted than planktonic developers?

The hypothesis that populations of direct developers exhibit greater geographic differentiation in life history features than populations of planktonic developers, was tested with two species of grazing snails of the genus Littorina from 1986 to 1987. Littorina sitkana (direct developer) and L. scutulata (planktonic developer) coexist on sun- and wave-sheltered beaches from Alaska to Oregon, USA. Seasonal patterns in growth, survival and reproduction were monitored for samples from four geographically separated populations of each species grown in population cages at a common site, Friday Harbor, Washington, USA. The environmental and population effects on growth in the two species were determined in a four-way reciprocal transplant experiment with the same populations. Both the direct and planktonic developers exhibited geographic differentiation in life history features. Differentiation in the direct developer occurred over distances shorter than 30 km, while differentiation in the planktonic developer occurred over the 500 km distance examined (greater than their larvae would likely travel).     

Population biology of the annual grass Triplasis purpurea in relation to distance from shore on Staten Island, New York

Four populations of the native annual grassTriplasis purpurea were surveyed on coastal beaches along the south shore of Staten Island, NY, to determine the potential of this species to colonize shoreline habitats mostly devoid of other vegetation. If the species can establish and maintain dense populations, it may have conservation value for urban beaches disturbed by human activities. For two populations, survivorship, growth, and reproduction were monitored at different distances from shore to determine the ability of this species to maintain viable populations. At three sites,T. purpurea occurred in >75% of all quadrats and the highest density was 1195 plants/m² at 74 m from shore in one recently disturbed site. Density generally increased with increasing distances from shore at low tide (from ca. 40–90 m). Plants showed the greatest growth and reproduction at close distances to shore (30–40 m); part of this effect was due to density in one population, but when density effects were removed statistically, there still remained a decline in growth and reproduction with increasing distance from shore. Improved vigor nearest to shore may be due to continual sand deposition. Survivorship showed a Type I pattern, with low mortality throughout the growing season. By colonizing newly-deposited and continually shifting sands,T. purpurea can contribute to the earliest stages of ecological succession along disturbed beaches in eastern North America and may be valuable to the development and management of urban coastal plant communities.     

Population biology of the annual grassTriplasis purpurea in relation to distance from shore on Staten Island, New York

Four populations of the native annual grassTriplasis purpurea were surveyed on coastal beaches along the south shore of Staten Island, NY, to determine the potential of this species to colonize shoreline habitats mostly devoid of other vegetation. If the species can establish and maintain dense populations, it may have conservation value for urban beaches disturbed by human activities. For two populations, survivorship, growth, and reproduction were monitored at different distances from shore to determine the ability of this species to maintain viable populations. At three sites,T. purpurea occurred in >75% of all quadrats and the highest density was 1195 plants/m² at 74 m from shore in one recently disturbed site. Density generally increased with increasing distances from shore at low tide (from ca. 40 – 90 m). Plants showed the greatest growth and reproduction at close distances to shore (30 – 40 m); part of this effect was due to density in one population, but when density effects were removed statistically, there still remained a decline in growth and reproduction with increasing distance from shore. Improved vigor nearest to shore may be due to continual sand deposition. Survivorship showed a Type I pattern, with low mortality throughout the growing season. By colonizing newly-deposited and continually shifting sands,T. purpurea can contribute to the earliest stages of ecological succession along disturbed beaches in eastern North America and may be valuable to the development and management of urban coastal plant communities.     

The impact of tidal inundation on salt marsh vegetation after de-embankment on Langeoog Island, Germany—six years time series of permanent plots

Salt marsh succession after de-embankment was monitored on the East Frisian barrier island Langeoog by investigating permanent plots. Seventy years after embankment salt marsh plants were once again influenced mainly by the tidal regime. From 2002 to 2004 the former high marsh and glycophytic vegetation died out and was replaced by species of lower salt marsh zones. Nitrophytic halophytes like Suaeda maritima, Atriplex prostrata and Artemisia maritima established first because of the high nutrient content in the soil, a direct result of former vegetation decay. With decreasing nitrogen afterwards other species became more competitive. Until 2007 Atriplex portulacoides became more dominant in the lower marsh and Elymus athericus reached dominance in areas where grazing has been abandoned in the high marsh. The dynamics in the study area is much lower than in natural marshes due to the still existing drainage system. Therefore vegetation units with low species diversity are widespread.     

Non-host organisms affect transmission processes in two common trematode parasites of rocky shores

The transmission of free-living trematode stages is mediated by various environmental factors, of which the presence of ambient organisms within the host space is a potential major determinant. In two laboratory mesocosm experiments, we investigated the influence of four intertidal rocky shore species on transmission success of cercariae of the digenean trematodes Echinostephilla patellae (encysting in the tissue of blue mussels Mytilus edulis) and Parorchis acanthus (encysting on mussel shells). Encystment success of both parasite species was significantly lower in the presence of test organisms when compared to controls. Observations revealed that barnacles Austrominius modestus actively filtered cercariae, whereas the larvae were obstructed by the seaweeds Corallina officinalis and Fucus serratus. Anemones Actinia equina both physically disturbed and consumed cercariae. In a further laboratory experiment, grazing gastropods (Littorina littorea, Patella vulgata, and Gibbula umbilicalis) were found to significantly reduce the numbers of P. acanthus metacercariae in artificially prepared dishes by ingestion of cysts. Our results suggest that non-host organisms may play a key role in regulating the transmission of free-living trematode stages in rocky shore ecosystems, which is especially important with regard to the relative diversity and density of species in these habitats. The findings also emphasize the need to include parasites into marine food webs, since cercariae seem to be consumed by certain organisms to a considerable extent and could possibly represent an important energy source.     

Effects of a larval stage on isolation of populations of the trochid<Emphasis Type="Italic"> Austrocochlea constricta</Emphasis> in tidal ponds

To test the importance of special environments for local genetic subdivision in species with a larval phase, we examined allozyme variation among populations of the intertidal snail Austrocochlea constricta, in 18 tidal ponds in the Houtman Abrolhos Islands, Western Australia. Levels of genetic divergence between pond populations were correlated with those of parallel analyses among adjacent shore populations. However, divergence among the isolated ponds, which lack surface connection to the sea, were generally substantially higher, with an overall F_ST of 0.408, compared with 0.274 among the shore sites. The pond populations had less genetic variation than their shore counterparts, and the reduction of heterozygosity was correlated with the isolation of the pond population, as measured by F_ST. Both the degree of isolation and the reduction of heterozygosity were greater in deeper ponds, where snails can produce a local pool of larvae. In contrast, ponds that dry out frequently are less likely to allow production of local recruits, and these appear to be better connected genetically to adjacent shore populations. These patterns contrast sharply with those previously documented at the same sites for the direct-developing snail Bembicium vittatum, which shows greater isolation in ponds that are often dry. The comparison between the two species shows significant interaction between intrinsic and extrinsic impediments to gene flow, and highlights the importance of characteristics of the life history in determining which circumstances favour isolation of local populations.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00277-005-1553-5.Communicated by G.F. Humphrey, Sydney     

Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.     

Heterogeneity in the photoprotective capacity of three Antarctic diatoms during short-term changes in salinity and temperature

The Antarctic marine ecosystem changes seasonally, forming a temporal continuum of specialised niche habitats including open ocean, sea ice and meltwater environments. The ability for phytoplankton to acclimate rapidly to the changed conditions of these environments depends on the species’ physiology and photosynthetic plasticity and may ultimately determine their long-term ecological niche adaptation. This study investigated the photophysiological plasticity and rapid acclimation response of three Antarctic diatoms—Fragilariopsis cylindrus, Pseudo-nitzschia subcurvata and Chaetoceros sp.—to a selected range of temperatures and salinities representative of the sea ice, meltwater and pelagic habitats in the Antarctic. Fragilariopsis cylindrus displayed physiological traits typical of adaptation to the sea ice environment. Equally, this species showed photosynthetic plasticity, acclimating to the range of environmental conditions, explaining the prevalence of this species in all Antarctic habitats. Pseudo-nitzschia subcurvata displayed a preference for the meltwater environment, but unlike F. cylindrus, photoprotective capacity was low and regulated via changes in PSII antenna size. Chaetoceros sp. had high plasticity in non-photochemical quenching, suggesting adaptation to variable light conditions experienced in the wind-mixed pelagic environment. While only capturing short-term responses, this study highlights the diversity in photoprotective capacity that exists amongst three dominant Antarctic diatom species and provides insight into links between ecological niche adaptation and species’ distribution.     

Seasonal variation in utilization of biogenic microhabitats by littorinid snails on tropical rocky shores

Mobile species may actively seek refuge from stressful conditions in biogenic habitats on rocky shores. In Hong Kong, the upper intertidal zone is extremely stressful, especially in summer when organisms are emersed for long periods in hot desiccating conditions. As a result, many species migrate downshore between winter and summer to reduce these stressful conditions. The littorinids Echinolittorina malaccana and E. vidua, for example, are found on open rock surfaces high on the shore in winter but the majority migrate downshore in summer to the same tidal height as a common barnacle, Tetraclita japonica. In the laboratory, where environmental conditions could be controlled to approximate those occurring on the shore, we tested whether the downshore migration allowed littorinids to select barnacles as biogenic habitats to reduce stress and if this behaviour varied between seasons. In summer, littorinids demonstrated a strong active preference for the barnacles, which was not observed in the cool winter conditions, when animals were found on open rock surfaces even when barnacles were present. Littorinids, therefore, only actively select biogenic habitats during the summer in Hong Kong when they migrate downshore, suggesting that such habitats may play an important, temporal, role in mitigating environmental stress on tropical shores.     

Desiccation enhanced nitrogen uptake rates in intertidal seaweeds

Desiccation increased nitrate and ammonium uptake rates upon resubmergence in late summer populations of the intertidal macroalgae Gigartina papillata (C.Ag.) J.Ag., Enteromorpha intestinalis (L.) Grev., Fucus distichus L., and Pelvetiopsis limitata (Setch) Gardn. The ratio of nitrogen uptake rates in desiccated plants to rates in hydrated plants (controls) was correlated with the position of the species in the intertidal zone. Gracilaria pacifica Abbott., the species occurring at the lowest shore level, showed no enhancement of nitrogen uptake following desiccation. The high intertidal species such as P. limitata and F. distichus showed a two-fold enhancement of nitrate and ammonium uptake following more extensive desiccation (>30%) and continued uptake even following severe desiccation (50 to 60%). After the plants had been desiccated, the increase in nitrate uptake rates upon submergence lasted much longer than a similar enhancement of ammonium uptake. The duration of the enhanced nitrate uptake was similar to the time required for total rehydration but the uptake rates were not related to the state of rehydration. The potential contribution that this enhanced nitrogen uptake following desiccation could make to total nitrogen procurement for growth is discussed. The experiments were carried out in 1979 or 1980 and repeated in 1981.This paper is dedicated to Dr. R. F. Scagel on the occasion of his retirement for his outstanding contribution to phycology     

Huddling up in a dry environment: the physiological benefits of aggregation in an intertidal gastropod

In many intertidal gastropods, the formation of aggregations and closing of the opercular opening are behaviors commonly assumed to be associated with water conservation and maintenance of body temperature during tidal emersion periods. In the laboratory, we quantified the relationship between these two behaviors in a littorinid snail common to the north-central shores of Chile, Echinolittorina peruviana, and evaluated any benefit of these behaviors during desiccating conditions. We predicted that solitary individuals would maintain their opercula open for less time than aggregated snails when exposed to drier conditions due, at least in part, to differences in evaporative water loss. In laboratory trials, where relative humidity was manipulated, we observed that aggregated snails maintained their opercula open for longer periods of time than solitary snails under increasingly drier conditions. These results, together with observations of body temperature, suggest that aggregated animals may able to maintain gaseous exchange with their environment for longer periods of time than solitary individuals in response to desiccation stress. Our results suggest an interactive effect of two behaviors that confer physiological benefits when confronted with extreme physical conditions experienced during periods of emersion.     

Desiccation as a factor in the intertidal zonation of barnacles

Four species of balanomorph barnacles, Balanus crenatus Brugière, B. balanoides (L.), Elminius modestus Darwin and Chthamalus stellatus (Poli), were studied to assess the susceptibility of intertidal barnacle species to desiccation. Known sized samples of barnacles were exposed to controlled desiccating conditions and subsequent survival and water loss were determined. It is clear that the ability to live high on the shore is dependent on a reduction of the overall permeability to water loss. Because of greater surface area to volume ratios, small stages are particularly prone to desiccation. In normal intertidal emersion periods, small stages of B. crenatus particularly, and also of B. balanoides and E. modestus which are similar in their desiccation resistance, would be susceptible to desiccation at normal temperatures and low humidities. Large barnacles would be more prone to death from high temperatures when the tide is out. The spat of C. stellatus, although surviving much longer than spat of larger dimensions of the other species, must also be prone to prolonged emersion conditions at high shore levels.     

Influence of occupant microhabitat on the composition of encrusting communities on gastropod shells

The exploitation of microhabitats is widely considered to increase biodiversity in marine ecosystems. Although intertidal hermit crabs and gastropods may inhabit the same shell type and shore level, their microhabitat may differ depending on the state of the tide. On the south coast of Wales the hermit crab Pagurus bernhardus mainly inhabits the shells of Nucella lapillus (84%). Hermit crab shells had a significantly different encrusting community compared with live N. lapillus shells. At low tide the live gastropods were found on exposed rock surfaces whereas hermit crabs were restricted to tidal pools. Communities encrusting live gastropod shells were characterised by lower species richness and abundance compared with shells inhabited by hermit crabs (12 species found in total). A greater abundance and richness of epibionts was recorded from both shell types during the summer compared with winter. Differences in community composition between shell occupant types were attributed to microhabitats used by gastropods and hermit crabs and the associated desiccation pressures, rather than competitive interactions or shell characteristics. This contradicts earlier studies of subtidal shells where biological processes were considered more important than physical factors in controlling species abundance and richness patterns. The use of rockpool microhabitats by hermit crabs increases the biodiversity of rocky shores, as some species commonly found on hermit-crab-inhabited shells were rare in other local habitats.Communicated by J.P. Thorpe, Port Erin     

Comparing the ability of a non-indigenous and a native seaweed to induce anti-herbivory defenses

Phenotypic plasticity could be a key mechanism of successful species invasion. Few studies, however, have evaluated whether a non-indigenous species induces anti-herbivore defenses after its introduction to new habitats. We tested how a non-indigenous (Mastocarpus stellatus) and a native red seaweed (Chondrus crispus) responded to grazing by a periwinkle (Littorina littorea) and an isopod (Idotea granulosa) in Helgoland, Germany. In the donor region of the Mastocarpus population, that is Iceland, the periwinkle is missing, whereas the isopod is present. Our results indicate that Mastocarpus experienced less consumption by Littorina (but not by Idotea) than Chondrus. Moreover, Mastocarpus induced defenses in response to Idotea (but not to Littorina) grazing, whereas Chondrus induced anti-herbivory responses to both species of herbivores. Grazed Mastocarpus grew equally to non-grazed conspecifics, while growth of grazed Chondrus was lower than of ungrazed conspecifics. Hence, the non-indigenous Mastocarpus responded differently to grazing stress than the native Chondrus with respect to anti-herbivory and compensatory responses, which perhaps supported the successful Mastocarpus establishment in a new range.     

Grazing rates of organic matter and living fungal biomass of decaying Spartina alterniflora by three species of salt-marsh invertebrates

 The pathway for the flow of salt-marsh grass production into marsh food-webs is still not well defined. We compared the abilities of three marsh macroinvertebrates [salt marsh periwinkles, Littoraria irrorata (Say) (=Littorina irrorata), salt-marsh coffee-bean snails, Melampus bidentatus (Say); and a talitrid amphipod, Uhlorchestia spartinophila Bounsfield and Heard] to access standing-dead leaves of smooth cordgrass (Spartina alterniflora Loisel). The invertebrates were incubated with naturally-decaying leaves, and the rates of removal of organic matter and living fungal biomass (ergosterol) were measured. The impact of invertebrate activity upon fungal growth rates was measured as rates of fungal-membrane synthesis (incorporation of radioacetate into ergosterol). The removal rates of organic leaf biomass per mg individual biomass were highest for amphipods (700 μg mg⁻¹ d⁻¹) and lowest for periwinkles (90 μg mg⁻¹ d⁻¹), but the relatively large biomass of the snails made their removal rates per individual greater than those of amphipods. Net removal of ergosterol by all three invertebrates was >50% for yellow-brown (early-decay) leaf blades. For fully-brown (advanced-decay) blades, >50% removal of ergosterol was found only for periwinkles; exposure to coffee-bean snails and amphipods resulted in a net ergosterol reduction of ≤20%. The lower net reduction of living fungal biomass by coffee-bean snails and amphipods may have been due to fungal-growth stimulation (2.3-fold stimulation in coffee-bean snails and 1.5-fold stimulation in amphipods). Grazing by periwinkles did not stimulate fungal growth, possibly because of its high intensity. Grazing by these three salt-marsh shredders may affect marsh-grass shoot-decay in different ways. Periwinkles may abbreviate the period of fungal production, and incorporate the decaying material relatively quickly into snail biomass and fecal-pellet rain to the sediments. Coffee-bean snails and amphipods may enhance and prolong fungal production, along with the formation of fecal-pellet rain. All three invertebrates fed preferentially on leaf blades rather than leaf sheaths, and feeding rates of gastropods were higher during the night than during the day. Received: 25 November 1998 / Accepted: 4 November 1999     

Efficacy of created and restored nesting sites for the conservation of colonial Laridae in the South of France

By rapidly modifying key habitat components, habitat restoration is at risk of producing attractive cues for animals without providing habitats of sufficient quality. As such, individual fitness components, such as reproduction, could be reduced and restored habitats could become ecological traps. This risk notably appears by using artificial constructions in restoration projects, yet few studies have evaluated their efficacy in a robust way. We investigated this by analyzing 154 islets that were created or restored to improve the conservation status of 7 colonial Laridae species in the South of France. From 2007 to 2016, we compared occupancy dynamics and breeding parameters of these species between the restored sites and 846 unmanaged nesting sites. We also explored species’ preference for different nesting site characteristics and their respective effect on breeding parameters. Restored nesting sites were 2–9 times as attractive as unmanaged sites for all species except the Black-headed Gull (Chroicocephalus ridibundus). Colonization probability was up to 100 times higher in sites already used by other species the previous year and increased with distance to the shore until >0.2 when distance was over 250 m. Abandonment probability was 29–70% lower when breeding was successful the previous year in all species except the Sandwich Tern (Thalasseus sandvicensis). Productivity and breeding success probability were 2 times higher on managed sites. Distance from the shore was an important attractive characteristic of artificial nesting sites in all species. Other nesting site characteristics had species-specific effects on colonization, abandonment, and breeding success. Our results indicate that managed nesting sites are successful conservation tools for colonial Laridae in the Mediterranean and do not act as ecological traps. Our study showed that testing the ecological trap hypothesis is a robust way to evaluate the success of restoration projects of breeding habitats.