Wintering Shorebird Assemblages and Behavior in Restored Tidal Wetlands in Southern California

Habitat restoration can partially compensate for the extensive loss of coastal wetlands, but creation of functional habitat and assessment of restoration success remain challenging tasks. To evaluate wintering shorebird use of restored coastal wetlands, we quantified shorebird assemblages and behavior of selected focal species at five restored sites and paired reference sites in Mugu Lagoon, southern California, United States. The Shannon–Wiener index of species diversity (for all birds in order Charadriiformes) was higher in the restored than in the reference portion of three of the five sites, higher in the reference portion of a fourth site, and similar between reference and restored areas of the fifth site. Species diversity was lower in sites closer to man‐made structures. The four most abundant species groups across the five sites were selected for detailed analysis of site use and behavior: Willets (Catoptrophorus semipalmatus), Marbled Godwits (Limosa fedoa), Dowitchers (Limnodromus spp.), and Sandpipers (Calidris spp.) (Western, Least, and Dunlin). Each focal species group exhibited distinct site preferences, and densities in restored sites were often as high or higher than in reference sites. Willets and Dowitchers preferred habitats with more extensive tidal flats, a characteristic of restored sites. Godwits and Sandpipers preferred heterogeneous habitats with a mix of water and tidal flats. Most birds were engaged in feeding activities during the ebb tides surveyed, and there were no apparent differences in behavior between reference and restored sites. Though not all restored sites were used equally by all species, the creation of multiple restored sites with varied habitat characteristics attracted a diverse assemblage of shorebirds and may have contributed to the integrity of the regional wetland landscape.     

Recruitment of shore crabsCarcinus maenas on tidal flats: Mussel clumps as an important refuge for juveniles

During the late summer and early fall, juvenile shore crabs (Carcinus maenas L.) occurred in high abundances in mussel clumps scattered on tidal flats of the Wadden Sea. Abundances were much lower on bare tidal flats without mussel clumps and decreased substantially from July to November, whereas numbers in mussel clumps remained high. Large crabs left the tidal flats in early fall, whereas juveniles undertook tidal migrations only in the late fall. In March very few shore crabs were found in the intertidal area. The size of juvenile shore crabs living between mussels did not increase significantly during fall. On the bare tidal flats surrounding the mussels, a size increase was observed. Mussel beds and mussel clumps serve as a spatial refuge for the early benthic phases of juvenile shore crabs. Between mussels they can hide effectively from their epibenthic predators. Juvenile shore crabs do not leave the intertidal area and the mussel habitats before their major predators have left the area. Mussel clumps scattered over the tidal flats may be a critical refuge for juvenile shore crabs settling on tidal flats. Intensified efforts in mussel culturing in the European Wadden Sea during recent decades may have caused an increased abundance of mussel clumps on tidal flats, thus enhancing habitat availability for some mussel-clump inhabitants.     

Lifetime reproductive output in a hermaphrodite cestode when reproducing alone or in pairs: a time cost of pairing

The cestode Schistocephalus solidus is a facultatively self-fertilising simultaneous hermaphrodite. Here we test for differences in the starting point, the rate, and the magnitude of egg production between individuals allowed to reproduce alone (only self-fertilisation possible) or in pairs (both self- and cross-fertilisation possible). Specifically, we want to distinguish between alternative processes responsible for the lower egg production in paired individuals observed in an earlier study (Wedekind et al., 1998). We designed an improved in vitro system, replacing the bird final host that allows us to measure, with high temporal resolution, the timing and magnitude of lifetime egg production of worms in these two social situations. We found that the experimental groups did not differ significantly in the starting point of egg production. However, the temporal pattern in egg production differed between them, in that paired individuals had a lower rate of egg production. This, however, did not lead to a significant reduction in lifetime egg production, as pairs compensated for the lower rate by producing eggs longer than single individuals. We argue that the lower rate of egg production may nevertheless lead to a time cost of pairing in the study species, and that this cost is likely to represent a cost of outcrossing due to sexual selection. This revised version was published online in July 2006 with corrections to the Cover Date.     

The influence of potential stressors on oviposition site selection and subsequent growth,survival and emergence of the non‐biting midge (Chironomus tepperi)

Theory predicts that animals should prefer habitats where their fitness is maximized but some mistakenly select habitats where their fitness is compromised, that is, ecological traps. Understanding why this happens requires knowledge of the habitat selection cues animals use, the habitats they prefer and why, and the fitness costs of habitat selection decisions. We conducted experiments with a freshwater insect, the non‐biting midge Chironomus tepperi to ask: (a) whether females respond to potential oviposition cues, (b) to explore whether oviposition is adaptive in relation to metal pollution and conductivity, and (c) whether individuals raised in poor quality sites are more likely to breed in similarly poor locations. We found the following: (a) females responded to some cues, especially conductivity and conspecifics, (b) females preferred sites with higher concentrations of bioavailable metals but suffered no consequences to egg/larval survival, (c) females showed some avoidance of high conductivities, but they still laid eggs resulting in reduced egg hatching, larval survival, and adult emergence, and (d) preferences were independent of natal environment. Our results show that C. tepperi is susceptible to ecological traps, depending on life stage and the relative differences in conductivities among potential oviposition sites. Our results highlight that (a) the fitness outcomes of habitat selection need to be assessed across the life cycle and (b) the relative differences in preference/suitability of habitats need to be considered in ecological trap research. This information can help determine why habitat preferences and their fitness consequences differ among species, which is critical for determining which species are susceptible to ecological traps.     

Empty habitat in coastal sediments for populations of macrozoobenthos

Species with wide dispersal are expected to have little empty habitat. This was tested by analysing habitat use in the Wadden Sea near the island of Sylt (North Sea). Sampling covered 222 intertidal sites spread across the K?nigshafen tidal flats (4.5 km², mapping approach conducted once) and 270 subtidal sites along 12 km of a tidal channel system (stratified random sampling in spring and autumn). For any single species, 'suitable habitat' was extrapolated from the ranges of water depth and sediment composition present at the ten sites with the highest frequency or abundance of the species. On average, macrobenthic species actually used less than half of the suitable sites within the scale of local populations; this was far less than expected from a lognormal distribution. In the subtidal, abundance of most species changed between the two sampling seasons. A 50% increase in overall abundance was accompanied by a decrease in empty habitat of only about 25%. Thus, a doubling of abundances would not fill the empty space but just of half of it. The polychaete Scoloplos armiger was an exception in occupying almost all of its suitable habitat in the intertidal and subtidal sediments. A distinct patch of high species richness occurred where flood waters persistently form a large gyre which may enhance larval settlement. We suggest that limitations to larval settlement and/or juvenile survival primarily cause the extent of the observed habitat emptiness. Electronic Publication     

The response of meiofauna and microphytobenthos to engineering effects of fiddler crabs on a subtropical intertidal sandflat

Fiddler crabs are key bioturbators on tidal flats. During their intense bioturbation process, they manipulate large amounts of sediment, altering the physical state of existing materials. We investigated whether different types of sediment bioturbation produced by fiddler crabs modulate meiofaunal assemblages and microphytobenthic content. We hypothesized that sedimentary structures produced by burrowing (the burrow itself and the excavation pellets) and feeding (feeding pellets) generate different microenvironments compared with areas without apparent signs of fiddler crab disturbance, affecting both meiofauna and microphytobenthos, independent of the sampling period. Our results indicate that the engineering effects of burrow construction and maintenance and the engineering effects of fiddler crab foraging modulate meiofaunal assemblages in different ways. Overall, meiofauna from burrows and excavation pellets was more abundant and diverse than at control sites, whereas feeding pellets contained poor meiofaunal assemblages. By contrast, only foraging effects were detected on microphytobenthos; independent of the sampling period, Chl a and phaeopigment content were higher in the feeding pellets, but similar among burrows, excavation pellets and control sites. The present study demonstrates that the different engineering effects of fiddler crabs are an important source of habitat heterogeneity and a structuring agent of meiofaunal assemblages on subtropical tidal flats.     

Spatial ecology of juvenile lemon sharks (<Emphasis Type="Italic">Negaprion brevirostris</Emphasis>) in tidal creeks and coastal waters of Eleuthera,The Bahamas

Fisheries exploitation and habitat alteration are threatening lemon shark (Negaprion bevirostris) populations because they use nearshore regions as nursery sites. As such, there is a need for information on the spatial ecology of juvenile lemon sharks to identify critical habitats that require protection, as well as to understand their basic ecology. The purpose of this study was to determine the habitat preferences and movement patterns of juvenile lemon sharks along a sub-section of coastline characterized by coastal flats and tidal creeks of Eleuthera, The Bahamas. Eleven juvenile lemon sharks (766 ± 127 mm total length; mean±SD) were captured from various tidal creeks within the 23 km study area and were surgically implanted with acoustic transmitters. A series of 27 hydrophone receivers acted as a passive monitoring array to detect tagged individuals as they moved among habitats. Findings suggest that juvenile lemon sharks tagged in this study prefer shallow water habitats within tidal creeks, and typically display high site fidelity with occasional forays to alternate habitats or creeks. In fact, more than 90% of tagged lemon sharks had the greatest percentage of detections located at a receiver at or close to the location where they were tagged. There was no evidence of differences in diel or seasonal movement and habitat use. Knowledge gained from this study will be useful for directing future conservation and management strategies including coastal development plans and marine protected areas.     

Multiple habitat use by declining migratory birds necessitates joined‐up conservation

Many species depend on multiple habitats at different points in space and time. Their effective conservation requires an understanding of how and when each habitat is used, coupled with adequate protection. Migratory shorebirds use intertidal and supratidal wetlands, both of which are affected by coastal landscape change. Yet the extent to which shorebirds use artificial supratidal habitats, particularly at highly developed stopover sites, remains poorly understood leading to potential deficiencies in habitat management. We surveyed shorebirds on their southward migration in southern Jiangsu, a critical stopover region in the East Asian Australasian Flyway (EAAF), to measure their use of artificial supratidal habitats and assess linkages between intertidal and supratidal habitat use. To inform management, we examined how biophysical features influenced occupancy of supratidal habitats, and whether these habitats were used for roosting or foraging. We found that shorebirds at four of five sites were limited to artificial supratidal habitats at high tide for ~11–25 days per month because natural intertidal flats were completely covered by seawater. Within the supratidal landscape, at least 37 shorebird species aggregated on artificial wetlands, and shorebirds were more abundant on larger ponds with less water cover, less vegetation, at least one unvegetated bund, and fewer built structures nearby. Artificial supratidal habitats were rarely used for foraging and rarely occupied when intertidal flats were available, underscoring the complementarity between supratidal roosting habitat and intertidal foraging habitat. Joined‐up artificial supratidal management and natural intertidal habitat conservation are clearly required at our study site given the simultaneous dependence by over 35,000 migrating shorebirds on both habitats. Guided by observed patterns of habitat use, there is a clear opportunity to improve habitat condition by working with local land custodians to consider shorebird habitat requirements when managing supratidal ponds. This approach is likely applicable to shorebird sites throughout the EAAF.     

HABITAT DIFFERENCES IN THE TIMING OF REPRODUCTION OF THE INVASIVE ALGA SARGASSUM MUTICUM (PHAEOPHYTA,SARGASSACEAE) OVER TIDAL AND LUNAR CYCLES1

Sargassum muticum (Yendo) Fensholt is an invasive species that is firmly established on intertidal and subtidal rocky shores of Europe and the Pacific coast of North America. Local success and spread of S. muticum is thought to rely on its reproductive potential that seems dependent on exogenous factors like tidal and lunar cycles. This study is the first to compare the reproductive patterns (periodicity of egg expulsion and embryo settlement) of this invader in two different habitats: the middle and low intertidal. The combination of monthly, daily, and tidal samples at triplicate sites within each habitat showed a semilunar periodicity of egg expulsion and embryo settlement coincident with increasing tidal amplitude just before full and new moons. In both habitats, duration of each egg expulsion event was ～1 week, and embryo settlement occurred during the first daily low tide and with the incoming high tide during spring tides. However, both expulsion and settlement started 1–2 d earlier, expulsion saturation was faster, and settlement was higher in the mid‐ compared to the low intertidal. Our results suggest that the exact timing of gamete expulsion and embryo release of S. muticum responds to local factors, including tidal cues, which result in differences between mid‐ and low‐intertidal habitats.     

At Palmyra Atoll,the fish-community environmental DNA signal changes across habitats but not with tides

At Palmyra Atoll, the environmental DNA (eDNA) signal on tidal sand flats was associated with fish biomass density and captured 98%–100% of the expected species diversity there. Although eDNA spilled over across habitats, species associated with reef habitat contributed more eDNA to reef sites than to sand-flat sites, and species associated with sand-flat habitat contributed more eDNA to sand-flat sites than to reef sites. Tides did not disrupt the sand-flat habitat signal. At least 25 samples give a coverage >97.5% at this diverse, tropical, marine system.     

Oviposition behavior and offspring performance in herbivorous insects: consequences of climatic and habitat heterogeneity

The preference–performance hypothesis predicts that when female herbivorous insects determine where to position offspring of low mobility, they will select sites that maximize development and survival of those offspring. How this critical relationship responds to variation in climatic and habitat conditions remains untested, however, despite its important consequences for population and evolutionary dynamics. Here we report on 13 years of data totaling 1348 egg clusters of the montane Gillette's checkerspot butterfly Euphydryas gillettii (Lepidoptera: Nymphalidae). We used these data to test the hypothesis that, in environments with climatic and habitat heterogeneity, the oviposition behavior–offspring performance relationship should vary in both space and time. Orientation of egg clusters for maximum morning sun exposure is known to affect developmental rate. We therefore predicted female preference for morning sun orientation to be variable and a function of climatic and habitat conditions. We found that preference for egg cluster orientation on the leaf tracked the phenology of the start of the female flight season but that seasonal temperatures drove most of the variation in egg cluster development time. The relationship between behavior and performance was also dependent upon the climatic effects on survival; sun‐oriented egg clusters had higher survivorship in the coldest year of the four years for which measurements were made. We also examined how conifer cover affected larval survival and female oviposition behavior in one year. Females selected oviposition sites in more open habitat. However, when egg clusters were oriented to intercept morning sun, conifer cover increased survivorship to diapause. Finally, we found that predator activity was lower for morning sun‐oriented egg clusters suggesting that predation patterns may further influence habitat selection for oviposition. This study exemplifies how the relationship between oviposition behavior and offspring performance is context‐dependent: habitat and climate interact to determine preference–performance outcomes.     

Density‐dependent habitat selection predicts fitness and abundance in a small lizard

Density‐dependent habitat selection has been used to predict and explain patterns of abundance of species between habitats. Thermal quality, a density‐independent component of habitat suitability, is often the most important factor for habitat selection in ectotherms which comprise the vast majority of animal species. Ectotherms may reach high densities such that individual fitness is reduced in a habitat due to increased competition for finite resources. Therefore, density and thermal quality may present conflicting information about which habitat will provide the highest fitness reward and ectotherm habitat selection may be density‐independent. Using ornate tree lizards Urosaurus ornatus at 10 sites each straddling two adjacent habitats (wash and upland), we tested the hypothesis that habitat selection is density‐dependent even when thermal quality differs between habitats. We first tested that fitness proxies decline with density in each habitat, indicating density‐dependent effects on habitat suitability. We also confirmed that the two habitats vary in suitability (quantified by food abundance and thermal quality). Next, we tested the predictions that habitat selection depends on density with isodar analyses and that fitness proxies are equal in the two habitats within a site. We found that monthly survival rates decreased with density, and that the wash habitat had more prey and higher thermal quality than the upland habitat. Lizards preferred the habitat with more food and higher thermal quality, lizard densities in the two habitats were positively correlated, and fitness proxies of lizards did not differ between habitats. These patterns are consistent with density‐dependent habitat selection, despite differences in thermal quality between habitats. We expect that density‐dependent habitat selection is widespread in terrestrial ectotherms when densities are high and temperatures are close to their optimal performance range. In areas where thermal quality is low, however, we expect that depletable resources, such as food, become less limiting because assimilating resources is more difficult.     

钦州湾越冬鸻鹬类对不同生境的选择分析

2008年12月至2009年1月及2011年12月至2012年1月对钦州湾的越冬鸻鹬类进行了调查,对越冬鸻鹬类退潮后在各类型生境的分布情况和原因进行了分析,以期为制定有效保护越冬鸟类及其生境的措施提供科学依据。冬季在钦州湾记录到鹬鸻类27种,结果表明,在不同的天气情况下退潮后各生境内记录到的鹬鸻类种数和数量均不同,钦州湾不同生境鹬鸻类结构特征随天气变化改变。相似性比较显示,晴天生境相似性指数最高的是盐田-养殖塘生境和农田生境,相似性指数为0.96;相似性指数最低的是红树林滩涂生境和光滩生境,相似性指数为0.49。阴冷天生境相似性指数最高的是光滩生境和盐田-养殖塘生境,相似性指数为0.79;最低是红树林滩涂生境和光滩生境,相似性指数为0.52。分析表明人类活动的干扰对越冬鹬鸻类栖息地选择的影响是巨大的。     

Environmental heterogeneity amplifies behavioural response to a temporal cycle

Resource acquisition is integral to maximise fitness, however in many ecosystems this requires adaptation to resource abundance and distributions that seldom stay constant. For predators, prey availability can vary at fine spatial and temporal scales as a result of changes in the physical environment, and therefore selection should favour individuals that can adapt their foraging behaviour accordingly. The tidal cycle is a short, yet predictable, temporal cycle, which can influence prey availability at temporal scales relevant to movement decisions. Here, we ask whether black‐legged kittiwakes Rissa tridactyla can adjust their foraging habitat selection according to the tidal cycle using GPS tracking studies at three sites of differing environmental heterogeneity. We used a hidden Markov model to classify kittiwake behaviour, and analysed habitat selection during foraging. As expected for a central‐place forager, we found that kittiwakes preferred to forage nearer to the breeding colony. However, we also show that habitat selection changed over the 12.4‐h tidal cycle, most likely because of changes in resource availability. Furthermore, we observed that environmental heterogeneity was associated with amplified changes in kittiwake habitat selection over the tidal cycle, potentially because environmental heterogeneity drives greater resource variation. Both predictable cycles and environmental heterogeneity are ubiquitous. Our results therefore suggest that, together, predictable cycles and environmental heterogeneity may shape predator behaviour across ecosystems.     

The Development of Vertical Migration Patterns in the Sediments of Estuaries as a Strategy for Algae to Resist Drift with Tidal Currents

The phenomenon of massive algae migrations to the surface of mud flats at low tide was quantitatively investigated. Differences were noted in the behavior of the individual species. This adaptation to the tidal rhythm is obviously effective for preventing the algae from drifting to the sea, and its development can shed much light on the natural history of the species involved. Euglena obtusa and Caloneis amphisbaena var. subsalina apparently arose by a natural selection process among ancestors from lentic freshwater bodies, while Navicula salinarum, Cylindrotheca signata, and several other diatoms probably invaded the freshwater habitat from marine mud flats, where they were preadapted to the tides.     

Migration stopover ecology of Cinnamon Teal in western North America

Identifying migration routes and fall stopover sites of Cinnamon Teal (Spatula cyanoptera septentrionalium) can provide a spatial guide to management and conservation efforts, and address vulnerabilities in wetland networks that support migratory waterbirds. Using high spatiotemporal resolution GPS‐GSM transmitters, we analyzed 61 fall migration tracks across western North America during our three‐year study (2017–2019). We marked Cinnamon Teal primarily during spring/summer in important breeding and molting regions across seven states (California, Oregon, Washington, Idaho, Utah, Colorado, and Nevada). We assessed fall migration routes and timing, detected 186 fall stopover sites, and identified specific North American ecoregions where sites were located. We classified underlying land cover for each stopover site and measured habitat selection for 12 land cover types within each ecoregion. Cinnamon Teal selected a variety of flooded habitats including natural, riparian, tidal, and managed wetlands; wet agriculture (including irrigation ditches, flooded fields, and stock ponds); wastewater sites; and golf and urban ponds. Wet agriculture was the most used habitat type (29.8% of stopover locations), and over 72% of stopover locations were on private land. Relatively scarce habitats such as wastewater ponds, tidal marsh, and golf and urban ponds were highly selected in specific ecoregions. In contrast, dry non‐habitat across all ecoregions, and dry agriculture in the Cold Deserts and Mediterranean California ecoregions, was consistently avoided. Resources used by Cinnamon Teal often reflected wetland availability across the west and emphasize their adaptability to dynamic resource conditions in arid landscapes. Our results provide much needed information on spatial and temporal resource use by Cinnamon Teal during migration and indicate important wetland habitats for migrating waterfowl in the western United States.     

丹江口库区滩涂与入库支流植被与土壤种子库: 水传播潜力探讨

为了探索库区滩涂与入库支流土壤种子库与植被的关系, 以及物种通过水传播扩散的潜力, 对丹江口库区内4条支流及其入库滩涂的地表植被和土壤种子库进行了调查。结果显示, 支流地表植被和种子库的物种多样性高于库区滩涂, 汉江库区滩涂和支流的地表植被与种子库均有较高的物种相似性。DCA排序和TWINSPAN聚类可以明显地将支流的植被和种子库样方与滩涂的植被和种子库样方分到不同的样方组, 结合物种排序图可以清楚地发现, 流入汉江库区支流的植被和种子库的物种组成与滩涂的植被和种子库的物种组成有密切联系。汉江库区滩涂0-15 cm的土层内都有种子萌发, 而丹江库区种子库中只有狗牙根(Cynodon dactylon)一种植物, 且仅存在于0-5 cm土层。研究结果表明, 繁殖体通过支流向滩涂的水传播作用对植被建立有积极贡献。     

Avian fitness consequences match habitat selection at the nest‐site and landscape scale in agriculturally fragmented landscapes

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Patterns of macro and microhabitat use of two rodent species in relation to agricultural practices

Habitat selection may reflect the location of the home ranges or the allocation of shelter and foraging sites within a given habitat. We studied seasonal patterns of habitat use by Akodon azarae and Calomys laucha at two spatial scales: between maize fields and their weedy edges (macrohabitats), and associations of rodents captures with vegetation variables at the trap site level (microhabitats). We evaluated if the different habitat uses were related to disturbances generated by practices associated to maize cycle. A. azarae used mainly field edges, but it showed an increased use of maize fields when the crop reached maturity in summer. Contrarily, C. laucha used maize fields in a higher proportion than edges in all seasons. C. laucha was more influenced by microhabitat characteristics than A. azarae. C. laucha was present in sites with abundant dicot weeds when maize was growing up, while it was associated to sites with weeds with scarce cover in stubble maize fields. Before harvesting, both species were segregated at the microscale within maize fields. A. azarae was related to sites with high availability of green plant cover and C. laucha occupied low-quality sites, probably attributed to differences in their diets. We conclude that the pattern of habitat use by both species is best predicted at the macrohabitat scale, and when they are impoverished and present internal heterogeneity, there is selection at microhabitat scale of those better sites. While A. azarae responds to changes in vegetation cover and habitat structure associated to agricultural practices, C. laucha uses cropfields in an opportunistic way, affected by interspecific competition.