Evolution of habitat use by deep-sea mussels

Previous phylogenetic studies proposed that symbiont-bearing mussels of the subfamily Bathymodiolinae (Bivalvia: Mytilidae) invaded progressively deeper marine environments and evolved from lineages that decomposed wood and bone to specialized lineages that invaded cold-water hydrocarbon seeps and finally deep-sea hydrothermal vents. To assess the validity of the hypotheses, we examined two nuclear (18S and 28S rRNA) and two mitochondrial genes (COI and ND4) from a broad array of bathymodiolin species that included several recently discovered species from shallow hydrothermal seamounts. Bayesian phylogenetic analysis and maximum-likelihood estimates of ancestral character states revealed that vent species evolved multiple times, and that reversals in vent and seep habitat use occurred within the sampled taxa. Previous hypotheses regarding evolution from wood/bone-to-seeps/vents are supported in that mid-ocean hydrothermal vent species may represent a monophyletic group with one noticeable reversal. Earlier hypotheses about progressive evolution from shallow-to-deep habitats appear to hold with a few instances of habitat reversals.     

Complementary habitat use by wild bees in agro-natural landscapes

Human activity causes abrupt changes in resource availability across the landscape. In order to persist in human-altered landscapes organisms need to shift their habitat use accordingly. Little is known about the mechanisms by which whole communities persist in human-altered landscapes, including the role of complementary habitat use. We define complementary habitat use as the use of different habitats at different times by the same group of species during the course of their activity period. We hypothesize that complementary habitat use is a mechanism through which native bee species persist in human-altered landscapes. To test this idea, we studied wild bee communities in agro-natural landscapes and explored their community-level patterns of habitat and resource use over space and time. The study was conducted in six agro-natural landscapes in the eastern United States, each containing three main bee habitat types (natural habitat, agricultural fields, and old fields). Each of the three habitats exhibited a unique seasonal pattern in amount, diversity, and composition of floral resources, and together they created phenological complementarity in foraging resources for bees. Individual bee species as well as the bee community responded to these spatiotemporal patterns in floral availability and exhibited a parallel pattern of complementary habitat use. The majority of wild bee species, including all the main crop visitors, used fallow areas within crops early in the season, shifted to crops in mid-season, and used old-field habitats later in the season. The natural-forest habitat supported very limited number of bees, mostly visitors of non-crop plants. Old fields are thus an important feature in these arable landscapes for maintaining crop pollination services. Our study provides a detailed examination of how shifts in habitat and resource use may enable bees to persist in highly dynamic agro-natural landscapes, and points to the need for a broad cross-habitat perspective in managing these landscapes.     

Spatiotemporal dynamics of habitat use by koalas: the checkerboard model

Animal movement patterns and use of space depend upon food and nonfood resources, as well as conspecific and heterospecific interactions, but models of habitat use often neglect to examine multiple factors and rarely include marsupials. We studied habitat use in an Australian population of koalas (Phascolarctos cinereus) over a 6-year period in order to determine how koalas navigate their environment and partition limited patchy food and nonfood resources. Tree selection among koalas appears to be mediated by folar chemistry, but nonfood tree selection exerts a major impact on home range use due to thermoregulatory constraints. Koalas moved on a daily basis, during both day and night, but daytime resting site was not necessarily in the same location as nighttime feeding site. Koalas had substantial home range overlap in the near absence of resource sharing with less than 1% of trees located in areas of overlap used by multiple koalas. We suggest that koala spatiotemporal distribution and habitat use are probably based upon a community structure of individuals, with a checkerboard model best describing overlap in home range area but not in resource use. Nonfood refugia and social networks should be incorporated into models of animal range and habitat use.     

Effects of habitat fragmentation and degradation on flocks of African ant-following birds.

Tropical rain forests are rapidly cleared, fragmented, and degraded in sub-Saharan Africa; however, little is known about the response of species and even of key ecological groups to these processes. One of the most intriguing (but often neglected) ecological phenomena in African rain forests is the interaction between swarm-raiding army ants and ant-following birds. Similar to their well-known Neotropical representatives, ant-following birds in Africa track the massive swarm raids of army ants and feed on arthropods flushed by the ants. In this study we analyzed the effect of habitat fragmentation and degradation of a mid-altitude Congo-Guinean rain forest in western Kenya on the structure of ant-following bird flocks. Significant numbers of swarm raids were located in all forest fragments and in both undegraded and degraded forest. Fifty-six different species of birds followed army ant raids, forming bird flocks of one to 15 species. We quantitatively differentiated the bird community into five species of specialized ant-followers and 51 species of opportunistic ant-followers. Species richness and size of bird flocks decreased with decreasing size of forest fragments and was higher in undegraded than in degraded habitat. This was caused by the decrease of the species richness and number of specialized ant-followers at swarms, while the group of opportunistic ant-followers was affected little by habitat fragmentation and degradation. The composition of bird flocks was more variable in small fragments and degraded forest, compared to undegraded habitat in large fragments. The effect of habitat fragmentation on flock structure was best explained by the strong decline of the abundance of specialized ant-followers in small forest fragments. To conserve the association of army ants and ant-following birds in its natural state, vast areas of unfragmented and undegraded tropical rain forest are necessary.     

Influence of fish on habitat choice of water birds: a whole system experiment

It is notoriously difficult to study population interactions among highly mobile animals that cannot be meaningfully confined to experimental plots of limited size. For example, migratory water birds are believed to suffer from competition with resident fish populations for shared food resources. While observational evidence in support of this hypothesis is accumulating, replicated experiments addressing this issue at the proper spatial scale are lacking. Here, we report from a replicated whole-system experiment in which we stocked large (0.07 km2), shallow (< or =2.5 m deep), highly eutrophic ponds in the bird protection area "Ismaninger Speichersee mit Fischteichen" with different densities of carp and assessed the responses of water birds and their food resources during summer over several years. In all years, the biomasses of benthic macroinvertebrates, macroalgae, and macrophytes as well as the densities of herbivorous, carnivorous, and omnivorous water birds were reduced in carp ponds compared to fishless ponds. The negative effects of carp on food resources and on the numbers of water birds feeding in carp ponds increased over the season (May-September) and were stronger at high than at low stocking densities of carp. Consequently, differences in resource densities between ponds with and without carp increased, and the ranking of ponds with respect to resource densities became more predictable over the season. These factors may have contributed to a seasonal improvement of the birds' abilities to track resource densities across ponds, as suggested by tight correlations of bird numbers on ponds with resource densities late, but not early, in the season.     

Spatio-temporal use of the urban habitat by feral pigeons (Columba livia )

Feral pigeons are descendants of wild rock pigeons that have adapted to the urban habitat. They have partially conserved the foraging behaviour of their wild ancestors (flights to agricultural areas) but have also developed new habits. Previous studies on the foraging strategies of feral pigeons have given various results, e.g. maximum distances reached by the pigeons (measured in a straight line from the resting places) differed between 0.3–0.5 km and 18–25 km. This study focuses on the spatio-temporal activity of feral pigeons in the urban habitat. We equipped 80 free-living feral pigeons from Basel, Switzerland with GPS receivers. We found three different foraging strategies for pigeons in Basel: (1) in the streets, squares and parks near the home loft, (2) in agricultural areas surrounding the city, (3) on docks and railway lines in harbours. The maximum distance reached by a pigeon was 5.29 km. More than 32% of the pigeons remained within 0.3 km of the home lofts and only 7.5% flew distances of more than 2 km. Females covered significantly longer distances than males, preferring to fly to more abundant and predictable food sources. Temporal activity patterns showed to be influenced by sex, breeding state and season. In contrast to wild rock pigeons and to feral pigeons in other cities, pigeons in Basel showed a clear bimodal activity pattern for breeding birds only. The differences between our results and those of other studies seem to be partly method-dependent, as the GPS-technique allows to record the pigeons’ localisations continuously in contrast to other methods. Other differences might be due to different kinds of food supply in the various cities. Our study shows that feral pigeons have individual foraging strategies and are flexible enough to adapt to different urban environments.Electronic supplementary material Supplementary material is available for this article at     

Foraging behaviour and habitat use by brown skuas Stercorarius lonnbergi breeding at South Georgia

Top predators are critical to ecosystem function, exerting a stabilising effect on the food web. Brown skuas are opportunistic predators and scavengers. Although skuas are often the dominant land-based predator at seabird colonies, this is the first detailed study of their movements and activity during breeding. The study was carried out at Bird Island, South Georgia (54°00′S, 38°03′W), in the austral summer of 2011/2012 and included GPS data from 33 breeding adults tracked during the late incubation and early chick-rearing periods. Brown skuas spent on average more than 80 % of time in the territory, and it was extremely rare for both partners to leave the territory simultaneously. Much more time was spent foraging at the coast than in penguin colonies and, based on saltwater immersion data, adults never foraged at sea. None of the tracked birds appeared to specialise in catching small petrels at night. Fewer foraging trips were made per day, and hence, more time was spent in the territory, during incubation than chick-rearing. Despite the pronounced sexual size dimorphism, there were no effects of sex on territorial attendance, foraging time or habitat use. Skuas at Bird Island show higher territorial attendance and are less likely to leave the territory unattended than those breeding elsewhere, suggesting closer proximity to more diverse or abundant food resources than at other colonies. The results tie in with previous diet studies, indicating that brown skuas at this site feed mostly on seal placentae and carrion and that birds may rely on a broader range of food resources as the season progresses.     

Foraging behavior and habitat use by the Southern Giant Petrel on the Patagonian Shelf

We explored the at-sea behavior and marine habitat use of the Southern Giant Petrel breeding in Patagonia, Argentina by means of satellite telemetry. Adult breeders showed a wide distribution over the Patagonian Shelf, using 74% of its surface. The maximum distance traveled from the colonies was 683 km, but on average birds moved no more than 200 km further away from their colony. Important marine areas were located in the shelf break, middle shelf and coastal waters. Areas of activity by sex overlap between 35 and 94%. Females foraged primarily away from the coast and males mainly on coastal areas. Both sexes were capable of flying up to 4,000 km but most of the foraging trips were of less than 200 km. Our results emphasize the importance of the Patagonian Shelf as foraging habitat for pelagic seabirds and contribute to international efforts to identify and protect a network of marine sites.     

Dynamic intra-seasonal habitat use by Antarctic fur seals suggests migratory hotspots near the Antarctic Peninsula

Antarctic fur seals (Arctocephalus gazella) are major secondary consumers in the Southern Ocean, placing them in potential competition with commercial fisheries and requiring research to understand their seasonal habitat use. Using the data obtained during 14 shipboard surveys sampled on a fixed grid (150 K km²) during mid- to late summer, I quantified the spatial distribution and intra-seasonal variability of fur seal sightings relative to distance to land and hydrographic boundaries. I test the hypothesis that fur seals display an increase in their at-sea abundance during mid- to late summer near the Antarctic Peninsula as they prepare to take up wintering grounds. I also test whether abundances of their potential prey, krill and myctophids, exhibit intra-seasonal variability. During midsummer, high-abundance areas are located near major breeding colonies; however, during late summer, there is an order-of-magnitude increase in fur seal abundance, coinciding with an increase in the number of high-abundance areas located in Bransfield Strait. Coincidently, abundance of Euphausia superba decreased and the myctophid Electrona antarctica increased between mid- and late-summer surveys. High-abundance areas of fur seals are not associated with the southern Antarctic Circumpolar Current front but are concentrated within 100 km from land, potentially indicating the location of haul out and important coastal habitat use areas. The dynamic increase in the number and location of high-abundance areas during late summer represents a considerable amount of mammalian predators entering the Antarctic Peninsula marine ecosystem. This information is important for understanding the seasonal impact of fur seals on regional marine food webs and their potential interaction with the autumn–winter krill fishery.     

Evidence of food-based competition among passerine migrants during stopover

Summary Local concentrations of migrating, fat-depleted birds with similar diets can lead to increased competition for food at a time when energy demand is high. Results of a predator-exclosure experiment indicate that intercontinental passerine migrants depress food abundance during stopover following migration across the Gulf of Mexico. Moreover, migrants that stop when a high number of potential competitors are present replenish energy reserves more slowly than migrants that stop under low density conditions. Competition increase the rate of food depletion and may decrease the probability that a migrant will meet its energetic requirements and complete a successful migration.     

A hierarchical,multi-scale,management-responsive model of Mediterranean vegetation dynamics

Ecosystems of the Mediterranean basin are characterized by a heterogeneous and dynamic landscape mosaic of vegetation formations. This landscape has been shaped over millenia by disturbances associated with agropastoral land use: clearing, grazing and burning, and by regeneration processes of the natural woody vegetation. The ability to predict the effects of management decisions on the structure and composition of the vegetation is essential for present and future land management. To improve this ability we developed a hierarchical multi-scale, management-responsive model of vegetation dynamics.     

Tidal influences on the habitat use of Indo-Pacific humpback dolphins in an estuary

This paper offers the first study of diurnal variations in the use of an estuarine habitat by Indo-Pacific humpback dolphins. Passive acoustic data loggers were deployed in the Xin Huwei River Estuary, Western Taiwan, from July 2009 to December 2010, to collect biosonar clicks. Acoustic encounter rates of humpback dolphins on the riverside of the estuary changed significantly over the four tidal phases, instead of the two diurnal phases based on the recordings from 268 days. Among the tidal phases, the encounter rates were lowest during ebb tides. Additionally, circling movements associated with the hunt for epipelagic fish significantly changed in temporal and spatial presence over the four tidal phases, matching the overall pattern of encounter rate changes in the focal estuary. Our findings suggest that the occurrence pattern and habitat utilization of humpback dolphins are likely to be influenced by the tidal-driven activity of their epipelagic prey.     

Spatial repartition and ontogenetic shifts in habitat use by coral reef fishes (Moorea,French Polynesia)

This study explores the extent to which ontogenetic habitat shifts modify spatial patterns of fish established at settlement in the Moorea Island lagoon (French Polynesia). The lagoon of Moorea Island was divided into 12 habitat zones (i.e. coral seascapes), which were distinct in terms of depth, wave exposure, and substratum composition. Eighty-two species of recently settled juveniles were recorded from March to June 2001. Visual censuses documented changes in the distribution of juveniles of each species over time among the 12 habitats. Two patterns of juvenile habitat use were found among species. Firstly, some species settled and remained in the same habitat until the adoption of the adult habitats (i.e. recruitment; e.g. Chaetodon citrinellus, Halichoeres hortulanus, Rhinecanthus aculeatus). Secondly, others settled to several habitats and then disappeared from some habitats through differential mortality and/or post-settlement movement (e.g. 65–70 mm size class for Ctenochaetus striatus, 40–45 mm size class for Epinephelus merra, 50–55 mm size class for Scarus sordidus). A comparison of the spatial distribution of juveniles to that of adults (61 species recorded at both stages) illustrated four patterns of subsequent recruitment in habitat use: (1) an increase in the number of habitats used during the adult stage (e.g. H. hortulanus, Mulloidichthys flavolineatus); (2) a decrease in the number of habitats adults used compared to recently settled juveniles (e.g. Chrysiptera leucopoma, Stethojulis bandanensis); (3) the use of different habitat types (e.g. Acanthurus triostegus, Caranx melampygus); and (4) no change in habitat use (e.g. Naso litturatus, Stegastes nigricans). Of the 20 most abundant species recorded in Moorea lagoon, 12 species modified the spatial patterns established at settlement by an ontogenetic habitat shift.Communicated by T. Ikeda, Hakodate     

Age-specific social dominance affects habitat use by breeding American redstarts (Setophaga ruticilla): a removal experiment

Summary By removing older males from their breeding territories, we tested the hypothesis that age-related dominance behavior influenced the pattern of habitat selection by breeding American redstarts Setophaga ruticilla (Aves: Parulinae). Fifteen older male redstarts removed in five experimental replicates during three breeding seasons were replaced by ten yearling and five older males; no redstart males of either age colonized the control sites during the same time periods, although two yearlings disappeared. Significantly more yearling males (67%, n=9) colonized the vacated areas than were present in the redstart population at large (26.8%, n=209). We reject the alternative hypothesis that yearling male redstarts occupy different habitats from older males because of age-related (innate) habitat preferences. Redstarts that colonized the territories made vacant by our removals (i.e., floaters) were a behaviorally heterogeneous group of animals. The presence of both yearling and older male floaters indicates that suitable habitat is limiting for this species and that intraspecific competitive interactions are important in habitat distribution, and potentially in population regulation.     

Tiger shark (Galeocerdo cuvier) movement patterns and habitat use determined by satellite tagging in eastern Australian waters

Partial migration is considered ubiquitous among vertebrates, but little is known about the movements of oceanodromous apex predators such as sharks, particularly at their range extents. PAT-Mk10 and SPOT5 electronic tags were used to investigate tiger shark (Galeocerdo cuvier) spatial dynamics, site fidelity and habitat use off eastern Australia between April 2007 and May 2013. Of the 18 tags deployed, 15 recorded information on depth and/or temperature, and horizontal movements. Tracking times ranged between four and 408 days, with two recovered pop-up archival tags allowing 63 days of high-resolution archived data to be analysed. Overall mean proportions of time-at-depth revealed that G. cuvier spent the majority of time-at-depths of <20 m, but undertook dives as deep as 920 m. Tagged sharks occupied ambient water temperatures from 29.5 °C at the surface to 5.9 °C at depth. Deep dives (>500 m) occurred mostly around dawn and dusk, but no definitive daily dive patterns were observed. Horizontal movements were characterised by combinations of resident and transient behaviour that coincided with seasonal changes in water temperature. While the majority of movement activity was focused around continental slope waters, large-scale migration was evident with one individual moving from offshore Sydney, Australia, to New Caledonia (c. 1,800 km) in 48 days. Periods of tiger shark residency outside of Australia’s fisheries management zones highlight the potential vulnerability of the species to unregulated fisheries and the importance of cross-jurisdictional arrangements for species’ management and conservation.     

Patch use as an indicator of habitat preference,predation risk,and competition

Summary A technique for using patch giving up densities to investigate habitat preferences, predation risk, and interspecific competitive relationships is theoretically analyzed and empirically investigated. Giving up densities, the density of resources within a patch at which an individual ceases foraging, provide considerably more information than simply the amount of resources harvested. The giving up density of a forager, which is behaving optimally, should correspond to a harvest rate that just balances the metabolic costs of foraging, the predation cost of foraging, and the missed opportunity cost of not engaging in alternative activities. In addition, changes in giving up densities in response to climatic factors, predation risk, and missed opportunities can be used to test the model and to examine the consistency of the foragers' behavior. The technique was applied to a community of four Arizonan granivorous rodents (Perognathus amplus, Dipodomys merriami, Ammospermophilus harrisii, and Spermophilus tereticaudus). Aluminum trays filled with 3 grams of millet seeds mixed into 3 liters of sifted soil provided resource patches. The seeds remaining following a night or day of foraging were used to determine the giving up density, and footprints in the sifted sand indicated the identity of the forager. Giving up densities consistently differed in response to forager species, microhabitat (bush versus open), data, and station. The data also provide useful information regarding the relative foraging efficiencies and microhabitat preferences of the coexisting rodent species.     

Sulfur isotopes link overwinter habitat use and breeding condition in Double-crested Cormorants.

North American Double-crested Cormorant (Phalacrocorax auritus) populations have increased greatly. Both breeding and overwintering ground factors have likely contributed to these increases. However, demonstrating how overwintering conditions may affect breeding birds has not been possible because of the difficulty in linking breeding birds to their wintering grounds. Here, we demonstrate the utility of stable sulfur isotopes to elucidate overwintering habitat use by cormorants breeding on Lake Erie. Sulfur isotopes in feathers grown on overwintering grounds provided insights into the degree to which birds used freshwater vs. marine environments. The proportion of birds utilizing freshwater habitats increased through time. This change may have reflected increases in freshwater aquaculture (i.e., catfish) in the U.S. south. Examination of body condition in birds returning to breed on Lake Erie indicated that those individuals that solely used marine habitats for at least a portion of the overwintering period were in poorer condition than birds using freshwater. Enhanced foraging opportunities at aquaculture facilities may improve the fitness of individuals that have returned to breed after overwintering at such locations. This study is the first to demonstrate a linkage between overwinter habitat use and breeding ground parameters in Double-crested Cormorants. These results underscore that factors throughout the Mississippi flyway are likely acting together to regulate cormorant populations.     

A general approach to the analysis of habitat selection

The investigation of animal habitat selection aims at the detection of selective usage of habitat types and the identification of covariates influencing their selection. The results not only allow for a better understanding of the habitat selection process but are also intended to help improve the conservation of animals. Usually, habitat selection by larger animals is assessed by radio-tracking or visual observation studies, where the chosen habitat is determined for some animals at a set of specific points in time. Hence the resulting data often have the following structure: a categorical variable indicating the habitat type selected by an animal at a specific point in time is repeatedly observed and will be explained by covariates. These may either describe properties of the habitat types currently available and/or properties of the animal. In this paper, we present a general approach to the analysis of such data in a categorical regression setup. The proposed model generalizes and improves upon several of the approaches previously discussed in the literature. In particular, it accounts for changing habitat availability due to the movement of animals within the observation area. It incorporates both habitat- and animal-specific covariates, and includes individual-specific random effects to account for correlations introduced by the repeated measurements on single animals. Furthermore, the assumption that the effects are linear can be dropped by including the effects in nonparametric manner based on a penalized spline approach. The methodology is implemented in a freely available software package. We demonstrate the general applicability and the potential of the proposed approach in two case studies: The analysis of a songbird community in South-America and a study on brown bears in Central Europe.     

Stingray life history trade-offs associated with nursery habitat use inferred from a bioenergetics model

Consumption rates of marine predators are vital to assessing their trophic impacts and potential consequences of fisheries removal and habitat alteration, yet are rarely estimated. Standard metabolic rates were estimated for juvenile brown stingrays, Dasyatis lata, and used as input parameters for a bioenergetics model to predict consumption rates. Temperature and mass had significant effects on metabolic rates. The energy budget of juvenile brown stingrays was heavily weighted toward metabolism, accounting for 66 % of consumed energy. Growth accounted for 7 % of the energy budget indicating very slow growth potentially due to limited food resources. Population consumption rates suggest potential for strong top-down effects on prey populations due to stingray predation. This study suggests the use of Kāne‘ohe Bay as a nursery habitat for juvenile brown stingrays is a trade-off between increased juvenile survival through predator avoidance and a late age at first maturity due to slow growth rates resulting from low prey availability.