GROWTH OF THE FILAMENTOUS GREEN ALGA CTENOCLADUS CIRCINNATUS (CHAETOPHORALES,CHLOROPHYCEAE) IN RELATION TO ENVIRONMENTAL SALINITY1

Clones of the filamentous green alga Ctenocladus circinnatus Borzi were isolated from algae collected at Abert Lake (Oregon) and Mono Lake (California). Stock cultures were exposed to varied salinities of natural lake water to examine the effects on growth rate, cell form, chlorophyll a, and water content. Growth rates were reduced in both clones with increased salinity over the range 25–100 g·L^?1 and were almost completely inhibited at 150 g·L^?1. Chlorophyll a increased between salinities of 25 and 100 g·L^?1, reflecting slower growth, higher proportions of akinetes, and smaller cell sizes as salinity increased. Tissue water content remained essentially constant from 25 to 100 g·L^?1 salinity. Shorter cell dimensions with increased salinity suggest that a lower surface-to-volume ratio may reduce the potential for passive loss of cell water. Prior acclimation of stock cultures to elevated salinity provided no enhancement of growth response at any salinity. The results indicate that environmental salinity can limit the productivity and distribution of Ctenocladus in nature.     

Effects of an epiphytic alga on abundances of fish and decapods associated with the seagrass Zostera capricorni

During a field experiment to test effects of changes to leaf height and density of Zostera capricorni on associated fish and decapods, more than one-third of the substratum at some sites was covered by the epiphytic brown alga Giffordia mitchelliae. This paper reports apparent effects of the alga on the 13 most abundant species of fish and decapods, and interactions between effects of the alga and those due to height and density of the seagrass leaves. Abundances of three species were significantly lower, and those of two species were significantly greater where the alga occurred. There were interactions between effects of the alga and those of leaf height and/or density for another four species. In all four cases, when the alga had an effect, it was to decrease abundance. The effects of Giffordia were generally opposite to those reported for drift algae in seagrass habitats elsewhere. The lower abundances of seven species where algal cover was great are best explained by an existing model for seagrass habitats that predicts reductions in abundance of individuals when the ratio of plant material to substratum is very high.     

Predation risk predicts use of a novel habitat

Predator–prey interactions are often highly co‐evolved, with selection over time for prey with morphological and behavioral traits that minimize predation risk. Consequently, in many environments prey choose among potential habitats according to their refuge value. It is unclear, however, when presented with new habitats, if prey are able to evaluate the predation risk of these relative to familiar habitats and utilize these in accordance with their value. We tested whether, along the east coast of the USA, native mud crabs Panopeus herbstii utilize the non‐native alga Gracilaria vermiculophylla according to its relative refuge value. Experiments examining predation by blue crabs Callinectes sapidus on mud crabs revealed that the non‐native alga had an intermediate refuge value relative to native oysters, which were the most protective, and unvegetated sediment, which was the least. In subsequent choice experiments, mud crabs selected oysters over alga over unvegetated sediment, in accordance with habitat refuge values. Further, in field experiments, the use of Gracilaria by mud crabs was inversely related to the proximity of the alga to the preferred habitat type, oysters, and was reduced by the presence of a blue crab predator. Consequently, mud crabs are utilizing the non‐native alga Gracilaria in accordance with its intermediate refuge value. The relative refuge value of non‐native vs native habitat‐forming species may provide a baseline expectation against which to measure the speed of learning and opportunism in the response of native prey to novel protective habitats.     

The Colonisation of Human-made Structures by the Invasive Alga Codium fragile ssp. tomentosoides in the North Adriatic Sea (NE Mediterranean)

Human-made structures, such as groynes, breakwaters, seawalls, pier pilings and floating pontoons, are becoming common features of the landscape in urbanised coastal and estuarine areas. Despite this tendency few studies have focused on their ecology or on their potential impacts on natural assemblages of organisms. When artificial structures are introduced in areas with little or no hard substrata, they not only provide novel habitats, which enables the colonisation of sandy areas by hard-bottom dwelling species, but they can also provide suitable habitats for exotic species. Along the north-east coast of Italy, sandy shores are protected from erosion by a line of breakwaters, which runs almost uninterrupted for about 300 km. These structures provide habitat for a variety of macroalgae and invertebrates and also for the invasive green alga Codium fragile ssp. tomentosoides. The aim of this study was, therefore, to investigate patterns of distribution of this alga on breakwaters in Cesenatico. In particular, we compared the density of thalli, biomass, length and degree of branching of C. fragile ssp. tomentosoides between the landward and the seaward sides of breakwaters, to test the hypothesis that sheltered habitats (landward) represent more suitable habitats than exposed habitats (seaward). In general, the landward side of breakwaters supported greater numbers of thalli of C. fragile ssp. tomentosoides than seaward sides. Thalli grew longer and more branched in sheltered habitats, leading to an overall larger biomass of the alga on the landward side of breakwaters. The presence of sheltered human-made hard substrata in the vicinity of major trading ports and sources of eutrophication could enhance the dispersal of invasive species across regional and geographic scales. Thus, the effects of artificial structures and introduced species on coastal assemblages cannot be evaluated separately, but their synergistic nature should be considered in planning strategies for conservation of biodiversity in coastal habitats.     

Ecological aspects of Sargassum muticum (Fucales,Phaeophyta) in Baja California,Mexico: reproductive phenology and epiphytes

The reproductive phenology and epiphytic macroalgae of Sargassum muticum were studied through an annual cycle (September 1987 to November 1988) at two sites on the northwestern coast of Baja California, Mexico, which were subjected to different degrees of wave exposure. Sargassum muticum is a brown alga of Japanese origin, now considered a permanent member of the marine flora of Baja California. A similar reproductive development was observed at both sites, with a maximum percentage of reproductive plants from May to July (spring–summer) and minimum from December to March (winter). Reproductive plants were found throughout the year. A total of 48 species of epiphytes were identified and seasonal variation in their diversity was observed. The greatest diversity was found at the more protected site.     

The dominant <Emphasis Type="Italic">Ulva</Emphasis> strain of the 2008 green algal bloom in the Yellow Sea was not detected in the coastal waters of Qingdao in the following winter

The region of Qingdao, China, experienced the world's largest green tide from May to July 2008. More than one million tons of fresh algal biomass of the green alga Ulva prolifera was harvested, while more was suspected to have sunk to the bottom. The original source of this seaweed was suspected to be from the south as revealed by satellite images. The floating biomass drifted with the water current northward and flourished in nearshore waters around Qingdao. However, direct biological evidence for “seed” source is lacking. It is still unclear whether this alga could survive the Qingdao local coastal environment and pose future danger of potential blooming. Systematic and seasonal sampling of waters in the intertidal zone at six collection sites along the Qingdao coast was conducted from December 2008 to April 2009. Forty-eight water samples were analyzed. From these, nine different morphotypes of Ulva were grown in the laboratory under standard temperature and light regimes. Growth of Ulva was observed in all water samples. However, molecular phylogenetic analyses revealed that the dominant U. prolifera strain of the 2008 bloom was absent in all the water-derived cultures during the sampling period. These results provide evidence that the dominant bloom-forming alga was unlikely able to survive the coastal waters (the minimal surface water temperature in February is 2°C) in winter conditions in Qingdao, even though all the sampling locations were heavily covered by this alga in June 2008.     

Littorina mariae — a factor structuring low shore communities?

The role of littorinids in structuring communities is discussed. On hard substrates preferential grazing of the dominant algae often enables competitively inferior species to utilize the rock substrate. Investigations of these systems has led to the proposal of a number of factors that should be analyzed in order to assess the effect of herbivores on community structure. Some of these factors have been investigated for Littorina mariae Sacchi & Rastelli. L. mariae is a micro-epiphytic grazer browsing the surface covering of epiphytes off the alga Fucus serratus (L.). The relationship between the host alga, F. serratus, and L. mariae is far more intricate than that between winkles and hard substrates as the alga itself is a dynamic resource. L. mariae is spatially and temporally linked to F. serratus. L. mariae is found almost exclusively on the alga and is positively attracted by extracts of the alga. The life history of the winkle is closely synchronized with that of the alga (the winter decrease of L. mariae populations is associated with the seasonal die-back of the host alga). On analysis of the factors considered important to assess herbivory, and examples of the effects of other epiphytic herbivores on algal success, it is suggested that L. mariae could potentially play an important role in structuring the community of it's host fucoid and that this may influence larger scale community structure. Experimental manipulations are proposed to evaluate these hypotheses.     

The morphology,distribution and ecology of Pseudococcomyxa simplex (Mainx) Fott (Chlorophyta,Chlorellaceae), a widespread terrestrial Antarctic alga

Summary Previous antarctic records of Pseudococcomyxa simplex are summarized and, together with new records, the alga is shown to have a circumpolar distribution in terrestrial habitats. It has been encountered over a latitudinal range of 54° to 77°S in sub-, maritime and coastal continental Antarctica, from sea-level to an altitude of 3600 m. The alga has been observed in grassland soils, as an epiphyte on a variety of mosses, in lithosols, in volcanic fumarolic soils and as a chasmoendolithophyte. A previous misidentification of this alga as Monodus subterraneus Boye Pet. is corrected. The morphology of antarctic specimens is deccribed using light and electron microscopy, the latter for the first time with this alga. Antarctic and european specimens are shown to be almost identical. It is concluded that culture studies of wide-ranging samples from antarctic terrestrial habitats will probably reveal additional cosmopolitan species.     

Temperature,Herbivory and Epibiont Acquisition as Factors Controlling the Distribution and Ecological Role of an Invasive Seaweed

The invasive canopy alga, Codium fragile ssp. tomentosoides, first observed at the Isles of Shoals in 1983, has become the dominant canopy species to 8 m throughout the islands. Codium populations are replacing themselves at most sites in what appears to be a new, climax, canopy species. However, Codium densities have declined in protected Gosport Harbor areas where it first became established. Codium has only slowly expanded its presence in adjacent nearshore subtidal habitats. Recent studies suggest a combination of factors that may be influencing the relative success of populations between habitats. The herbivorous sea slug, Placida dendritica, may be reducing populations in protected areas in spite of predators such as the green crab, Carcinus maenas, while surge may inhibit herbivore buildup in exposed habitats. Temperature instability due to localized, wind-driven upwelling may be slowing the buildup of subtidal Codium populations in nearshore sites. The combination of Codium dominance and the acquisition of increasing epibiont diversity are producing a new, potentially more complex community state than the previous kelp-dominated climax typical of the Gulf of Maine.     

Modification of Sediments and Macrofauna by an Invasive Marsh Plant

Invasive grasses have recently altered salt marsh ecosystems throughout the northern hemisphere. On the eastern seaboard of the USA, Phragmites australis has invaded both brackish and salt marsh habitats. Phragmites australis influence on sediments and fauna was investigated along a salinity and invasion-age gradient in marshes of the lower Connecticut River estuary. Typical salinities were about 19–24 ppt in Site I, 9–10 ppt in Site II and 5–7 ppt in Site III. Strongest effects were evident in the least saline settings (II and III) where Phragmites has been present the longest and exists in monoculture. Limited influence was evident in the most saline region (I) where Phragmites and native salt marsh plants co-occur. The vegetation within Phragmites stands in tidal regions of the Connecticut River generally exhibits taller, but less dense shoots, higher above-ground biomass, and lower below-ground biomass than does the un-invaded marsh flora. There were lower sediment organic content, greater litter accumulation and higher sediment chlorophyll a concentrations in Phragmites- invaded than un-invaded marsh habitat. Epifaunal gastropods (Succinea wilsoni and Stagnicola catascopium) were less abundant in habitats where Phragmites had invaded than in un-invaded marsh habitat. Macro-infaunal densities were lower in the Phragmites-invaded than un-invaded habitats at the two least saline sites (II and III). Phragmites stands supported more podurid insects, sabellid polychaetes, and peracarid crustaceans, fewer arachnids, midges, tubificid and enchytraeid oligochaetes, and greater habitat-wide taxon richness as measured by rarefaction, than did the un-invaded stands. The magnitude and significance of the compositional differences varied with season and with site; differences were generally greatest at the oldest, least saline sites (II and III) and during May, when faunal densities were higher than in September. However, experimental design and the 1-year study period precluded clear separation of salinity, age, and seasonal effects. Although structural effects of Phragmites on salt marsh faunas are evident, further investigation is required to determine the consequences of these effects for ecosystem function. This revised version was published online in July 2006 with corrections to the Cover Date.     

HABITAT DURATION,LENGTH OF LARVAL PERIOD,AND THE EVOLUTION OF A COMPLEX LIFE CYCLE OF A SALAMANDER,AMBYSTOMA TEXANUM

In many organisms, genotypic selection may be a less effective means of adapting to unpredictable environments than is selection for phenotypic plasticity. To determine whether genotypic selection is important in the evolution of complex life cycles of amphibians that breed in seasonally ephemeral habitats, we examined whether mortality risk from habitat drying in natural populations of small-mouthed salamanders (Ambystoma texanum) corresponded to length of larval period when larvae from the same populations were grown in a common laboratory environment. Comparisons were made at two levels of organization within the species: 1) among geographic races that are under strongly divergent selection regimes associated with the use of pond and stream habitats and 2) among populations within races that use the same types of breeding habitats. Morphological evidence indicates that stream-breeding A. texanum evolved from pond-breeding populations that recently colonized streams. Larvae in streams incur heavy mortality from stream drying, so the upper bound on length of larval period is currently set by the seasonal duration of breeding sites. We hypothesized that selection would reduce length of larval period of pond-breeders that colonize streams if their larval periods are inherently longer than those of stream-breeders. The results of laboratory experiments support this hypothesis. When grown individually in a common environment, larvae from stream populations had significantly shorter larval periods than larvae from pond populations. Within races, however, length of larval period did not correlate significantly with seasonal duration of breeding sites. When males of both races were crossed to a single pond female, offspring of stream males had significantly shorter larval periods than offspring of pond males. Collectively, these data suggest that differences in complex life cycles among pond and stream-breeders are due to genotypic selection related to mortality from habitat drying. Stream larvae in the common-environment experiment were significantly smaller at metamorphosis than pond larvae. Yet, the evolution of metamorphic size cannot be explained readily by direct selection: there are no intuitively obvious advantages of being relatively small at metamorphosis in streams. A positive phenotypic correlation was observed between size at metamorphosis and length of larval period in most laboratory populations. A positive additive genetic correlation between these traits was demonstrated recently in another amphibian. Thus, we suspect that metamorphic size of stream-breeders evolved indirectly as a consequence of selection to shorten length of larval period.     

Tibetan sacred sites conserve old growth trees and cover in the eastern Himalayas

Khawa Karpo, in the eastern Himalayas, is a mountain considered sacred throughout Tibet, and is internationally recognized as a global biodiversity hotspot. Numerous areas within this landscape are considered ‘sacred’ by the indigenous Tibetans of the region, who interact with these sites in ways potentially beneficial to conservation. Our previous remote sensing study indicated that sacred sites are found in habitats with greater species richness, diversity, and endemism than randomly selected non-sacred sites. This study examines the role of sanctity in biodiversity conservation within habitats in the Khawa Karpo region by pairing plots within the same habitats in sacred and non-sacred areas. Understory richness, diversity, cover, and number of useful species are measured; for trees, richness, diversity, cover, and density are measured. Results indicate that within habitats sanctity does not affect understory plant communities; however, within sacred areas trees are larger (p = 0.003) and forests have greater cover (p = 0.003) than non-sacred areas. Our results indicate that, whereas placement of sacred areas and preservation of vegetation cover affects useful plants, biodiversity and endemism, within habitats sacred sites preserve old growth trees and forest structure. In sum, Tibetan sacred sites are ecologically unique and important for conservation on varying scales of landscape, community, and species.     

Survival types of high mountain plants under extreme temperatures

Extreme temperatures are a main factor limiting plant growth in high mountain habitats. During winter, the risk of frost damage is highest at windblown and often snow-free sites. During summer, actively growing plants are particularly endangered by episodic cold spells, but also by short-term overheating. The current review gives an overview of extreme temperatures in the European Alps and observations of temperature damage on plants in their natural habitats. Furthermore, seasonal time courses of frost and heat resistance derived from laboratory tests on different plant growth forms are presented. Study species were the cushion plants Silene acaulis, Minuartia sedoides, Saxifraga oppositifolia and Carex firma collected on wind-exposed ridges; the rosette plant Soldanella alpina collected on snow-protected sites, and three Sempervivum species collected in xerothermic habitats.     

Colonization of UK coastal realignment sites by mosquitoes: implications for design,management, and public health

Coastal realignment is now widely instituted in the UK as part of local flood risk management plans to compensate for the loss of European protected habitat and to mitigate the effects of sea‐level rise and coastal squeeze. Coastal aquatic habitats have long been known to provide suitable habitats for brackish‐water mosquitoes and historically, coastal marshes were considered to support anopheline mosquito populations that were responsible for local malaria transmission. This study surveyed the eight largest managed realignment (MRA) sites in England (Essex and the Humber) for mosquito habitats. The apparent absence of anopheline mosquitoes exploiting aquatic habitats at all of these sites suggests that the risk of malaria associated with MRA sites is currently negligible. However, three of the eight sites supported populations of two nuisance and potential arboviral vector species, Aedes detritus and Aedes caspius. The aquatic habitats that supported mosquitoes resulted from a) specific design aspects of the new sea wall (ballast to mitigate wave action and constructed saline borrow ditches) that could be designed out or managed or b) isolated pools created through silt accretion or expansion of flooded zones to neighbouring pasture. The public health risks and recommendations for management are discussed in this report. This report highlights the need for pro‐active public health impact assessments prior to MRA development in consultation with the Health Protection Agency, as well as the need for a case‐by‐case approach to design and management to mitigate mosquito or mosquito‐borne disease issues now and in the future.     

New evidence for the potential role of Culex pipiens mosquitoes in the transmission cycle of West Nile virus in Tunisia

Physiological and molecular characteristics of natural populations of Culex pipiens Linnaeus, 1758 (Diptera: Culicidae) were investigated to elucidate how this species is potentially involved in the transmission of West Nile virus in Tunisia. A total of 215 Cx. pipiens females from 11 breeding habitats were analysed in the laboratory to estimate autogeny and stenogamy rates. They were tested individually for the locus CQ11 to distinguish between the two Cx. pipiens forms, pipiens and molestus. All tested Cx. pipiens populations were stenogamous. Females from underground breeding sites were all autogeneous, whereas females from above‐ground habitats were mostly anautogeneous. Of all the females tested, 59.7% were identified as pipiens, 22.4% as molestus, and 17.9% as hybrid pipiens/molestus. Furthermore, both Cx. pipiens forms and their hybrids were found to co‐occur in sympatry in all sites. The results of this study represent the first evidence that both Cx. pipiens forms and their hybrids are present in Tunisia. Because hybrids able to act as bridge vectors are present in all studied habitats, Tunisia can be considered to have a high degree of receptivity for the establishment of West Nile virus zoonotic cycles.     

Facultative mutualism between an herbivorous crab and a coralline alga: advantages of eating noxious seaweeds

Because encrusting coralline algae rely on herbivory or low light levels to prevent being overgrown by competitively superior fleshy algae, corallines are relatively rare in shallow areas with low rates of herbivory. In contrast to this general trend, the branching coralline alga Neogoniolithon strictum occurs primarily in shallow seagrass beds and along the margins of shallow reef flats where herbivory on macrophytes is low. This alga apparently persists in these habitats by providing refuge to the herbivorous crab Mithrax sculptus at mean densities of 1 crab per 75 g of algal wet mass. When crabs were removed from some host corallines, hosts without crabs supported 9 times the epiphytic growth of hosts with crabs after only 30 days. Crabs without access to a coralline alga were rapidly consumed by reef fishes, while most of those tethered near a host alga survived. These results suggest that the crabs clean their algal host of fouling seaweeds and associate with the host to minimize predation. However, to effectively clean the host, the crab must consume the wide array of macroalgae that commonly co-occur with coralline algae in these habitats, including chemically defended species in the genera Halimeda, Dictyota, and Laurencia. Crabs did readily consume these seaweeds, which were avoided by, and are chemically defended from, herbivorous fishes. Even though crabs readily consumed both Halimeda and Dictyota in whole-plant feeding assays, chemical extracts from these species significantly reduced crab feeding, suggesting that factors other than secondary chemistry (e.g., food value, protein, energy content), may determine whole-plant palatability. Having the ability to use a wide variety of foods, and choosing the most profitable rather than the least defended foods, would diminish foraging time, increase site fidelity, and allow the crab to function mutualistically with the host alga. Despite the obvious benefit of associating with N. strictum, M. sculptus did not prefer it over other habitats offering a structurally similar refuge, suggesting that these crabs are not N. strictum specialists, but rather occupy multiple habitats that provide protection from predators. Structurally complex organisms like N. strictum may commonly suppress competitors by harboring protective symbionts like M. sculptus. It is possible that diffuse coevolution has occurred between these two groups; however, this seems unlikely because both herbivore and host appear to respond most strongly to selective pressures from predators and competitors outside this association.     

Seasonality and distribution of epilithic diatoms,macroalgae and macrophytes in a spring-fed stream system in Ontario,Canada

A study of the epilithic diatom, macroalgal and macrophyte communities from a spring-fed stream in Ontario, Canada was undertaken from September 1996 to July 1997. The relative abundance of the epilithic diatom flora, percent cover of macroalgal and macrophyte taxa, and several physical and chemical stream conditions were monitored along a 20-m stretch at each of four sites, approximately every 2 months. Several stream conditions were relatively constant over the sampling period (pH, maximum width and maximum depth), while others exhibited a distinct seasonal pattern (water temperature, specific conductance and daylength) and some fluctuated strongly with no discernable seasonal pattern (turbidity, current velocity). A total of 124 taxa were identified from the four sites, including 79 epilithic diatoms, three macroalgal diatom species (large gelatinous masses), one cyanobacterium, two red algae, eight green algae, one chrysophyte alga, one tribophyte alga, three mosses, three horsetails and 23 angiosperm taxa. Species richness was positively correlated to stream channel maximum width and depth, indicating that the total number of species tends to increase in a downstream direction. Distribution of several diatom and macroalgal species was significantly correlated to stream conditions (e.g. Gomphonema parvulum and Phormidium subfuscum with current velocity); however, the vast majority of species did not display seasonal variation in abundance that could be explained by changes in stream conditions. Many of the taxa identified from Blue Springs Creek are common elsewhere in North America.     

Site specific distribution of the mud shore crab <Emphasis Type="Italic">Helograpsus haswellianus</Emphasis> in temperate wetlands

An important aspect in determining the ecology of a species is an understanding of its abundance and distribution. The population structure and distribution of Helograpsus haswellianus (Brachyura, Grapsidae) were evaluated over 12 months at four intertidal wetland sites that differed in vegetation and inundation. Surface activity of H. haswellianus varied over time with no consistent seasonal pattern. Sex ratios were always significantly skewed towards males, and crabs of both sexes were distributed evenly across habitats at all sites. Recruitment patterns and size frequency distributions varied significantly between sites, independent of habitats. Crabs were smaller in wetter, more sheltered areas, but size ranges overlapped between habitats. The site-specific difference in population structure and distribution of H. haswellianus indicates that the functional role of this burrowing crab may vary across wetlands.     

Seasonal dynamics of macrophyte communities from a stream flowing over granite flatrock in North Carolina,USA

The seasonal distribution and abundance of benthic macrophytes were characterized from second- and third-order segments of a stream flowing over granite flatrock in the southeastern United States. Eighteen genera were identified over two annual cycles including macroalgae (60% of the total), angiosperms (30%), and bryophytes (10%). Light availability as affected by riparian shading was a major factor influencing community structure. Based on strong agreement among two-way indicator species analysis, detrended correspondence analysis and cluster analysis, we identified four communities characteristic of distinct light regimes and seasons. In shaded sites the red alga Lemanea australis was dominant during cool seasons, and the aquatic moss Fontinalis sp. was dominant during warm seasons. By contrast, in open sites L. australis and the angiosperm Podostemum ceratophyllum were co-dominant during cool seasons, and P. ceratophyllum was also dominant in warm seasons. The prolific macrophyte communities followed a pattern of broad seasonal maxima for dominant species along with rapid fluctuations in ephemerals. The community dynamics suggest that competitive interactions control space partitioning among macrophytes on the granite flatrock.     

Winter sites and cold-hardiness of two gastropod species in a boreal river

Summary The overwintering sites and cold-hardiness of Gyraulus acronicus and Valvata piscinalis were investigated in a northern Swedish river subject to wide seasonal fluctuations in water-level and ice-cover. More than 95% of the population of young and older G. acronicus overwintered in the frozen hydrolittoral zone. No specimens of V. piscinalis were found in the frozen zone. The cold-hardiness in G. acronicus in unfrozen sites was maximal in November and early December. V. piscinalis was not cold-hardy at any time of the year. These interspecific differences in cold-hardiness accord with the differences in overwintering habitats.