Genetic and historic evidence for climate-driven population fragmentation in a top cetacean predator: the harbour porpoises in European water

Recent climate change has triggered profound reorganization in northeast Atlantic ecosystems, with substantial impact on the distribution of marine assemblages from plankton to fishes. However, assessing the repercussions on apex marine predators remains a challenging issue, especially for pelagic species. In this study, we use Bayesian coalescent modelling of microsatellite variation to track the population demographic history of one of the smallest temperate cetaceans, the harbour porpoise (Phocoena phocoena) in European waters. Combining genetic inferences with palaeo-oceanographic and historical records provides strong evidence that populations of harbour porpoises have responded markedly to the recent climate-driven reorganization in the eastern North Atlantic food web. This response includes the isolation of porpoises in Iberian waters from those further north only approximately 300 years ago with a predominant northward migration, contemporaneous with the warming trend underway since the ‘Little Ice Age’ period and with the ongoing retreat of cold-water fishes from the Bay of Biscay. The extinction or exodus of harbour porpoises from the Mediterranean Sea (leaving an isolated relict population in the Black Sea) has lacked a coherent explanation. The present results suggest that the fragmentation of harbour distribution range in the Mediterranean Sea was triggered during the warm ‘Mid-Holocene Optimum’ period (approx. 5000 years ago), by the end of the post-glacial nutrient-rich ‘Sapropel’ conditions that prevailed before that time.     

Predictive Modelling to Identify Near-Shore,Fine-Scale Seabird Distributions during the Breeding Season

During the breeding season seabirds are constrained to coastal areas and are restricted in their movements, spending much of their time in near-shore waters either loafing or foraging. However, in using these areas they may be threatened by anthropogenic activities such as fishing, watersports and coastal developments including marine renewable energy installations. Although many studies describe large scale interactions between seabirds and the environment, the drivers behind near-shore, fine-scale distributions are not well understood. For example, Alderney is an important breeding ground for many species of seabird and has a diversity of human uses of the marine environment, thus providing an ideal location to investigate the near-shore fine-scale interactions between seabirds and the environment. We used vantage point observations of seabird distribution, collected during the 2013 breeding season in order to identify and quantify some of the environmental variables affecting the near-shore, fine-scale distribution of seabirds in Alderney’s coastal waters. We validate the models with observation data collected in 2014 and show that water depth, distance to the intertidal zone, and distance to the nearest seabird nest are key predictors in the distribution of Alderney’s seabirds. AUC values for each species suggest that these models perform well, although the model for shags performed better than those for auks and gulls. While further unexplained underlying localised variation in the environmental conditions will undoubtedly effect the fine-scale distribution of seabirds in near-shore waters we demonstrate the potential of this approach in marine planning and decision making.     

The distribution and abundance of planktonic penaeid larvae in Tudor creek,Mombasa, Kenya

The temporal distribution and abundance of penaeid prawn larvae in surface waters of Tudor creek were studied. Only late mysis and early post mysis stages of development were encountered in the creek waters with a descending gradient from the mouth station (station 1) to the innermost station 5. Larval encounter incidence (% incidence) was highest in plankton tows taken during night spring tides (59%) followed by night neap tides (45.4%), day spring tides (34.9%) and day neap tides (21.4%). This encounter incidence was more skewed towards the seaward stations (1 and 2) during the neap tides than during spring tides. Likewise, larval density (mean catch/standard 5 min. tow) decreased less steeply up the creek during spring tides than it did during neap tides. Higher catch rates were in general obtained in night samples than in day samples at each station and in each neap/spring phase. The diel cycle had a greater effect on both incidence and abundance of larvae than did the tidal cycle.Intensified larval incursion into the creek was observed between March and June.     

Populations of Anaerobic Phototrophic Bacteria in a Spartina alterniflora Salt Marsh

Habitat-simulating media were used with the Hungate anaerobic roll tube technique to enumerate culturable anaerobic photosynthetic bacteria in sediment, tidal waters, and Spartina alterniflora plant samples collected from the salt marsh at Sapelo Island, Ga. No phototrophs were detected in samples of creekside (low marsh) sediment or in tidal waters in creekside regions. In the high marsh region, 90% of anaerobic phototrophic bacteria occurred in the top 5 mm of sediment and none were detected below 6 mm. There was a seasonal variation, with maximal populations occurring in summer and fall (mean, 4.4 × 10⁵ phototrophs g of dry sediment⁻¹) and minimal numbers occurring in winter (mean, 3.9 × 10³ phototrophs g of dry sediment⁻¹). During winter and late spring, phototrophs had a patchy distribution over the high marsh sediment surface. In contrast, during late summer they had a random uniform distribution. Tidal water collected over high marsh sediment contained an average of 8.7 × 10² phototrophs ml⁻¹, with no significant seasonal variation. Anaerobic phototrophic bacteria were also cultured from the lower stem tissue of S. alterniflora growing in both the high (4.3 × 10⁴ phototrophs g of dry tissue⁻¹) and creekside (4.9 × 10⁴ phototrophs g of dry tissue⁻¹) marsh regions. Chromatium buderi, Chromatium vinosum, Thiospirillum sanguineum, Rhodospirillum molischianum, and Chlorobium phaeobacteroides were the predominant anaerobic phototrophic species cultured from high marsh sediment. The two Chromatium species were dominant.     

Feeding rhythms of the green-lipped mussel, Perna viridis (Linnaeus, 1758) (Bivalvia: Mytilidae) during spring and neap tidal cycles

Food availability and feeding responses of the green mussel Perna viridis were investigated for two complete tidal cycles during both spring and neap tides. Temporal changes in total particulate matter (TPM), particulate inorganic matter (PIM) and particulate organic matter (POM), were smaller during neap than spring tides. Seston characteristics at different times of a tidal cycle were compared for both spring and neap tides. Only during spring tides were TPM and PIM significantly higher at high tides while POM remained relatively constant (P>0.05). The clearance rate of the mussels underwent temporal variations with tides, and was a negative power function of TPM and a positive linear function of f (organic content), during both spring and neap tides. f was the key factor influencing filtration rate, organic ingestion rate, absorption rate and absorption efficiency. All feeding rates increased linearly with increases in organic content. Pseudofaeces were produced only during spring but not neap tides. Feeding rates and absorption efficiency were highest at low and lowest at high tides (P<0.01). There was no significant temporal change in the wet weight and protein content of the crystalline style with the tidal regime. For the digestive gland, alpha-amylase activity was higher at spring than at neap tides, and higher during high tides in a tidal cycle. The digestive gland cellulase activity did not change significantly with the tides. For the crystalline style, both the activity of cellulase and alpha-amylase were not significantly different (P>0.05) between spring and neap tides. Tidal rhythms in feeding and digestion in this species were likely controlled by temporal variations in food availability in the seawater. By adjusting feeding rates and enzymatic activities, absorption in Perna viridis remained constant, irrespective of the changes in food availability.     

Population structure of nuclear and mitochondrial DNA variation among South American Burmeister’s porpoises (<Emphasis Type="Italic">Phocoena spinipinnis</Emphasis>)

Little is known about the biology of Burmeister’s porpoises (Phocoena spinipinnis), a small cetacean species endemic to South American waters. Information on stock structure, however, is urgently needed, as the species suffers from considerable mortality due to local fishery activities throughout its distribution range. Using mitochondrial control region sequences and 11 species-specific microsatellite loci, we assessed the genetic differentiation among 118 stranded, incidentally or directly-caught Burmeister’s porpoises from different localities in Peruvian, Chilean, and Argentine waters. F-statistics and Bayesian clustering analyses indicate a major population differentiation along the South American Pacific coast, separating Peruvian from both Chilean and Argentine individuals. Interestingly, this population boundary is consistent with the population structure found in another sympatrically-occurring cetacean species: the dusky dolphin (Lagenorhynchus obscurus). Given that vulnerability to local depletion for South American coastal porpoises and dolphins is probably highest in the Peruvian population (due to high exploitation levels and recurrent El Niño events), the genetic data reported here considerably strengthen the need for conservation efforts focused on regulation of catches in local waters. Moreover, we discuss possible genetic differentiation among Burmeister’s porpoises (i) from the Atlantic and Pacific Ocean and (ii) from different Peruvian harbors. Finally, cross-species amplifications suggest that our newly-developed microsatellite markers will be useful in population genetic studies in the five other extant porpoise species.     

Breed Locally,Disperse Globally: Fine-Scale Genetic Structure Despite Landscape-Scale Panmixia in a Fire-Specialist

An exciting advance in the understanding of metapopulation dynamics has been the investigation of how populations respond to ephemeral patches that go ‘extinct’ during the lifetime of an individual. Previous research has shown that this scenario leads to genetic homogenization across large spatial scales. However, little is known about fine-scale genetic structuring or how this changes over time in ephemeral patches. We predicted that species that specialize on ephemeral habitats will delay dispersal to exploit natal habitat patches while resources are plentiful and thus display fine-scale structure. To investigate this idea, we evaluated the effect of frequent colonization of ephemeral habitats on the fine-scale genetic structure of a fire specialist, the black-backed woodpecker (Picoides arcticus) and found a pattern of fine-scale genetic structure. We then tested for differences in spatial structure between sexes and detected a pattern consistent with male-biased dispersal. We also detected a temporal increase in relatedness among individuals within newly burned forest patches. Our results indicate that specialist species that outlive their ephemeral patches can accrue significant fine-scale spatial structure that does not necessarily affect spatial structure at larger scales. This highlights the importance of both spatial and temporal scale considerations in both sampling and data interpretation of molecular genetic results.     

Climate and pH Predict the Potential Range of the Invasive Apple Snail (Pomacea insularum) in the Southeastern United States

Predicting the potential range of invasive species is essential for risk assessment, monitoring, and management, and it can also inform us about a species’ overall potential invasiveness. However, modeling the distribution of invasive species that have not reached their equilibrium distribution can be problematic for many predictive approaches. We apply the modeling approach of maximum entropy (MaxEnt) that is effective with incomplete, presence-only datasets to predict the distribution of the invasive island apple snail, Pomacea insularum. This freshwater snail is native to South America and has been spreading in the USA over the last decade from its initial introductions in Texas and Florida. It has now been documented throughout eight southeastern states. The snail’s extensive consumption of aquatic vegetation and ability to accumulate and transmit algal toxins through the food web heighten concerns about its spread. Our model shows that under current climate conditions the snail should remain mostly confined to the coastal plain of the southeastern USA where it is limited by minimum temperature in the coldest month and precipitation in the warmest quarter. Furthermore, low pH waters (pH <5.5) are detrimental to the snail’s survival and persistence. Of particular note are low-pH blackwater swamps, especially Okefenokee Swamp in southern Georgia (with a pH below 4 in many areas), which are predicted to preclude the snail’s establishment even though many of these areas are well matched climatically. Our results elucidate the factors that affect the regional distribution of P. insularum, while simultaneously presenting a spatial basis for the prediction of its future spread. Furthermore, the model for this species exemplifies that combining climatic and habitat variables is a powerful way to model distributions of invasive species.     

Predicting greater sage‐grouse habitat selection at the southern periphery of their range

Mapping suitable habitat is an important process in wildlife conservation planning. Species distribution reflects habitat selection processes occurring across multiple spatio‐temporal scales. Because habitat selection may be driven by different factors at different scales, conservation planners require information at the scale of the intervention to plan effective management actions. Previous research has described habitat selection processes shaping the distribution of greater sage‐grouse (Centrocercus urophasianus; sage‐grouse) at the range‐wide scale. Finer‐scale information for applications within jurisdictional units inside the species range is lacking, yet necessary, because state wildlife agencies are the management authority for sage‐grouse in the United States. We quantified seasonal second‐order habitat selection for sage‐grouse across the state of Utah to produce spatio‐temporal predictions of their distribution at the southern periphery of the species range. We used location data obtained from sage‐grouse marked with very‐high‐frequency radio‐transmitters and lek location data collected between 1998 and 2013 to quantify species habitat selection in relation to a suite of topographic, edaphic, climatic, and anthropogenic variables using random forest algorithms. Sage‐grouse selected for greater sagebrush (Artemisia spp.) cover, higher elevations, and gentler slopes and avoided lower precipitations and higher temperatures. The strength of responses to habitat variables varied across seasons. Anthropogenic variables previously reported as affecting their range‐wide distribution (i.e., roads, powerlines, communication towers, and agricultural development) were not ranked as top predictors at our focal scale. Other than strong selection for sagebrush cover, the responses we observed differed from what has been reported at the range‐wide scale. These differences likely reflect the unique climatic, geographic, and topographic context found in the southern peripheral area of the species distribution compared to range‐wide environmental gradients. Our results highlight the importance of considering appropriateness of scale when planning conservation actions for wide‐ranging species.     

Acoustic monitoring reveals the times and tides of harbor porpoise (Phocoena phocoena) distribution off central Oregon,U.S.A.

Harbor porpoises (Phocoena phocoena) are commonly observed in Oregon's nearshore marine environment yet knowledge of their ecosystem use and behavior remains limited, generating concerns for potential impacts on this species from future coastal development. Passive acoustic monitoring was used to investigate spatial and temporal variations in the presence and foraging activity of harbor porpoises off the Oregon coast from May through October 2014. Digital monitoring devices (DMONs) were deployed to record acoustic data (320 kHz sample rate) in two neighboring but bathymetrically different locations off the Oregon coast: (1) a site on the 30 m isobath in close proximity (<50 m) to a rocky reef, and (2) a site on the 60 m isobath in an open sandy environment. Data were analyzed with respect to two dynamic cyclic variables: diel and tidal phase. Porpoise presence at the rocky reef site was aligned with the ebb phase of the tidal forcing, while, harbor porpoise presence and foraging at the offshore, sandy bottom site was associated with night‐time foraging. The spatial and temporal patterns identified in this study suggest harbor porpoise habitat use is modulated by specific environmental conditions particular to each site that maximize foraging efficiency.     

Techniques for the objective detection of pattern in chlorophyll samples at different temporal scales: an example using data from Lough Hyne, a seasonally stratified inlet

Chlorophyll concentrations in coastal systems are frequently variable to the extent that identifying the scales where pattern occurs is very difficult. Judgements on the temporal structure of data sets are frequently rather subjective. By examining the temporal structure of chlorophyll variation in Lough Hyne with hierarchical techniques, it was possible to identify the important temporal scales objectively. Suggestions can also be made about appropriate sampling programmes for similar coastal systems. There was significant variation in measured chlorophyll concentrations between seasons and between months within seasons. High chlorophyll concentrations were more likely during spring and autumn, as would be predicted from the seasonal cycle of stratification in the lough. Seasonality could also be detected in the tidal inflow to the lough from adjacent coastal waters. More intensive sampling during the summer did not reveal any temporal structure in surface water samples. However, there were 14 day periodicities associated with measurements of depth integrated chlorophyll, oxygen, salinity, water column stability and attenuation coefficient. It is suggested that these periodicities are consistent with spring neap tidal forcing. No interannual variation in chlorophyll concentrations was detected. Examination of the power in the analysis of variance suggested that the monthly sampling frequency was unlikely to have detected differences of less than 145% between the means of pairs of years. A sampling interval of less than a week would be needed to have confidence in detecting differences of 50% between annual means.     

Landscape Variation in Tree Species Richness in Northern Iran Forests

Mapping landscape variation in tree species richness (SR) is essential to the long term management and conservation of forest ecosystems. The current study examines the prospect of mapping field assessments of SR in a high-elevation, deciduous forest in northern Iran as a function of 16 biophysical variables representative of the area’s unique physiography, including topography and coastal placement, biophysical environment, and forests. Basic to this study is the development of moderate-resolution biophysical surfaces and associated plot-estimates for 202 permanent sampling plots. The biophysical variables include: (i) three topographic variables generated directly from the area’s digital terrain model; (ii) four ecophysiologically-relevant variables derived from process models or from first principles; and (iii) seven variables of Landsat-8-acquired surface reflectance and two, of surface radiance. With symbolic regression, it was shown that only four of the 16 variables were needed to explain 85% of observed plot-level variation in SR (i.e., wind velocity, surface reflectance of blue light, and topographic wetness indices representative of soil water content), yielding mean-absolute and root-mean-squared error of 0.50 and 0.78, respectively. Overall, localised calculations of wind velocity and surface reflectance of blue light explained about 63% of observed variation in SR, with wind velocity accounting for 51% of that variation. The remaining 22% was explained by linear combinations of soil-water-related topographic indices and associated thresholds. In general, SR and diversity tended to be greatest for plots dominated by Carpinus betulus (involving ≥ 33% of all trees in a plot), than by Fagus orientalis (median difference of one species). This study provides a significant step towards describing landscape variation in SR as a function of modelled and satellite-based information and symbolic regression. Methods in this study are sufficiently general to be applicable to the characterisation of SR in other forested regions of the world, providing plot-scale data are available for model generation.     

Temporal scales of variation in settlement and recruitment of the mussel Perna perna (Linnaeus, 1758)

Population dynamics of many intertidal organisms are strongly affected by the abundance and distribution of larvae arriving on the shore. In particular, not only absolute numbers of settlers but also the degree of synchronisation of settlement can have a strong influence on whether density-dependent or density-independent processes shape adult shape populations. Temporal variation in rates of settlement and recruitment of the mussel Perna perna on the south coast of South Africa was investigated using a nested spatial design at different temporal scales. Variability in settlement at spring tides was examined at two temporal scales: lunar (to investigate the effect of state of the moon on settlement) and tidal (to investigate the influence of state of the tide on mussel settlement). Recruitment over neap tides was examined at one temporal scale, fortnight (to investigate the effect of date on mussel recruitment).Strong temporal variation was evident for both settlement and recruitment, but not at all time scales. Distinct peaks of settler/recruit abundance were observed during the lunar and neap tide studies. Recruitment intensity differed over the course of the year, and pulsing of recruitment was generally synchronised among locations. However, the strength of pulsing differed dramatically among locations, giving a significant interaction between fortnight and location. The finest temporal scale, investigated in the tidal study, did not reveal a significant effect of the state of the tide on settlement. The state of the moon (new or full) was not significant as a main factor (p = 0.052), although generally more settlers arrived on the shore during new moon. Phase of the moon appeared to have an effect on settler abundances, but only when and where densities were high.     

Carbon flux between an estuary and the ocean: a case for outwelling

The classical outwelling hypothesis states that small coastal embayments (e.g. estuaries, wetlands) export their excess production to inshore marine waters. In line with this notion, the present study tested whether the Swartkops estuary acts as source or sink for carbon. To this end, concentrations of dissolved inorganic carbon (DIC), dissolved organic carbon (DOC) and particulate organic carbon (POC) were determined hourly during the first monthly spring and neap tides over one year in the tidal waters entering and leaving the estuary. Each sampling session spanned a full tidal cycle, yielding a total of 936 concentration estimates. Carbon fluxes were calculated by integrating concentrations with water flow rates derived from a hydrodynamic model calibrated for each sampling datum. Over the year, exports to marine waters markedly exceeded imports to the estuary for all carbon species: on the basis of total spring tidal drainage area, 1083 g m^–2 of DIC, 103 g m^–2 of DOC, and 123 g m^–2 of POC left the estuary annually. Total carbon export from the estuary to the ocean amounted to 4755 tonnes, of which 83% was in the inorganic form (DIC). Thus, the bulk of carbon moving in the water column is inorganic - yet, DIC seems to be measured only rarely in most flux studies of this nature. Salt marshes cover extensive areas in this estuary and produce some carbon, particularly DOC, but productivity of the local Spartina species is low (P:B=1.1). Consequently, the bulk of carbon exported from the estuary appears to originate from the highly productive macroinvertebrate and the phytoplankton component and not from the salt marsh plants.     

Foraging strategy of the mudskipper Periophthalmus sobrinus Eggert in a Kenyan mangrove

The feeding behaviour of Periophthalmus sobrinus was studied in a population living along the channels of the Tana river delta. In November 1993 mudskippers were sampled every 2 tidal h for two consecutive spring and neap tides both during the day and night. The fishes were sexed and their stomach contents analysed. The results indicated that the mudskipper has a carnivorous diet and forages during both day and night although feeding greatly decreases after dark. Quantitative and qualitative differences in diet were found during the day and night of the two synodic and tidal phases. These differences were also demonstrable in the two sexes. The study showed that females of P. sobrinus are more selective than males and that differences between the two sexes may be related to reproductive condition. Furthermore, the mudskippers showed a tidal rhythm in feeding behaviour with peaks of activity around low tide (LT) on spring tides and around high tide (HT) on neap tides. Foraging activity seems to be influenced by the different spatial and temporal distributions of the mudskipper's prey caused by synodic, tidal and diel changes in the environment.     

Crossover Heterogeneity in the Absence of Hotspots in Caenorhabditis elegans

Crossovers play mechanical roles in meiotic chromosome segregation, generate genetic diversity by producing new allelic combinations, and facilitate evolution by decoupling linked alleles. In almost every species studied to date, crossover distributions are dramatically nonuniform, differing among sexes and across genomes, with spatial variation in crossover rates on scales from whole chromosomes to subkilobase hotspots. To understand the regulatory forces dictating these heterogeneous distributions a crucial first step is the fine-scale characterization of crossover distributions. Here we define the wild-type distribution of crossovers along a region of the C. elegans chromosome II at unprecedented resolution, using recombinant chromosomes of 243 hermaphrodites and 226 males. We find that well-characterized large-scale domains, with little fine-scale rate heterogeneity, dominate this region’s crossover landscape. Using the Gini coefficient as a summary statistic, we find that this region of the C. elegans genome has the least heterogeneous fine-scale crossover distribution yet observed among model organisms, and we show by simulation that the data are incompatible with a mammalian-type hotspot-rich landscape. The large-scale structural domains—the low-recombination center and the high-recombination arm—have a discrete boundary that we localize to a small region. This boundary coincides with the arm-center boundary defined both by nuclear-envelope attachment of DNA in somatic cells and GC content, consistent with proposals that these features of chromosome organization may be mechanical causes and evolutionary consequences of crossover recombination.     

Temporal and spatial patterns in the abundance of wintering Red-breasted Mergansers Mergus serrator in an estuary

Tidal, temporal and spatial aspects in the abundance of wintering Red-breasted Mergansers in the Ythan estuary, northeast Scotland, were investigated. The number and location of birds were examined in relation to four variables: tidal cycle and tidal range, and month and time of day. A significant increase in the numbers of this piscivorous bird occurred as the winter progressed. Numbers were highest in early morning and then decreased significantly during the day. Spring tides were favoured over neap tides and a significant increase in numbers was recorded between low tide and high tide. The lower part of the estuary was preferred over the upper part with no detectable shift in the preference occurring in relation to any of the four variables.     

Long-Term Data Reveal a Population Decline of the Tropical Lizard Anolis apletophallus,and a Negative Affect of El Nino Years on Population Growth Rate

Climate change threatens biodiversity worldwide, however predicting how particular species will respond is difficult because climate varies spatially, complex factors regulate population abundance, and species vary in their susceptibility to climate change. Studies need to incorporate these factors with long-term data in order to link climate change to population abundance. We used 40 years of lizard abundance data and local climate data from Barro Colorado Island to ask how climate, total lizard abundance and cohort-specific abundance have changed over time, and how total and cohort-specific abundance relate to climate variables including those predicted to make the species vulnerable to climate change (i.e. temperatures exceeding preferred body temperature). We documented a decrease in lizard abundance over the last 40 years, and changes in the local climate. Population growth rate was related to the previous years’ southern oscillation index; increasing following cooler-wetter, la niña years, decreasing following warmer-drier, el nino years. Within-year recruitment was negatively related to rainfall and minimum temperature. This study simultaneously identified climatic factors driving long-term population fluctuations and climate variables influencing short-term annual recruitment, both of which may be contributing to the population decline and influence the population’s future persistence.     

Environmental heterogeneity, spatial segregation of prey, and the utilization of southwest Atlantic mudflats by migratory shorebirds

We studied the relationships between the habitat use of migratory shorebirds and the spatial distributions of the Southwestern Atlantic Fiddler Crab Uca uruguayensis , polychaetes, sediment characteristics and tidal levels in the Río de La Plata estuary, Argentina, where U. uruguayensis is one of the most important intertidal species. Crabs have a well-defined patchy distribution that is segregated spatially from that of polychaetes. Crab density on the surface varied across the tidal cycle, reaching maximum values during low tide. Polychaete density decreased with depth but showed no change through the tidal cycle; however, given that sediment penetrability did change during the same period, their availability to probing shorebirds is expected to change. Habitat use by shorebirds followed the spatial distribution of prey; shorebirds that foraged on polychaetes (White-rumped Sandpiper Calidris fuscicollis , Two-banded Plover Charadrius falklandicus and Hudsonian Godwit Limosa haemastica ) focused their attention on the areas with the highest densities of polychaetes, whereas species that preyed mostly on crabs (Ruddy Turnstone Arenaria interpres , Whimbrel Numenius phaeopus and Grey Plover Pluvialis squatarola ) predominantly used areas with crabs. This segregation occurred particularly during low tide, a period in which polychaetes became fully available. Results show that the spatial and temporal heterogeneity of mudflats in relation to the types and availability of prey has a strong effect on shorebird habitat use.     

Brown Adipose Tissue in Cetacean Blubber

Brown adipose tissue (BAT) plays an important role in thermoregulation in species living in cold environments, given heat can be generated from its chemical energy reserves. Here we investigate the existence of BAT in blubber in four species of delphinoid cetacean, the Pacific white-sided and bottlenose dolphins, Lagenorhynchus obliquidens and Tursiops truncates, and Dall’s and harbour porpoises, Phocoenoides dalli and Phocoena phocoena. Histology revealed adipocytes with small unilocular fat droplets and a large eosinophilic cytoplasm intermingled with connective tissue in the innermost layers of blubber. Chemistry revealed a brown adipocyte-specific mitochondrial protein, uncoupling protein 1 (UCP1), within these same adipocytes, but not those distributed elsewhere throughout the blubber. Western blot analysis of extracts from the inner blubber layer confirmed that the immunohistochemical positive reaction was specific to UCP1 and that this adipose tissue was BAT. To better understand the distribution of BAT throughout the entire cetacean body, cadavers were subjected to computed tomography (CT) scanning. Resulting imagery, coupled with histological corroboration of fine tissue structure, revealed adipocytes intermingled with connective tissue in the lowest layer of blubber were distributed within a thin, highly dense layer that extended the length of the body, with the exception of the rostrum, fin and fluke regions. As such, we describe BAT effectively enveloping the cetacean body. Our results suggest that delphinoid blubber could serve a role additional to those frequently attributed to it: simple insulation blanket, energy storage, hydrodynamic streamlining or contributor to positive buoyancy. We believe delphinoid BAT might also function like an electric blanket, enabling animals to frequent waters cooler than blubber as an insulator alone might otherwise allow an animal to withstand, or allow animals to maintain body temperature in cool waters during sustained periods of physical inactivity.