Satellite imagery can be used to detect variation in abundance of Weddell seals (Leptonychotes weddellii) in Erebus Bay, Antarctica

The Weddell seal population in Erebus Bay, Antarctica, represents one of the best-studied marine mammal populations in the world, providing an ideal test for the efficacy of satellite imagery to inform about seal abundance and population trends. Using high-resolution (0.6 m) satellite imagery, we compared counts from imagery to ground counts of adult Weddell seals and determined temporal trends in Erebus Bay during November 2004–2006 and 2009, and December 2007. Seals were counted from QuickBird-2 and WorldView-1 images, and these counts were compared with ground counts at overlapping locations within Erebus Bay during the same time. Counts were compared across years and within individual haul-out locations. We counted a total of 1,000 adult Weddell seals from five images across all years (for a total of 21 satellite-to-ground count comparisons), approximately 72% of the total counted on the ground at overlapping locations. We accurately detected an increase in abundance during 2004–2009. There was a strong, positive correlation (r = 0.98, df = 3, P < 0.003) between ground counts and counts derived from the imagery. The correlation between counts at individual haul-out locations was also strong (r = 0.80, df = 19, P < 0.001). Detection rates ranged from 30 to 88%. Overall, our results showed the utility of high-resolution imagery to provide an accurate way to detect the presence and variation in abundance of Weddell seals. Our methods may be applied to other species in polar regions, such as walruses or polar bears, particularly in areas where little is known about population status.     

Harbor seal population decline in the Aleutian Archipelago

Populations of Steller sea lions, northern fur seals, and northern sea otters declined substantially during recent decades in the Bering Sea and Aleutian Islands region, yet the population status of harbor seals has not been assessed adequately. We determined that counts obtained during skiff‐based surveys conducted in 1977–1982 represent the earliest estimate of harbor seal abundance throughout the Aleutian Islands. By comparing counts from 106 islands surveyed in 1977–1982 (8,601 seals) with counts from the same islands during a 1999 aerial survey (2,859 seals), we observed a 67% decline over the ～20‐yr period. Regionally, the largest decline of 86% was in the western Aleutians (n= 7 islands), followed by 66% in the central Aleutians (n= 64 islands), and 45% in the eastern Aleutians (n= 35 islands). Harbor seal counts decreased at the majority of islands in each region, the number of islands with >100 seals decreased ～70%, and the number of islands with no seals counted increased ～80%, indicating that harbor seal abundance throughout the Aleutian Islands was substantially lower in the late 1990s than in the 1970s and 1980s.     

DIVING BEHAVIOR OF THE PACIFIC HARBOR SEAL (PHOCA VITULINA RICHARDII) IN MONTEREY BAY, CALIFORNIA

Physical environment and physiological characteristics of marine mammals potentially affect the duration and depth of diving. Härkönen (1987b) proposed a hypothesis that the harbor seal would gain maximum energy by foraging at intermediate depths. To investigate this hypothesis, we studied diving behavior of the Pacific harbor seal (Phoca vitulina ricbardii) during 1995 through 1997 in Monterey Bay, California. Dive depths (n = 13,063 dives) were recorded via time‐depth recorders. Approximately 80% of recorded dives were classified as square dives (type I), which typically were associated with foraging in pinnipeds. Approximately 11% of dives were V dives (type II; 1,402 dives), and the remainder (1,225 dives) were skewed dives (type III and IV). The deepest recorded dive was 481 m, while the greatest duration was 35.25 min. Body mass explained the variability of durations of long dives for females (95th percentile; D₉₅♂=‐5.47 + 0.18 × (mass♀), r²= 0.91, 95% CI for slope = [0.08, 0.28], n= 5) and for males (D₉₅♂=‐5.86 + 0.18 × (mass♀), r²= 0.83, 95% CI for slope = [0.12, 0.24], n= 11). The large proportion of variability in deep dives, however, was explained by body mass only for males (95th percentile; Z₉₅♂=‐363.9 + 6.05 × (mass♀), r²= 0.83, 95% CI for slope = [3.93, 8.17], n= 11) and not for females (Z₉₅,♂=?148.1 +3.11 × (mass♀), r²= 0.58, 95% CI for slope = [‐1.7, 7.9], n= 5, 95% CI for slope= [?1.7, 7.9]). Median depths of presumed foraging dives of harbor seals in the Monterey Bay area were between 5 and 100 m, which were within the range of the previously reported depths for other areas (< 100 m). Our findings generally supported Härkönen's hypothesis that harbor seals forage in the intermediate depth in their environment.     

Postrelease dive ability in rehabilitated harbor seals

The efficacy of seal rehabilitation is examined in a postrelease study of dive ability in harbor seal pups (Phoca vitulina) in the Wash, United Kingdom. Six rehabilitated seals were fitted with Sea Mammal Research Unit (SMRU) Argos Satellite Relay Data Logger tags and their individual dive behavior was monitored for an average of 122 d. The upper 90 percentile edge of dive behavior (dive duration [DD₉₀] and percentage of time at‐sea spent in a dive [PD₉₀]), in 7 d bins, was used as a proxy for physiological dive ability. The results are compared with data from five wild adult harbor seals. There was no statistically significant difference between (1) the mean track duration of rehabilitated seals (126.20 ± 27.48 [SD] d) and adult seals (150.2 ± 24.62 d) (P= 0.108), indicating no evidence that short‐term survival was less in the rehabilitated group; (2) the mean mass‐scaled DD₉₀ of rehabilitated seals (3.95 ± 0.37 min) and adult seals (4.09 ± 0.55 min) (P= 0.632); and (3) the mean PD₉₀ of rehabilitated seals (81.62 ± 1.21%) and adult seals (81.48 ± 3.93%) (P= 0.943). These three results all suggest the success of the rehabilitation program in terms of short‐term survival and dive ability.     

Resource partitioning by sympatric pagophilic seals in Alaska: monitoring effects of climate variation with fatty acids

Four species of pagophilic phocid seals in the Western Arctic—bearded (Erignathus barbatus), ringed (Phoca hispida), ribbon (Phoca fasciata), and spotted (Phoca largha)—are particularly vulnerable to arctic warming trends. Documenting diet composition over time is one way in which the flexibility of a species, in the face of broad ecosystem changes, can be measured. Because the fatty acid (FA) composition of depot lipids has long been known to reflect diet, we analysed the FA composition of blubber samples collected from bearded (n = 30), ringed (n = 15), ribbon (n = 32) and spotted seals (n = 24). All animals were taken near Little Diomede Island in May and June 2003, providing a unique opportunity to study interspecies differences with minimal effects of spatial and temporal variation. Bearded seal FA composition was significantly different from all other seals (P < 0.001 in all cases), ringed seal FA composition was significantly different from that of both ribbon and spotted seals (P < 0.001), but ribbon and spotted seal FA compositions could not be distinguished from each other. Overall, the blubber FA compositions of ribbon, ringed and spotted seals implied diets dominated by pelagic prey. Inferences we made about current diets of bearded and ringed seals suggest that certain prey important to them in this area in the 1970s remain important today, despite notable changes in the ecosystem. We believe that blubber FA analysis will be a useful tool in assessing the response of ice-associated pinnipeds and their ecosystem to changes associated with arctic warming.     

MONITORING THE TREND OF HARBOR SEALS IN PRINCE WILLIAM SOUND,ALASKA, AFTER THE EXXON VALDEZ OIL SPILL

We used aerial counts to monitor the trend in numbers of harbor seals, Phoca vitulina richardsi, in Prince William Sound, Alaska, following the 1989 Exxon Valdez oil spill. Repetitive counts were made at 25 haul-out sites during the annual molt period each year from 1990 through 1997. A generalized linear model indicated that time of day, date, and time relative to low tide significantly affected seal counts. When Poisson regression was used to adjust counts to a standardized set of survey conditions, results showed a highly significant decline of 4.6% per year. Unadjusted counts indicated a slight, but not statistically significant, decline in the number of seals. The number of harbor seals on the trend-count route in eastern and central PWS has been declining since at least 1984, with an overall population reduction of 63% through 1997. Programs to monitor long-term changes in animal population sizes should account for factors that can cause short-term variations in indices of abundance. The inclusion of such factors as covariates in models can improve the accuracy of monitoring programs.     

Reaction of Harbor Seals to Cruise Ships

Abstract: The largest aggregations of harbor seals (Phoca vitulina) in Alaska, USA, haul out on floating ice in tidewater glacial fjords. Seals use these fjords in peak numbers during the critical periods of pupping, breeding, and molting when visits by tour ships also peak. Documented and suspected declines of harbor seals in fjords with rising vessel traffic underscore the need to better understand possible impacts, particularly in areas where ship visits have risen substantially in the past 2 decades. We examined the interruption of haul-out bouts of harbor seals due to approaching cruise ships in Disenchantment Bay, Alaska. We conducted observations from cruise ships and focused on disturbance of seals as evidenced by seals flushing into the water from the floating ice on which they rested. We investigated rate of flushing in relation to vessel distance, approach angle, group size, and seal type (mother, pup, or other). Using a survival-regression analysis, we found that the risk of disturbing harbor seals increased when ships approached within 500 m; seals approached as close as 100 m were 25 times more likely to enter the water than seals 500 m from a ship. Seals were 4 times more prone to enter the water when ships were approaching directly rather than passing abeam. Seals responded similarly regardless of group size or seal type. Energetic models indicated a potential to disrupt energy balance and cause thermal stress in disturbed pups if they spent >50% of their time in ice-chilled water. Studies at non-glacial sites suggest that pups spend 40–70% of their time in the water. Voluntary guidelines for approaching seals in Alaska recommend that cruise ships approach ≥91 m (100 yards), a distance at which we show 90% of seals would flush into the water. Our findings indicate a need to develop regulations to maintain a 500-m separation between cruise ships and seals in all Alaskan glacial fjords.     

MOVEMENTS OF SATELLITE-TAGGED SUBADULT AND ADULT HARBOR SEALS IN PRINCE WILLIAM SOUND, ALASKA

Satellite-linked tags were attached to 49 subadult and adult harbor seals captured in Prince William Sound (PWS), Alaska, and their movements were monitored during 1992–1997. Seals were tracked for a total of 5,517 seal-days and were located on about 80% of the days that tags transmitted. Most locations were in or near PWS, but some juvenile seals moved 300–500 km east and west into the Gulf of Alaska. While several seals travelled to 50–100 km offshore, virtually all locations were in water <200 m deep. Overall, juvenile seals moved more than adults and had larger home ranges. Movements were significantly affected by month, and age by month and sex by month interactions. In all months, mean distances between successively used haulouts were <10 km for adults and <20 km for juveniles. Mean monthly home ranges varied from <100 km² to >1,500 km², and were smallest during June-July. Mean haul-out to at-sea distance was 5–10 km for adults and generally 10–25 km for juveniles. Satellite-linked tags provided an effective means of monitoring and describing the full range of harbor seal movements in this region, with the exception of late summer when tags were shed during the molt.     

Interannual differences in postrelease movements of rehabilitated harbor seal pups (Phoca vitulina richardii) in the Salish Sea

Despite the large number of harbor seals (Phoca vitulina richardii) rehabilitated worldwide, few studies have been conducted on postrelease movement and behavior of rehabilitated harbor seal pups. We compared interannual differences in movements and survival of 24 rehabilitated seal pups released in the Salish Sea in 2010 (n = 10), 2012 (n = 5), 2013 (n = 5), and 2014 (n = 4). We also compared the postrelease movement of these seals to the movement of 10 wild seal pups tracked in the same ecosystem in 2010. Transmission duration, total cumulative distance, and average daily distance varied annually. Maximum linear distance traveled from the release site was similar for the rehabilitated seal groups. Compared to wild seals (n = 10), and consistent with prior studies, rehabilitated pups (n = 24) traveled significantly farther daily and cumulatively than wild weaned pups. Unlike in a prior study in this ecosystem, we found no significant difference between transmission duration in wild and rehabilitated pups.     

DIEL HAUL-OUT PATTERNS AND SITE FIDELITY OF HARBOR SEALS (PHOCA VITULINA RICHARDSI) ON SAN MIGUEL ISLAND, CALIFORNIA, IN AUTUMN

We studied the haul-out patterns and movements of harbor seals ( Phoca vitulina richardsi ) on San Miguel Island, California, from 23 October through 6 December 1982 by attaching a radio transmitter to each of 18 seals and monitoring their presence ashore with continuously scanning receivers. Seals hauled out at all hours although, on average, the largest proportion of tagged seals was ashore between 1300 and 1500 h. Median durations of haul-out bouts of individual seals ranged from 4.7 to 21.8 h; 81% of all haul-out bouts were less than 12 h and 3% were longer than 24 h. Eighty-one percent of the seals that were resighted at least twice used only the sites where they were tagged; two seals used two sites and one seal used three. Most seals were hauled out on fewer than 51% of the days sampled. On average, about 41% of tagged seals hauled out each day whereas an average of about 19% was hauled out during peak afternoon hours. Using telemetry data to correct a count of 412 seals made during an aerial survey, we estimated absolute abundance at about 2,168 seals; a modified Peterson mark-recapture model produced an estimate of about 1,445 seals.     

The effect of a large Danish offshore wind farm on harbor and gray seal haul-out behavior

This study investigates the effects of the construction and operation of a large Danish offshore wind farm on harbor and gray seal haul-out behavior within a nearby (4 km) seal sanctuary. Time-lapse photography, visual monitoring, and aerial surveys were used to monitor the number of seals on land in daylight hours. Seals were monitored during two preconstruction periods (19 June–31 August 2001 and April–August 2002), a construction period of the wind farm (August 2002–December 2003), and a period of operation of the wind farm (December 2003–December 2004). Monthly aerial surveys were conducted to estimate the proportion of seals in the sanctuary relative to neighboring haul-out sites. From preconstruction to construction and through the first year of operation the number of harbor seals in the sanctuary increased at the same rate as the number of seals at the neighboring haul-out sites. No long-term effects on haul-out behavior were found due to construction and operation of the wind farm. However, a significant short-term decrease was seen in the number of seals present on land during sheet pile driving in or near the wind farm. Acoustic deterrents were utilized simultaneously to avoid hearing damage.     

Fine-Scale Variability in Harbor Seal Foraging Behavior

Understanding the variability of foraging behavior within a population of predators is important for determining their role in the ecosystem and how they may respond to future ecosystem changes. However, such variability has seldom been studied in harbor seals on a fine spatial scale (<30 km). We used a combination of standard and Bayesian generalized linear mixed models to explore how environmental variables influenced the dive behavior of harbor seals. Time-depth recorders were deployed on harbor seals from two haul-out sites in the Salish Sea in 2007 (n = 18) and 2008 (n = 11). Three behavioral bout types were classified from six dive types within each bout; however, one of these bout types was related to haul-out activity and was excluded from analyses. Deep foraging bouts (Type I) were the predominant type used throughout the study; however, variation in the use of bout types was observed relative to haul-out site, season, sex, and light (day/night). The proportional use of Type I and Type II (shallow foraging/traveling) bouts differed dramatically between haul-out sites, seasons, sexes, and whether it was day or night; individual variability between seals also contributed to the observed differences. We hypothesize that this variation in dive behavior was related to habitat or prey specialization by seals from different haul-out sites, or individual variability between seals in the study area. The results highlight the potential influence of habitat and specialization on the foraging behavior of harbor seals, and may help explain the variability in diet that is observed between different haul-out site groups in this population.     

Contrasting behavior between two populations of an ice‐obligate predator in East Antarctica

The Austral autumn–winter is a critical period for capital breeders such as Weddell seals that must optimize resource acquisition and storage to provision breeding in the subsequent spring. However, how Weddell seals find food in the winter months remains poorly documented. We equipped adult Weddell seals after their annual molt with satellite‐relayed data loggers at two sites in East Antarctica: Dumont D'Urville (n = 12, DDU) and Davis (n = 20). We used binomial generalized mixed‐effect models to investigate Weddell seals’ behavioral response (i.e., “hunting” vs. “transit”) to physical aspects of their environment (e.g., ice concentration). Weddell seal foraging was concentrated to within 5 km of a breathing hole, and they appear to move between holes as local food is depleted. There were regional differences in behavior so that seals at Davis traveled greater distances (three times more) and spent less time in hunting mode (half the time) than seals at DDU. Despite these differences, hunting dives at both locations were pelagic, concentrated in areas of high ice concentration, and over areas of complex bathymetry. There was also a seasonal change in diving behavior from transiting early in the season to more hunting during winter. Our observations suggest that Weddell seal foraging behavior is plastic and that they respond behaviorally to changes in their environment to maximize food acquisition and storage. Such plasticity is a hallmark of animals that live in very dynamic environments such as the high Antarctic where resources are unpredictable.     

Determining a correction factor for aerial surveys of harbor seals in California

Counts of pinnipeds provide a minimal estimate of population size because some unknown proportion of individuals is in the water during surveys. We determined a correction factor (CF) for Pacific harbor seals (Phoca vitulina richardii) by estimating the proportion ashore of 180 seals tagged with flipper‐mounted radio tags throughout California. The mean proportions of tagged individuals ashore during four complete surveys in 2004 were not different between central and northern California (F= 1.85, P= 0.18) or between sexes (F= 0.57, P= 0.45), but a lesser proportion of weaners was ashore than subadults or adults (F= 7.97, P= 0.001), especially in northern California. The CF calculated for the statewide census of harbor seals was 1.65, using transmitters operating during the survey (n= 114). Using a mark‐recapture estimator for tag survival (phi) and the four telemetry surveys the mean CF for central and northern California was 1.54 ± 0.38 (95% CI). A CF for southern California of 2.86 was based on a single survey. Using the mean CF of 1.54 and a statewide count in 2009 we estimated 30,196 (95% CI = 22,745–37,647) harbor seals in California.     

Factors Affecting Haul-Out Behavior of Harbor Seals (Phoca vitulina) in Tidewater Glacier Inlets in Alaska: Can Tourism Vessels and Seals Coexist?

Large numbers of harbor seals (Phoca vitulina) use habitat in tidewater glaciers in Alaska for pupping, breeding, and molting. Glacial fjords are also popular tourist destinations; however, visitation by numerous vessels can result in disturbance of seals during critical life-history phases. We explored factors affecting haul-out behavior of harbor seals at a glacial site frequented by tourism vessels. In 2008-10, we deployed VHF transmitters on 107 seals in Endicott Arm, Alaska. We remotely monitored presence and haul-out behavior of tagged seals and documented vessel presence with time-lapse cameras. We evaluated the influence of environmental and physical factors on the probability of being hauled out, duration of haul-out bouts, and as factors associated with the start and end of a haulout. Location, season, hour, and interactions of location by year, season, hour, and sex significantly influenced haul-out probability, as did ice, weather, and vessels. Seals were more likely to be hauled out with greater ice availability during the middle of the day, and less likely to be hauled out if vessels were present. Cruise ships had the strongest negative effect; however, most vessel types negatively affected haul-out probability. Haul-out duration was longest in association with starting on incoming tides, clear skies, no precipitation, occurring in the middle of the day, and ending in the late afternoon or evening. End of haulouts was associated with increasing cloud cover, low ice availability, and vessel presence; large-sized tourism vessels or all-vessel-types combined were significant predictors of ending a haul-out bout. Probability of being hauled out was highest in June, during pupping season. Potential disturbances of harbor seals could be reduced, enabling longer resting times for seals and fewer interruptions for nursing pups, if vessels focused the majority of visits to glacial habitat to before or after the hours of 08:00-17:00 or, less optimally, 09:00-16:00.     

Breeding habitat and lair structure of the ringed seal (Phoca hispida ladogensis) in northern Lake Ladoga in Russia

The breeding habitat and lair structures of the ringed seal (Phoca hispida ladogensis) were studied by snowmobile expeditions in northern Lake Ladoga, Russia, during 1996 to 1999. Mostly flat ice and no large pack ice existed in northern Lake Ladoga during this study. All birth lairs and 88% of haul-out lairs were located in snowdrifts in the shorelines of islands or islets and 12% of the haul-out lairs occurred in pressured ice ridges. Some large haul-out lairs had been used communally by the seals. A total of 15% of the lairs observed had been attacked or marked by red fox (Vulpes vulpes), wolf (Canis lupus) or unidentified canines. The colour of the lanugo fur of the Ladoga ringed seal pup appeared to be dark grey. We conclude that the northern part of Lake Ladoga is an active breeding area for the ringed seal, which attests conservation needs for this area. Accepted: 18 September 2000     

Harbour seal grouping as an anti-predator strategy

Terrestrial grouping by western Atlantic harbour seals, Phoca vitulina concolor, was investigated to determine whether it has a functional significance and, if so, what the function might be. Two nofunctional hypotheses were tested: (1) groups arise from a shortage of haul-out sites or space on sites; and (2) terrestrial groups are extensions of groups formed in the water for reasons not related to hauling-out. Aerial surveys of haul-out site use by seals indicated that sites were not in short supply. Observations of the use of space on one haul-out site suggested that space on sites was not limiting. Arrivals and departures of seals at one study site showed that groups formed at the haul-out site and disbanded after the haul-out period. The hypothesis that grouping functions to increase the probability of predator detection was tested by measuring individual and group vigilance relative to group size, and simulating predator attacks in the field. Although individual vigilance decreased with increasing group size, group vigilance increased. Larger groups detected ‘attacks’ at greater distances and a solitary seal failed to detect an ‘attack’. It is proposed that an individual increases its probability of detecting a predator by joining a group.     

Haul-out behavior of harbor seals (Phoca vitulina) in Hood Canal, Washington

The goal of this study was to model haul-out behavior of harbor seals (Phoca vitulina) in the Hood Canal region of Washington State with respect to changes in physiological, environmental, and temporal covariates. Previous research has provided a solid understanding of seal haul-out behavior. Here, we expand on that work using a generalized linear mixed model (GLMM) with temporal autocorrelation and a large dataset. Our dataset included behavioral haul-out records from archival and VHF radio tag deployments on 25 individual seals representing 61,430 seal hours. A novel application for increased computational efficiency allowed us to examine this large dataset with a GLMM that appropriately accounts for temporal autocorellation. We found significant relationships with the covariates hour of day, day of year, minutes from high tide and year. Additionally, there was a significant effect of the interaction term hour of day : day of year. This interaction term demonstrated that seals are more likely to haul out during nighttime hours in August and September, but then switch to predominantly daylight haul-out patterns in October and November. We attribute this change in behavior to an effect of human disturbance levels. This study also examined a unique ecological event to determine the role of increased killer whale (Orcinus orca) predation on haul-out behavior. In 2003 and 2005 these harbor seals were exposed to unprecedented levels of killer whale predation and results show an overall increase in haul-out probability after exposure to killer whales. The outcome of this study will be integral to understanding any changes in population abundance as a result of increased killer whale predation.     

Habitat use by harbour seals (Phoca vitulina) in a seasonally ice-covered region, the western Hudson Bay

Over the last few decades, the period of ice cover in Hudson Bay has decreased, owing to climate warming, with breakup occurring approximately 3 weeks earlier than it did 30 years ago. The trend towards lengthening of the open water season has led to speculation that ringed seal numbers would decline, but then harbour seals might become numerous enough to replace ringed seals in the diet of polar bears. The movement patterns of 18 harbour seals equipped with satellite-linked transmitters in the Churchill River estuary (western Hudson Bay) were examined, as well as the dive behaviour of 11 of these seals. During the ice-free period, seals followed a general central place-foraging strategy, making repeated trips between their haul-out site in the Churchill River estuary and nearshore areas (<20 km) near the river mouth and estuary. Seal behaviour changed significantly as ice started to form along the coast of western Hudson Bay: animals remained significantly farther from the Churchill River haul-out site and from the coast and performed longer and deeper dives. However, throughout the entire tracking period, whether ice was present or not, all animals restricted their movements to a narrow band of shallow coastal waters (<50 m depth) along a 600-km stretch of the western Hudson Bay coastline, centred on the Churchill River estuary haul-out site. This natural self-limitation to nearshore shallow waters could restrict the potential for the population to increase in size and replace ringed seals as a primary energy resource for polar bears.     

Mark-resight estimates of seasonal variation in harbor seal abundance and site fidelity

Monitoring trends in abundance of pinnipeds typically involves counting seals at terrestrial haul-out sites during the breeding season. Counts of seals made at other times of the year are typically lower; however, it is often unknown whether this is because fewer animals are present or whether lower counts simply reflect a reduction in haul-out probability. Here we illustrate how photo-identification data from an individual-based study of harbor seals (Phoca vitulina) can be used to estimate seasonal variation in abundance and site fidelity. Monthly data collected over a two-year period were analyzed using a mark-recapture mark-resight model accounting for individuals transitioning between observable and unobservable states. Levels of site fidelity were high throughout the year and abundance estimates showed no seasonal pattern. This suggests that individual seals used haul-out sites to the same extent throughout the year, and that peaks in counts during the breeding season are a result of seasonal changes in haul-out probability. The results of this study have implications for understanding population sub-structuring, gene flow and disease spread.