Effects of Tidal and Diel Cycles on Dugong Habitat Use

ABSTRACT Quantifying the factors influencing behaviors of aquatic mammalian grazers may enhance the generic understanding of grazer ecology. We investigated diel and tidal patterns in movements of the dugong (Dugong dugon) by Global Positioning System—tracking 12 animals in 5 inshore—intertidal and 3 offshore—subtidal habitats along the coast of Queensland, Australia. We examined effects of tide height and time of day on the dugong's distance from 1) the nearest coast, 2) water >3 m deep, 3) actual water depth (bathymetry + tide ht) experienced, and 4) distribution of the directions of movements. Both tidal and diel cycles influenced dugong movement. Tracked dugongs tended to be closer to shore at high tide than at low tide and closer to shore at night than during the day. Onshore movement was more prevalent on incoming tides and in the afternoon and evening. Offshore movement was more prevalent on outgoing tides and from midnight through the morning until midday. Tidal and diel variation in water depths used by the inshore—intertidal dugongs was small, but probably underestimated, hidden by a sampling bias in the telemetry equipment. Onshore movement at high tide allowed dugongs to exploit intertidal seagrass beds. Dugongs are closer to shore in afternoons and evenings than in mornings. This behavior may be related to the avoidance of predators or watercraft. Our findings can be used to predict spatial patterns of dugongs within areas of conservation management significance and to assess, avoid, and mitigate adverse effects of anthropogenic disturbance.     

SHORT-TERM EFFECTS OF BOAT TRAFFIC ON BOTTLENOSE DOLPHINS, TURSIOPS TRUNCATUS, IN SARASOTA BAY, FLORIDA

Coastal cetaceans are subject to potential injury or disturbance from vessels. In Sarasota, Florida, where about 120 resident bottlenose dolphins, Tursiops truncatus , share the inshore waters with over 34,000 registered boats, disturbance potential is high. We assessed specific behavioral responses of individual dolphins to boat traffic. We conducted focal animal behavioral observations during opportunistic and experimental boat approaches involving 33 well-known identifiable individual bottlenose dolphins. Dolphins had longer interbreath intervals (IBI) during boat approaches compared to control periods (no boats within 100 m). Treatment IBI length was inversely correlated with distance to the nearest boat in opportunistic observations. During 58 experimental approaches to 18 individuals, a video system suspended from a tethered airship was used to observe subsurface responses of focal dolphins as boats under our control, operating at specified speeds, were directed near dolphins. Dolphins decreased interanimal distance, changed heading, and increased swimming speed significantly more often in response to an approaching vessel than during control periods. Probability of change for both interanimal distance and heading increased when dolphins were approached while in shallow water. Our findings provide additional support for the need to consider disturbance in management plans for cetacean conservation.     

CURRENT STATUS AND CONSERVATION NEEDS OF DUGONGS IN SOUTHERN JAPAN

We conducted aerial surveys of dugongs ( Dugong dugon ) using the line-transect method and snorkeling surveys of dugong feeding trails in 1998 and 1999 around Okinawa Island (26°30'N, 128°00'E) and the Sakishima Islands, southern Japan. A total of ten dugongs were sighted and feeding trails were confirmed in the sea grass beds off the east coast of Okinawa Island. In the Sakishima Islands, however, no dugongs were observed, and there was no evidence of feeding trails despite the existence of apparently suitable sea grass beds for feeding. The results of these surveys and other available information suggest that Okinawan dugongs represent a small, geographically isolated population. Our sightings of dugongs during the daytime, offshore of sea grass beds where feeding trails were recorded, suggest that Okinawan dugongs principally feed at night when human activities are limited. Survival of this remnant dugong population is threatened by habitat degradation and occasional entanglement mortality in fishing nets.     

Seagrass Herbivory Levels Sustain Site-Fidelity in a Remnant Dugong Population

Herds of dugong, a largely tropical marine megaherbivore, are known to undertake long-distance movements, sequentially overgrazing seagrass meadows in their path. Given their drastic declines in many regions, it is unclear whether at lower densities, their grazing is less intense, reducing their need to travel between meadows. We studied the effect of the feeding behaviour of a small dugong population in the Andaman and Nicobar archipelago, India to understand how small isolated populations graze seagrasses. In the seven years of our observation, all recorded dugongs travelled either solitarily or in pairs, and their use of seagrasses was limited to 8 meadows, some of which were persistently grazed. These meadows were relatively large, contiguous and dominated by short-lived seagrasses species. Dugongs consumed approximately 15% of meadow primary production, but there was a large variation (3–40% of total meadow production) in consumption patterns between meadows. The impact of herbivory was relatively high, with shoot densities c. 50% higher inside herbivore exclosures than in areas exposed to repeated grazing. Our results indicate that dugongs in the study area repeatedly graze the same meadows probably because the proportion of primary production consumed reduces shoot density to levels that are still above values that can trigger meadow abandonment. This ability of seagrasses to cope perhaps explains the long-term site fidelity shown by individual dugongs in these meadows. The fact that seagrass meadows in the archipelago are able to support dugong foraging requirements allows us to clearly identify locations where this remnant population persists, and where urgent management efforts can be directed.     

Waterbird responses to regular passage of a birdwatching tour boat: Implications for wetland management

Participation in outdoor recreation can increase support for wildlife conservation, but may also disturb wildlife. We examined responses of wintering waterbirds to the regular passage of a small boat specifically dedicated to birdwatching tours in a coastal Ramsar site in northern Spain. Disturbances were measured during two separate periods: 2006–2008 and 2012–2015. Incidence and magnitude of disturbance events were compared by grouping species based on their interest to birders (target vs. non-target) and compared across sectors of the tour route. Flight-initiation distances (FID) were used to estimate species-specific buffer zones, which can be used to manage recreational disturbance to waterbirds. We further examined relationships between species-specific traits and FID, time flying, and distance flying following disturbance. A single boat tour disturbed on average 0.3% of non-target and 2.8% of target wintering bird populations within the wetland, with the effect being more pronounced on target species due to their smaller populations. Wing loading was positively associated with distance flying after disturbance. Based on measured FIDs, we calculated an overall buffer zone for all species of 100 m, and species-specific buffer zones ranging from 41 to 211 m. Disturbance incidence and the number of birds disturbed per tour were both greatest in narrow tidal channels (<200 m), where boats were forced to pass within 100 m of waterbirds. We urge caution in allowing boat passage through tidal channels in which boat operators cannot effectively maintain recommended buffer zones between their boat and waterbirds.     

A review of research on the interactions between dugongs (<Emphasis Type="Italic">Dugong dugon</Emphasis> Müller 1776) and intertidal seagrass beds in Indonesia

Research on dugong–seagrass interactions in Indonesia was done during the period 1990 until 2005 in respectively East Aru, Maluku Province and East Kalimantan, Indonesia. This research investigated intensive rotational grazing by dugongs in intertidal inshore Halodule univervis seagrass meadows, dugong movements in relation to these grazing swards, and analyses of parameters explaining the temporal and spatial patterns of grazing in these meadows. In this paper, we report the findings of this long-term study. The patterns of movement and the results of snorkelling surveys confirmed a practice of regular recropping of restricted grazing swards by small feeding assemblages of dugongs. Dugong grazing showed a significant correlation with carbohydrate content of the below-ground biomass and no significant relation with total N. The timing of dugong grazing in these intertidal meadows coincides with high below-ground biomass and high carbohydrate content in the rhizomes of H. uninervis in the upper 0–4 cm sediment layer. Our findings support the hypothesis that temporal dugong grazing is ruled by carbohydrate content in below-ground biomass. The mechanisms of rotational grazing in restricted grazing swards are not yet well understood, and the maximisation of carbohydrates does not fully explain this phenomenon. Our research confirms that intertidal H. univervis seagrass meadows form a crucial resource for dugong survival. These relatively unknown biotopes need therefore more attention in research and conservation programmes.     

Distribution and abundance of the dugong in New Caledonia, southwest Pacific

New Caledonia is at the eastern limit of the dugong's range. In June 2003 standardized dugong aerial survey methodology was used to estimate the abundance and distribution of dugongs in the coastal waters of New Caledonia, resulting in a population estimate of 1,814 ± SE 332. This represents the largest concentration of dugongs in Melanesia and one of the largest populations in the world, outside Australia and the Arabian region. Calves comprised 7.2% of the population. The observed density of dugongs was highest in the center and southern part of the west coast of the island but not significantly different from the density on the north west and north east. In the central west region, sightings were associated with a pass in the barrier reef and up to a third of the on‐survey sightings were outside of the barrier reef. The dugong distribution we observed during June 2003 differs from the reported locations of historical hunts in several respects.     

Dugong habitat use in relation to seagrass nutrients,tides, and diel cycles

The dugong is the only herbivorous mammal that is strictly marine and a seagrass community specialist. The pasture available to the dugong varies with the tides because seagrass occurs in both intertidal and subtidal areas. We GPS-tracked seven dugongs within a 24 km², intensively used seagrass habitat in subtropical Australia in winter. We modeled resource selection within the habitat by comparing the dugongs’ use of space with the distribution of seagrass in an area defined using the combined space-use of the tracked animals. Selection by dugongs for seagrass quantity (biomass) and quality (nutrients) was analyzed within six time/tide combinations to examine the influences of tidal periodicity and the diel cycle on resource selection. Dugong habitat use was consistently centered over seagrass patches with high nitrogen concentrations, except during the day at low tides when the animals had fewer habitat choices and their space use was centered over high seagrass biomass. The association of dugongs with seagrass high in starch was positive during both day and night high tides when the animals could access the intertidal areas where seagrass biomass was generally low. Associations between dugongs and seagrass species were less definite, reflecting the potential for dugongs to exploit several species. Our model of dugong resource selection suggests that nitrogen is the primary limiting nutrient for dugong populations and also confirms the preference of dugongs for high-energy foods.     

Diving behaviour of dugongs, Dugong dugon

The diving behaviour of 15 dugongs (Dugong dugon) was documented using time-depth recorders (TDRs), which logged a total of 39,507 dives. The TDRs were deployed on dugongs caught at three study sites in northern Australia: Shark Bay, the Gulf of Carpentaria and Shoalwater Bay. The average time for which the dive data were collected per dugong was 10.4±1.1 (S.E.) days. Overall, these dugongs spent 47% of their daily activities within 1.5 m of the sea surface and 72% less than 3 m from the sea surface. Their mean maximum dive depth was 4.8±0.4 m (S.E.), mean dive duration was 2.7±0.17 min and the number of dives per hour averaged 11.8±1.2. The maximum dive depth recorded was 20.5 m; the maximum dive time in water >1.5 m deep was 12.3 min. The effects of dugong sex, location (study site), time of day and tidal cycle on diving rates (dives per hour), mean maximum dive depths, durations of dives, and time spent ≤1.5 m from the surface were investigated using weighted split-plot analysis of variance. The dugongs exhibited substantial interindividual variation in all dive parameters. The interaction between location and time of day was significant for diving rates, mean maximum dive depths and time spent within 1.5 m of the surface. In all these cases, there was substantial variation among individuals within locations among times of day. Thus, it was the variation among individuals that dominated all other effects. Dives were categorised into five types based on the shape of the time-depth profile. Of these, 67% of dives were interpreted as feeding dives (square and U-shaped), 8% as exploratory dives (V-shaped), 22% as travelling dives (shallow-erratic) and 3% as shallow resting dives. There was systematic variation in the distribution of dive types among the factors examined. Most of this variation was among individuals, but this differed across both time of day and tidal state. Not surprisingly, there was a positive relationship between dive duration and depth and a negative relationship between the number of dives per hour and the time spent within 1.5 m of the surface after a dive.     

Pleistocene sea level fluctuations and the phylogeography of the dugong in Australian waters

We investigated phylogeography, demography, and population connectivity of the dugong (Dugong dugon) in Australian waters using mitochondrial control region DNA sequences from 177 Australian dugongs and 11 from elsewhere. The dugong is widespread in shallow Indo‐West Pacific waters suitable for growth of its main food, seagrass. We hypothesized that the loss of habitat and creation of a land barrier (the Torres Strait landbridge) during low sea level stands associated with Pleistocene glacial cycles have left a persisting genetic signature in the dugong. The landbridge was most recently flooded about 7,000 yr ago. Individual dugongs are capable of traveling long distances, suggesting an alternative hypothesis that there might now be little genetic differentiation across the dugong's Australian range. We demonstrated that Australian dugongs fall into two distinct maternal lineages and exhibit a phylogeographic pattern reflecting Pleistocene sea‐level fluctuations. Within each lineage, genetic structure exists, albeit at large spatial scales. We suggest that these lineages diverged following the last emergence of the Torres Strait landbridge (ca. 115 kya) and remained geographically separated until after 7 kya when passage through Torres Strait again became possible for marine animals. Evidence for population growth in the widespread lineage, especially after the last glacial maximum, was detected.     

海南省儒艮资源现状调查

为拟在海南省建立儒艮自然保护区提供科学依据,于2001 年11 月至2002 年10 月在海南省西部沿岸海域进行了为期一年的海洋环境综合调查和儒艮海上目视调查。此外还在海南省及广西壮族自治区、广东省进行了有关访问调查。调查结果表明,在广西沿岸合浦海域仍有儒艮存在。但在海南省西海域近岸未发现儒艮,且原有儒艮栖息地的海草均被破坏消失,认为目前建立海南儒艮自然保护区的条件尚不成熟。儒艮资源减少的主要原因是受人类活动加剧的影响,以及生态环境不断恶化,威胁着本种的生存。为此,亟待采取有效的管理措施和健全管理机制,加强对儒艮资源及其栖息地的保护。     

Movement heterogeneity of dugongs, Dugong dugon (Müller), over large spatial scales

Seventy dugongs were fitted with satellite PTTs and/or GPS transmitters in sub-tropical and tropical waters of Queensland and the Northern Territory, Australia. Twenty-eight of the 70 dugongs were also fitted with time-depth recorders. The dugongs were tracked for periods ranging from 15 to 551 days and exhibited a large range of individualistic movement behaviours; 26 individuals were relatively sedentary (moving < 15 km) while 44 made large-scale movements (> 15 km) of up to 560 km from their capture sites. Male and female animals, including cows with calves, exhibited large-scale movements (LSM; > 15 km). Body length of travelling dugongs ranged from 1.9 to 3 m. At least some of the movements were return movements to the capture location, suggesting that such movements were ranging rather than dispersal movements. LSMs included macro-scale regional movements (> 100 km) and meso-scale inter-patch local movements (15 to < 100 km) and were qualitatively different from tidally-driven micro-scale commuting movements between and within seagrass beds (< 15 km). The mean ± S.E. macro-scale movement distance per individual was 243.8 ± 35.4 km (N = 14 individuals that travelled > 100 km), with a mean ± S.E. travel time of 179.8 ± 29.0 h. The mean ± S.E. meso-scale movement distance per individual was 49.7 ± 3.3 km (N = 28 individuals that made movements of 15-100 km), with a mean ± S.E. travel time of 52.3 ± 7.1 h. LSMs were rapid and apparently directed (mean ± S.E. travel speeds for GPS tagged animals; meso-scale movements = 1.3 ± 0.11 km/h, min = 0.3, max = 3.0; macro-scale movements = 1.6 ± 0.16 km/h, min = 0.8, max = 1.3). Tracked dugongs rarely travelled far from the coast (mean ± S.E. max distance = 12.8 ± 1.3 km). Dive profiles from the time-depth recorders suggest that dugongs make repeated deep dives while travelling rather than remaining at the surface, increasing their likelihood of capture in bottom set gill nets. Some animals caught in the high latitude limits of the dugongs' range on the Australian east coast in winter apparently undertook long distance movements in response to low water temperatures, similar to migrational movements by Florida manatees. Our findings that dugongs frequently undertake macro-scale movements have implications for management at a range of scales, and strengthen the aerial survey and genetic evidence for management and monitoring at ecological scales that cross jurisdictions.     

Long-Term Occupancy Trends in a Data-Poor Dugong Population in the Andaman and Nicobar Archipelago

Prioritizing efforts for conserving rare and threatened species with limited past data and lacking population estimates is predicated on robust assessments of their occupancy rates. This is particularly challenging for elusive, long-lived and wide-ranging marine mammals. In this paper we estimate trends in long-term (over 50years) occupancy, persistence and extinction of a vulnerable and data-poor dugong (Dugong dugon) population across multiple seagrass meadows in the Andaman and Nicobar archipelago (India). For this we use hierarchical Bayesian dynamic occupancy models accounting for false negatives (detection probability<1), persistence and extinction, to two datasets: a) fragmentary long-term occurrence records from multiple sources (1959–2004, n = 40 locations), and b) systematic detection/non-detection data from current surveys (2010–2012, n = 57). Dugong occupancy across the archipelago declined by 60% (from 0.45 to 0.18) over the last 20 years and present distribution was largely restricted to sheltered bays and channels with seagrass meadows dominated by Halophila and Halodule sp. Dugongs were not found in patchy meadows with low seagrass cover. In general, seagrass habitat availability was not limiting for dugong occupancy, suggesting that anthropogenic factors such as entanglement in gillnets and direct hunting may have led to local extinction of dugongs from locations where extensive seagrass meadows still thrive. Effective management of these remnant dugong populations will require a multi-pronged approach, involving 1) protection of areas where dugongs still persist, 2) monitoring of seagrass habitats that dugongs could recolonize, 3) reducing gillnet use in areas used by dugongs, and 4) engaging with indigenous/settler communities to reduce impacts of hunting.     

Impacts of dugong foraging on benthic animal communities in a Thailand seagrass bed

We studied the community structure and abundance of benthic animals in intertidal seagrass beds in Thailand, where dugongs (Dugong dugon) feed on the seagrass Halophila ovalis. Our objective was to examine whether the benthic community was affected by the foraging activity of the dugongs, which can cause disturbance to the sediment. We collected a total of 4156 benthic animals from three study sites and classified them into 77 different taxa. A lucinid bivalve, Pillucina sp., was the most abundant taxa (approximately 80% of individuals). Taxa richness of the benthic animals was higher in intact H.ovalis vegetation compared to dugong trails at all sites. In general, the densities of five functional groups (classified by position on/in the sediment and motility) and five dominant taxa were higher in intact seagrass vegetation compared to dugong trails, although the magnitude of this difference varied among functional groups and taxa. Densities were at least threefold higher in the seagrass vegetation than in the dugong trails for epifauna and shallow infauna, but were at most twofold higher for deep infauna. Taxa richness and abundance of the benthic animals in the dugong trails were similar between the day following, and 7–9days after, the creation of the trails. Thus, delayed and/or indirect effects of dugong herbivory on benthic communities were not detected in this study. These findings demonstrate that the foraging activity of dugongs in seagrass can have considerable impacts on the abundance and diversity of benthic organisms, although this effect varies among benthic animals with different lifestyles.     

Effects of nautical traffic and noise on foraging patterns of Mediterranean damselfish (Chromis chromis)

Chromis chromis is a key species in the Mediterranean marine coastal ecosystems where, in summer, recreational boating and its associated noise overlap. Anthropogenic noise could induce behavioural modifications in marine organisms, thereby affecting population dynamics. In the case of an important species for the ecosystem like C. chromis, this could rebound on the community structure. Here, we measured nautical traffic during the summer of 2007 in a Southern Mediterranean Marine Protected Area (MPA) and simultaneously the feeding behaviour of C. chromis was video-recorded, within both the no-take A-zone and the B-zone where recreational use is allowed. Feeding frequencies, escape reaction and school density were analysed. C. chromis specimens were also collected from 2007 to 2008 to evaluate their physiological state using the Body Condition Index as a proxy of feeding efficiency. The MPA was more exploited by nautical tourism during holidays than on weekdays, particularly in the middle of the day. Greater traffic volume corresponded with lower feeding frequencies. The escape reaction was longer in duration (>1 min) when boat passed nearby, while moored boats did not induce an escape response. We found no differences in density between schools in the A- and B-zones and worse body conditions among those individuals inhabiting the B-zone in one area only. Overall, our findings revealed a significant modification of the daily foraging habits of C. chromis due to boat noise, which was slightly buffered by no-take zones established within the MPA.     

Unmanned Aerial Vehicles (UAVs) for Surveying Marine Fauna: A Dugong Case Study

Aerial surveys of marine mammals are routinely conducted to assess and monitor species’ habitat use and population status. In Australia, dugongs (Dugong dugon) are regularly surveyed and long-term datasets have formed the basis for defining habitat of high conservation value and risk assessments of human impacts. Unmanned aerial vehicles (UAVs) may facilitate more accurate, human-risk free, and cheaper aerial surveys. We undertook the first Australian UAV survey trial in Shark Bay, western Australia. We conducted seven flights of the ScanEagle UAV, mounted with a digital SLR camera payload. During each flight, ten transects covering a 1.3 km² area frequently used by dugongs, were flown at 500, 750 and 1000 ft. Image (photograph) capture was controlled via the Ground Control Station and the capture rate was scheduled to achieve a prescribed 10% overlap between images along transect lines. Images were manually reviewed post hoc for animals and scored according to sun glitter, Beaufort Sea state and turbidity. We captured 6243 images, 627 containing dugongs. We also identified whales, dolphins, turtles and a range of other fauna. Of all possible dugong sightings, 95% (CI = 90%, 98%) were subjectively classed as ‘certain’ (unmistakably dugongs). Neither our dugong sighting rate, nor our ability to identify dugongs with certainty, were affected by UAV altitude. Turbidity was the only environmental variable significantly affecting the dugong sighting rate. Our results suggest that UAV systems may not be limited by sea state conditions in the same manner as sightings from manned surveys. The overlap between images proved valuable for detecting animals that were masked by sun glitter in the corners of images, and identifying animals initially captured at awkward body angles. This initial trial of a basic camera system has successfully demonstrated that the ScanEagle UAV has great potential as a tool for marine mammal aerial surveys.     

Mechanical disruption of seagrass in the digestive tract of the dugong

The cheek teeth in dugongs are considered to be largely non-functional whereas the oral horny pads are important both in mechanical disruption of the diet and in conveying seagrass through the mouth. Particle size distributions of digesta from 41 dead stranded dugongs were examined to investigate the relationship between degree of food breakdown, gut region and functional surface area of the mouthparts. The in vitro ease of fracture of major dietary seagrass species were compared. The rate of food breakdown through the gut appears to be more closely linked to fibre level of the diet than to size or age of the dugong and its mouthparts. Low fibre seagrass, for example Halophila ovalis , breaks down at a faster rate than high fibre seagrass, for example Zostera capricorni both in dugong guts and in vitro . Several structural characteristics of seagrass, including level and arrangement of fibre, and water content, make it particularly amenable to mechanical breakdown. The soft mouthparts of the dugong are highly modified so that the entire oral cavity functions to crush low fibre seagrasses. Thus, the dugong has developed an efficient method of food ingestion and mastication that is suited to processing large quantities of soft seagrass during short dive times. The potential cost to the dugong in having lost its hard dental surfaces is that it has become restricted to a low fibre diet.     

Improving population estimates by quantifying diving and surfacing patterns: A dugong example

Diving animals are available for detection from above the water when environmental conditions are favorable and the animals are near the surface. The number of animals that are unavailable for detection needs to be estimated to obtain unbiased population estimates. The current availability correction factors used in aerial surveys for the dugong (Dugong dugon) allow for variation in environmental conditions but use the average time dugongs spend near the surface (i.e., constant availability corrections). To improve availability estimates, we examined location and dive data from nine dugongs fitted with satellite telemetry units and time‐depth recorders (TDRs) in eastern Australia. The effects of water depth, tidal conditions, and habitat types on dugong surfacing time were examined using generalized linear mixed models (GLMMs). We found that availability for detection differed with water depth, and depth‐specific availability estimates were often lower than the constant estimates. The habitat effect was less influential, and there was no tidal effect. The number of dugongs estimated using depth‐specific availabilities were higher than those obtained using constant availabilities across water depth. Hence, information on water depth can refine availability estimates and subsequent abundance estimates from dugong aerial surveys. The methodology may be applicable to other aquatic wildlife.     

Behavioral responses to encounter of fishing boats in wandering albatrosses

Animals are attracted to human food subsidies worldwide. The behavioral response of individuals to these resources is rarely described in detail, beyond chances of encounters. Seabirds for instance scavenge in large numbers at fishing boats, triggering crucial conservation issues, but how the response to boats varies across encounters is poorly known. Here we examine the behavioral response of wandering albatrosses (Diomedea exulans), equipped with GPS tags, to longline fishing boats operating near their colony for which we had access to vessel monitoring system data. We distinguish between encounters (flying within 30 km of a boat) and attendance behavior (sitting on the sea within 3 km of a boat), and examine factors affecting each. In particular, we test hypotheses that the response to encountered boats should vary with sex and age in this long‐lived dimorphic species. Among the 60% trips that encountered boats at least once, 80% of them contained attendance (but attendance followed only 60% of each single encounter). Birds were more attracted and remained attending longer when boats were hauling lines, despite the measures enforced by this fleet to limit food availability during operations. Sex and age of birds had low influence on the response to boats, except the year when fewer boats came fishing in the area, and younger birds were attending further from boats compared to older birds. Net mass gain of birds was similar across sex and not affected by time spent attending boats. Our results indicate albatrosses extensively attend this fishery, with no clear advantages, questioning impacts on foraging time budgets. Factors responsible for sex foraging segregation at larger scale seem not to operate at this fleet near the colony and are not consistent with predictions of optimal foraging theory on potential individual dominance asymmetries. This approach complements studies of large‐scale overlap of animals with human subsidies.     

Spatial distribution of soil penetration resistance as affected by soil compaction: The fractal approach

Using semivariograms and fractal dimension (D), we identified the spatial variation of penetration resistance (PR) in variously compacted silty loam in the Lublin region, South-East Poland. Four compaction treatments were as follows: zero traffic (0p), one pass of tractor (1p), three passes of tractor (3p), and (8p) eight passes. Penetration resistance was measured in a square net of 0.6 m × 0.6 m with grid density of 0.05 m, in horizontal planes at depths of 0.05, 0.15, 0.25, 0.35, 0.45 and 0.55 m in each traffic treatment. The data were analysed in 6 horizontal planes and 12 vertical planes. The vertical planes were obtained by transforming the measured data along one side of the square at every 0.05 m. Total number of penetrations was 864. Fractal dimension (D) was estimated from the slope of the log–log semivariogram plots. The semivariograms showed spatial autocorrelation of penetration resistance in the horizontal and vertical planes. Direction in space was important in this study. In the horizontal planes the differentiation of penetration resistance semivariance at different depths was considerable and not clearly related with traffic intensity. In the vertical planes the semivariograms showed spatial dependence of the PR and evident decreasing of semivariance with increasing traffic intensity. Kriging-interpolated maps revealed that the differentiation of penetration resistance was higher in the vertical than in the horizontal planes. The vertical differentiation was higher in 0p than in the remaining treatments. The overall mean fractal dimensions in the vertical planes increased with increasing compaction levels and can be a useful indicator of the compaction level. The opposite courses of fractal dimension in the vertical and horizontal planes indicate spatial anisotropy in distribution of penetration resistance.