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(11)Department of Ecology and Evolutionary Biology, University of California, (​16)Department of Marine Sciences, University of South Alabama, Mobile, AL (39)Biosecurity South Australia, Primary Industries and Regions South Movement patterns were primarily explained by the interaction.


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Tsong kha pa's Commentary on the Yogacara Doctrine of Mind Tson-kha-pa Blo-​bzan-grags-pa says the source of Tsong kha pa' s explanation is the Guhyartha​-vyakhya. This verse is identical to the version of the verse quoted from the MSam 39). Guhyartha-vyakhyaP MSU Pb7;cp.


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Endnote 4—Amendment history An Act relating to the amended Hague Rules has the meaning given in section 7. Australia, when used in (b) the operation of section 11 or 16 of the Carriage of Goods by Sea Act ” 22 Regulations.


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Endnote 4—Amendment history An Act relating to the amended Hague Rules has the meaning given in section 7. Australia, when used in (b) the operation of section 11 or 16 of the Carriage of Goods by Sea Act ” 22 Regulations.


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However, the importance of this instability in the context of ocean waves is not well interactions and associated modulational instabilities inherent to the excess kurtosis is smaller since it is of (see also Methods section).


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Tsong kha pa's Commentary on the Yogacara Doctrine of Mind Tson-kha-pa Blo-​bzan-grags-pa says the source of Tsong kha pa' s explanation is the Guhyartha​-vyakhya. This verse is identical to the version of the verse quoted from the MSam 39). Guhyartha-vyakhyaP MSU Pb7;cp.


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Tracks are coloured according to sex, with locations of inferred resident behavioural state shown as circles. Future population assessments considering feeding ground density dependence have also been recommended It is unknown to what extent prey availability, as a consequence of habitat quality in the Southern Ocean, may provide a constraint on HBW population numbers. The spatial distribution of whaling catches Fig. It seems likely that these three important elements are together potentially facilitating the strong recovery of the WA population 15 , 22 : the availability of suitable breeding, calving and resting areas along the WA coast 47 , a relatively direct migration compared to other HBW populations , and temporally persistent high-quality prey in a productive boundary-current ecosystem. Humpback whale populations migrate extensively between winter breeding grounds and summer feeding grounds, however known links to remote Antarctic feeding grounds remain limited in many cases. Satellite telemetry is helping to clarify migratory routes, providing detailed information about key corridors, important habitats and potential for overlap with human activities 26 , 27 , 28 , 29 , 30 , 31 , 32 , We show directed movements towards the southern Kerguelen plateau, a known area of high biological productivity 34 , 35 , 36 , including a focal usage of the western boundary current 37 , 38 , 39 by both male and female HBWs. Detailed oceanographic fronts 37 , as surveyed the summer prior during the KAXIS marine ecosystem survey, are shown as bold black lines. Recovery patterns following intensive industrial whaling vary across populations, with differing levels of uncertainty, but the recent comprehensive assessment by the IWC Scientific Committee projected the Western Australian WA population to have the highest abundance, in the range of 18,—24, Western breeding stock D and Eastern Australian breeding stock E1 populations have amongst the highest increase rates estimated, in the order of 9— In contrast, the neighbouring Oceania population comprising breeding stocks E2, E3, F1 and F2 remains listed as endangered by the International Union for Conservation of Nature 17 and show no apparent recovery trend 18 but see also However, improved understanding of HBW feeding grounds may elucidate the role of factors such as migration length and energetic requirements in regulating recovery 22 , and provide an improved context for understanding likely impacts of future ecosystem changes 23 , 24 , Links between specific breeding grounds and remote, relatively unsurveyed Antarctic feeding grounds remain poorly understood in many cases e. It is important to note that we did not tag mother-calf pairs which tend to move more slowly 33 , Hence, the difference in movement rates might be simply explained by size differences with the larger body size of females enabling faster travel relative to males. A single Southern Ocean migration available from a female WA whale ID tagged in 41 tracked during an earlier deployment is shown in yellow. Reported whaling catches west of the SKP are comparatively low until the Weddell gyre. For Southern Hemisphere populations, early knowledge of humpback whale HBW movements were derived from commercial catch distributions and basic Discovery tag mark-recaptures 4 , 5. With respect to the spatial extent of the krill biome, an empirical relationship derived for krill growth rate in relation to chlorophyll and temperature has recently been applied across the Southern Ocean 55 , This high-growth habitat contracts farther southward towards the Antarctic continent by January The SKP western boundary current may therefore represent one of the locations where winter ice extent overwintering krill larvae coincides with high adult potential growth rates during summer, enabling the krill lifecycle to be completed successfully 55 , their Fig. Background shading shows bathymetry Here, the topographic steering turns these fronts sharply to the north before they retroflect southward again and continue meandering eastward 35 , 37 , This important western boundary current 39 is known to advect subpolar waters and sea ice northward, extending sea-ice extent and duration in this region Migration pathways for satellite-tracked Western Australian humpback whales detailing the focus at the southern Kerguelen plateau. Humpback whale Megaptera novaeangilae migrations, between low-latitude winter breeding grounds and high-latitude summer feeding areas, represent one of the most extensive 1 , 2 , 3 and consistent mammalian migrations known. The remaining tags commenced reporting once offshore from WA. We consider this new information within the historical context of the spatial distribution of whaling catches, and what is known of other HBW movements in the region. Tracks from whales which continued into January IDs , , and show movements southward from Banzare Bank, into the Princess Elizabeth Trough and the north of Prydz Bay. Ship-based survey sightings highlight cetacean densities around the southern and eastern flanks of the SKP Both catch and survey data show generally high cetacean densities extend farther northward i. HBWs are known for the variety and complexity of their feeding behaviours, with capture techniques that include bubble nets, tail slaps, lunges and bottom feeding The distribution T. Summarised dive data available for ID in the vicinity of the northern Kerguelen plateau area The four whales with the longest tag life records that continued past mid-December all logged locations here. However, regional-scale contact between, and interchange of, individuals is assumed to take place primarily on shared feeding areas in the Southern Ocean 10 , Discovery mark-recaptures imply that HBWs from WA stock D are likely to mix with individuals from eastern Australia stock E1 , particularly via east-to-west movements on the feeding grounds 5 and 42 , their Fig. Bathymetry contour 91 shows the m isobath. Early work proposed the Southern Boundary of the ACC to be of circumpolar ecological importance to the productivity of Southern Ocean ecosystems and the baleen whales which depend upon them {/INSERTKEYS}{/PARAGRAPH} The topographic steering of currents dictates both the southern extent of the ACC and the northern limit of subpolar waters, which likely influences the distribution and productivity of marine ecosystem at multiple trophic levels Examining KAXIS dissolved iron measurements within the broader context of in situ and satellite chlorophyll- a concentrations, Schallenberg and co-authors 36 concluded the regional ocean circulation plays an important role in supplying the iron necessary to sustain these persistent blooms. With respect to the question regarding likely alternative prey, for bulk-feeding baleen whales swarming or schooling organisms are the most lucrative food resource Globally, the known prey of HBWs include a wide assortment of schooling fish including herring Clupea spp. This new research clarifying biophysical linkages provides a deepened understanding of the importance of this feeding ground to migrating HBWs, where the conditions support predictable and persistent high biological productivity. However the krill net tow data 57 necessary to confirm this hypothesis are only typically available from scientific research voyages conducted during mid-summer January—February, as in KAXIS and closer to the Antarctic continent 58 , Since whaling operations favour low ice conditions, spatial distributions drawn from historical whale catches may also bias towards the Antarctic continent in summer. The detailed migration pathways presented here enrich our knowledge of the long-distance migrations undertaken by WA humpback whales and their usage of important Southern Ocean seasonal feeding grounds. These highlight a focal feeding area during austral spring and early summer at the southern Kerguelen plateau, in a western boundary current where a sharp northward turn and retroflection of ocean fronts occurs along the eastern plateau edge. These cetacean distributions reflect the overall patterns of regional biological productivity and oceanographic processes 43 , 44 , The regional surface ocean circulation and frontal structure, including sharp northward turn and retroflection of the fronts in a western boundary current along the eastern plateau margin Fig. Relatively short and directed migratory pathways overall, together with high-quality, reliable forage resources may provide a partial explanation for the ongoing strong recovery demonstrated by this population. Once south of Australia i. Movement modelling discriminates sex-related behaviours, with females moving faster during both transit and resident periods, which may be a consequence of size or indicate differential energetic requirements. A Nature Research Journal. The northward horizontal advection of subsurface iron, likely originating from Antarctic shelf sediments in the south, and subsequent upwelling — along the eastern margin of the SKP and in association with frontal processes — combine to create a biological hotspot. Alternatively, with high pregnancy rates recently reported in southern hemisphere HBWs 73 it is possible these females were pregnant, driving high energetic requirements and hence greater foraging effort. This individual showed a northernmost terminus turning point at Overall, these investigations suggest that WA breeding ground movements are likely to be of a similar, or somewhat smaller, scale than the comparable movements of Oceania HBWs. Our analysis of movements on the forage grounds, showing significantly higher movement rates by females during both transit and residency, further raises interesting questions regarding the energetic requirements and benefits between the sexes. Spatial distributions derived from catches are also confounded by many other conditions e. New satellite tracks detail humpback whale migration pathways from Western Australia into the Southern Ocean. In comparison, isotopic profiles of 13 East Australian stock E1 animals showed this population was more likely to feed outside the traditional Antarctic zone; for example, with supplementary temperate feeding as has been regularly reported 51 , 52 , Recent satellite tracking analyses also indicate supplemental feeding during residencies in south-eastern Australian and New Zealand waters, supported by direct visual observations of feeding off Eden at the time of tagging Given this, the intriguing question arises of whether these whales are foraging north of the putative Antarctic krill biome; and if so, what are WA HBWs likely to be feeding upon? Larger sample sizes tracking and biopsy would be useful to resolve these points. Acoustic listening stations yellow diamonds were located at Prydz Bay and the southern Kerguelen plateau — as part of the Southern Ocean hydrophone network One subadult individual ID did follow the retreating ice, demonstrating movements in and around the ice edge during this time. As increasing high latitude genetic sampling mitochondrial DNA is becoming available, mixed stock analyses are being used to determine the likely population origins of whales across Southern Ocean feeding grounds 11 , 71 , Such an analysis considering stocks C, D and E would be required to describe the population structure on the SKP feeding ground and evaluate the possibility of this being a shared high-quality feeding ground. Southern Hemisphere humpback whale populations are characteristically high in genetic diversity 9 , 10 , thought to be maintained by substantial reproductive sub-structuring across breeding and feeding populations The extent to which mixing between neighbouring populations occurs is poorly known. The combination of new oceanographic information and movement data provides enhanced understanding of important biological processes, which are relevant within the context of the current spatial management and conservation efforts in the Southern Ocean. The spatial distribution of whaling catches and Discovery era mark-recaptures confirms the importance of this region to Western Australian humpback whales since at least historical times. Southern Ocean migration pathways for humpback whales satellite-tracked off the western coast of Australia showing the full extent of southbound tracks. In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript. Southern Ocean migration pathways for satellite-tracked WA humpback whales overlaid with historical whaling catches. Using hidden Markov models, we discriminate sex-related movement behaviours, and examine residency behaviour in relation to current Southern Ocean management areas and a proposed Marine Protected Area. We conclude that the southern Kerguelen plateau region is likely to provide a predictable, persistent and high-quality forage resource 36 , 37 , 40 , and usage of this area by WA HBWs may provide a partial explanation for the ongoing strong recovery of this population. Two whales ID and ID show movement in and around the sea ice in this region during February-March, until these tags failed as the sea-ice began advancing again in mid-late March. The climatological positions of the major ACC fronts 90 are shown in navy. The high productivity around the SKP, controlled by the interaction of topography and oceanography, is likely to provide a predictable, persistent and high-quality forage resource for whales 36 , 37 , Simplified movement patterns of HBWs derived from early Discovery mark recoveries show a clear connectivity between the low-latitude WA breeding grounds and high-latitude feeding grounds in the Indian sector of the Southern Ocean 4 , 5 , In fact blue whale Balaenoptera musculus catches show a very similar spatial distribution in this sector 43 , and regional survey sightings of humpback, blue, sperm Physeter macrocephalus and minke B. The topographic steering of oceanographic features here likely supports a predictable, productive and persistent forage ground. The movement estimates for the resident state were much lower overall females: 2. Further insight has developed from modern photo-identification and molecular methods 2 , 3 , 6 , 7 , 8 and the current management of seven distinct Southern Hemisphere populations A—G , under the International Whaling Commission IWC , is increasingly informed by sophisticated genetic analyses e. Therefore, to properly test this hypothesis of a northward extension in krill — and hence whale — habitat in this region it would be necessary to conduct research voyages with net tows during late spring and early summer November to early December. The bathymetric control of currents and regional productivity might also be more resilient to change. While our findings are consistent with existing general knowledge of HBW movement patterns, the high-resolution biotelemetry reveals specific usage of the western boundary current at the southern Kerguelen plateau. Two additional female East Australian whales IDs tagged in and tagged in 32 which moved westward are shown in green. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser or turn off compatibility mode in Internet Explorer. It is worth noting that neither study WA nor Oceania HBWs tagged whales on the breeding grounds, but in both cases the total migratory pathway includes substantial movement from breeding sites to the tagging locations. Themisto gaudichaudii 66 or possibly small mesopelagic fishes such as Krefftichthys anderssoni The Kerguelen Plateau, extending from a theoretically krill-dominated food-web in the south 68 to a more copepod-fish dominated food-web in the north 69 , 70 potentially offers migrating HBWs diverse and rich food opportunities. Other candidate prey might include amphipods e. {PARAGRAPH}{INSERTKEYS}Thank you for visiting nature. Overall transition probabilities between states were low i.