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Lucy A Hawkes

Bangor University
Brambell Laboratories
Deiniol Road
Bangor
LL57 2UW, UK
l.hawkes@bangor.ac.uk
Lucy Hawkes is a physiological ecologist, whose work focuses on the costs and drivers of migration in vertebrates using techniques such as satellite telemetry, heart rate recording and metabolic rate measurements. Her work has also investigated the impact of external forcing factors, such as climate change, on migration and breeding ecology.

Lucy has an h factor of 7 and a total of 151 cites on first author papers.

Journal articles

InPress
L A Hawkes, J Tomas, O Revuelta, Y M Leon, J M Blumenthal, A C Broderick, M Fish, J A Raga, M J Witt, B J Godley (InPress)  Migratory patterns in hawksbill turtles described by satellite tracking   Marine Ecology Progress Series  
Abstract: The advent of telemetry has improved knowledge of the spatio-temporal distribution of marine species of conservation concern. Among the sea turtles, the movements of the hawksbill turtle (Eretmochelys imbricata) are among the least well described. We tracked 10 adult female hawksbill turtles by satellite after nesting in the Dominican Republic and describe a dichotomy in patterns of movement: (i) some (n=2) turtles remained in the DR, while others migrated to waters off Honduras and Nicaragua (n=5) and the Bahamas (n=1). Two transmitters failed before turtles reached their foraging grounds, although they had left DR waters. We present results from long tracking durations for three turtles, including three entire remigration intervals, highlighting foraging ground and nest-site fidelity. Threats to hawksbill turtles are not well documented for Nicaragua, or neighbouring Honduras, and represent a major information gap. We suggest that directing conservation effort to regionally important foraging areas, such as those in Nicaragua, as well as strengthening national conservation in each nation hosting significant hawksbill nesting offers a clear way forward for the conservation of hawksbill turtles in the region.
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2012
R T Graham, M J Witt, D W Castellanos, F Remolina, S Maxwell, B J Godley, L A Hawkes (2012)  Satellite tracking of manta rays highlights challenges to their conservation   PLoS One 7: 5. e36834  
Abstract: We describe the real-time movements of the last of the marine mega-vertebrate taxa to be satellite tracked – the giant manta ray (or devil fish, Manta birostris), the world's largest ray at over 6 m disc width. Almost nothing is known about manta ray movements and their environmental preferences, making them one of the least understood of the marine mega-vertebrates. Red listed by the International Union for the Conservation of Nature as ‘Vulnerable’ to extinction, manta rays are known to be subject to direct and incidental capture and some populations are declining. Satellite-tracked manta rays associated with seasonal upwelling events and thermal fronts off the Yucatan peninsula, Mexico, and made short-range shuttling movements, foraging along and between them. The majority of locations were received from waters shallower than 50 m deep, representing thermally dynamic and productive waters. Manta rays remained in the Mexican Exclusive Economic Zone for the duration of tracking but only 12% of tracking locations were received from within Marine Protected Areas (MPAs). Our results on the spatio-temporal distribution of these enigmatic rays highlight opportunities and challenges to management efforts.
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F G Moncada, L A Hawkes, M R Fish, B J Godley, S C Manolis, Y Medina, G Nodarse, G J W Webb (2012)  Patterns of dispersal of hawksbill turtles from the Cuban shelf inform scale of conservation and management   Biological Conservation 148: 191-199 April  
Abstract: Conserving and managing populations of marine vertebrates can be complex when they occupy the waters of multiple nations, crossing heterogeneous legal and management landscapes. Hawksbill turtles (Eretmochelys imbricata) are subject to varying levels of use in Caribbean countries and their conservation and management is complicated by the extent to which they are a ‘shared’ resource. In 1997 and 2000, Cuba attempted to ‘downlist’ hawksbills from Cuban waters to CITES Appendix II to allow limited international trade. The research on movement and dispersal of hawksbills reported here was undertaken to better inform discussion about the impacts of their harvest. Flipper tagging and satellite tracking demonstrate that the majority of study turtles remained in Cuban territorial waters. Of 1,170 hawksbills tagged (525 adults and 606 juveniles), 12% (n = 143) were recaptured. All recaptured adults (n = 16 males, 38 nesting females and 30 adult females in-water) were in Cuban waters. Of the 59 juveniles recaptured, only four recaptures were outside Cuban waters (Nicaragua = 2, Colombia = 1, USA = 1). Fourteen hawksbills tagged in the waters of other nations were recaptured in Cuban waters. We also satellite tracked 21 turtles (one adult male, ten nesting females and ten non-nesting adult females), of which five tags failed, 11 stayed in Cuban waters for the duration of transmissions (1 to 809 days) and five foraged in the waters of other nations (Mexico n=1 an adult female; Honduras n=2, both post-nesting turtles; Colombia n=1 an adult female; and the eastern Lesser Antilles n=1, an adult male), with differences for nesting and non-nesting turtles. Our results, demonstrating extended site fidelity within Cuban waters, suggest that strengthening national management within national jurisdictions that host hawksbill turtles is fundamental to improving regional conservation as a whole.
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2011
J Patino-Martinez, A Marco, L Quiñones, L A Hawkes (2011)  A Potential Tool to Mitigate the Impacts of Climate Change to the Caribbean Leatherback Sea Turtle   Global Change Biology 18: 1.  
Abstract: It is now well understood that climate change has the potential to dramatically affect biodiversity, with effects to spatio-temporal distribution patterns, trophic relationships and survivorship. In the marine turtles, sex is determined by incubation temperature, such that warming temperatures could lead to a higher production of female hatchlings. By measuring nest temperature, and using a model to relate the incubation temperature to sex ratio, we estimate that Caribbean Colombian leatherback sea turtles currently produce approximately 92% female hatchlings. We modelled the relationship between incubation, sand and air temperature, and under all future climate change scenarios (0.4 to 6.0 °C warming over the next 100 years) complete feminisation could occur, as soon as the next decade. However, male producing refugia exist in the periphery of smaller nests (0.7°C cooler at the bottom than at the centre), within beaches (0.3°C cooler in the vegetation line and inter-tidal zone) and between beaches (0.4°C higher on dark beaches) and these natural refugia could be assigned preferential conservation status. However, there exists a need to develop strategies that may ameliorate deleterious effects of climate-induced temperature changes in the future. We experimentally shaded clutches using screening material, and found it was effective in reducing nest temperature, producing a higher proportion of male hatchlings, without compromising the fitness or hatching success. Artificial shade in hatcheries is a very useful and simple tool in years or periods of high environmental temperatures. Nevertheless, this is only an emergency response to the severe impacts that will eventually have to be reversed if we are to guarantee the stability of the populations.
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L A Hawkes, S Balachandran, N Batbayar, P J Butler, P B Frappell, W K Milsom, N Tseveenmyadag, S H Newman, G R Scott, P Sathiyaselvam, J Y Takekawa, M Wikelski, C M Bishop (2011)  The trans-Himalayan flights of bar-headed geese (Anser indicus).   Proc Natl Acad Sci U S A 108: 23. 9516-9519 Jun  
Abstract: Birds that fly over mountain barriers must be capable of meeting the increased energetic cost of climbing in low-density air, even though less oxygen may be available to support their metabolism. This challenge is magnified by the reduction in maximum sustained climbing rates in large birds. Bar-headed geese (Anser indicus) make one of the highest and most iconic transmountain migrations in the world. We show that those populations of geese that winter at sea level in India are capable of passing over the Himalayas in 1 d, typically climbing between 4,000 and 6,000 m in 7-8 h. Surprisingly, these birds do not rely on the assistance of upslope tailwinds that usually occur during the day and can support minimum climb rates of 0.8-2.2 km·h(-1), even in the relative stillness of the night. They appear to strategically avoid higher speed winds during the afternoon, thus maximizing safety and control during flight. It would seem, therefore, that bar-headed geese are capable of sustained climbing flight over the passes of the Himalaya under their own aerobic power.
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L A Hawkes, M J Witt, A C Broderick, J W Coker, M S Coyne, M Dodd, M G Frick, M H Godfrey, D B Griffin, S R Murphy, T M Murphy, K L Williams, B J Godley (2011)  Home on the range: spatial ecology of loggerhead turtles in Atlantic waters of the USA   Diversity and Distributions 17: 4. 624-640 July  
Abstract: Aim  Although satellite tracking has yielded much information regarding the migrations and habitat use of threatened marine species, relatively little has been published about the environmental niche for loggerhead sea turtles Caretta caretta in north-west Atlantic waters. Location  North Carolina, South Carolina and Georgia, USA. Methods  We tracked 68 adult female turtles between 1998 and 2008, one of the largest sample sizes to date, for 372.2 ± 210.4 days (mean ± SD). Results  We identified two strategies: (1) ‘seasonal’ migrations between summer and winter coastal areas (n = 47), although some turtles made oceanic excursions (n = 4) and (2) occupation of more southerly ‘year-round’ ranges (n = 18). Seasonal turtles occupied summer home ranges of 645.1 km2 (median, n = 42; using α-hulls) predominantly north of 35 ° latitude and winter home ranges of 339.0 km2 (n = 24) in a relatively small area on the narrow shelf off North Carolina. We tracked some of these turtles through successive summer (n = 8) and winter (n = 3) seasons, showing inter-annual home range repeatability to within 14.5 km of summer areas and 10.3 km of winter areas. For year-round turtles, home ranges were 1889.9 km2. Turtles should be tracked for at least 80 days to reliably estimate the home range size in seasonal habitats. The equivalent minimum duration for ‘year-round’ turtles is more complex to derive. We define an environmental envelope of the distribution of North American loggerhead turtles: warm waters (between 18.2 and 29.2 °C) on the coastal shelf (in depths of 3.0–89.0 m). Main conclusions  Our findings show that adult female loggerhead turtles show predictable, repeatable home range behaviour and do not generally leave waters of the USA, nor the continental shelf (< 200m depth). These data offer insights for future marine management, particularly if they were combined with those from the other management units in the USA.
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2010
J Y Takekawa, S R Heath, D C Douglas, W M Perry, S Javed, S H Newman, R N Suwal, A R Rahmani, B C Choudhury, D J Prosser, B Yan, Y Hou, N Batbayar, T Natsagdorj, C M Bishop, P J Butler, P B Frappell, W K Milsom, G R Scott, L A Hawkes, M Wikelski (2010)  Geographic variation in Bar-headed Geese Anser indicus: connectivity of wintering areas and breeding grounds across a broad front   Wildfowl 59: 100-123  
Abstract: The connectivity and frequency of exchange between sub-populations of migratory birds is integral to understanding population dynamics over the entire species’ range. True geese are highly philopatric and acquire lifetime mates during the winter, suggesting that the number of distinct sub-populations may be related to the number of distinct wintering areas. In the Bar-headed Goose Anser indicus, a species found exclusively in Central Asia, the connectivity between breeding and wintering areas is not well known. Their migration includes crossing a broad front of the Himalaya Cordillera, a significant barrier to migration for most birds. Many Bar-headed Geese fly to breeding areas on the Tibetan-Qinghai Plateau (TQP), the highest plateau in the world. From 2005–2008, 60 Bar-headed Geese were captured and marked with satellite transmitters in Nepal (n = 2), India (n = 6), China (n = 29), and Mongolia (n = 23) to examine their migration and distribution. Distinct differences were observed in their migration corridors and timing of movements, including an apparent leap-frog migration pattern for geese from Mongolia. Measurements of geese from Mongolia were larger than their counterparts from China, providing some evidence of morphological differences. Alteration of habitats in China, including the warming effects of climate change on glaciers increasing runoff to TQP wetlands, may be changing goose migration patterns and timing. With the exception of one individual, all geese from Qinghai Lake, China wintered in the southern TQP near Lhasa, and their increasing numbers in that region may be related to the effects of climate change and agricultural development. Thus, our findings document both morphological and geographical variation in sub-populations of Bar-headed Geese, but their resilience to environmental change may be lost if migratory short-stopping results in larger congregations restricted to a smaller number of wintering areas.
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F Moncada, F A Abreu-Grobois, D Bagley, K A Bjorndal, A B Bolten, J A Caminas, L Ehrhart, A Muhlia-Melo, G Nodarse, B A Schroeder, J Zurita, L A Hawkes (2010)  Movement patterns of loggerhead turtles Caretta caretta in Cuban waters inferred from flipper tag recaptures   Endangered Species Research 11: 61-68 March  
Abstract: Understanding the spatial movements of threatened marine species, such as sea turtles, is essential as a means of informing appropriate conservation management. Although novel techniques for tracking spatial movements are becoming more widely available (such as satellite tracking), simple techniques such as mark-release-recapture remain effective. A flipper tagging and recovery program in Cuba tagged 210 loggerhead turtles over 14 yr and recovered 7% of the tags between 2 d and 3 yr later (mean = 296 d). All but one turtle was recaptured in Cuban waters, and data showed limited movement of turtles between northern and southern coasts. A further 50 turtles were recovered that had been tagged in foreign projects, the majority of which were from the USA (but also Mexico, The Bahamas, Canary Islands and Spain). A range of life stages of loggerhead turtles are found in Cuban waters year-round, and given that Cuba has the second largest reef in the Caribbean, it likely provides foraging habitat for significant numbers of loggerhead turtles from at least 6 different countries.
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L Bourouiba, Jianhong Wu, S Newman, J Takekawa, T Natdorj, N Batbayar, C M Bishop, L A Hawkes, P J Butler, M Wikelski (2010)  Spatial dynamics of bar-headed geese migration in the context of H5N1.   J R Soc Interface 7: 52. 1627-1639 Nov  
Abstract: Virulent outbreaks of highly pathogenic avian influenza (HPAI) since 2005 have raised the question about the roles of migratory and wild birds in the transmission of HPAI. Despite increased monitoring, the role of wild waterfowl as the primary source of the highly pathogenic H5N1 has not been clearly established. The impact of outbreaks of HPAI among species of wild birds which are already endangered can nevertheless have devastating consequences for the local and non-local ecology where migratory species are established. Understanding the entangled dynamics of migration and the disease dynamics will be key to prevention and control measures for humans, migratory birds and poultry. Here, we present a spatial dynamic model of seasonal migration derived from first principles and linking the local dynamics during migratory stopovers to the larger scale migratory routes. We discuss the effect of repeated epizootic at specific migratory stopovers for bar-headed geese (Anser indicus). We find that repeated deadly outbreaks of H5N1 on stopovers during the autumn migration of bar-headed geese could lead to a larger reduction in the size of the equilibrium bird population compared with that obtained after repeated outbreaks during the spring migration. However, the opposite is true during the first few years of transition to such an equilibrium. The age-maturation process of juvenile birds which are more susceptible to H5N1 reinforces this result.
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M J Witt, L A Hawkes, M H Godfrey, B J Godley, A C Broderick (2010)  Predicting the impacts of climate change on a globally distributed species: the case of the loggerhead turtle.   J Exp Biol 213: 6. 901-911 Mar  
Abstract: Marine turtles utilise terrestrial and marine habitats and several aspects of their life history are tied to environmental features that are altering due to rapid climate change. We overview the likely impacts of climate change on the biology of these species, which are likely centred upon the thermal ecology of this taxonomic group. Then, focusing in detail on three decades of research on the loggerhead turtle (Caretta caretta L.), we describe how much progress has been made to date and how future experimental and ecological focus should be directed. Key questions include: what are the current hatchling sex ratios from which to measure future climate-induced changes? What are wild adult sex ratios and how many males are necessary to maintain a fertile and productive population? How will climate change affect turtles in terms of their distribution?
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2009
L A Hawkes, A C Broderick, M H Godfrey, B J Godley (2009)  Climate change and marine turtles   Endangered Species Research 7: 154 May  
Abstract: Marine turtles occupy a wide range of terrestrial and marine habitats, and many aspects of their life history have been demonstrated to be closely tied to climatic variables such as ambient temperature and storminess. As a group, therefore, marine turtles may be good indicators of climate change effects on coastal and marine habitats. Despite the small number of species in the taxon and a growing body of research in the field, the evidence base to predict resultant impacts of climate change remains relatively poor. We review the data from peer-reviewed publications to assess the likely impacts of climate change on marine turtles and highlight the types of data that would be most useful for an accurate assessment of future effects. The cumulative indications from these previous studies indicate that future research should focus on: (1) climate change effects on key habitats upon which turtles depend; (2) factors that influence nest site selection; (3) the consequences of skewed primary sex ratios; and (4) the effect of climate change on turtles at sea, for example range shifts and dietary breadth. Although it is too early to give detailed management recommendations, careful protection of coastlines along which turtles nest should be considered, as should the protection of beaches that produce male hatchlings, which may be of increased importance in the future. More active management approaches, for example translocation of eggs to suitable yet vacant nesting beaches, may be necessary to consider under worst-case scenarios.
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2008
B J Godley, J M Blumenthal, A C Broderick, M S Coyne, M H Godfrey, L A Hawkes, M J Witt (2008)  Satellite tracking of sea turtles: Where have we been and where do we go next?   Endangered Species Research 10: 3-22  
Abstract: The use of satellite tracking for the fundamental and applied study of marine turtles began in the 1980s but has undergone rapid growth in recent years. To provide a background against which to judge the past success and future directions of these research efforts we carried out a comprehensive review of over 130 scientific papers on the use of this technique in this taxon. We show how satellite tracking has changed over time as well as outlining biases in spatial, species and lifestage coverage. Descriptions of migration routes and other habitats have offered novel insights into the basic life history patterns of some species, highlighted focal areas for conservation and reinforced the multi-national nature of the stakeholders of many populations. In foraging areas, knowledge is growing as to how animals move within dynamic seascapes, thus facilitating our understanding of 3-dimensional habitat use and seasonal patterns of behaviour. More experimental approaches have elucidated navigational capabilities and post-release survival following fisheries interaction and long-term captivity. In addition, through the Internet and other media, satellite tracking appears to have been effective in engaging public attention in many countries. Finally, we discuss why the use of the technique has increased so markedly over time and point out key areas of concern that we feel should be addressed by the community of researchers and donors who focus on sea turtles.
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2007
L A Hawkes, A C Broderick, M H Godfrey, B J Godley (2007)  Investigating the potential impacts of climate change on a marine turtle population   Global Change Biology 13: 1-10  
Abstract: Recent increases in global temperatures have affected the phenology and survival of many species of plants and animals. We investigated a case study of the effects of potential climate change on a thermally sensitive species, the loggerhead sea turtle, at a breeding location at the northerly extent of the range of regular nesting in the United States. In addition to the physical limits imposed by temperature on this ectothermic species, sea turtle primary sex ratio is determined by the temperature experienced by eggs during the middle third of incubation. We recorded sand temperatures and used historical air temperatures (ATs) at Bald Head Island, NC, to examine past and predict future sex ratios under scenarios of warming. There were no significant temporal trends in primary sex ratio evident in recent years and estimated mean annual sex ratio was 58% female. Similarly, there were no temporal trends in phenology but earlier nesting and longer nesting seasons were correlated with warmer sea surface temperature. We modelled the effects of incremental increases in mean AT of up to 7.5 1C, the maximum predicted increase under modelled scenarios, which would lead to 100% female hatchling production and lethally high incubation temperatures, causing reduction in hatchling production. Populations of turtles in more southern parts of the United States are currently highly female biased and are likely to become ultra-biased with as little as 1 1C of warming and experience extreme levels of mortality if warming exceeds 31C. The lack of a demonstrable increase in AT in North Carolina in recent decades coupled with primary sex ratios that are not highly biased means that the male offspring from North Carolina could play an increasingly important role in the future viability of the loggerhead turtle in the Western Atlantic.
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L A Hawkes, A C Broderick, M S Coyne, M H Godfrey, B J Godley (2007)  Only some like it hot — quantifying the environmental niche of the loggerhead sea turtle   Diversity and Distributions 13: 447-457  
Abstract: Although the Atlantic waters of North America support hundreds of thousands of loggerhead sea turtles (Caretta caretta), remarkably little is known regarding their migratory ecology and habitat use. We integrate satellite tracking with remotely sensed oceanographic data to uncover two different migratory strategies used by loggerhead turtles at the northern part of their range. Most turtles travelled from the nesting beach to forage at higher latitudes in summer, before migrating south to wintering grounds in the autumn. Others moved south after nesting to forage for up to 514 days and did not make an autumn migration. Both groups utilized warm waters at the very edge of the Gulf Stream during winter: for southerly turtles obviating seasonal migration, and for northerly turtles minimizing the distance, time and energy required to reach northern areas for subsequent foraging seasons, avoiding lethally cold winter temperatures in inshore waters at the same latitude, and reducing energy costs that would be incurred within the fast-flowing Gulf Stream. Females made long resting dives of up to 7 h 24 min, effectively hibernating during the colder months. Offshore federal waters of the USA constitute a more important habitat for both foraging and wintering turtles than previously appreciated. These areas are potential hotspots for interaction with fisheries and proposed US military training activities and should receive special monitoring efforts to fully assess the extent of overlap.
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2006
L A Hawkes, A C Broderick, M S Coyne, M H Godfrey, L F Lopez-Jurado, P Lopez-Suarez, S E Merino, N Varo-Cruz, B J Godley (2006)  Phenotypically linked dichotomy in sea turtle foraging requires multiple conservation approaches.   Curr Biol 16: 10. 990-995 May  
Abstract: Marine turtles undergo dramatic ontogenic changes in body size and behavior, with the loggerhead sea turtle, Caretta caretta, typically switching from an initial oceanic juvenile stage to one in the neritic, where maturation is reached and breeding migrations are subsequently undertaken every 2-3 years. Using satellite tracking, we investigated the migratory movements of adult females from one of the world's largest nesting aggregations at Cape Verde, West Africa. In direct contrast with the accepted life-history model for this species, results reveal two distinct adult foraging strategies that appear to be linked to body size. The larger turtles (n = 3) foraged in coastal waters, whereas smaller individuals (n = 7) foraged oceanically. The conservation implications of these findings are profound, with the population compartmentalized into habitats that may be differentially impacted by fishery threats in what is a global fishing hotspot. Although the protection of discrete areas containing coastal individuals may be attainable, the more numerous pelagic individuals are widely dispersed with individuals roaming over more than half a million square kilometers. Therefore, mitigation of fisheries by-catch for sea turtles in the east Atlantic will likely require complex and regionally tailored actions to account for this dichotomous behavior.
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2005
L A Hawkes, A C Broderick, M H Godfrey, B J Godley (2005)  Status of nesting loggerhead turtles Caretta caretta at Bald Head Island (North Carolina, USA) after 24 years of intensive monitoring and conservation   Oryx 39: 1. 65-72 January  
Abstract: A 24-year set of data from monitoring of a nesting beach at Bald Head Island, North Carolina, USA, was analysed in parallel with limited data from nearby rookeries to investigate trends in loggerhead turtle Caretta caretta nesting numbers. There was no statistical evidence of an increasing or decreasing trend in numbers of clutches laid per year, although a significant decrease in the number of turtles nesting and number of clutches laid per year was found from 1991. Remigrating turtles were larger and had larger annual clutch frequencies than neophyte turtles. Annual levels of nesting at beaches within the Cape Fear area were significantly correlated. The stable trend in number of clutches laid across more than two decades is discussed in relation to other factors affecting marine turtles in North American waters.
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Technical reports

2011
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