Juvenile Southern Bluefin Tuna Research Articles

Migration dynamics of juvenile southern bluefin tuna

Large scale migrations are a key component of the life history of many marine species. We quantified the annual migration cycle of juvenile southern bluefin tuna (Thunnus maccoyii; SBT) and spatiotemporal variability in this cycle, based on a multi-decadal electronic tagging dataset. Behaviour-switching models allowed for the identification of cohesive areas of residency and classified the temporal sequence of movements within a migration cycle from austral summer foraging grounds in the Great Australian Bight (GAB) to winter foraging grounds in the Indian Ocean and Tasman Sea and back to the GAB. Although specific regions within the Indian Ocean were frequented, individuals did not always return to the same area in consecutive years. Outward migrations from the GAB were typically longer than return migrations back to the GAB. The timing of individual arrivals to the GAB, which may be driven by seasonality in prey availability, was more cohesive than the timing of departures from the GAB, which may be subject to the physiological condition of SBT. A valuable fishery for SBT operates in the GAB, as do a number of scientific research programs designed to monitor SBT for management purposes; thus, understanding SBT migration to and from the area is of high importance to a number of stakeholders.

Southern bluefin tuna habitat use and residence patterns in the Great Australia Bight

Juvenile southern bluefin tuna (SBT) undertake seasonal migrations to and from the coastal waters of southern Australia until around age 5. In particular, they are present in the waters of the Great Australia Bight (GAB) over the summer months, where they are targeted by a commercial purse seine fishery. Arrival to and departure from the GAB represent important constraints on the valuable fishery, and this information is also now seen as essential for managing other activities within the GAB. As such, understanding the general patterns of the distribution of juvenile SBT and the environmental factors might be driving their distribution is valuable for establishing a baseline against which any future changes to population arrival or departure may be assessed. As part of a large investigation into the ecosystem of the GAB, data from a multi-year archival tagging program focused on juvenile SBT were used to construct statistical models of spatial usage within the residence areas of the GAB. Climatological data describing mixed-layer-depth, sea-surface temperatures, along with distance to the shelf break and coast, spatial position and time of year were used as predictor variables. Generalized additive models were fit which accounted for the seasonal progression of SBT into the GAB throughout their annual cycle and predicted the broad scale environmental features associated with their average distribution. The statistical models predicted habitat use by juvenile SBT in the proximity of the continental shelf break, with a preference for warmer waters in conjunction with deeper mixed layers. The model-generated annual distribution climatology can be used to plan the timing for a range of activities such as fishing, shipping, oil and gas exploration, and also inform development of forecasting applications. The predictions are also useful for determining large scale variation from the typical usage patterns of SBT in the GAB into the future.

A summary of oil and gas exploration in the Great Australian Bight with particular reference to southern bluefin tuna

Large numbers of juvenile southern bluefin tuna (SBT; Thunnus maccoyii) migrate into the warm shelf waters of the Great Australian Bight (GAB) each austral summer. Whilst in the GAB, they are the focus of a commercially important Australian fishery. Recent expansion of activities associated with oil and gas exploration in the GAB over the last five years has raised concerns that noise associated with geophysical surveys and exploratory drilling activities may impact on the migration and behaviour of SBT, with potential flow-on effects on the commercial fishery and current monitoring of the population for management purposes.As a first step in identifying the potential impacts of expansion of oil and gas exploration activities in the GAB on SBT, we establish the historical extent of these activities in the GAB with a primary focus on geophysical surveys. Potential overlap with the distribution of juvenile SBT in the GAB determined from an archival tag dataset (1998–2011) was then explored. Geophysical surveys have occurred throughout the vast majority of the region since 1960, with the size of energy sources used in surveys and the extent of areas surveyed increasing through time. The use of more complex, higher density 3D surveys has increased gradually since 2000. The timing and distribution of exploration activity has had varying degrees of overlap with SBT occurrence and distribution as described by tagged fish.Determining robust measures of the responses to surveys however, is difficult due to a paucity of data on the hearing capabilities of SBT, uncertainties in the true extent of spatial overlap of activities and tagged fish resulting from geolocation methods used for position estimation from archival tag data, complexities in the drivers of behaviour in SBT, and limitations with using observational data to determine cause-and-effect relationships. These difficulties are typical not just of SBT and the GAB region, but apply to investigations of the responses of marine animals to geophysical surveys more broadly. The data presented in this study, however, represent a first step towards understanding oil and gas exploration within a defined region of the Australian marine environment, and provide important context for exploring the potential impact of such activities on a commercially important top order predator of the GAB ecosystem.

Modelling surfacing behaviour of southern bluefin tuna in the Great Australian Bight

Large numbers of juvenile southern bluefin tuna (SBT; Thunnus maccoyii) migrate into the warm shelf waters of the Great Australian Bight (GAB) each austral summer. Whilst in the GAB, they aggregate in schools that spend substantial periods in the surface layer of the water column. In this study we investigate biological, temporal and environmental factors influencing this surfacing phenomena using an extensive archival tagging dataset collected between 1998 and 2011. High frequency data on the vertical movement of SBT collected by these tags were used to calculate the proportion of time fish spent in the shallowest 20 m during each day and night period. Estimates of fish location derived from light sensor data on the tags allowed us to investigate the influence that local environmental conditions had on a fish's surfacing behaviour. Although there is high variability in surfacing behaviour within and between individuals, some general patterns emerge. There are clear diel differences in surfacing, with the proportion of time fish spend at the surface tending to be high during most days and either very high (>90%) or very low (<10%) during most nights. Complex relationships were found between surfacing behaviour and the environmental variables considered (sea surface temperature, chlorophyll, salinity and wind speed). The results from this study have relevance to the commercial purse-seine fishery targeting surface schools of SBT in the GAB during the austral summer, as well as the scientific aerial survey conducted each summer that collects data on sightings of surface schools of SBT in order to derive a relative abundance index used directly in management.

Evidence of separate subgroups of juvenile southern bluefin tuna.

Archival tagging studies of southern bluefin tuna (SBT , Thunnus maccoyii) have revealed that juveniles residing in the Great Australian Bight (GAB) over the austral summer undertake seasonal cyclic migrations to the southeast Indian Ocean and the Tasman Sea during winter. However, there remains disagreement about the extent of mixing between juvenile SBT regularly caught by longline fleets south of Africa and those observed in the GAB. Some researchers have argued that archival tag recoveries indicate most juveniles reside in the GAB over the austral summer. Others have suggested that recoveries of conventional and archival tags are better explained by a juvenile population consisting of separate groups on the eastern and western sides of the Indian Ocean with limited intermixing. We present analyses of catch and tag recovery data and re‐examine archival tagging studies. The evidence provided strongly favors the hypothesis of separate juvenile subgroups, or contingents, with limited intermixing. We draw some tentative conclusions about the nature of the putative contingents and discuss some implications of these findings for the interpretation of existing datasets and future research priorities. We also provide the first evidence that the migration choices of juveniles that summer in the GAB are influenced by fidelity to winter feeding grounds and suggest this helps explain the collapse of the surface fishery off New South Wales in the 1980s.

A standardised abundance index from commercial spotting data of southern bluefin tuna (Thunnus maccoyii): random effects to the rescue.

Commercial aerial spotting of surface schools of juvenile southern bluefin tuna (SBT), Thunnus maccoyii, is conducted as part of fishing operations in the Great Australian Bight in summer. This provides the opportunity to efficiently collect large amounts of data on sightings of SBT. The data can potentially be used to construct a time-series index of relative abundance by standardising the data for issues such as weather, spotter ability and ocean conditions. Unlike a statistically designed survey, the commercial spotting is governed by business considerations and fishing operations. The SBT dataset is therefore highly unbalanced with regard to spotters operating in each season. This complicates the standardisation of the data, particularly with regard to interactions between covariates. We show how a generalized additive model with random effects can simplify both the fitting of the model and the construction of an index, while also avoiding the need to leave out strata or interaction terms that are important. The approach is applicable to standardisation of more traditional catch and effort data.

The complete mitochondrial genome of the Southern bluefin tuna Thunnus maccoyii

The complete mitochondrial genome of the juvenile Southern bluefin tuna Thunnus maccoyii collected from Sea of Japan was determined by next-generation sequencing. The mitogenome is a circular molecule 16,529 bp in length, including the typical structure of 13 protein-coding genes, 2 ribosomal RNA genes, 22 transfer RNA genes and a control region. The TAS, central CSB and CSB were detected in the control region. The gene contents of the mitogenome are identical to those observed in most bony fishes.

Using electronic tag data to improve mortality and movement estimates in a tag-based spatial fisheries assessment model

Despite increased deployment of archival tags on exploited fish species, analytical methods for including archival tag data in fishery assessment models are lacking. We present a method for integrating archival tag data into a spatial tag–recapture model for estimating natural mortality, fishing mortality, abundance, and movement. Archival tags provide important information on fish movement not available from conventional tags that facilitates separation of movement from mortality estimates. Using simulations, we evaluate the benefit of including archival tag data in the model using two model formulations: one with a general spatial structure and one with movement and fishery dynamics based on juvenile southern bluefin tuna (SBT; Thunnus maccoyii ). If fish are not tagged in all regions and time periods, then including archival tag data can substantially improve the precision of the fishing mortality and movement estimates. For example, with the general spatial structure and specific scenario presented, standard errors of the fishing mortality estimates decreased by an average of 34% (21%) when 25 archival tags were released in addition to 500 (2500) conventional tags in each region and period of tagging, respectively. Furthermore, with the SBT spatial structure, archival tag data were necessary for all parameters to be estimable.

Departure behaviour of juvenile southern bluefin tuna (Thunnus maccoyii) from southern Western Australia temperate waters in relation to the Leeuwin Current

The arrival of the warm tropical Leeuwin Current (LC) into southern Western Australia (SWA) may influence the movement timing and foraging habitat of juvenile southern bluefin tuna (SBT). Seasonal and interannual changes in the strength of the LC lead to thermal differences and potential changes in food availability between tropical and temperate waters in SWA. This phenomenon could influence the habitat utilization of SBT in these summer grounds. Movement characteristics determined from SBT tagged with acoustic transmitters (N = 244) using cross-shelf lines of automated receivers for three summer-autumn seasons (2004/2005, 2005/2006, 2006/2007) in SWA revealed interannual variability. Each year, the eastward movements of tagged fish increased as temperatures increased, and fish left the region when temperatures exceeded 20.0°C, a temperature indicative of the leading edge of the LC in SWA waters. Interannual fluctuations in the timing of movements were detected. When the LC was narrow and restricted to the shelf edge in 2004/2005, the distribution of SBT in shelf waters did not change before or after LC intrusion. In contrast, long distance eastward movements frequently occurred when the LC intrusion was spread wide over the continental shelf in 2005/2006 and 2006/2007. This suggests that, off SWA, juvenile SBT move quickly out of local foraging habitats defined by cool sub-tropical temperate waters ahead of the tropical LC intrusion, despite these waters not being physiologically limiting. These results suggest that the behavioural response of SBT may be related to changes in prey availability as a result of changes in oceanographic conditions.

Diet of young southern bluefin tuna Thunnus maccoyii in the southwestern coastal waters of Australia in summer

The diet of juvenile (predominantly age 1) southern bluefin tuna Thunnus maccoyii (SBT, N = 720), caught over 11 years of the recruitment monitoring survey off southern Western Australia during summer, consisted overwhelmingly of teleosts (97.4% by volume). Pilchard Sardinops sagax (27.4%V), blue mackerel Scomber australasicus (16.7%V), and jack mackerel Trachurus declivis (14.2%V) were the major taxa, with pilchard more abundant in coastal waters and jack mackerel more frequently encountered in fish caught closer to the shelf-edge. Prey size varied from 5 to 240 mm, with 67% of ingested items measuring between 30 and 50 mm. Pilchard dominated the prey size category 130–190 mm (84% by number), but the overall contribution of this species to the diet of juvenile SBT was much lower than previously reported. Future research in relation to the feeding ecology of juvenile SBT should focus on the biology and ecology of the young lifestages of the main prey species in this area and on prey distribution and dynamics as a key factor linking environmental change and SBT distribution.

Interannual variation in summer habitat utilization by juvenile southern bluefin tuna (Thunnus maccoyii) in southern Western Australia

AbstractThe spatial habitat utilization of juvenile southern bluefin tuna in southern Western Australia was investigated using automated acoustic receivers with acoustic transmitters implanted in tagged fish during three austral summers (2004/2005, N = 79 fish, 2005/2006, N = 81, 2006/2007, N = 84). Seventy acoustic receivers were deployed at three cross‐shelf lines and three coastal topographic features (lumps) between December and May. We observed markedly different patterns of habitat utilization between the three seasons: (i) aggregation at lumps in 2004/2005 and 2006/2007, and (ii) wide distribution over the continental shelf (i.e., few occurring at lumps) in 2005/2006. Vertical profile by conductivity‐temperature‐depth casts showed these spatial shifts were caused by interannual changes in the presence of sub‐Antarctic water. The sub‐Antarctic water was present in the subsurface layer close to the continental slope only during 2005/2006, and the area had higher chlorophyll‐a concentrations than the coastal areas, including at the lumps. These environmental characters, related to the nutrient rich sub‐Antarctic water, appear to have a strong influence on fish distributions in 2005/2006, and may occur generally during La Niña events. Interannual fluctuations in habitat utilization will influence detection of fish in recruitment monitoring surveys and thus bias the resulting juvenile abundance indices.

Spatial and temporal variation in the distribution of juvenile southern bluefin tuna Thunnus maccoyii: implication for precise estimation of recruitment abundance indices

Acoustic tags were used to examine the spatial and temporal distribution of southern bluefin tuna (SBT) in southern Western Australia, which is in a region where fishery-independent acoustic surveys of the recruitment abundance index of SBT have been historically undertaken. We investigated patterns of SBT distribution within and inshore of the acoustic survey area during three summer seasons. Annual differences in distribution patterns were characterized by two distinctive migration pathways. An inshore-migrating pathway was observed in two seasons (2004/2005 and 2006/2007), with a relatively high proportion of tagged SBT (84.5, 65.0%) migrating inshore of the acoustic survey area. The other pathway was concentrated along the shelf (2005/2006 season), with an estimated 63.3% of tagged SBT moving within the survey area. These variable migration patterns may bias the interannual fluctuations in abundance indices. Current survey methods can be modified to include both inshore and continental shelf areas. This contribution shows that the accuracy of acoustic surveys can be improved by including ecological patterns.

Age‐dependent distribution of juvenile southern bluefin tuna (Thunnus maccoyii) on the continental shelf off southwest Australia determined by acoustic monitoring

AbstractJuvenile southern bluefin tuna (Thunnus maccoyii, SBT) were monitored in nearshore waters off southwest Australia using acoustic tagging and monitoring over five austral summers (2002/2003–2006/2007) to determine patterns in age‐based distribution of SBT. A total of 20–70 receivers were deployed in early December along one to three cross‐shelf transects and at three inshore topographic features (lumps) where SBT are known to occur; and a total of 59–84 juvenile SBT (41–90 cm fork length) were tagged and released each year. After several months, receivers were recovered, and data extracted. In 2002/2003 and 2003/2004, 2‐yr‐old SBT were detected more frequently in nearshore areas than did 1‐yr old fish. Similarly in 2004/2005 and 2006/2007, it was the larger 1‐yr‐old SBT that were detected more frequently at inshore lumps (&gt;85% of detections) while small 1‐yr‐old SBT were detected more widely over the study area. In 2005/2006, although large 1‐yr‐old SBT were still detected more frequently at inshore lumps than small 1‐yr old SBT, the percentage of all detections at lumps was lower at 31.5%, indicating wider distribution of both small and large age‐1 fish. The observed age‐dependent distribution pattern may be enhanced by the distribution of prey, which disperse or concentrate depending on the local oceanography.

Plasticity in vertical behaviour of migrating juvenile southern bluefin tuna (Thunnus maccoyii) in relation to oceanography of the south Indian Ocean

AbstractElectronic tagging provides unprecedented information on the habitat use and behaviour of highly migratory marine predators, but few analyses have developed quantitative links between animal behaviour and their oceanographic context. In this paper we use archival tag data from juvenile southern bluefin tuna (Thunnus maccoyii, SBT) to (i) develop a novel approach characterising the oceanographic habitats used throughout an annual migration cycle on the basis of water column structure (i.e., temperature‐at‐depth data from tags), and (ii) model how the vertical behaviour of SBT altered in relation to habitat type and other factors. Using this approach, we identified eight habitat types occupied by juvenile SBT between the southern margin of the subtropical gyre and the northern edge of the Subantarctic Front in the south Indian Ocean. Although a high degree of variability was evident both within and between fish, mixed‐effect models identified consistent behavioural responses to habitat, lunar phase, migration status and diel period. Our results indicate SBT do not act to maintain preferred depth or temperature ranges, but rather show highly plastic behaviours in response to changes in their environment. This plasticity is discussed in terms of the potential proximate causes (physiological, ecological) and with reference to the challenges posed for habitat‐based standardisation of fishery data used in stock assessments.

Feeding ecology of wild migratory tunas revealed by archival tag records of visceral warming

1. Seasonal long-distance migrations are often expected to be related to resource distribution, and foraging theory predicts that animals should spend more time in areas with relatively richer resources. Yet for highly migratory marine species, data on feeding success are difficult to obtain. We analysed the temporal feeding patterns of wild juvenile southern bluefin tuna from visceral warming patterns recorded by archival tags implanted within the body cavity. 2. Data collected during 1998-2000 totalled 6221 days, with individual time series (n = 19) varying from 141 to 496 days. These data span an annual migration circuit including a coastal summer residency within Australian waters and subsequent migration into the temperate south Indian Ocean. 3. Individual fish recommenced feeding between 5 and 38 days after tagging, and feeding events (n = 5194) were subsequently identified on 76.3 +/- 5.8% of days giving a mean estimated daily intake of 0.75 +/- 0.05 kg. 4. The number of feeding events varied significantly with time of day with the greatest number occurring around dawn (58.2 +/- 8.0%). Night feeding, although rare (5.7 +/- 1.3%), was linked to the full moon quarter. Southern bluefin tuna foraged in ambient water temperatures ranging from 4.9 degrees C to 22.9 degrees C and depths ranging from the surface to 672 m, with different targeting strategies evident between seasons. 5. No clear relationship was found between feeding success and time spent within an area. This was primarily due to high individual variability, with both positive and negative relationships observed at all spatial scales examined (grid ranges of 2 x 2 degrees to 10 x 10 degrees ). Assuming feeding success is proportional to forage density, our data do not support the hypothesis that these predators concentrate their activity in areas of higher resource availability. 6. Multiple-day fasting periods were recorded by most individuals. The majority of these (87.8%) occurred during periods of apparent residency within warmer waters (sea surface temperature > 15 degrees C) at the northern edge of the observed migratory range. These previously undocumented nonfeeding periods may indicate alternative motivations for residency. 7. Our results demonstrate the importance of obtaining information on feeding when interpreting habitat utilization from individual animal tracks.

Moving with the beat: heart rate and visceral temperature of free-swimming and feeding bluefin tuna

Owing to the inherent difficulties of studying bluefin tuna, nothing is known of the cardiovascular function of free-swimming fish. Here, we surgically implanted newly designed data loggers into the visceral cavity of juvenile southern bluefin tuna (Thunnus maccoyii) to measure changes in the heart rate (fH) and visceral temperature (TV) during a two-week feeding regime in sea pens at Port Lincoln, Australia. Fish ranged in body mass from 10 to 21 kg, and water temperature remained at 18-19 degrees C. Pre-feeding fH typically ranged from 20 to 50 beats min(-1). Each feeding bout (meal sizes 2-7% of tuna body mass) was characterized by increased levels of activity and fH (up to 130 beats min(-1)), and a decrease in TV from approximately 20 to 18 degrees C as cold sardines were consumed. The feeding bout was promptly followed by a rapid increase in TV, which signified the beginning of the heat increment of feeding (HIF). The time interval between meal consumption and the completion of HIF ranged from 10 to 24 hours and was strongly correlated with ration size. Although fH generally decreased after its peak during the feeding bout, it remained elevated during the digestive period and returned to routine levels on a similar, but slightly earlier, temporal scale to TV. These data imply a large contribution of fH to the increase in circulatory oxygen transport that is required for digestion. Furthermore, these data oppose the contention that maximum fH is exceptional in bluefin tuna compared with other fishes, and so it is likely that enhanced cardiac stroke volume and blood oxygen carrying capacity are the principal factors allowing superior rates of circulatory oxygen transport in tuna.

School-based indicators of tuna population status

Abstract Dell, J. T. and Hobday, A. J. 2008. School-based indicators of tuna population status. – ICES Journal of Marine Science, 65: 612–622. Theory and limited observation suggest that fish schools consist of more individuals of similar size when populations are large than when they are small. The hypothesis that population size might be indicated by school structure is tested for southern bluefin tuna (SBT), a commercially important large pelagic species, which has undergone an estimated 60% reduction in juvenile biomass since 1960. Fish size data are used to determine whether there have been changes in schooling behaviour that can be used as simple indicators of abundance. During tagging studies, juvenile SBT are removed sequentially from a school, measured, tagged, and released. These sequential size measurements are used here to describe school composition from different years in two locations using simple school metrics (including mean fish size, variance in size, and mean difference in size between sequential fish). Trends were significant in most metrics over the 40-year period analysed, and were inversely related to independent estimates of population size. Simple school metrics are cost-efficient and easily interpreted by stakeholders. Monitoring population trends in near real time through school composition metrics may indicate further decline or recovery of SBT and, therefore, assist future management of tuna and other schooling species.

Historical changes in juvenile southern bluefin tuna Thunnus maccoyii growth rates based on otolith measurements

Suspected historic changes in juvenile southern bluefin tuna Thunnus maccoyii growth rates were investigated using otolith increment width data. Four hundred and ninety otoliths were selected from fish estimated to be between 1 and 41 years‐old. The distance between the first five annuli were measured on the otoliths, giving estimates of otolith growth for age classes 1+ to 4+ years for fish spawned from the early 1960s to mid 1990s. The data showed that growth rates of juveniles (age 1+ and 2+ years) started to increase at around 1979–1980, and that growth continued to increase throughout the 1980s and early 1990s. Lee’s phenomenon was not observed in the data. Correlation tests did not reveal clear relationships between annual otolith growth and regional environmental variables such as sea surface temperature or Southern Oscillation Index. The increase in otolith growth, however, was consistent with juvenile growth estimates obtained from other sources, and correlated with large‐scale trends in population size and environmental conditions.

Influence of upwelling on movement of southern bluefin tuna (Thunnus maccoyii) in the Great Australian Bight

Large pelagic predators move quickly in and out of local ecosystems that may be separated by long distances: their trophic effects are determined by their behaviour while present. To investigate movement and local residence times of one such predator we implanted 29 acoustic tags into juvenile southern bluefin tuna (Thunnus maccoyii) (SBT) in the Great Australian Bight. We used acoustic detectors at a reef known to attract tuna and detected fifteen SBT on 941 occasions over 62 days. SBT were tagged at the reef, 40 km, and 120 km distant. A total of 100% of local and 60% of SBT tagged 40 km away were subsequently recorded at the reef. Presence and absence was related to an upwelling event. Water temperature decreased just after SBT departure from the monitoring region. The immediate area was aerially surveyed 22 times for SBT schools during the experiment. We combined aerial survey observations with computer simulation, calibrated against field studies of SBT movement, to test the hypothesis that tuna could be well simulated by a correlated random walk throughout the area of known occurrence. The most plausible explanation for the observed behaviour was short-term (hours) fidelity to schools combined with medium-term (weeks) fidelity to bathymetric features. The present study illustrates how dynamic models aid interpretation of experiments designed to understand trophic effects of large pelagic predators.

ターゲットストレングス推定に必要なミナミマグロ（Ｔｈｕｎｎｕｓ ｍａｃｃｏｙｉｉ）幼魚の魚肉内音速測定

The ratio between the speed of sound in animals and that of the surrounding water (the sound-speed contrast), particularly for fish lacking any gas-filled organs or for zooplankton, is an important parameter in a theoretical scattering model for estimating target strength (TS). Since the swimbladder of a juvenile southern bluefin tuna (SBT) is not developed or gas-filled, the accurate and precise measurement of the sound-speed inside the body is required for TS estimation by the scattering model. The sound-speed of juvenile SBT tissue was measured using a FURUNO medical bone density-measuring instrument, the CM-100, during acoustic surveys conducted in Western Australia. This instrument measures the ultrasound travel time in tissues (and converts sound-speed to bone density). Since it has been recognized that sound-speed in tissues changes as time progresses and that it varies between fresh and defrosted tissues, the measurement should be taken immediately after the fish is caught. The sound-speed of juvenile SBT tissue ranged from 1610 to 1655 m/s and showed a tendency to depend on body temperature. On the assumption that the muscle temperature of a living SBT was 5 degrees higher than the surrounding water, the sound-speed contrast ranged from 1.06 to 1.09.