Ryan Johnson’s research while affiliated with Oceans and other places

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Publications (17)


Satellite tracks of 33 white sharks Carcharodon carcharias tagged at five locations along the South African coast in 2012. Each colour represents an individual shark’s track between 2012 and 2014. These are time-regular predicted locations (one location every 12 hours) from a correlated random walk model fitted to the original data, accounting for the estimated CLS Argos location error, using the foieGras package in R (Jonsen and Patterson, 2020).
Estimated state-dependent distribution of step length (km) and turning angle (radians). The resident state (purple) is made up of smaller steps and bigger turns (slow, undirected movement), while the transient state (sea green) is made up of bigger steps and smaller turns (fast, directed movement). The marginal distribution (lime) shows the sum of the two densities, while the data appear as a grey histogram.
Predicted shark locations coloured by Viterbi-decoded states. The tracks are separated by state (resident in purple: left column, transient in sea green: right column) and sex (females: top row, males: bottom row). Country codes are shown on the map: SA, South Africa; NA, Namibia; BW, Botswana; ZW, Zimbabwe; MZ, Mozambique; MG, Madagascar.
Predicted shark locations are coloured by the probability of a given location belonging to the transient state, where lighter locations indicate a high probability of being transient. This illustrates the level of uncertainty in state assignment; the closer the colour to either end of the colour scale, the greater the certainty in state assignment. The tracks are separated here by life stage based on size (juvenile: left column, subadult: middle column, adult: right column) and by sex (females: top row, males: bottom row). Country codes are shown on the map: SA, South Africa; NA, Namibia; BW, Botswana; ZW, Zimbabwe; MZ, Mozambique; MG, Madagascar.
The effect of day of the year on the probability of finding a shark in a given state, shown here by sex (females: left column, males: right column) and life stage (adult: top row, subadult: middle row, juvenile: bottom row). The shading around the state line represents the 95% confidence interval around the mean effect.

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Sex and Size Influence the Spatiotemporal Distribution of White Sharks, With Implications for Interactions With Fisheries and Spatial Management in the Southwest Indian Ocean
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April 2022

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734 Reads

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20 Citations

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Amanda T. Lombard

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Human activities in the oceans increase the extinction risk of marine megafauna. Interventions require an understanding of movement patterns and the spatiotemporal overlap with threats. We analysed the movement patterns of 33 white sharks (Carcharodon carcharias) satellite-tagged in South Africa between 2012 and 2014 to investigate the influence of size, sex and season on movement patterns and the spatial and temporal overlap with longline and gillnet fisheries and marine protected areas (MPAs). We used a hidden Markov model to identify ‘resident’ and ‘transient’ movement states and investigate the effect of covariates on the transition probabilities between states. A model with sex, total length and season had the most support. Tagged sharks were more likely to be in a resident state near the coast and a transient state away from the coast, while the probability of finding a shark in the transient state increased with size. White sharks moved across vast areas of the southwest Indian Ocean, emphasising the need for a regional management plan. White sharks overlapped with longline and gillnet fisheries within 25% of South Africa’s Exclusive Economic Zone and spent 15% of their time exposed to these fisheries during the study period. The demersal shark longline fishery had the highest relative spatial and temporal overlap, followed by the pelagic longline fishery and the KwaZulu-Natal (KZN) shark nets and drumlines. However, the KZN shark nets and drumlines reported the highest white shark catches, emphasising the need to combine shark movement and fishing effort with reliable catch records to assess risks to shark populations accurately. White shark exposure to shark nets and drumlines, by movement state, sex and maturity status, corresponded with the catch composition of the fishery, providing support for a meaningful exposure risk estimate. White sharks spent significantly more time in MPAs than expected by chance, likely due to increased prey abundance or less disturbance, suggesting that MPAs can benefit large, mobile marine megafauna. Conservation of white sharks in Southern Africa can be improved by implementing non-lethal solutions to beach safety, increasing the observer coverage in fisheries, and continued monitoring of movement patterns and existing and emerging threats.

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Comparing shark exposure risk between AIS longline fishing effort datasets
a–d, Estimated exposure risk of sharks to capture by GFW AIS longline fishing effort across ocean regions for Queiroz et al.¹ (a) compared with three improved data releases since the paper was published (b–d). The plots show minor effects of any changes on estimates of shark exposure risk from AIS longline fishing effort and confirm the global results and conclusions of our paper. a, Data from Queiroz et al.¹. b, Data from GWF 2012–2016. c, Data from GWF 2012–2018. d, Data from GWF 2018.
Reply to: Caution over the use of ecological big data for conservation

July 2021

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1,412 Reads

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11 Citations

Nature

106,107 ✉ replying to A. V. Harry & J. M. Braccini Nature https://doi.org/10.1038/s41586-021-03463-w (2021) Our global analysis 1 estimated the overlap and fishing exposure risk (FEI) using the space use of satellite-tracked sharks and longline fishing effort monitored by the automatic identification system (AIS). In the accompanying Comment, Harry and Braccini 2 draw attention to two localized shark-longline vessel overlap hotspots in Australian waters, stating that 47 fishing vessels were misclassified as longline and purse seine vessels in the Global Fishing Watch (GFW) 3 2012-2016 AIS fishing effort data product that we used. This, they propose 2 , results in misi-dentifications that highlight fishing exposure hotspots that are subject to an unexpected level of sensitivity in the analysis and they suggest that misidentifications could broadly affect the calculations of fishing exposure and the central conclusions of our study 1. We acknowledged in our previously published paper 1 that gear reclassifications were likely to occur for a small percentage of the more than 70,000 vessels studied, however, here we demonstrate that even using much larger numbers of vessel reclassifications than those proposed by Harry and Braccini 2 , the central results and conclusions of our paper 1 do not change. In our use of a third-party dataset such as GFW 3 , we stated clearly 1 that the dataset is undergoing continuous refinement to correct for acknowledged contamination of some gear types with others in some regions (for example, drifting longlines with bottom-set longlines off New Zealand 1). The characterization of GFW vessels (gear) is under-taken using two convolutional neural networks that were trained 3 on 45,441 marine vessels (fishing and non-fishing) that identified six Published online: xx xx xxxx Check for updates Q1 Q2 Q3 Q4 Q5 A list of affiliations appears at the end of the paper.




Figure 1: Capture locations for white sharks (Carcharodon carcharias) sampled in this study.
using antibiotic disc diffusion tests according to the Kirby-Bauer methodology outlined in the guidelines of the US Clinical and Laboratory Standards Institute. 11Classes and types of antibiotics used in disc diffusion tests to assess the antibiotic sensitivity of bacterial flora isolated from Carcharodon carcharias oral cavities
Antibiotic sensitivity of bacteria isolated from the oral cavities of live white sharks (Carcharodon carcharias) in South African waters

November 2019

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984 Reads

South African Journal of Science

The white shark (Carcharodon carcharias) is responsible for 49% of shark-related injuries in South Africa, yet no information currently exists on the composition or antibiotic resistance of bacteria hosted by these apex predators in South African waters. This study aimed to address this gap by sampling the bacteria present in the oral cavities of 28 live C. carcharias along South Africa’s southern coastline. The antibiotic resistance of the range of microbiota was also assessed using antibiotic disc diffusion tests. A total of 51 strains from at least 20 species of bacteria were isolated from the oral cavities of C. carcharias. Of these strains, the most common bacteria present were Serratia spp., Proteus vulgaris and Vibrio alginolyticus. The overall antibiotic resistance was relatively higher in this study than that reported for bacterial microbiota sampled from other shark species. Results indicate that the combination therapy of imipenem (carbapenem antibiotic) and vancomycin (glycopeptide antibiotic) might be the most parsimonious option to effectively treat infections resulting from white shark bites, particularly in South Africa. It is hoped that, in addition to assisting medical professionals to treat shark bite victims, these findings enhance the understanding of the microbial communities present in large coastal predators and their surrounding environments. Significance: • Overall antibiotic resistance of bacteria in the oral cavities of C. carcharias was relatively high. • Combination therapy of imipenem (carbapenem antibiotic) and vancomycin (glycopeptide antibiotic) is recommended for the treatment of white shark bites, particularly in South Africa. • The findings add to understanding of the microbial communities present in large coastal predators and their surrounding environments.


Global spatial risk assessment of sharks under the footprint of fisheries

July 2019

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4,352 Reads

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360 Citations

Nature

Effective ocean management and the conservation of highly migratory species depend on resolving the overlap between animal movements and distributions, and fishing effort. However, this information is lacking at a global scale. Here we show, using a big-data approach that combines satellite-tracked movements of pelagic sharks and global fishing fleets, that 24% of the mean monthly space used by sharks falls under the footprint of pelagic longline fisheries. Space-use hotspots of commercially valuable sharks and of internationally protected species had the highest overlap with longlines (up to 76% and 64%, respectively), and were also associated with significant increases in fishing effort. We conclude that pelagic sharks have limited spatial refuge from current levels of fishing effort in marine areas beyond national jurisdictions (the high seas). Our results demonstrate an urgent need for conservation and management measures at high-seas hotspots of shark space use, and highlight the potential of simultaneous satellite surveillance of megafauna and fishers as a tool for near-real-time, dynamic management.


Ten-second-averaged distance-to-signal strength results from the six detection range test experiments conducted in coastal waters off Mossel Bay (estimated by additive models using gam package and span = 0.9; the dotted lines represent the 95% confidence intervals)
Occurrence of depth and temperature false detections per signal strength bins in an experiment with a transmitter position underneath a fixed buoy (0.05 threshold line is added)
Reliability test of tracking data from white sharks (n = 6) actively tracked in Mossel Bay. The absolute error values (telemetered data compared to a 1-h averaging moving window), absolute mean error values (second row), and absolute maximum error values (third row) for water temperature (Tw, left column) and stomach temperature (Ts, right column) were plotted against signal strength (proxy for distance from the animal) row values (first row) and binned values (second and third row)
Weighted occurrence of outliers for each speed (3D-ROM) bin of the white sharks (n = 6) actively tracked in Mossel Bay
Performance and reliability of active acoustic biotelemetry to best track marine pelagic species in temperate coastal waters

July 2018

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73 Reads

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4 Citations

Marine Biology

Acoustic tracking is a commonly used method to study the movement ecology of marine species. The characteristics of the data collected are not simply functions of the location of the tagged animal as they are also influenced by the method of data collection and its sampling frequency. In particular, since the data are acoustically driven, the significance of any result hinges not only on the accuracy of the equipment but also on the reliability of the information received. While passive acoustic telemetry requires an evaluation of the detection efficiency of receivers moored in different habitats (or sites) to obtain reliable data, active acoustic telemetry requires more field-related information on how to best track tagged animals without impacting on their natural behaviour. This study aimed to evaluate the reliability of active acoustic telemetry data in a temperate coastal environment. This was achieved by conducting a series of field experiments to assess (i) the use of signal strength-to-infer distance to the tracked animal under different tracking conditions, (ii) what signal strength threshold can be used to obtain reliable bio-telemetered data, (iii) whether the behaviour of a tracked animal would be a main concern in terms of data reliability, and (iv) the best filtering option to reduce the data to be analysed. The findings yielded a signal strength-to-distance relationship to improve the accuracy of the positional fixes of actively tracked animals, which can be used as a practical reference for future tracking studies on pelagic species in temperate coastal environments.



Table 1 (continued) 
Table 2 Level of overlap from one sharks home range to each of the others in the study as determined by IOSS 
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Fine scale movements and activity areas of white sharks (Carcharodon carcharias) in Mossel Bay, South Africa

July 2013

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2,018 Reads

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45 Citations

Environmental Biology of Fishes

Previous work on white sharks indicate the species show seasonally limited movement patters, at certain aggregation sites small areas may play vital roles in the life history of a large amount of the population. Acoustic telemetry was used to estimate habitat use of white sharks, Carcharodon carcharias, while aggregating at Mossel Bay, South Africa. Total range of all shark tracks combined accumulated 782 h and covered an area of 93.5 km2 however, within this range, sharks were found to highly utilise a core habitat (50 % Kernel, K50) of just 1.05 km2 over a reef system adjacent to a river mouth. Individual tracks revealed additional core habitats, some of which were previously undocumented and one adjacent to a commercial harbor. Much was found to be dependent on the size of the shark, with larger sharks (>400 cm) occupying smaller activity areas than subadult (300–399 cm) and juvenile (<300 cm) conspecifics, while Index of Reuse (IOR) and Index of Shared Space (IOSS) were both found to increase with shark size. Such results provide evidence that larger white sharks are more selective in habitat use, which indicates they have greater experience within aggregation sites. Furthermore, the focused nature of foraging means spatially restricted management strategies would offer a powerful tool to aid enforcement of current protective legislation for the white shark in similar environments of limited resources and capacity.


Values are means ± SE (n = 60 replicates).
Values are means ± SE (n = 22 replicates).
Accuracy of Using Visual Identification of White Sharks to Estimate Residency Patterns

April 2012

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91 Reads

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14 Citations

Determining the residency of an aquatic species is important but challenging and it remains unclear what is the best sampling methodology. Photo-identification has been used extensively to estimate patterns of animals' residency and is arguably the most common approach, but it may not be the most effective approach in marine environments. To examine this, in 2005, we deployed acoustic transmitters on 22 white sharks (Carcharodon carcharias) in Mossel Bay, South Africa to quantify the probability of detecting these tagged sharks by photo-identification and different deployment strategies of acoustic telemetry equipment. Using the data collected by the different sampling approaches (detections from an acoustic listening station deployed under a chumming vessel versus those from visual sightings and photo-identification), we quantified the methodologies' probability of detection and determined if the sampling approaches, also including an acoustic telemetry array, produce comparable results for patterns of residency. Photo-identification had the lowest probability of detection and underestimated residency. The underestimation is driven by various factors primarily that acoustic telemetry monitors a large area and this reduces the occurrence of false negatives. Therefore, we propose that researchers need to use acoustic telemetry and also continue to develop new sampling approaches as photo-identification techniques are inadequate to determine residency. Using the methods presented in this paper will allow researchers to further refine sampling approaches that enable them to collect more accurate data that will result in better research and more informed management efforts and policy decisions.


Citations (12)


... As upper-trophic-level predators, they play an important ecological role (Carrier et al., 2010), primarily feeding on small sharks and rays, squid, and benthic fishes as small juveniles (< 2.5 m, Estrada et al., 2006;Clark et al., 2023) and incorporating marine mammals into their diet as they grow (Tricas and McCosker, 1984;Hussey et al., 2012). Large juvenile, subadult, and adult white sharks seasonally aggregate near pinniped colonies often when water temperatures correspond to their preferred range (Klimley et al., 2001;Bruce and Bradford, 2015;Hewitt et al., 2018;Kock et al., 2022;Winton et al., 2023). Much of our understanding of white shark movement ecology comes from studies where animals are tracked with acoustic or satellite tags at these sites (e.g., Neptune Islands, South Australia; Guadalupe Island, Mexico; California and Cape Cod, U.S.A), which have documented philopatric behaviors, such as high site fidelity, seasonal residency as well as long-distance return migrations (Bonfil et al., 2005;Jorgensen et al., 2010;Bruce and Bradford, 2015;Skomal et al., 2017;Huveneers et al., 2018;Bastien et al., 2020). ...

Reference:

First evidence of white sharks, Carcharodon carcharias, in the tongue of the ocean, central Bahamas
Sex and Size Influence the Spatiotemporal Distribution of White Sharks, With Implications for Interactions With Fisheries and Spatial Management in the Southwest Indian Ocean

... Oceanic shark and ray populations are estimated to have decreased by as much as 71% since 1970 , and approximately 32% of elasmobranch species are currently assessed as being at risk of extinction . A few regions of high space use are free from industrial fishing pressures, with many species exposed to significant capture risk for extended periods of time across ranges (Queiroz et al., 2016;Queiroz et al., 2019;Queiroz et al., 2021). The typically slow growth and maturation rates, coupled with low fecundity, make elasmobranchs particularly susceptible to population declines after large-scale removals and limit the ability of many stocks to recover (Cortes et al., 2010;Myers & Worm, 2005;Smith et al., 1998). ...

Matters arising: Reply to: Shark mortality cannot be assessed by fishery overlap alone
  • Citing Article
  • July 2021

Nature

... Other studies have indeed shown that consensus models can produce reliable macro patterns (Coro et al., 2024a,d;Coro, 2024). The biases on the individual cells can indeed fade away when macroscopic patterns are produced through spatial aggregation (Weber et al., 2004;Queiroz et al., 2021). ...

Reply to: Caution over the use of ecological big data for conservation

Nature

... While there is a growing knowledge base of information on how threats like overfishing, habitat loss, and climate change-induced shifts in prey availability impact shark species in shallow coastal and open ocean regions, there is comparatively limited information available from the deep sea (Field et al., 2009;Queiroz et al., 2019). Additionally, the extensive exploitation of oil, gas, and fisheries biomass presents substantial, and potentially increasing, challenges to deep-sea environments (Howell et al., 2020). ...

Global spatial risk assessment of sharks under the footprint of fisheries

Nature

... A combination of sensor technologies can form a wireless communication system to monitor large scale aquaculture activities, and increases the accuracy of data, processing and interpretation (Kessel et al., 2014). These data provide the industry with information needed to facilitate performance, health and welfare and to optimise management practices. ...

Performance and reliability of active acoustic biotelemetry to best track marine pelagic species in temperate coastal waters

Marine Biology

... It is noteworthy that as sharks increased in age, the individual variability of area use increased, where a portion of 3-and 4-year-old sharks (the largest sharks analyzed here) used smaller amounts of space than 1-and 2-year olds (see Figure 3). Using active acoustic tracking, Jewell et al. (2013) found that large white sharks (>400 cm TL) occupied smaller activity areas (~2 km 2 ) than both subadult and juveniles (8-10 km 2 ) in South Africa, suggesting that larger sharks may competitively exclude smaller sharks from optimum habitat. Similarly, on a small scale like the present study, individual variation in movement patterns and habitat use has also been shown across ontogeny in juvenile bull sharks (Carcharhinus leucas), where a portion of older individuals spent a greater proportion of time in the most productive habitats, while others did not (Matich & Heithaus, 2015); however, on larger scales bull sharks may partition space more predictably in regard to size (Matich et al., 2020). ...

Fine scale movements and activity areas of white sharks (Carcharodon carcharias) in Mossel Bay, South Africa

Environmental Biology of Fishes

... Most studies examining the effects of wildlife tourism have largely been on focal species, with only 7% exploring non-focal species or ecosystemwide effects (Trave et al., 2017. Similar impacts to those observed in focal species have also been observed in non-focal species , Vignon et al., 2010, providing some of the only evidence of non-focal impacts. Due to the lack of studies examining non-focal impacts the need for further research has been highlighted in several studies and reviews (Gallagher et al., 2015, Trave et al., 2017, Gallagher and Huveneers, 2018, Patroni et al., 2018, Meyer et al., 2021. ...

Impact of shark-feeding tourism on surrounding fish populations off Moorea Island (French Polynesia)

Marine and Freshwater Research

... Sharks have been demonstrated to aggregate around sites of known prey productivity (Mourier et al. 2016;Schilds et al. 2019). For C. carcharias, adults have been known to aggregate around pinniped colonies (Johnson et al. 2009;Schilds et al. 2019) and neonate to subadults have been known to aggregate in areas with an abundance of bottom dwelling fishes and elasmobranchs (White et al. 2019;Grainger et al. 2020). Similar to C. Carcharias nursery areas that have been characterized as having abundant prey resources (e.g., Oñate-González et al. 2017;Santana-Morales et al. 2012) and as a result of their high metabolic rates (Semmens et al. 2013), it can be inferred that parturition sites should exhibit high prey availability to maximize survival. ...

Coastal swimming patterns of white sharks (Carcharodon carcharias) at Mossel Bay, South Africa

Environmental Biology of Fishes

... One of the main conditions for photo ID is that morphological features used for identification must be both discernible and stable through time, and several species of elasmobranchs exhibit natural patterns conducive to ID (Armstrong et al., 2020;Arzoumanian et al., 2005;Harty et al., 2022). However, only a few studies have attempted to validate photo ID through multi-modal methodologies or by convergent evidence from different markers, including sex and external tags (Bansemer & Bennett, 2008;Dudgeon et al., 2008;Gubili et al., 2009;Winton et al., 2023). For example, white sharks are identified by the trailing edge of their dorsal fin, which may seem quite similar among individuals, but on closer examination it carries substantial individuality (Andreotti et al., 2014). ...

Concordance of genetic and fin photo identification in the great white shark, Carcharodon carcharias, off Mossel Bay, South Africa

Marine Biology

... The relatively straightforward integration of georeferenced data sources, such as telemetry data, is an additional advantage of the spatial nature of our framework. Tagging studies are often conducted in conjunction with photo-ID surveys at aggregation sites (Delaney et al. 2012), and the data that are collected are used indirectly to aid in interpretation of the resulting abundance estimates and to assess the validity of model assumptions , Burgess et al. 2014. Telemetry data have been directly integrated via additional likelihood components in closed SCR models for terrestrial carnivores to inform relationships with environmental covariates and the scale of the movement process via σ (Royle et al. 2013, Sollmann et al. 2013, Tenan et al. 2017, Linden et al. 2018) but have not previously been used to inform parameters related to movements into and out of the state space in an open SCR model. ...

Accuracy of Using Visual Identification of White Sharks to Estimate Residency Patterns