Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

Contents lists available at ScienceDirect

Journal of Experimental Marine Biology and Ecology

journal homepage: www.elsevier.com/locate/jembe

Monograph

Global tracking of shark movements, behaviour and ecology: A review of

the renaissance years of satellite tagging studies, 2010–2020
Samantha Renshaw a, f, *, 1, Neil Hammerschlag b, Austin J. Gallagher c, Nicolas Lubitz d,
David W. Sims e, f
a
Dalhousie University, Faculty of Science, Marine Affairs, Halifax, NS, Canada
b
Rosenstiel School of Marine and Atmospheric Science, University of Miami, 4600 Rickenbacker Causeway, Miami, FL 33149, USA
c
Beneath the Waves, PO Box 126, Herndon, VA 2017, USA
d
College of Science and Engineering, James Cook University, Australia
e
Marine Biological Association of the UK, The Laboratory, Citadel Hill, Plymouth PL1 2PB, UK
f
Ocean and Earth Science, National Oceanography Centre Southampton, University of Southampton, European Way, Southampton, UK

A R T I C L E I N F O A B S T R A C T

Keywords: Satellite telemetry as a tool in marine ecological research continues to adapt and grow and has become
Satellite telemetry increasingly popular in recent years to study shark species on a global scale. A review of satellite tag application
Sharks to shark research was published in 2010, provided insight to the advancements in satellite shark tagging, as well
Tracking
as highlighting areas for improvement. In the years since, satellite technology has continued to advance, creating
Habitat use
smaller, longer lasting, and more innovative tags, capable of expanding the field. Here we review satellite shark
Conservation
tagging studies to identify early successes and areas for rethinking moving forward. Triple the amount of shark
satellite tagging studies have been conducted during the decade from 2010 to 2020 than ever before, tracking
double the number of species previously tagged. Satellite telemetry has offered increased capacity to unravel
ecological questions including predator and prey interactions, migration patterns, habitat use, in addition to
monitoring species for global assessments. However, ω17% of the total reviewed studies directly produced re-
sults with management or conservation outcomes. Telemetry studies with defined goals and objectives produced
the most relevant findings for shark conservation, most often in tandem with secondary metrics such as fishing
overlap or management regimes. To leverage the power of telemetry for the benefit of shark species, it remains
imperative to continue improvements to tag function and maximize the outputs of tagging efforts including
increasing data sharing capacity and standardization across the field, as well as spatial and species coverage.
Ultimately, this review offers a status report of shark satellite tagging and the ways in which the field can
continue to progress.

1. Introduction movement extents, and spatial overlap with fisheries (Hammerschlag

et al., 2011b). Insights from tracking studies have led to several tangible
Tracking technologies have provided unparalleled insights into use management and conservation outcomes, such as providing supporting
of the three-dimensional ocean environment by marine animals and evidence for the designation of marine protected areas (MPAs) and
contributed invaluable information to ecological, biological, and policy motivating the protection of at-risk elasmobranch species (Chapman
disciplines, particularly for apex predators like sharks (Hussey et al., et al., 2005; Espinoza et al., 2015; Byrne et al., 2017).
2015; Hays et al., 2019). Elasmobranchs (sharks, skates and rays) have Hammerschlag et al. (2011b) provided a review of studies using
been studied using various telemetry or bio-logging technologies for satellite telemetry to monitor shark movements and behaviour from
several decades, providing better understanding of species migrations, 1984 to 2010, and described the state of the field, identified knowledge
population connectivity, habitat preferences, horizontal and vertical gaps, and suggested numerous avenues for future research. A key

* Corresponding author at: Dalhousie University, Faculty of Science, Marine Affairs, Halifax, NS, Canada.
E-mail address: [email protected] (S. Renshaw).
1
Permanent address: 1475 Lower Water St., B3J 0B5, Halifax, NS

https://doi.org/10.1016/j.jembe.2022.151841
Received 26 September 2021; Received in revised form 6 November 2022; Accepted 18 November 2022
Available online 3 January 2023
0022-0981/© 2022 Published by Elsevier B.V.
S. Renshaw et al. Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

research priority identified was the need to go beyond simply using failures and premature releases of pop-off satellite archival transmitter
satellite transmitters to describe shark movements and environmental (PSAT) tags were recorded when the information was available. In cases
preferences but to extend studies by testing competing hypotheses and where the minimum tracking duration was zero, this was considered tag
attempting to evaluate explicit drivers underlying movement and failure, so the next shortest tracking duration was used to calculate
habitat use patterns. averages. Premature tag release was not considered tag failure, unless
In the decade since the review by Hammerschlag et al. (2011b), the tracking time was zero days. If minimum or maximum tracking
spanning 2010–2020, there has been a substantial expansion in shark duration, premature release or tag failure rate was not provided, the
satellite tagging studies (henceforth, tagging will refer to satellite study was removed from respective analyses. Each study's purpose(s)
technologies unless otherwise specified), adding a range of new species wasrecorded to identify common themes throughout the reviewed
and size classes, with new monitoring occurring in every ocean basin. In literature. In some cases, studies had multiple purposes and they were
addition, several large tagging initiatives have been completed, for each identified accordingly. Purpose(s) was identified from stated ob-
example, the Tagging of Pacific Predators Program (TOPP, Block et al., jectives, goals, or explicit hypotheses within the introductory para-
2011), which tracked several thousand marine vertebrates, including graphs of a study. Instances where satellite telemetry was carried out in
sharks, in the Eastern Pacific Ocean between 2000 and 2010. The past service of a testable hypothesis were recorded. Statistical significance
decade has also seen the emergence of collaborative networks synthe- was determined between quantitative values including tagging dura-
sizing the increased volume of shark satellite tracking data from across tion, using simple two-tailed t-test analyses, performed using R Studio
the globe in order to address ocean-wide conservation issues, for (Version 1.3.1093) with significance determined at α → 0.05. A hotspot
example the Global Shark Movement Project (GSMP; www.glob analysis calculating the Getis-Ord Gi* statistic was used to determine
alsharkmovement.org;Queiroz et al., 2019). Furthermore, new in- binned locations where a high or low number of sharks were tagged
novations in tagging technology have allowed researchers to better spatially, based on geolocations provided by each study. This process
describe the drivers of habitat use patterns for numerous shark species. works by looking at each feature within the context of neighboring
Spanning nearly 40 years of use, shark tagging studies have evolved in features. Hotspots represent areas where the z-score is high and p-value
intellectual design, tag innovation, and output. Therefore, an updated is small, resulting in a 99% confidence interval. Cold-spots indicate areas
review of this most recent ‘renaissance’ decade of shark tagging is of low shark tagging and thus low z-scores and high p-values. Where
warranted to identify progress made and to propose new questions and studies aggregated tagging resulted from several different tagging sites
research priorities, as we approach nearly a half-century of development (ex. Queiroz et al., 2019 and Block et al., 2011), every effort was taken to
and implementation. account for each tagging location used. Where exact latitude or longi-
The purpose of this study is to generate a contemporary review of tude of tagging locations was not provided, an approximate location
shark satellite tagging studies from 2010 to 2020, comparing the current based on the study's description was chosen. Geo-spatial analysis was
state of the field with the results since Hammerschlag et al. (2011b), performed using QGIS (3.18.3).
hereafter referred to as the “original review”) and highlighting ad-
vancements and shortcomings over the past decade. In order to compare 3. Results and discussion
to the original review, we address specific topics including why, where
and how shark tagging studies have been performed, the types of sat- Our search identified a total of 176 studies available for examination
ellite tags used and what types of data they have provided, as well as in this review (Table 1), resulting in a ω 3.6 increase in the volume of
how satellite tracking data has supported shark conservation and man- studies published on the topic since 2010 (n → 48). Similar to the find-
agement decisions. Through these topics, this study aims to showcase ings of Hammerschlag et al. (2011b), most studies utilized satellite
novel applications of satellite technology for tracking sharks, assess the telemetry to describe the movement of shark species. Of these, just 26%
extent to which knowledge gaps identified in the original review have (n → 46) were in direct service of conservation or management. Average
been addressed, understand how the field of shark tagging and tracking sample size across all studies was 37 tagged individuals (ranging from 1
has developed in the last 10 years, and explore the field's future tra- to 1804; mode → 1, median → 15; Fig. 1). The longest track recorded a
jectory for improving our understanding of shark behaviour, ecology, white shark tagged in 2008 at Guadalupe Island, Mexico with a fin
and conservation. mounted Argos-linked satellite transmitter (SAT) (model used: SPOT –
smart position-only transmitter – tag, Wildlife Computers, Redmond
2. Methods WA, USA; Domeier and Nasby-Lucas, 2012) where data transmission
ended in 2016, however the total distance traveled is not presently re-
Database searches of Web of Science and Google Scholar were con- ported (Nasby-Lucas and Domeier, 2020). Whale shark tagging has
ducted as per Hammerschlag et al. (2011b) and limited to primary consistently pushed satellite tracking boundaries, providing some of the
literature published from July 2010 to July 2020. The words ‘satellite furthest and deepest tracking records to date. Guzman et al. (2018)
(s)’, ‘shark(s)’, ‘elasmobranch(s)’, and/or ‘Argos’ were used in combi- recorded a whale shark travelling from Panama to the Marianas Trench,
nation to search through key words, titles, and abstracts to find appli- a journey totaling over 20,140 km (or over 13,800 km in a straight path).
cable sources. Additional papers were identified from the works cited of Similarly, Tyminski et al. (2015) recorded one of the deepest tracked
the reviewed literature. For the purpose of this review, studies using dives by a whale shark 1928 m in the Northern Gulf of Mexico, limited
only bio-logging devices without satellite connectivity, or where satel- by triggered tag release due to extreme pressure. With each tagging
lite connectivity was only used for a tag's physical retrieval at sea, were milestone comes novel ecological data. For example, the longest known
excluded. tracks for tiger sharks (1101 days; Lea et al., 2015a) revealed extensive
Following the methodology from the original review, the informa- seasonal migrations and return movements to core habitats demon-
tion recorded from each study included: (1) study location, (2) species strating philopatric behaviour (SPOT5, Wildlife Computers, Redmond,
tagged, (3) study objective(s), (4) satellite tag type, (5) any additional Washington, USA).
methods used (i.e., acoustic telemetry), (6) sample size; (7) catch and
release methods; (8) tag attachment and deployment; (9) minimum and 3.1. Study objectives and common themes
maximum tracking days (duration); (10) tag performance; (11) shark
behaviour; and (12) any study limitations. Consideration of a studies The original review found that the primary objectives of most shark
contribution to conservation was also considered where applicable. tracking studies were to determine movement patterns and/or evaluate
Recording procedures were replicated as outlined by Hammerschlag the depth and temperature preferences of sharks, which mirrors the
et al. (2011b), including tag performance, whereby the percentage of tag studies reviewed here (98%). The remaining studies measured the

2
 S. Renshaw et al.
Table 1
Published studies between 2010 and 2020 using satellite telemetry on sharks. Study objectives are abbreviated and provided to reveal common themes across studies as follows: (i) ‘Fisheries/PRM’ - fisheries interaction/
post-release survival; (ii) ‘Tag effects’ - effects of tags on sharks' wellbeing; (iii) “MPA’ – tagging to determine MPA effectiveness; (iv) “Enviro’ - environmental drivers of movements; and (vi) ‘Pred/Prey’ - predator/prey
interactions. An asterisks (*) notes that calculated failure rate was inclusive of premature-pop off tags. In cases where estimates of tracking times, premature pop-offs, and tag failures were reported together across all tag
types, for instance, when sharks were double tagged, it is noted as ‘ALL’.
Author(s) Study Ocean Region/Country Species Tagged Satellite Total Tag Min. Max. Premature Tag
Objectives Basin Tag Type Sample Attachment Tracking Tracking Pop-off Failures
Size Methods Days days

Tiger shark (Galeocerdo cuvier), 100

PSAT, Fin mount, 3 (PSAT),
1 Meyer et al., 2010 Pacific Hawaii, USA Galapagos shark (Carcharhinus 8 (PSAT), 88% (PSAT) 0% (ALL)
SAT dart 22 (SAT)
galapagensis) 196 (SAT)
Sulikowski et al.,
2 Atlantic Northwest Atlantic Spiny dogfish (Squalus acanthias) PSAT 3 Fin loop 65 189 100% 0%
2010
Heberer et al., Fisheries/
3 Pacific Southern California Common thresher shark (Alopias vulpinus) PSAT 18 Dart 0 10 NA 6%
2010 PRM
da Silva et al.,
4 Atlantic South Africa Blue sharks (Prionace glauca) PSAT 3 NA NA NA NA NA
2010
Otway and Ellis, New South Wales,
5 Pacific Sand tiger (Carcharias taurus) PSAT 15 Dart 38 180 27% 13%
2011 Australia
Saunders et al.,
6 Enviro Atlantic Ireland Porbeagle sharks (Lamna nasus) PSAT 3 Dart NA 122 0% 0%
2011
Hammerschlag
7 Atlantic Florida Keys, USA Great Hammerhead (Sphyrna mokarran) SAT 1 Fin mount NA 62 NA 0%
et al., 2011a
Campana et al.,
8 Enviro Atlantic Canada Blue sharks (Prionace glauca) PSAT 40 Dart 40 210 43% 8%
2011
Abascal et al.,
9 Pacific Northern Chile Shortfin mako (Isurus oxyrinchus) PSAT 9 Dart 3 159 77% 0%
2011
Barnett et al., Broadnose sevengill shark (Notorynchus
10 MPA Pacific Tasmania, Australia PSAT 5 Dart 34 156 100% 0%
3

2011 cepedianus)
Blue sharks (Prionace glauca), Porbeagle
sharks (Lamna nasus), White sharks
Hoolihan et al., Fisheries/ (Carcharodon carcharias), Oceanic
11 Various Various PSAT 53 Dart 26 358 NA NA
2011 PRM whitetip sharks (Carcharhinus
longimanus), Silky sharks (Carcharhinus
falciformis)
Shortfin mako (Isurus oxyrinchus), White

Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

sharks (Carcharodon carcharias), Blue
Several locations - see PSAT, Dart, Fin
12 Block et al., 2011 Enviro Pacific sharks (Prionace glauca), Salmon shark 321 NA 1380 NA NA
study for full list. SAT mount
(Lamna ditropis), Common thresher shark
(Alopias vulpinus)
Blue shark (Prionace glauca), Shortfin
mako (Isurus oxyrinchus), Silky shark
Fisheries/
13 Musyl et al., 2011 Pacific Hawaii, USA (Carcharhinus falciformis), Oceanic PSAT 71 Fin loop 0 247 NA 38%
PRM
whitetip shark (Carcharhinus lorigimanus),
Big eye thresher (Alopias superciliosus).
14 Jewell et al., 2011 Tag effects Indian Gansbaai, South Africa White sharks (Carcharodon carcharias) SAT 15 Fin mount NA NA NA NA
Bessudo et al., Malpelo and Cocos Scallopped hammerhead sharks (Sphyrna
15 Pacific PSAT 12 Dart 1 148 100% 25%
2011 Islands lewini)
Brunnschweiler
16 Indian Mozambique Whale shark (Rhincodon typus) PSAT 1 NA NA 87 100% NA
and Sims, 2011
Hammerschlag Florida USA &
17 Atlantic Tiger shark (Galeocerdo cuvier) SAT 25 Fin mount 26 297 NA 1%
et al., 2012a Bahamas
18 Wang et al., 2012 Pacific South China Sea Whale shark (Rhincodon typus) PSAT 1 NA NA 74 100% 0%
Wegner and Fisheries/
19 Pacific California, USA Shortfin mako (Isurus oxyrinchus) PSAT 1 Dart NA 54 NA 0%
Cartamil, 2012 PRM
Domeier et al., Guadalupe Island, PSAT, Dart, Fin
20 Pacific White sharks (Carcharodon carcharias) 73 15 (PSAT) 239 (PSAT) NA NA
2012 Mexico SAT mount
(continued on next page)
 S. Renshaw et al.
Table 1 (continued )
Author(s) Study Ocean Region/Country Species Tagged Satellite Total Tag Min. Max. Premature Tag
Objectives Basin Tag Type Sample Attachment Tracking Tracking Pop-off Failures
Size Methods Days days

Chatham Islands & 274

PSAT, Dart, Fin
21 Duffy et al., 2012 Pacific Stewart Islands, New White sharks (Carcharodon carcharias) 25 4 (PSAT) (PSAT), 23 16% (PSAT) 0% (ALL)
SAT Mount
Zealand (SPOT)
Jorgensen et al.,
22 Pacific Central California White sharks (Carcharodon carcharias) PSAT 97 Dart 0 362 NA 30%
2012
Queiroz et al., Fisheries/
23 Atlantic England & Portugal Blue sharks (Prionace glauca) PSAT 16 Dart 13 105 100% 19%
2012 PRM, Enviro
Hammerschlag
24 Pred/Prey Atlantic Florida USA Bull shark (Carcharhinus leucas) SAT 18 Fin mount 17 244 NA 11%
et al., 2012b
Greenland sharks (Somniosus
25 Fisk et al., 2012 Arctic Norway PSAT 20 Dart 8 196 15% 30%
microcephalus)
Fitzpatrick et al., Great Barrier Reef
26 Pred/Prey Pacific Tiger shark (Galeocerdo cuvier) SAT 10 Fin mount 16 356 NA 0%
2012 Marine Park
Guadalupe Island,
Domeier and
27 Pacific Mexico & Southeast White sharks (Carcharodon carcharias) SAT 17 Fin mount 356 722 NA NA
Nasby-Lucas, 2012
Farallon Island, USA
Filmalter et al., Fisheries/ Fin mount,
28 Indian Seychelles Silky sharks (Carcharhinus falciformis) PSAT 43 4 94 NA NA
2013 PRM Fin loop
Great Australian Bight,
29 Rogers et al., 2013 Indian Dusky shark (Carcharhinus obscurus) PSAT 3 Dart 176 182 0% 0%
Southern Australia
PSAT, Fin Clamp, 30 (PSAT),
30 Thums et al., 2013 Indian Western Australia Whale shark (Rhincodon typus) 4 6 (PSAT) NA NA
SAT Dart 88 (SAT)
Howey-Jordan Oceanic whitetip sharks (Carcharhinus
31 Enviro, MPA Atlantic Bahamas PSAT 12 Dart 31 245 0% 8%
et al., 2013 longimanus)
Hoffmayer et al., Scallopped hammerhead sharks (Sphyrna
4

32 Atlantic Mississippi Delta PSAT 1 Dart NA 27 0% NA

2013 lewini)
Domeier and Guadalupe Island,
33 Pacific White sharks (Carcharodon carcharias) SAT 4 Fin mount NA 510 NA 0%
Nasby-Lucas, 2013 Mexico
34 Hearn et al., 2013 Pacific Galapagos Islands Whale shark (Rhincodon typus) SAT 6 Dart 31 167 0% 0%
190
Yucatan Peninsula, PSAT, Dart, Fin 20%
35 Hueter et al., 2013 Atlantic Whale shark (Rhincodon typus) 35 2 (PAT) (PSAT), 30 NA
Mexico SAT mount (PSAT)
(SAT)

Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

36 Hazin et al., 2013 Atlantic Pernambuco, Brazil Tiger shark (Galeocerdo cuvier) PSAT 5 Fin loop 4 74 20% 0%
Del Raye et al.,
37 Pacific Central California, USA White sharks (Carcharodon carcharias) PSAT 97 Dart NA NA NA NA
2013
Fisheries/ SAT,
38 Lyons et al., 2013 Pacific Southern California White sharks (Carcharodon carcharias) 17 Fin mount 10 (SAT) 149 (SAT) NA 12% (SAT)
PRM PSAT
US Virgin Islands,
39 Vaudo et al., 2014 Enviro Atlantic Tiger shark (Galeocerdo cuvier) PSAT 14 Dart 9 184 NA 29%
Bermuda
18%
PSAT, Fin mount, 4 (PSAT), 38 (SAT),
40 Werry et al., 2014 Pacific Coral Sea Tiger shark (Galeocerdo cuvier) 14 73% (PSAT) (PSAT),
SAT Dart 7 (SAT) 210 (PSAT)
0% (SAT)
Stehfest et al., Broadnose sevengill shark (Notorynchus
41 Pacific Tasmania, Australia PSAT 10 Dart 13 153 60% 0%
2014 cepedianus)
Great Hammerhead (Sphyrna mokarran),
Gallagher et al., Fisheries/ Florida USA &
42 Atlantic Bull sharks (Carcharhinus leucas), Tiger SAT 83 Fin mount NA NA NA NA
2014 PRM Bahamas
shark (Galeocerdo cuvier)
Hoffmayer et al.,
43 Enviro Atlantic Mississippi Delta Dusky shark (Carcharhinus obscurus) PSAT 10 Dart 6 124 80% 0%
2014
Afonso and Hazin, Fisheries/
44 Atlantic Recife, Brazil Tiger shark (Galeocerdo cuvier) PSAT 21 Fin loop 4 159 57% 10%
2014 PRM
Kneebone et al.,
45 Atlantic Massachusetts, USA Sand tiger (Carcharias taurus) PSAT 15 Dart 0 56 13% 80%
2014
(continued on next page)
 S. Renshaw et al.
Table 1 (continued )
Author(s) Study Ocean Region/Country Species Tagged Satellite Total Tag Min. Max. Premature Tag
Objectives Basin Tag Type Sample Attachment Tracking Tracking Pop-off Failures
Size Methods Days days

Poisson et al., Fisheries/

46 Indian Western Indian Ocean Silky sharks (Carcharhinus falciformis) PSAT 31 Dart 2 100 NA 31%
2014 PRM
Dart, Dorsal
47 Cuevas et al., 2014 Atlantic Northern Patagonia Tope sharks (Galeorhinus galeus) PSAT 5 8 22 60% 40%
dart anchor
SAT, 5 (SAT), 42 45 (SAT),
48 Curtis et al., 2014 Enviro Atlantic Massachusetts, USA Basking sharks (Cetorhinus maximus) 17 Dart NA 0% (ALL)
PSAT (PSAT) 98 (PSAT)
Carlson et al.,
49 Atlantic New England, USA Spiny dogfish (Squalus acanthias) PSAT 40 Fin loop 10 369 79% 15%
2014a
315
Berumen et al., PSAT, 11 (PSAT),
50 Indian Saudi Arabian Red Sea Whale shark (Rhincodon typus) 59 Dart (PSAT), NA 20% (ALL)
2014 SAT 16 (SAT)
156 (SAT)
273
Vandeperre et al., SAT, Dart, Fin 3 (PSAT),
51 Atlantic Azores Archipelago Blue sharks (Prionace glauca) 37 (PSAT), 67% (PSAT) 0% (ALL)
2014 PSAT mount 33 (SAT)
877 (SAT)
Rodríguez-Cabello and S!anchez, El Cachucho (Le Danois Leafscale gulper sharks (Centrophorus
52 Atlantic PSAT 5 Dart 45 120 40% 20%
2014 Bank), Spain squamosus)
Holmes et al., Queensland & New SAT, Fin mount, 100%
53 Enviro Pacific Tiger shark (Galeocerdo cuvier) 18 4 408 17% (ALL)
2014 South Wales, Australia PSAT Dart (PSAT)
Hammerschlag Florida USA &
54 Pred/Prey Atlantic Tiger shark (Galeocerdo cuvier) SAT 31 Fin mount 8 844 NA NA
et al., 2015 Bahamas
55 Lea et al., 2015a Indian Seychelles Bull shark (Carcharhinus leucas) PSAT 1 Fin loop NA 133 0% 0%
Meekan et al., Christmas Island,
56 Indian Whale shark (Rhincodon typus) SAT 1 Fin Clamp NA 88 NA NA
2015 Australia
57 Rogers et al., 2015 Enviro Indian Southern Australia Shortfin mako (Isurus oxyrinchus) SAT 10 Fin mount 49 672 NA 0%
58 Teter et al., 2015 Atlantic Delaware Bay, USA Sand tiger (Carcharias taurus) PSAT 13 Dart 64 154 27% 15%
5

Sepulveda et al., Fisheries/ Southern California

59 Pacific Common thresher shark (Alopias vulpinus) PSAT 17 Dart 1 90 NA 6%
2015 PRM Bight
Carlisle et al.,
60 Pred/Prey Pacific Alaska, USA Salmon shark (Lamna ditropis) SAT 44 NA 365 NA NA NA
2015
Afonso and Hazin,
61 Enviro Atlantic Recife, Brazil Tiger shark (Galeocerdo cuvier) PSAT 20 Fin loop 7 159 60% 10%
2015
Ferreira et al.,
62 Enviro Indian Western Australia Tiger shark (Galeocerdo cuvier) SAT 11 Fin mount 7 517 NA 27%

Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

2015
Hutchinson et al., Fisheries/ Western & Central
63 Pacific Silky sharks (Carcharhinus falciformis) PSAT 28 NA 3 129 32% 7%
2015 PRM Pacific
Bonfil and O'Brien, Guadalupe Island,
64 Pacific White sharks (Carcharodon carcharias) SAT 1 NA NA 288 NA 0%
2015 Mexico
Campana et al., Greenland sharks (Somniosus
65 Atlantic Northeast Canada PSAT 15 Dart 48 350 67% 7%
2015 microcephalus)
Comfort and
66 Enviro Pacific Hawaii, USA Bluntnose sixgill (Hexanchus griseus) PSAT 6 NA 53 97 0% 17%
Weng, 2015
67 Bond et al., 2015 Pacific Astrolabe Reef, Fiji Silvertip (Carcharhinus albimarginatus) PSAT 1 Dart NA 12 100% NA
Carvalho et al., Fisheries/
68 Atlantic South Atlantic Blue sharks (Prionace glauca) PSAT 16 NA 40 205 NA 0%
2015 PRM
69 Lea et al., 2015b Enviro Atlantic Bermuda Tiger shark (Galeocerdo cuvier) SAT 24 NA 41 1101 0% 0%
PSAT, Dorsal loop, 70 (PSAT), 120(PSAT),
70 Doyle et al., 2015 Atlantic Ireland Blue sharks (Prionace glauca) 5 0% 0%
SAT Fin mount 74 (SAT) 314 (SAT)
Francis et al., Stewart Island, New PSAT, Dart, Fin
71 Pacific White sharks (Carcharodon carcharias) 31 4 (PSAT) 365 (PSAT) 10% (PSAT) NA
2015a Zealand SAT Mount
Fisheries/ Oceanic whitetip sharks (Carcharhinus
72 Tolotti et al., 2015 Atlantic Brazil PSAT 11 Fin loop 60 178 9% 27%
PRM longimanus)
Francis et al., Fisheries/
73 Pacific New Zealand Porbeagle sharks (Lamna nasus) PSAT 12 Dart 72 300 17% 17%
2015b PRM
(continued on next page)
 S. Renshaw et al.
Table 1 (continued )
Author(s) Study Ocean Region/Country Species Tagged Satellite Total Tag Min. Max. Premature Tag
Objectives Basin Tag Type Sample Attachment Tracking Tracking Pop-off Failures
Size Methods Days days

Jorgensen et al.,
74 Pacific California, USA White sharks (Carcharodon carcharias) PSAT 8 NA NA NA NA NA
2015
Filmalter et al., Fisheries/ Dart, Dorsal
75 Indian Western Indian Ocean Silky sharks (Carcharhinus falciformis) PSAT 13 NA NA NA NA
2015 PRM dart anchor
Tyminski et al., Yucatan Peninsula, Dart, Fin
76 Enviro Atlantic Whale shark (Rhincodon typus) PSAT 38 2 190 NA 18%
2015 Mexico & Florida USA mount
Tropical Northeast
77 Coelho et al., 2015 Atlantic Big eye thresher (Alopias superciliosus) PSAT 15 Dart 30 182 NA 20%
Atlantic
Comezzi et al.,
78 Pacific Maldives Whale shark (Rhincodon typus) SAT 2 Fin clamp 0 25 NA 50%
2016
Escalle et al., Fisheries/ Tropical eastern
79 Atlantic Whale shark (Rhincodon typus) PSAT 6 Dart 21 71 33% 17%
2016a PRM Atlantic
Bull shark (Carcharhinus leucas), Great
Graham et al., Florida USA & Fin mount,
80 MPA Atlantic hammerhead (Sphyrna mokarran), Tiger SAT 86 2 828 NA 0%
2016 Bahamas Dart
sharks (Galeocerdo cuvier)
Oceanic whitetip sharks (Carcharhinus
81 Howey et al., 2016 Enviro Atlantic Bahamas PSAT 16 Dart 22 335 NA 0%
longimanus)
Vandeperre et al., SAT, Dart, Fin
82 Enviro Atlantic Azores Archipelago Blue sharks (Prionace glauca) 31 NA NA NA NA
2016 PSAT mount
193
Sea of Hebrides, PSAT, 34 (SAT),
83 Witt et al., 2016 MPA Atlantic Basking sharks (Cetorhinus maximus) 83 Dart (PSAT), NA NA
Scotland SAT 64 (PSAT)
197 (SAT)
Blue sharks (Prionace glauca), Porbeagle
Campana et al., Fisheries/
84 Atlantic Nova Scotia, Canada sharks (Lamna nasus), Shortfin mako PSAT 131 Dart 0 356 NA 17%
2016 PRM
(Isurus oxyrinchus)
6

Fisheries/ Silky sharks (Carcharhinus falciformis),

85 Eddy et al., 2016 Pacific Ecuador PSAT 16 Dart 4 28 100% NA
PRM Scalloped Hammerheads (Sphyrna lewini)
Blue shark (Prionace glauca), Shortfin
Dart, Fin 179
Queiroz et al., Fisheries/ mako (Isurus oxyrinchus), Tiger shark PSAT, 4 (SAT), 7
86 Atlantic North Atlantic 113 mount, Fin (PSAT), NA 12% (ALL)
2016 PRM, Enviro (Galeocerdo cuvier), Great hammerhead SAT (PSAT)
loop 551 (SAT)
(Sphyrna mokarran)
Massachusetts, USA &

Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

New York, USA & Dart, Fin
87 Vaudo et al., 2016 Enviro Atlantic Shortfin mako (Isurus oxyrinchus) PSAT 8 7 181 63% 0%
Yucatan Peninsula, mount
Mexico
Cartamil et al., Southern California,
88 Pacific Common thresher shark (Alopias vulpinus) PSAT 23 Dart 6 373 NA 9%
2016 USA & Mexico
Norman et al., Dart, Fin
89 Indian Western Australia Whale shark (Rhincodon typus) SAT 14 9 261 NA 7%
2016 clamp
Fisheries/ New South Wales, Dusky shark (Carcharhinus obscurus),
90 Barnes et al., 2016 Pacific PSAT 16 Dart 0.1 59.7 75% 0%
PRM Australia Sandbar shark (Carcharhinus plumbeus)
Rodríguez-Cabello El Cachucho (Le Danois Leafscale gulper sharks (Centrophorus
91 Atlantic PSAT 9 NA 45 120 22% 11%
et al., 2016 Bank), Spain squamosus)
30%
SAT, (SAT),
92 Hearn et al., 2016 Pacific Galapagos Islands Whale shark (Rhincodon typus) 39 Dart 7 (SAT) 176 (SAT) NA
PSAT 50%
(PSAT)
Hammerschlag Great Barrier Reef
93 Pred/Prey Pacific Tiger shark (Galeocerdo cuvier) SAT 11 Fin mount 16 408 NA 18%
et al., 2016 Marine Park
Guttridge et al., Florida USA &
94 Atlantic Great Hammerhead (Sphyrna mokarran) PSAT 5 Dart 3 31 NA 0%
2017 Bahamas
Hammerschlag Western Cape, South
95 Pred/Prey Atlantic White shark (Carcharodon carcharias) SAT 37 Fin mount NA NA NA 8%
et al., 2017 Africa
(continued on next page)
 S. Renshaw et al.
Table 1 (continued )
Author(s) Study Ocean Region/Country Species Tagged Satellite Total Tag Min. Max. Premature Tag
Objectives Basin Tag Type Sample Attachment Tracking Tracking Pop-off Failures
Size Methods Days days

New Zealand and

96 Horton et al., 2017 Enviro Pacific White sharks (Carcharodon carcharias) SAT 5 Fin mount NA NA NA NA
Guadalupe
Queiroz et al., English Channel, & Basking sharks (Cetorhinus maximus),
97 Enviro Atlantic PSAT 68 Fin loop, Dart 7 544 NA NA
2017 Scotland Blue sharks (Prionace glauca)
Gulf of Mexico &
98 Vaudo et al., 2017 Enviro Atlantic Shortfin mako (Isurus oxyrinchus) SAT 32 Fin mount 78 527 NA 19%
Caribbean Sea
Shipley et al., Fisheries/
99 Atlantic Bahamas Cuban dogfish (Squalus cubensis) PSAT 8 Fin loop 5 14 63% 13%
2017a PRM
100 Lynch et al., 2017 Tag effects Atlantic Virginia, USA Sandbar shark (Carcharhinus plumbeus) PSAT 18 Dorsal loop NA NA NA NA
Shipley et al., Caribbean Reef shark (Carcharhinus
101 MPA Atlantic Bahamas PSAT 11 Fin loop 30 243 NA 0%
2017b perezi)
Doherty et al., PSAT,
102 Atlantic Isle of Man, Scotland Basking sharks (Cetorhinus maximus) 70 NA 365 (ALL) NA NA NA
2017a SAT
U.S. Palmyra Atoll
Grey reef shark (Carcharhinus
103 White et al., 2017 MPA Pacific National Wildlife SAT 11 Fin mount 32 375 NA 45%
amblyrhynchos)
Refuge
Fisheries/ Great Australian Bight,
104 Rogers et al., 2017 Indian School shark (Galeorhinus galeus) PSAT 10 Dart 5 44 100% 0%
PRM Southern Australia
105 Hueter et al., 2017 Enviro Atlantic Gulf of Mexico, Cuba Longfin mako (Isurus paucus) PSAT 2 Dart 8 21 0% 0%
106 Biais et al., 2017 Atlantic Bay of Biscay Porbeagle sharks (Lamna nasus) PSAT 13 Dart 128 365 15% 8%
Doherty et al., Sea of Hebrides, PSAT,
107 MPA Atlantic Basking sharks (Cetorhinus maximus) 36 Dart NA NA NA NA
2017b Scotland SAT
Ramírez-Macías Gulf of California,
108 Enviro Pacific Whale shark (Rhincodon typus) PSAT 17 Dart 14 134 100% 53%
et al., 2017 Mexico
Indian, Mozambique & Oceanic whitetip sharks (Carcharhinus
7

109 Tolotti et al., 2017 Enviro PSAT 6 Fin loop, Dart 100 178 0% 0%
Atlantic Atlantic Equator longimanus)
110 Ryan et al., 2017 Enviro Pacific Galapagos Islands Whale shark (Rhincodon typus) SAT 27 Dart 7 176 NA NA
Yucatan Peninsula,
Fisheries/
111 Byrne et al., 2017 Atlantic Mexico & Maryland, Shortfin mako (Isurus oxyrinchus) SAT 46 Fin mount 81 754 NA 13%
PRM
USA.
Acu!na-Marrero MPA, Pred/ Galapagos Marine
112 Pacific Tiger shark (Galeocerdo cuvier) SAT 20 Fin mount 14 333 NA 11%
et al., 2017 Prey Reserve

Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

Scallopped hammerhead sharks (Sphyrna
113 Spaet et al., 2017 Indian Red Sea PSAT 1 Dart NA 183 0% NA
lewini)
125
Fernando de Noronha PSAT, Fin loop, Fin
114 Afonso et al., 2017 Atlantic Tiger shark (Galeocerdo cuvier) 10 10 (PSAT) (PSAT), 50% (PSAT) 33% (SAT)
Archipelago, Brazil SAT mount
207 (SAT)
180
Robinson et al., PSAT, 1 (SAT), 10 23%
115 Indian Arabian Gulf, Qatar Whale shark (Rhincodon typus) 59 Dart (PSAT), 28% (PSAT)
2017 SAT (PSAT) (PSAT)
227 (SAT)
116 Niella et al., 2017 Enviro Atlantic Recife, Brazil Bull shark (Carcharhinus leucas) PSAT 1 Fin loop NA 75 0% 0%
330 31%
Skomal et al., PSAT, Dart, Fin 5 (PSAT),
117 Atlantic Massachusetts, USA White sharks (Carcharodon carcharias) 32 (PSAT), NA (PSAT),
2017 SAT mount 987 (SAT)
1567 (SAT) 0% (SAT)
Reynolds et al.,
118 MPA, Enviro Indian Western Australia Whale shark (Rhincodon typus) SAT 29 Dart 7 457 NA 0%
2017
SAT, Fin mount,
119 Coffey et al., 2017 Enviro Pacific Alaska, USA Salmon shark (Lamna ditropis) 96 5 (ALL) 366 (ALL) NA 30% (ALL)
PSAT Dart
Massachusetts, USA,
120 Howey et al., 2017 Enviro Atlantic Blue sharks (Prionace glauca) PSAT 33 Dart 4 273 NA 30%
Bahamas
King and Surry, British Columbia,
121 Enviro Pacific Bluntnose sixgill (Hexanchus griseus) PSAT 8 Dart 6 274 25% 25%
2017 Canada
122 Atlantic Gulf of Mexico Great hammerheads (Sphyrna mokarran) 3 66% (PSAT)
(continued on next page)
 S. Renshaw et al.
Table 1 (continued )
Author(s) Study Ocean Region/Country Species Tagged Satellite Total Tag Min. Max. Premature Tag
Objectives Basin Tag Type Sample Attachment Tracking Tracking Pop-off Failures
Size Methods Days days

0%
Drymon and Fisheries/ PSAT, Dart, Fin 0 (SAT), 20 30 (PSAT),
(PSAT),
Wells, 2017 PRM SAT mount (PSAT) 41 (SAT)
66% (SAT)
Fisheries/
123 Coelho et al., 2017 Atlantic Northeast Atlantic Shortfin mako (Isurus oxyrinchus) PSAT 10 NA NA NA NA NA
PRM
Fisheries/ Southern California, PSAT,
124 Benson et al., 2018 Pacific White sharks (Carcharodon carcharias) 37 NA 2 (PSAT) 270 (PSAT) NA 0% (PSAT)
PRM USA & Baja Mexico SAT
125 Braun et al., 2018 Enviro Atlantic Massachusetts, USA Basking sharks (Cetorhinus maximus) PSAT 57 Dart 79 424 14% 21%
Corrigan et al.,
126 Indian Southern Australia Shortfin mako (Isurus oxyrinchus) SAT 13 Fin mount 249 672 NA 0%
2018
127 Dewar et al., 2018 Pacific California, USA Basking sharks (Cetorhinus maximus) PSAT 4 Dart 9 240 50% 0%
Port MacDonnell, Blue sharks (Prionace glauca), Common
128 Heard et al., 2018 Indian PSAT 5 Dart 50 184 33% 0%
South Australia thresher shark (Alopias vulpinus)
Scallopped hammerhead sharks (Sphyrna
129 Wells et al., 2018 Enviro Atlantic Gulf of Mexico SAT 33 Fin mount 5 479 NA 0%
lewini)
Rohner et al.,
130 Enviro Indian Mozambique Whale shark (Rhincodon typus) SAT 15 Dart 2 88 NA 0%
2018
131 Fontes et al., 2018 Atlantic Azores Archipelago Blue sharks (Prionace glauca) PSAT 1 Fin Clamp NA NA NA NA
PSAT, Dart, Fin 34 (PSAT),
132 Hueter et al., 2018 Atlantic Cuba Silky sharks (Carcharhinus falciformis) 3 23 (PAT) 67% (PSAT) 0% (ALL)
SAT mount 240 (SAT)
Greenland sharks (Somniosus
133 Hussey et al., 2018 Arctic Nunavut, Canada PSAT 5 Fin mount 34 45 NA 0%
microcephalus)
Guzman et al.,
134 Pacific Panama Whale shark (Rhincondon typus) SAT 1 Dart NA 841 0% 0%
2018
South Africa,
8

135 Daly et al., 2018 MPA Indian Tiger shark (Galeocerdo cuvier) SAT 26 Fin mount 10 375 NA 3%
Mozembique
Diamant et al.,
136 Indian Madagascar Whale shark (Rhincodon typus) SAT 8 Dart 9 199 NA 0%
2018
Santos and Fisheries/
137 Atlantic Northeast Atlantic Smooth Hammerhead (Sphyrna zygaena) PSAT 8 Dart 6 159 50% 13%
Coelho, 2018 PRM
Fisheries/
138 Lea et al., 2018 PRM, MPA, Atlantic Bermuda Tiger shark (Galeocerdo cuvier) SAT 24 NA 41 1101 NA NA

Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

Enviro
Sulu and Bohol Seas,
139 Araujo et al., 2018 Pacific Whale shark (Rhincodon typus) SAT 17 Dart 6 126 NA 0%
Philippines
Great Hammerhead (Sphyrna mokarran),
Fisheries/ Florida USA & Dart, Fin
140 Calich et al., 2018 Atlantic Bull sharks (Carcharhinus leucas), Tiger SAT 96 NA NA NA NA
PRM, Enviro Bahamas mount
shark (Galeocerdo cuvier)
Musyl and Gilman, Fisheries/ Marshall Islands & Blue sharks (Prionace glauca), Silky sharks
141 Pacific PSAT 83 Dart NA NA NA NA
2018 PRM Palau (Carcharhinus falciformis)
Rodríguez-Cabello
142 Atlantic Cantabrian Sea, Spain Bluntnose sixgill (Hexanchus griseus) PSAT 1 NA NA 84 100% 0%
et al., 2018
Nyk!anen et al.,
143 Atlantic Cork, Ireland Blue sharks (Prionace glauca) PSAT 2 Fin loop NA NA NA NA
2018
Andrzejaczek Oceanic whitetip sharks (Carcharhinus
144 Enviro Atlantic Bahamas PSAT 16 Dart 22 335 NA NA
et al., 2018 longimanus)
SAT, Fin mount,
145 Curtis et al., 2018 Atlantic New York, USA White sharks (Carcharodon carcharias) 10 6 (ALL) 305 (ALL) NA NA
PSAT Dart
SAT, Fin mount,
146 Gaube et al., 2018 Enviro Atlantic Massachusetts, USA White sharks (Carcharodon carcharias) 2 NA NA NA NA
PSAT Dart
147 Meyer et al., 2018 Pacific Hawaii, USA Tiger shark (Galeocerdo cuvier) SAT 28 Fin mount 7 595 NA 11%
Fisheries/ Tropical eastern
148 Escalle et al., 2018 Atlantic Whale shark (Rhincodon typus) PSAT 11 Dart 3 150 27% 9%
PRM Atlantic
(continued on next page)
 S. Renshaw et al.
Table 1 (continued )
Author(s) Study Ocean Region/Country Species Tagged Satellite Total Tag Min. Max. Premature Tag
Objectives Basin Tag Type Sample Attachment Tracking Tracking Pop-off Failures
Size Methods Days days

149 Santos et al., 2018 Atlantic Various Shortfin mako (Isurus oxyrinchus) PSAT 34 Dart 1 121 NA 6%
Barkley et al.,
150 MPA Indian Kenya Tiger shark (Galeocerdo cuvier) SAT 5 Fin mount 35 235 NA 20%
2019
Bradley et al., Fisheries/ Gray reef sharks (Carcharhinus
151 Pacific Marshall Islands PSAT 15 NA NA NA NA NA
2019 PRM, MPA amblyrhynchos)
Yucatan Peninsula,
152 Byrne et al., 2019 Enviro Atlantic Mexico & Maryland, Shortfin mako (Isurus oxyrinchus) SAT 39 Fin mount 49 527 NA 0%
USA.
Johnston et al.,
153 Atlantic Northern Ireland Basking sharks (Cetorhinus maximus) SAT 1 NA NA 33 NA 0%
2019
California,
Maxwell et al., Washington, USA &
154 Enviro Pacific Blue sharks (Prionace glauca) SAT 54 Fin mount 25 614 NA NA
2019 British Columbia,
Canada
Thorburn et al.,
155 Atlantic Scotland Tope sharks (Galeorhinus galeus) PSAT 1 Dart NA 180 0% 0%
2019
156 Francis et al., 2019 Enviro Pacific New Zealand Shortfin mako (Isurus oxyrinchus) SAT 14 Fin mount 34 588 0% 0%
Western & Central
Hutchinson et al., Fisheries/ PSAT, Dart, Fin
157 Pacific Pacific, Easter Tropical Silky sharks (Carcharhinus falciformis) 29 2 (PSAT) 129 (PSAT) NA NA
2019 PRM SAT mount
Pacific
Whale shark (Rhincodon typus), Tiger
shark (Galeocerdo cuvier), Shortfin mako SAT,
158 Rooker et al., 2019 Atlantic Gulf of Mexico 133 NA NA NA NA NA
(Isurus oxyrinchus), Scallopped PSAT
hammerhead sharks (Sphyrna lewini)
White sharks (Carcharodon carcharias),
9

Fisheries/ Shortfin mako (Isurus oxyrinchus), Blue PSAT,

159 White et al., 2019a Pacific See Block et al., 2011. 309 NA NA NA NA NA
PRM, MPA sharks (Prionace glauca), Salmon shark SAT
(Lamna ditropis)
SAT,
160 White et al., 2019b Enviro Pacific California Bight, USA White sharks (Carcharodon carcharias) 18 Fin mount 4 (SAT) 544 (SAT) NA 5% (SAT)
PSAT
Silky Sharks (Carcharhinus falciformis),
Carlisle et al., British Indian Ocean
161 MPA Indian Silverstip sharks (Carcharhinus PSAT 9 Dart 86 269 67% NA
2019 Territory

Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

albimarginatus)
New South Wales &
162 Bruce et al., 2019 Pacific White sharks (Carcharodon carcharias) SAT 43 Fin mount 1 275 NA 0%
Victoria, Australia
Schaefer et al., Fisheries/
163 Pacific Costa Rica & Ecuador Silky sharks (Carcharhinus falciformis) PSAT 40 Dart 1 180 55% 5%
2019 PRM
McMillan et al.,
164 Enviro Pacific Great Australian Bight School shark (Galeorhinus galeus) PSAT 13 Dart 3 120 46% 8%
2019
Southern California Blue shark (Prionace glauca), Shortfin
165 Nosal et al., 2019 Pacific PSAT 10 Dart 13 86 30% 0%
Bight mako (Isurus oxyrinchus)
Atlantic,
Queiroz et al., Fisheries/ PSAT, Fin mount,
166 Pacific, Various 23 shark species (see study for full list) 1804 NA NA NA 7% (ALL)*
2019 PRM, Enviro SAT Dart
Indian
180 20%
PSAT, Dart, Fin 16 (PSAT),
167 Braun et al., 2019 Enviro Atlantic New England, USA Blue sharks (Prionace glauca) 15 (PSAT), NA (PSAT),
SAT mount 54 (SAT)
288 (SAT) 0% (SAT)
Cochran et al., SAT,
168 Indian Saudi Arabia Whale shark (Rhincodon typus) 38 Dart 16 (ALL) 281(ALL) NA 18% (ALL)
2019 PSAT
Salinas-de-Le!
on Pred/Prey,
169 Pacific Galapagos Islands Tiger shark (Galeocerdo cuvier) SAT 23 Fin mount NA NA NA 9%
et al., 2019 MPA
Doherty et al., PSAT,
170 Atlantic Scotland Basking sharks (Cetorhinus maximus) 32 Dart 10 (ALL) 292 (ALL) NA NA
2019 SAT
(continued on next page)
 S. Renshaw et al.
Table 1 (continued )
Author(s) Study Ocean Region/Country Species Tagged Satellite Total Tag Min. Max. Premature Tag
Objectives Basin Tag Type Sample Attachment Tracking Tracking Pop-off Failures
Size Methods Days days

0%
Saint Peter and Saint
Bezerra et al., Scallopped hammerhead sharks (Sphyrna PSAT, (PSAT),
171 MPA Atlantic Paul Archipelago, 7 Dart 5 (PSAT) 120 (PSAT) 33% (PSAT)
2019 lewini) SAT 100%
Brazil
(SAT)
Sepulveda et al., Fisheries/ Southern California
172 Pacific Big eye thresher (Alopias superciliosus) PSAT 14 Dart 2 32 0% 0%
2019 PRM Bight
173 Araujo et al., 2019 Pacific Honda Bay, Philippines Whale shark (Rhincodon typus) PSAT 10 Dart 61 200 NA 50%
Sulikowski et al., Fisheries/
174 Atlantic North Carolina, USA Dusky shark (Carcharhinus obscurus) PSAT 50 Dart 1 28 62% 14%
2020 PRM
3%
Fisheries/ Cape Verde Blue shark (Prionace glauca), Shortfin PSAT, Dart, Fin 3 (PSAT), 88 (SAT),
175 Coelho et al., 2020 Atlantic 30 4%(PSAT) (PSAT),
PRM Archipelago mako (Isurus oxyrinchus) SAT mount 22 (SAT) 121 (PSAT)
0% (SAT)
10

Guadalupe Island,
Nasby-Lucas and
176 Tag effects Pacific Mexico & Southeast White sharks (Carcharodon carcharias) SAT 12 Fin mount 0 2706 NA NA
Domeier, 2020
Farallon Island, USA

Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

S. Renshaw et al. Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

Fig. 1. Fig. 1 Distribution of sample sizes across all studies. The maximum sample size from reviewed studies was 1804, however, this outlier was removed as the
study used data from multiple sources (denoted by *).

impact of satellite tags on welfare (i.e., fin damage) and to test tag (Hammerschlag et al., 2014).
application methods (n → 3; 2%). Tags were administered to investigate
subtopics including environmental or oceanographic drivers of move- 3.1.2. Environmental and oceanographic drivers of movement
ment (n → 38; 22%), species interactions with fisheries and in- To better understand shark habitat use, researchers have coupled
vestigations of post-release mortality/survival (n → 33; 19%), marine satellite telemetry data with a suite of environmental parameters to
protected area (MPA) effectiveness (n → 17; 10%), predator and prey investigate potential drivers of movements. Sea surface temperature
interactions (n → 9; 5%), movement energetics and thermoregulation (n (SST; see Lea et al., 2018; Coffey et al., 2017; Tolotti et al., 2017;
→ 5; 3%). Testable hypotheses were explored in 36 reviewed studies, 10 Howey-Jordan et al., 2013; Thums et al., 2013), chlorophyll-a concen-
(28%) of which were directly related to conservation or fisheries man- trations (i.e.., areas of high primary production; see Rohner et al., 2018),
agement. Hypotheses were tested with satellite telemetry most often in depth or pressure (see Nyk! anen et al., 2018; Wells et al., 2018) and
relation to shark's potential association with fishing gear (Hutchinson bathymetric slope (see Rogers et al., 2015) have been identified as
et al., 2019), survival outcomes (Musyl and Gilman, 2018), and deter- important extrinsic predictors for shark habitat use. Tagging studies
mining critical habitat use in relation to conservation (Domeier and using environmental and oceanographic data to provide context to
Nasby-Lucas, 2013). Tagging was used in service of informing conser- observed movement patterns were able to inform successful conserva-
vation and management objectives (including fisheries management tion recommendations. In these instances, studies determined drivers of
procedures) in 26% of all studies (n → 46). Recommendations for con- shark movements using habitat models and generated advice based on
servation or management of tagged species resulted from 23% of all the outcomes. Shark habitat modelling has expanded in recent decades
studies (n → 40), while just 13% (n → 23) were able to directly evaluate thus increasing researchers' ability to unravel complex relationships
the efficacy of conservation or fisheries management strategies using between covariates and telemetry data. Details on the types of models
satellite telemetry. used is available in the Supplementary Material.

3.1.1. Tag effects on wellbeing 3.1.3. Predator-prey interactions

Prior to 2010, no published reports had investigated the impacts of Although infrequent (Table 1), shark predator-prey studies have
satellite tags on the histology and vitality of dorsal fins of sharks helped to address shark distribution and behaviour in relation to prey
(Hammerschlag et al., 2011b). In the last decade three studies investi- movements and availability (Hammerschlag et al., 2015; Acu” na-Marrero
gated the impacts of satellite tagging on sharks. Two monitored the ef- et al., 2017; Lea et al., 2015a; Carlisle et al., 2015). For example, Lea
fects of dorsal fin mounted SPOT tags on white sharks (Jewell et al., et al. (2015a) found roughly ~37% overlap of tiger sharks tracks with
2011; Nasby-Lucas and Domeier, 2020), each concluding that despite satellite-tagged juvenile loggerhead turtles (Caretta caretta; McClellan
evidence of physical trauma, scars, and biofouling (as a result of satellite and Read, 2007) and validated the observed patterns with tiger shark
tag application), the sharks' behaviour was unaltered and their surviv- stomach contents sampled from commercial longlines. Further, preda-
ability likely unimpaired. A third study concluded that dorsal looped tion events (Horning and Mellish, 2014) and relative predation risk to
PSAT tags did not significantly increase the metabolic costs or alter the prey (Hammerschlag et al., 2017) have been documented using satellite
swimming kinematics of sandbar sharks (Lynch et al., 2017). These telemetry. Satellite tagging studies can expand knowledge of predator
studies were not considered to have produced conservation or man- and prey relationships, anti-predator behaviour (Hammerschlag et al.,
agement outcomes given the evidence that tags have little impact on the 2012b) and stress responses, as well as hunting and foraging strategies
survivability of species. The trade-offs between satellite tag data, public when coupled with additional ecological and physiological information,
perception, and animal welfare have since been discussed such as stable isotope analysis (Salinas-de-Le# on et al., 2019; Carlisle

11
S. Renshaw et al. Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

et al., 2015), hormone analysis, energetic models, and population met- attachment methods, (i.e., tag anchor failure), early triggered detach-
rics (Barnett et al., 2017). Despite these opportunities, there remains a ment (i.e., constant pressure readings from mortality events; Eddy et al.,
limited body of work investigating sharks' role in ocean food webs as 2016; Bezerra et al., 2019; Holmes et al., 2014; Rodríguez-Cabello et al.,
both predator and prey. 2018; Bond et al., 2015; Hoffmayer et al., 2014; Rogers et al., 2017).
Musyl et al. (2011) found that tag retention rates were affected by shark
behaviours, environment, biofouling, and infection, as well as tag
3.2. Tag types, duration and deployment characteristics such as manufacturer and year. One reviewed study
noted that female sand tiger shark behaviour may increase their rate of
Tracking durations ranged from 0.1 (2.4 h) to 2706 days, more than tag shedding compared to their tagged male counterparts (Teter et al.,
twice the length recorded prior to 2010 (1335 days). Overall, tracking 2015).
times have stayed relatively the same as before, however minimum time Most studies reviewed used PSAT tags (74%, n → 130) while less than
has increased (minimum was 43 compared to 25 days previously; p ω half used SAT tags (48%, n → 84), both of which were applied more
0.05, whereas maximum was 269 compared to 201 days; p ε 0.05). On frequently than during previous review (Table 1; Fig. 2). Similar to the
average, SAT tags provided longer minimum and maximum tracks previous review, many studies opted to use SPOT tags from Wildlife
compared to PSAT tags (p ω 0.05). A SPOT tag was responsible for the Computers Inc (see Hammerschlag et al., 2011a for more information).
overall longest tracking duration of 2706 days (Nasby-Lucas and Of the reviewed studies, 22% (n → 39) deployed both PSAT and SAT tags
Domeier, 2020). In all cases, tracking durations currently reviewed were matching the previous review's findings. Similar proportions of tag types
longer than those previously reported, suggesting tag technology and were used in service of conservation objectives (see Supplementary
refined application improved the longevity of individual deployments. Materials). In some cases, both tag types were affixed to an animal,
In service of conservation, across both PSAT and SAT tags the minimum referred to as ‘double tagging’ (Coffey et al., 2017; Drymon and Wells,
tracking time was 19 days on average while all other statistics are 2017), thus combining the advantages of both technologies by collecting
reflective of the overall findings. More details can be found in supple- complementary data of depth and temperature (PSAT tag) and more
mentary materials. accurate spatial locations (SAT tag). Sharks tagged with both SAT and
In total, 74 studies reported premature tag detachment. The average PSAT tags have allowed researchers to test the accuracy of location es-
rate of early pop-off was 45%, outperforming estimates of 65% prema- timates between tags, providing better data quality and larger data
ture detachment of shark tags previously determined by Musyl et al. quantities (Hueter et al., 2013). Several studies (41%, n → 73) utilized
(2011). Of the reviewed studies for which tag failure rates could be alternative methods to complement satellite telemetry data, such as
determined (n → 136), 57% (n → 77) reported malfunctions resulting in conventional identification tags (n → 27) for mark-recapture, photo-
no data transmission from at least one deployed tag. Average failure rate graphs (n → 20) for resighting, as well as acoustic telemetry (n → 25) for
of 12% across all studies was consistent with previous review findings further movement data, and archival tags (n → 3) for environmental
(10%). These rates are likely attributedto novel experimental tag types, information storage. Use of anti-biofouling materials was noted in 19
application methods, or new shark species being tagged, thus deviating studies. Disclosure of these materials and their application methods is
from more refined tagging methods. Premature tag release and tag inconsistent across studies. For transparency, we suggest that best
failure may occur for several reasons, including improper or insecure tag

Fig. 2. Fig. 2 Most common reported satellite tag types used for shark tracking (not to scale). Wildlife Computers Inc. provided most tags used across studies
including the MK10, miniPAT, mark-report (mrPAT) tags (Box A); and SPOT and SPLASH tags with Fastloc technology incorporated.(Box C) (from www.wildl
ifecomputers.com/ accessed 27 March 2020). X-tags, made by Microwave Telemetry (Box B), were designed for smaller marine species with smaller dimensions
and as such, were used to tag all of the small-bodied shark studied since 2010 (Squalidae; Carlson et al., 2014a; Sulikowski et al., 2010; Shipley et al., 2017a; from
www.microwavetelemetry.com/x_tag, accessed 27 March 2020). Photos 1 and 2 show fin-mounted and towed (dart) style tags respectively.

12
S. Renshaw et al. Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

practice for future telemetry surveys is to list their anti-biofouling ma- (IUCN, 2020). While a wider representation of tagged species has been
terials to better understand its worth and the impacts specific materials achieved, 50% of satellite tagging efforts are still focused on just 5
have (if any) on the animals themselves. Additional details about tag species (Fig. 3).
application are provided in the Supplementary Material. Satellite telemetry has been most frequently used to study wide-
Within the last 10 years the application of Fastloc GPS technology (i. ranging, highly mobile species, including whale sharks (n → 25
e., www.wildlifecomputers.com) in shark tagging has emerged (n → 3), studies), tiger sharks (n → 25), white sharks (n → 24), and blue sharks (n
with its absence and potential noted in the original review. The potential → 24; Prionace glauca) (Fig. 3). White sharks continue to be a primary
of Fastloc GPS for pelagic fish tracking was first explored by Sims et al. tagging subject with the total number of studies on white sharks having
(2009) documenting large scale movements of ocean sunfish (Mola more than doubled from the original review (n → 10 previously). In the
mola), and the ability of Fastloc GPS to produce nearly four times the last decade, these studies have facilitated investigations of regional
amount of location estimates compared to Argos transmitters. Fastloc movements and long-distance migrations (Francis et al., 2015a; Bonfil
therefore, may help overcome some of the traditional disadvantages of and O'Brien, 2015; Duffy et al., 2012) with studies identifing essential
satellite tags by requiring less surface time (as little as 10 ms) to generate habitats for conservation including mating, pupping (Domeier and
accurate position fixes (Ferreira et al., 2015; Hammerschlag et al., Nasby-Lucas, 2012, 2013; Domeier et al., 2012), and nursery grounds
2011b; Dujon et al., 2014). Additional details about the application of (Curtis et al., 2018; White et al., 2019b; Lyons et al., 2013), including the
Fastloc GPS is provided in the Supplementary Materials. first attempt at studying white shark energetics using satellite tags (Del
Raye et al., 2013).
3.3. Species Pelagic species such as blue sharks, shortfin makos (Isurus oxy-
rinchus), silky sharks (Carcharhinus falciformis, n → 13) and threshers
Since 2010, 35 shark species have been satellite tagged, from 12 (bigeye and common; Alopias superciliosus and Alopias vulpinus, respec-
families and 5 orders (Table 1; Fig. 3), more than double the species tively, n → 9 together) constituted a large proportion of tagged sharks
recorded originally(n → 17). Some of the newly tagged species are of (Fig. 3). Most notably we found a substantial increase in studies con-
high conservation priority, including the great hammerhead (Sphyrna ducted on shortfin makos (n → 20, versus n → 2 previously). Despite their
mokarran, n → 8 studies) and pelagic thresher shark (Alopias pelagicus, n Endangered status, longfin mako sharks (Isurus paucus, n → 2) remain
→ 1 study). Over half of the tagged species (62%, Fig. 3) were catego- poorly studied in comparison to the closely related shortfin mako shark
rized by the IUCN Red List as threatened, comprising 34% Vulnerable (n (Hueter et al., 2017). Fisheries management outcomes were provided
→ 12), 17% Endangered (n → 6), and 11% Critically Endangered (n → 4) most often for silky sharks (n → 9 studies) and blue sharks (n → 8)

Fig. 3. Fig. 3 Number of studies by species tagged between 2010 and 2020. Each species current global status per the IUCN RedList (IUCN, 2020, accessed August 14,
2020) is indicated by colour, with the percentage of studies that tracked species under each listing displayed in the upper right corner. An asterisk (*) denotes the
species for which tracking studies' outcomes have directly contributed to conservation and management.

13
S. Renshaw et al. Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

frequently in relation to their spatial overlap and risk of interactions the aforementioned obstacles for affected species. These methods
with fisheries and fish aggregating devices (FADs) as a means of deter- coupled with the increased availability of smaller tag technology have
mining best practice for mitigating bycatch (Schaefer et al., 2019; resulted in early successes with increased post-release survival,
Queiroz et al., 2012, 2016, 2017). providing preliminary insights into vertical and horizontal distributions
Species rarely tagged were also present in reviewed studies, under- of a small subset of deep-water species (see referenced studies for further
scored by the adaptation of satellite tags for tracking small bodied and details).
deep-water sharks like dogfish (n → 3; family Squalidae; Shipley et al., Studies directly informed conservation and management of 21 tag-
2017a; Carlson et al., 2014a; Sulikowski et al., 2010); these species, and ged species. Overall, 48% of publications resulting in direct conserva-
others like them, were largely absent from the collection of studies tion/management outcomes focused on just four species: silky sharks,
previously reviewed. For example, Hussey et al. (2018) provided the tiger sharks, blue sharks, and great hammerheads. Other study species
first near-real time tracks over extended periods for Greenland sharks for conservation driven tracking included large, frequently tagged,
(Somniosus microcephalus, n → 3), a large bodied, deep-dwelling shark. charismatic species such as whale sharks and basking sharks. Silky
There are several challenges for tagging and tracking deep-water species sharks and blue sharks were also tagged due to conservation concern due
such as their vulnerability to capture/handling stress, rapid temperature to their occurrence as bycatch in commercial fisheries.
changes, post-release predation, and sampling gear limitations. How-
ever, improvements to tagging procedures such as body size selectivity
to maintain suitable tag to body weight ratio (American Society of 3.4. Geographic location
Ichthyologists and Herpetologists, 2003; Carlson et al., 2014a), and
release cages or free diver assisted swimming to prevent predation Reviewed tagging studies span nearly every ocean basin, including
during initial descent (Shipley et al., 2017a) have been used to overcome the Arctic Ocean (n → 2; 1%; Fig. 4) which was absent in the original
review. These studies took place in Norway and Canada tracking

Fig. 4. Fig. 4 Global distribution of tagging hotspots. Top panel shows tagging location and sample sizes from the presently reviewed literature, using clustered
analysis. Bottom left panel shows circle sizes which denote the number of tagged individuals at each tagging location, as well as a coloured scale to show hotspots
globally. Coloured scale indicated results from hotspot analysis whereby red indicates areas of high numbers of sharks tagged (with 99% confidence) and blue
indicated areas of low numbers of sharks tagged (with 99% confidence). Areas of no significance are transparent. Bar graph in the bottom right panel shows the
number of studies taken place in each ocean basin. Studies with tagging locations in multiple locations, across oceans were included in the count for each ocean and
marked in respective locations on the map. Map constructed using QCIS (3.18.3). (For interpretation of the references to colour in this figure legend, the reader is
referred to the web version of this article.)

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S. Renshaw et al. Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

Greenland sharks (Fisk et al., 2012; Hussey et al., 2018). Challenges of is often not enough to change policy, and perhaps more importantly,
accessibility, poor weather conditions, and fewer shark species present human behaviour. Although most studies mention the importance of
at extreme latitudes contribute to the relative absence of studies from tagging for conservation, putting findings directly into conservation
these areas. However, tagging initiatives have demonstrated movements action has been less frequent.
of pelagic species, such as white sharks, into sub-Antarctic waters and The most direct investigation of shark conservation efficacy using
their affinity for travel along the circumpolar front (Queiroz et al., satellite tags was the evaluation of MPAs. Proposal, designation, and
2019), thus highlighting the need for broadened spatial coverage of evaluation of various MPAs has been informed by shark space use pat-
fisheries-independent data like tagging. terns determined using satellite telemetry (e.g., Doherty et al., 2017b;
Most reviewed studies took place in the Atlantic Ocean, accounting Graham et al., 2016; Lea et al., 2018; Gallagher et al., 2020; Daly et al.,
for 48% (n → 85) of tagging locations, a substantial increase in number 2018). In total, 10 different protected areas were evaluated, which
of studies than previously reported (n → 19) (Fig. 4). In the original included both MPAs as well as EEZs with shark fishing restrictions.
review nearly 50% of studies originated in the Pacific Ocean (n → 23), However, several studies found that conservation measures such as
compared to 36% reviewed here (n → 64). Despite the shift from Pacific MPAs and regulations within geo-political boundaries (i.e., state waters
to Atlantic predominance among tagging locations, these ocean basins or EEZs) did not cover a significant portion of shark distribution (Gra-
together continue to account for ω80% of total shark tagging sites. Some ham et al., 2016) or provide sufficient protections from high-risk ac-
areas were identified as ‘tagging hotspots’, where a significant number tivities such as commercial fishing (Queiroz et al., 2012, 2017),
of individuals were tagged relative to other areas (Fig. 4) and include particularly to highly mobile, pelagic species. Generally, EEZs in the
both the central Atlantic and Pacific coasts of North America (99% Southern USA and Caribbean were found to provide inad-
confidence interval). Frequent tagging locations included inshore areas equateprotection to sharks, highlighting the need for transboundary
off southern California and northern Mexico, as well as southern Florida management to ensure sharks are protected throughout their entire
and the Bahamas. Overall, the diversity in satellite tagging locations has range both in and outside of these areas (Lea et al., 2015a; Graham et al.,
expanded, however tagging studies remain scarce in several areas. 2016). MPAs that provided effective protection to sharks were large, old,
Notably there are a lack of published tagging literature throughout the and well protected (Carlisle et al., 2019; Acu!na-Marrero et al., 2017). In
Northwest Pacific (with the exception of tracking data from Japan 2015, it was determined that 29% of all ocean conservations areas had
included in Queiroz et al., 2019), particularly the east China Sea, as well been established primarily for the protection of shark and ray species
as northeast Indian Ocean in the Bay of Bengal despite a major fishing (Davidson and Dulvy, 2017). In order to understand the true efficacy of
presence throughout these regions. It was suggested by Queiroz et al. MPAs, Carlisle et al. (2019) recommended comprehensive tagging of
(2019) that fisheries independent data in these areas, such as satellite species and individuals across: “seasons, years, and ontogenetic stages,
tracking data, may exist but is simply inaccessible or unpublished, in order to fully characterize their spatial ecology”.
however, further investigation was beyond the scope of this review. Satellite tags are advantageous for measuring the efficacy of large,
Studies addressing shark conservation and management were restricted remote protected areas as data can be recorded independently of geo-
to the Atlantic (n → 21), Pacific (n → 15), and Indian (n → 9) oceans. political zones, and in areas beyond national jurisdiction (White et al.,
However, throughout these regions conservation and management were 2017; MacKeracher et al., 2019; Sequeira et al., 2018). Emphasis in
determined for a range of locations, concentrating most heavily on the recent years has been placed on amalgamating sources of tracking data
Galapagos (n → 4), and other islands and archipelagos (i.e., Hawaii, in order to optimize tagging outcomes and produce results on a global
Bahamas, Madagascar, Seychelles, Cabo Verde, etc.). In most cases, scale (see Sequeira et al., 2018; Harcourt et al., 2019, and Hays et al.,
tagging locations were chosen in order to determine the level of pro- 2019). Multi-year, continuous studies are beneficial for overcoming the
tection afforded to sharks within a defined area such as a country's often-limited duration of tagging studies (usually less than one year), by
Exclusive Economic Zone (EEZ) or an MPAs. This demonstrates that creating pooled resources that can be used to generate long-term studies
conservation efforts have been biased towards locations with existing of seasonality and species movements, migrations, residency and/or
spatial protections. While this presents opportunities to evaluate their aggregations (‘hotspots’) on a global scale (Queiroz et al., 2019), whilst
effectiveness, it leaves information gaps regarding shark distribution reducing tagging bias based on the tagging location (Maxwell et al.,
patterns in areas without formal conservation or management struc- 2019; Francis et al., 2019; Queiroz et al., 2016), irregular post-release
tures, particularly in areas known to be heavily fished (i.e.., Southeast behaviour (Shipley et al., 2017a), and sex and age of tagged sharks
Asia). (Andrzejaczek et al., 2018; Howey et al., 2016; Howey-Jordan et al.,
2013; Wells et al., 2018).
4. Satellite tagging as a tool for shark conservation Many studies provided conservation support by tracking species of
conservation concern in novel or understudied areas where known risks
Satellite telemetry has long been used to inform shark conservation occur (i.e., Fishing in EEZs; Barkley et al., 2019). In fact, one study
initiatives (see Hays et al., 2019). Over the past decade research has concluded that illegal fishing had occurred based on the satellite track of
expanded in this field, providing novel information of at-risk species a tagged shark (Bradley et al., 2019). This type of analysis could be
habitat use, for instance scalloped hammerhead movements in Atlantic replicated throughout other suspicious tag recordings in order to iden-
equatorial oceanic waters (Bezerra et al., 2019), or range expansion of tify these events, potentially aiding regulating and monitoring bodies in
great hammerheads in the North Atlantic (Hammerschlag et al., 2011a). fisheries observations. Combining telemetry studies with fisheries
Evaluations of sharks' core habitat and activity space were determined dependent data (i.e., Logbooks) can help to pull apart the overlap be-
with methods like kernel density distributions (e.g., Doherty et al., tween fisheries and shark species (Coelho et al., 2020) and maximize
2017b), residency behaviours and indices (e.g., Acu! na-Marrero et al., fisheries management for the conservation of highly migratory species.
2017) and quantifications of site fidelity (e.g., Guttridge et al., 2017; Studies of post release survival also provided recommendations of best
Barnett et al., 2011), often to identify and prioritize areas for conser- practice for whale shark release when caught in commercial fishing nets,
vation (Queiroz et al., 2012; Queiroz et al., 2017; Rohner et al., 2018; while having little to no effect on fishers catch (Escalle et al., 2016a,b).
Bradley et al., 2019; Reynolds et al., 2017; Daly et al., 2018).
While most studies used shark conservation as a motive for satellite 4.1. Interactions with fisheries
tagging, only 45 studies used telemetry to answer questions directly
related to a conservation or fisheries management problem, rather than Investigations of fisheries interactions in reviewed studies included
simply expanding the literature regarding species habitat use and post-release mortality (PRM), species overlap with fishing activity, and
movement patterns. While valuable, movement information on its own assistance in stock assessments, constituting 31 total studies. Estimates

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S. Renshaw et al. Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

of PRM from 6 different gear types (longlines, tuna purse seines, remains ample opportunity to improve and expand satellite telemetry
drumlines, deep set buoys, gillnets, rod and reel) were made in 22 results into fisheries management or conservation planning, this tech-
studies, monitoring 792 individuals from 16 species across commercial, nology has proved to be a compelling tool that can provide impactful
recreational, and fishery-independent (i.e., research) fishing. Motivation outcomes.
for PRM studies included identifying safe and effective handling
methods for tagging and/or release (Escalle et al., 2016a; Escalle et al., 5. Conclusions and future directions
2018; Gallagher et al., 2014; Schaefer et al., 2019), bycatch survivability
(Benson et al., 2018; Campana et al., 2016), and gear associated risks This review has examined what can now be described as the ‘re-
(Sepulveda et al., 2015; Sulikowski et al., 2020). Longlining gear was naissance’ period of shark satellite tagging where focus has been paid to
most often monitored for instances of shark PRM (n → 9 studies), fol- expanding the geographical and ecological scope of tagging studies, as
lowed by tuna purse seines (n → 6 studies) often regarding FADs. PRM well as increased capacity for collaboration and data sharing. These
estimates varied widely depending on gear and species, most notably opportunities have been leveraged to support conservation and syn-
100% of tagged scalloped hammerheads (Sphyrna lewini) released from thesis driven research whereas previously, satellite telemetry studies
purse seines did not survive (Eddy et al., 2016), whereas silky sharks were often exploratory and lacking well-defined a priori research
released from tuna purse seines, suffered PRM ranging from 48 to 84% questions and therefore, appropriate designs to answer such questions.
(estimates from Poisson et al., 2014 and Hutchinson et al., 2015 The review shows that progress throughout the field has been made in
respectively). Lower instances of PRM were determined for several the past decade and the recent uptake of satellite tag derived data in
pelagic sharks captured by longlines (i.e., 0% PRM for tiger sharks, marine management is a clear bright spot, with plenty of opportunity for
Afonso and Hazin, 2014). continued implementation. Yet, despite satellite tag proliferation in
Ten additional studies investigated patterns between shark habitat shark research, tag innovation has not progressed as rapidly throughout
use and fisheries operations, all of which showed some evidence of this renaissance period compared to years prior. It is likely that the
overlap between sharks and fishing effort (n → 10 species total; Queiroz coming decade may offer substantially more technological innovation.
et al., 2016; Calich et al., 2018; Queiroz et al., 2019; White et al., 2019a; It remains imperative to continue tag innovation and development to
Hutchinson et al., 2019;). Estimates of shark overlap with fisheries was produce smaller tags (currently the smallest available prototype satellite
highly dependent on species, ocean region, and season (Queiroz et al., tag is the mark-report satellite tag (mrPAT), 121 mm long, 23 mm
2019). For example, Queiroz et al. (2019) assessed spatial risk to sharks diameter and 26 g in weight; Hussey et al., 2018), with strengthened or
using a fishing exposure index determined from satellite telemetry data enhanced functions like lower failure rates, more reliable data trans-
(for sharks and longline vessels) and found that on average, 49% of blue mission, longer battery life, and added sensor capabilities (Hammers-
shark movements overlapped with longline fisheries across all ocean chlag et al., 2014.) For example, the International Cooperation for
regions globally, whereas this estimate was only 1.3% for salmon sharks Animal Research Using Space receiver was deployed on the Interna-
(Lamna ditropis). Further, blue sharks overlapped with fishing effort tional Space Station in 2018 and its lower Earth's orbit means less energy
across 79% of their space use in the North Atlantic, versus just 14% in is required for tag transmissions and so batteries can become smaller to
the Eastern Pacific. Species vulnerability to fisheries interactions was meet the same duration times thus enabling tagging of smaller bodied
determined using covariates such as level of fisheries management species.or remain the same size as now to provide extended tracking
(Graham et al., 2016; Calich et al., 2018; Rooker et al., 2019), logbook durations (Sequeira et al., 2019b). Similarly, new Argos ground-based
records and catch per unit effort (Lyons et al., 2013), onboard observer listening stations (‘motes’ with 55 km range) and greater satellite area
or fisher recapture data (Tolotti et al., 2015; Afonso and Hazin, 2014), coverage will also contribute to shark tracking over longer temporal
gear characteristics (i.e., set depth; Santos and Coelho, 2018), and duration and/or across a broader geographic range, where there remains
locational information supplied by GPS, vessel monitoring systems spatial gaps in data collection (Sequeira et al., 2019b). As a community
(VMS) and automatic identification systems (AIS). Co-application of of researchers tagging sharks, it is also imperative that we learn from our
these satellite technologies for vessel monitoring (i.e., VMS/AIS data; mistakes with regards to the relationship between tag performance and
Queiroz et al., 2016, Queiroz et al., 2019; White et al., 2019a) was ab- animal welfare and survival; indeed, it is becoming clear that
sent from the previous review. While these studies provide substantial morphology, growth rates, and behaviour render some species as suit-
evidence for fisheries management, those whose results span large able candidates for certain tag types, while others are clearly poor
oceanographic regions will likely be slower to be taken into manage- candidates (Hussey et al., 2018). Indeed, as this review demonstrates
ment decisions due to the complexity of issues on the high seas and the from the few available studies on tag and tagging effects, research
need for transnational and transboundary co-management of fisheries projects should continue to monitor the impacts of tags on a broader
resources. Further research on uptake of telemetry studies into man- array of species as tagging studies proliferate.
agement decision making at an international level should be considered. Sequeira et al. (2019a) addressed the issue of appropriate sample
Poor knowledge of shark distribution (particularly for pelagic spe- sizes for animal tagging studies by evaluating the use of small (ω10) to
cies) in relation to active fisheries can make fisheries results difficult to very large (1000s) sample sizes and their applicability in marine
interpret, especially when fisheries data is scarce or incomplete (e.g., megafauna tracking research. Tag quantity and study design should be
poor observer coverage, unreported catch, etc.) (Queiroz et al., 2019, determined by the ability to draw meaningful conclusions and based on
2021a, 2021b). Recently, there has been increasing integration of fish- well-developed hypotheses, and, ideally, not simply availability or op-
eries independent information from electronic tagging studies into portunity. Further improvements can be made by pooling information
species stock assessment models, which has particular use for highly from across studies to generate larger data sets, made possible from the
migratory stocks by providing fine-scale resolution and long-term establishment and collaboration of multi-institutional programs (e.g.,
movement data of populations across geo-political boundaries (Sippel TOPP, see Block et al., 2011; GSMP, see Queiroz et al., 2019), open data
et al., 2015; Carvalho et al., 2015; Coelho et al., 2020). Population as- policies, and online repositories (i.e.., Integrated Marine Observing
sessments can benefit from information generated by satellite telemetry System [IMOS] and Ocean Tracking Network [OTN]) that continue to be
including utilization distribution, home range, and spatial dynamics that vital to increasing outputs from tagging events (Sequeira et al., 2019b;
extend beyond the reaches of fisheries to provide an improved under- Harcourt et al., 2019; Sequeira et al., 2021). Ongoing telemetry net-
standing of where and when sharks overlap with fisheries (Carvalho works like the Animal Tracking Network (ATN) hosted by the Integrated
et al., 2015; Coelho et al., 2020) and the influences of climate-driven Ocean Observing Systems (IOOS) have been responsible for large ac-
changes, such as ocean deoxygenation that can potentially increase cumulations of shark tracking data (Coffey et al., 2017; Queiroz et al.,
shark vulnerability to fishing (Vedor et al., 2021). Thus, while there 2019; White et al., 2019b; Bruce et al., 2019). Many existing programs

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S. Renshaw et al. Journal of Experimental Marine Biology and Ecology 560 (2023) 151841

have developed the required infrastructure needed for large data re- governance, human population, shark consumption levels, etc.) to better
positories that would include elasmobranch tracking data, such as understand the avenues for impacting change. Similarly, telemetry
MoveBank (www.movebank.org). Initiatives including the Spatial studies would do well to quantify shark movement and residency across
Ecological Analysis of Mega-vertebrate Population (OBIS-SEAMAP), a range of management regimes and to understand how this may impact
Migratory Connectivity (MiCO), Marine Megafauna Movement Analyt- populations globally. This work has been performed with other mea-
ical Program (MMMAP), and the Global Shark Movement Project sures of shark and ray abundance (see MacNeil et al., 2020), where the
(GSMP) have also begun to address persistent challenges of data sharing authors were able to produce a suite of policy recommendations based
pertaining to ownership rights (Sequeira et al., 2019b, 2021). However, on their findings. Doherty et al. (2017) is directly referenced to support
as Sequeira et al. (2019b) and Harcourt et al. (2019) pointed out, a lack the MPAs designation and its protection of basking shark populations,
of universal data standards, misinterpretations of data, limited funding, designated in December of 2020. Studies like this provide useful, prac-
opaque resource sovereignty, and analytical difficulties of pooled data tical, solutions that provide the evidence needed to support proper MPA
continue to create hurdles for data collection and require innovative boundaries and implementation. Evidence from telemetry studies may
solutions (see Sequeira et al., 2019b, 2021 for recommendations and not be utilized for several years after publication or may never end up in
Harcourt et al., 2019 for a list of marine data repositories). primary literature available for review.
The majority of originally reviewed tracking studies were inherently In conclusion, the data summarised here provides a refreshed over-
foundational, as they described some of the first spatio-temporal pat- view of telemetry endeavours to date, as well as pointing a way forward
terns of species-specific shark movement. While a thorough under- in shark tracking studies to further our understanding of species
standing of when and where sharks move is imperative, our review movement ecology. Ultimately, satellite telemetry will always provide a
revealed that, despite an additional 10 years of expanded work using relative snapshot of the greater life history of tagged animals, and as
satellite telemetry on sharks, investigations that explicitly formulate such will always need to be integrated with other tools in the shark
hypotheses or management objectives a priori are still in the minority. science toolbox to understand the behaviour of sharks in an ever-
This finding is likely not taxon specific. Nguyen et al. (2017) claimed changing ocean.
the “lack of explicit management objectives designed into telemetry
projects delays conservation impact of animal-borne telemetry in-
vestments.” Additionally, the mechanistic drivers of shark movement Declaration of Competing Interest
and intra-specific variability of movement patterns remain poorly un-
derstood in many species and regions. Thus, there is a continuing need to The authors declare that they have no known competing financial
integrate satellite telemetry with individual physiology, metabolism, interests or personal relationships that could have appeared to influence
gene expression, and reproductive strategy to better understand the the work reported in this paper.
context of movement decisions (Hays et al., 2016). Investigating the
ecological and economic consequences of seasonal shark presence/ Data availability
absence (tourism, fisheries, indirect/direct predation effects; see Ham-
merschlag et al., 2019), overlap of potentially dangerous shark species No data was used for the research described in the article.
with humans, climate change effects on shark movement and distribu-
tion, and population connectivity will be essential to create holistic and Acknowledgements
effective management decisions across scales (e.g. from local and
regional to international) (Queiroz et al., 2019; Hays et al., 2019). The The authors would like to thank Cassidy Walker and Sarah Hughes
effects of climate-driven environmental changes on shark movements for the support throughout the writing process and for providing feed-
and distribution are comparatively understudied, particularly with back on sections they read. Further, the guidance of the Marine Affairs
respect to future predicted changes (Vedor et al., 2021). This area of faculty at Dalhousie University is greatly appreciated. Finally, to Fred
shark tracking research promises much in the decade ahead as both Whoriskey whose continued mentorship and trust was paramount to this
shark spatial datasets and predictive ocean climate models mature publication.
further.
While telemetry in general, including acoustic methods, accounts for Appendix A. Supplementary data
many more elasmobranch species than reviewed here, satellite telem-
etry has only been used to track ω8% of shark species (with respect to Supplementary data to this article can be found online at https://doi.
over 500→ shark species known worldwide). Geographic gaps in shark org/10.1016/j.jembe.2022.151841.
tracking persist primarily off East and West Africa, South America and
Asia. Much of the world's shark catch is generated off these continents, References
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