Received: 21 July 2021 | Revised: 20 January 2022 | Accepted: 27 January 2022

DOI: 10.1111/faf.12648

ORIGINAL ARTICLE

Global assessment of shark strandings

Natascha Wosnick1 | Renata D. Leite1 | Eloísa P. Giareta1 | Danny Morick2,3 |

Michael Musyl4

1
Programa de Pós-­G raduação em
Zoologia, Universidade Federal do Paraná, Abstract
Curitiba, Brasil
Marine wildlife stranding is a global phenomenon, and for some taxonomic groups,
2
Morris Kahn Marine Research Station,
University of Haifa, Haifa, Israel
these events are well monitored and documented. Although sharks are among the
3
Hong Kong Branch of Southern Marine most threatened vertebrates, strandings have historically been neglected, with little
Science and Engineering Guangdong information on this topic currently available. To address this knowledge gap, a sys-
Laboratory, Guangzhou, China
4 tematic review on scientific publications from indexed databases, multimedia material
Pelagic Research Group LLC, Honolulu,
Hawaii, USA and citizen science databases was performed. A total of 3150 reports were recov-
ered, with strandings dating back to 1880 including 89 species of sharks from 22
Correspondence
Natascha Wosnick, Programa de Pós-­ families. Both single/small-­scale and mass stranding events were observed. Species
Graduação em Zoologia, Universidade
ranged from coastal to oceanic, with reports for small-­, medium-­and large-­sized
Federal do Paraná –­Centro Politécnico,
Curitiba, Paraná, Brasil. sharks. Leopard (Triakis semifasciata, Triakidae), brown smooth-­hound (Mustelus hen-
Email: n.wosnick@gmail.com
lei, Triakidae) and salmon sharks (Lamna ditropis, Laminidae) were the species with
Funding information most records. Females stranded significantly more than males, but juveniles and
Coordenação de Aperfeiçoamento de
adults were affected in the same proportion. Shark strandings were reported for 47
Pessoal de Nível Superior (CAPES)
countries, and the United States was the most representative location, highlighting
the states of California and Oregon. Survival rates were extremely low, indicating
that stranded sharks are more vulnerable that other taxonomic groups. Potential and
putative causes were identified, being similar to those suggested for marine mam-
mal strandings. Our results indicate that although they occur to a lesser extent when
compared to mammals, shark strandings are neglected, and urgent measurements are
necessary to better understand, document and to properly respond to these events.

KEYWORDS
beached, conservation, shark, threats, wash-­up

1 | I NTRO D U C TI O N
Marine wildlife stranding is a global phenomenon that affects
both invertebrates and vertebrates (Chan et al., 2017; Cockcroft,
2001). Many definitions are available, being guided mostly by the
taxonomic group affected. The National Oceanic and Atmospheric
Administration (NOAA) recognizes three main taxonomical catego-
ries of strandings: cetaceans, pinnipeds and sea turtles. Stranded
animals may be found dead (either on land or floating) or alive but
unable to return to sea due to injury or illness. Broader definitions
are also available and do not consider taxonomic groups, but simply
the presence of an aquatic animal on land, whether dead or alive (but
physiologically/physically unable to return to the sea) (Branco et al.,
2015; Brusius et al., 2020; Geraci & Lounsbury, 2005).
Whereas single or small-­scale strandings are more common,
mass strandings are of great concern as they are usually associated
with disease outbreaks, climatic events or as a result of anthropo-
genic disturbances (Moore et al., 2018). Most stranding reports
are of marine mammals, turtles and seabirds with cetaceans most
commonly reported (Bogomolni et al., 2010; Fahmi et al., 2014;
Geraci, 1978). The frequency of marine mammal strandings led to
the implementation of initiatives at a global level, given the need

Fish and Fisheries. 2022;00:1–14. wileyonlinelibrary.com/journal/faf© 2022 John Wiley & Sons Ltd | 1
2 | WOSNICK et al.

to understand not only why these events are affecting marine spe-
cies, but also as a promising way to assess population size, mortality
1. INTRODUCTION 1
rates, movement patterns and the impacts of strandings on pop-
2. METHODS 2
ulation dynamics (Geraci & Lounsbury, 2005). Along with rescue,
rehabilitation centres and monitoring initiatives carry out necrop- 2.1. Scientific publications and citizen science 2

sies and collect relevant data to assess the causes of strandings. 2.2. Complementary data 3

However, to date, most strand monitoring initiatives are focused 3. RESULTS 3

on charismatic marine megafauna, with few data available for other 3.1. Stranding reports 3
taxonomic groups. 3.2. Shark species 3
Sharks are among the vertebrates most affected by anthro- 3.3. Sex and Maturity 3
pogenic activities, and 31.2% of currently described species are 3.4. Geographic distribution 4
now threatened with extinction (Dulvy et al., 2021). Overfishing 3.5. Stranding trends across the United States 4
is undoubtedly the main contributing factor to the observed pop-
3.6. Historical trends of shark strandings 4
ulation declines (Dulvy et al., 2014, 2021). Sharks are also affected
3.7. Survival rates 5
by habitat degradation, pollution and climate change (Dulvy et al.,
3.8. Potential causes 5
2014; Pacoureau et al., 2021). Stranding reports offer important
4. DISCUSSION 6
opportunities to assess additional stressors that could impact
4.1. Shark species 6
survival and/or population-­level processes, and should be moni-
tored closely. For marine mammals, stranding refers to both live 4.2. Sex and Maturity 9

and dead animals, but for sharks, the term stranding is used only 4.3. Geographic distribution 9
when the animal is found alive. For dead animals, the term wash-­up 4.4. Historical trends on shark strandings 10
is prevalent without making a distinction as to whether the ani- 4.5. Survival rates 10
mal stranded alive and died, or it was already dead when found 4.6. Potential causes 10
(personal observation). Although superficial, this operational dif- 4.6.1. Infectious Disease 10
ference in definition highlights the little importance that has been 4.6.2. Abrupt environmental alterations 12
given to this phenomenon on sharks. Moreover, stranding reports
4.6.3. Negative interaction with fisheries 12
in the literature are scarce for these animals, with most data used
4.6.4. Other trauma 12
as occurrence records.
4.6.5. Foraging, sandbanks and tide pools 12
As little attention has been given to shark strandings, many
5. CONCLUSIONS AND FUTURE DIRECTIONS 12
knowledge gaps exist. To shed light on this matter, a systematic
ACKNOWLEDGEMENTS 12
review on the subject was performed, focusing on answering
specific questions. Firstly, for what species are stranding re- DATA AVAILABILITY STATEMENT 12

ports currently available? What species exhibit the highest rates REFERENCES 12

of strandings? Are strandings sex-­specific, ontogenic-­d ependent

or size dependent? Are there regions where sharks strand more
often? Are there spatial–­temporal patterns in stranding reports?
How much and what sort of data are available in traditional da-
tabases (peer-­reviewed papers, grey literature and books)? How
much does informal records and citizen science contribute to the
data currently available? How many sharks are stranded alive, and
of these, how many survive after rescue attempts? Finally, what
are the putative causes and prognostic factors related to reported
strandings?
2 | M E TH O D S
2.1 | Scientific publications and citizen science
Stranding reports were searched primarily in scientific publications
through indexed databases, including, Scopus and Web of Science.
A structured Boolean search (AND, OR, +) was performed using
the following search terms: "shark," "stranded," "strandings," "wash
up," "massive strandings." Then, a snowball method was applied
(Greenhalgh & Peacock, 2005), to search for additional publica-
tions from the reference lists. As some published reports focused
on the pathologies involved in strandings (and therefore were not
identified through the search terms), a second structured Boolean
search using the terms “shark,” “stranded,” “pathology” was per-
formed, and papers not found in the initial search were manually
added by the authors (Paul & Criado, 2020). A search for confer-
ence abstracts (grey literature) was also performed, applying the
same search terms.
Reports from iNaturalist (https://www.inatu​
ralist.org/) were
also considered, as a source of citizen science. Data from this spe-
cific database were found by adding the filter ‘elasmobranch’, and a
search was made manually from the first report in 2008 (data added,
but some reports date back to the 1990s) to the last one added in
January 2021. To avoid any bias in the recovery process, three au-
thors examined each record independently.
WOSNICK et al.
2.2 | Complementary data
An additional search for videos and photographic records on multi-
media platforms was performed using the search terms (Facebook
and YouTube) or hashtags (Instagram and Twitter) “shark,” “stranded,”
“standings,” “wash up” and “news,” according to Boldrocchi and
Storai (2021). Open-­
access database reports from rescuing pro-
grams websites and stranding monitoring Facebook groups (e.g.
California Department of Fish and Wildlife, UK Cetacean Strandings
Investigation Programme, Cornish Wildlife Trust Marine Strandings
Network, Sanctuary Integrated Monitoring Network, Pelagic Shark
Research Foundation) were also added in this search phase. It is im-
portant to mention that we cannot be sure to have captured all data
available or in a systematic fashion when it comes to media (e.g. news-
papers and websites), public platforms (i.e. YouTube) or social media
(i.e. Facebook, Instagram and Twitter). Moreover, it was beyond the
scope of this work to collect data from every oceanographic institute/
aquarium/state or national agency, as most databases are of restricted
access. Therefore, the reports recovered during this search phase are
treated as complementary and were not added to the quantitative
analysis performed. While this is a weakness, the data gathered in-
sights into the critical topic of shark stranding from two points of view,
namely science and happenstance (multimedia material).
Lastly, a visual examination of raw data was carried out to vali-
date records and reject replicated entries for both multimedia material
(TV news, newspapers and online platforms) and iNaturalist reports
(Toivonen et al. 2019). For six mass events, it was not possible to recover
the exact number of individuals affected, so these events were excluded
from the analyses (i.e. leopard sharks in California). For all data sources,
only reports for which it was possible to extract date (year), location
(country) and taxonomic identification at the species level (confirmed by
a specialist—­i.e. Gadig, O. and Duffy, C.) were considered. Lastly, to stan-
dardize data collection, only entries/reports in English were considered.
Qualitative analyses were performed to evaluate the most com-
monly stranded species, strandings per sex, as well as to evaluate
the countries with the highest number of reports, and reports over
time. Moreover, the possible causes of strandings were also evalu-
ated. To test the significance of relationships, an analysis of deviance
was applied. Quantitative analyses were also performed for number
of strandings by sex using chi-­squared (χ2) tests, and for species and
location (countries and States of the United States) separately, using
Kruskal–­Wallis tests. Statistical significance was set at p < .05. All
analyses were performed employing the R software version 1.4.1103
(R Development Core Team, 2021), using the MASS and car packages.
3 | R E S U LT S
3.1 | Stranding reports
A total of 3150 individual reports were retrieved. Concerning tra-
ditional scientific sources (both research articles and book chap-
ters), 42 publications were recovered (Table S1). An additional
| 3
12 reports from grey literature were found, totalizing 54 scien-
tific documents and 2073 records of shark strandings (65.8% of
total)—­m ost of them from mass events in California, USA. As for
citizen science, a total of 913 reports were recovered (28.9% of
total), being the second-­m ost relevant data source of global shark
strandings.
The additional search performed on multimedia platforms (web-
sites and social media) retrieved 70 reports, and along with data re-
covered from open-­access databases of rescuing programs websites
and monitoring groups (94 reports), accounted for 5.2% of all reports
found (n = 164). It is noteworthy that 57% of all stranding reports
for the basking shark, 32% of all reports for the great white shark
and 20% of all reports for the salmon shark are from complementary
entries, and although these represent the most recent and also sig-
nificant sources of information, the data were not considered for the
statistical analyses presented below.
3.2 | Shark species
Of the total, 89 species belonging to 22 families were identi-
fied (Table S2). Both large and small-­sized species were repre-
sented. The largest individual reported was a 9.45-­m whale shark
(Rhincodon typus, Rhincodontidae) and the smallest was a 20-­
cm Chilean catshark (Schroederichthys chilensis, Scyliorhinidae).
Species’ habitats ranged from coastal to pelagic, with some
reports for deep-­
s ea sharks (e.g. velvet belly lanternshark,
Etmopterus spinax, Etmopteridae; velvet dogfish, Zameus squa-
mulosus, Somniosidae and the angular roughshark, Oxynotus
centrina, Oxynotidae). Records were recovered for both benthic
and pelagic sharks. Some mass strandings were reported, being
relatively common for the leopard shark, and the brown smooth-­
hound, and isolated mass events were reported for the blue shark
(Prionace glauca, Carcharhinidae) and the dusky smooth-­h ound
shark (Mustelus canis, Triakidae).
The number of stranded individuals considered differed signifi-
cantly between species (Kruskal–­Wallis; H = 188.88, df = 87, p-­value
< .05 Figure 1). Among the ten most commonly stranded species
for which reports are available, the leopard shark stands out (n =
1153), followed by brown smooth-­hound (n = 531) and salmon shark
(n = 403, being 83 reports from complementary sources). The whale
shark also stands out, being the largest-­sized species with the high-
est number of records (n = 114, being 17 records from complemen-
tary sources) (Figure 1).
3.3 | Sex and maturity
For most considered records (n = 1837; 61.5%), sex was not identi-
fied. For individuals for which sex was identified (n = 1149, 38.5%),
848 (73.8%) were females and 301 (26.2%) were males (Chi-­sq; χ²
= 260.41, df = 1, p-­value <.05). Pregnancy (1.7%) was detected
in stranded females for the blacktip shark (Carcharhinus limbatus,
4 | WOSNICK et al.

Global reports of shark strandings (1880-2021)

Leopard shark
(Triakis semifasciata, Triakidae)
Brown smooth−hound shark
(Mustelus henlei, Triakidae)
Salmon shark
(Lamna ditropis, Laminidae)
Whale shark
(Rhincodon typus, Rhincodontidae)
Starry smooth hound shark
(Mustelus asterias, Triakidae)
Small spotted catshark
(Scyliorhinus canicula, Scyliorhinidae)
Dusky smooth−hound shark
(Mustelus canis, Triakidae))
Blue shark
(Prionace glauca, Carcharhinidae)
Bonnethead shark
(Sphyrna tiburo, Sphyrnidae)
Great White shark
(Carcharodon carcharias, Lamnidae)

0 300 600 900 1200

Reports

F I G U R E 1 Global reports of shark strandings, highlighting the ten species with the greatest number of reports. Date range: 1880 to 2021

Carcharhinidae), the draughtsboard shark (Cephaloscyllium isabel-
lum, Scyliorhinidae) and the salmon shark. Moreover, an abortion
event was recorded in a pregnant shortspine spurdog shark (Squalus
mitsukurii, Squalidae) stranded in a tide pool. Neonates of bonnet-
head (Sphyrna tiburo, Sphyrnidae), smooth hammerhead (Sphyrna
zygaena, Sphyrnidae) and scalloped hammerhead sharks, (Sphyrna
lewini, Sphyrnidae) were also sporadically reported, as well as one
bull (Carcharhinus leucas, Carcharhinidae) and one dusky shark
(Carcharhinu obscurus, Carcharhinidae). For most of the sharks re-
corded (78.2%), there was no information on total length, and thus
trends in size and (or) maturity could not be analysed in detail. For
records with shark size available, 7.1% presented exact total length
(TL), while 2.6% presented estimated TL. Considering the sharks for
which life stage was available, there appears to be no prevalence of
adults (5.4%) or juveniles (6.7%).
3.4 | Geographic distribution
strandings (22/41) also occurred in the United States. For both starry
smooth-­hound and small spotted catsharks reports were predom-
inantly from the United Kingdom (53/84 and 38/60, respectively).
Whale shark (45/110) and blue shark strandings (21/57) occurred
primarily in South Africa (Figure 2b). A list of the remaining species
recorded in the four main hotspots is presented as Table S4.
3.5 | Stranding trends across the United States
As the United States was the country with the highest rates of re-
ported strandings, data were analysed at the state level, to assess
possible patterns regarding specific locations. Among the 2409 re-
ports for the country (53 complementary entries excluded), 58.9%
are from California (n = 1419) (Kruskal–­Wallis; H = 41.976, df = 21,
p-­value <.05). The second state with most reports is Oregon (6%; n =
146) (Figure 3) followed by Florida (n = 54), Texas (n = 49), and North
(n = 28) and South Carolina (n = 17) (Figure 3).

Shark strandings were reported for 47 countries of 148 countries

with a coast (31.7%) (Table S3), with statistically significant differ-
ences between locations (Kruskal–­Wallis; H = 17.091, df = 9, p-­value
<.05). The United States stands out in the number of strandings
(n = 2462), followed by New Zealand (n = 121), the United Kingdom
(n = 118) and South Africa (n = 114) (Figure 2a).
As for the top 10 stranded species, all leopard shark and brown
smooth-­
hound stranding reports occurred in the United States,
where they are native. The majority of salmon shark (376/403),
dusky smooth-­hound (58/60), bonnethead (48/49) and great white
3.6 | Historical trends of shark strandings
Overall, shark stranding reports were more common in the years
2011–­2021 than in 2000–­2010 and were less commonly available, in
general, before 1999 (Figure 4). Sixty-­nine per cent of all stranding
reports were reported in the last 10 years (2011–­2021), configuring
the period with the highest number of reports. In addition to sev-
eral unique reports, a massive stranding of more than 1000 sharks
in 2017 was identified. Even excluding this specific mass stranding
WOSNICK et al. | 5

F I G U R E 2 (a) Global hotspots for

shark strandings. The gradual colour (a)
scale represents the locations with the
highest (dark blue) and lowest (light blue)
number of strandings. (b) Top 10 species,
indicating most representative countries
regarding the number of stranding reports

(b) Leopard shark Dusky smooth−hound shark

United States United States

1154 reports 58 reports

Brown smooth−hound shark Small spotted catshark

United Kingdom
United States
38 reports
532 reports
France
9 reports

Salmon shark United States Blue shark South Africa

376 reports 21 reports

Canada United States

24 reports 11 reports

Whale shark South Africa Bonnethead shark

45 reports United States
48 reports
Mexico
29 reports

Starry smooth hound shark Great White shark

United Kingdom United States
53 reports 21 reports

Ireland Canada
29 reports 10 reports

event, 2017 to 2020 accounted for 83.6% of the strandings reported
for that time period. Apart from two mass stranding events in 1967
(n = 505), only 164 strandings were reported in free-­access data-
bases between 1880 and 1999. Of these, 70 were reported in 1982,
being the most representative year in terms of single and (or) small-­
scale stranding reports. A detailed temporal distribution of strand-
ing for the top 10 species is presented in Figure 5.
delayed mortality occurred for most of the moribund subjects, and
future studies could better quantify post-­stranding survival rates
through reflex action mortality predictors (RAMPs) and condition
scores. Condition codes from PSAT tags are also efficient in estimat-
ing survival rates in sharks (Musyl & Gilman, 2018), being promising
tools to stranded sharks returned to the sea.

3.8 | Potential causes

3.7 | Survival rates
Considering all reports, only 9.9% of sharks were found alive
(Figure 6). Of these, 94% did not survive, even when rescue attempts
were made. For the remaining, 1% died soon after being returned to
the sea. That said, of the total (n = 3150), only 0.49% of sharks found
ostensibly alive were returned to the sea; however, it is likely that
For most reports, stranding causes were not mentioned and (or)
investigated. Moreover, most stranded individuals were found in
advanced stages of decomposition, preventing laboratory analyses
from being carried out. Despite these difficulties, some interesting
patterns could be identified (Figure 7). For many strandings with de-
tailed information (especially mass strandings), the main cause was
6 | WOSNICK et al.

United States Shark Standings (1880-2021) F I G U R E 3 Shark stranding across the

United States considering the states with
California at least one report. Data range: 1880–­
Oregon 2021

Florida

Texas

Massachusetts

South Carolina

Washington

New York

New Jersey

Georgia

0 250 500 750 1000 1250

Reports

0.6% 40.8%
0.1% 99.3% 5% 54.2%

669 2185

1928-1999 2000-2010 2011-2021

Total
Publications
Media 277
Citizen science
22.7%
7.2% 70.1%

F I G U R E 4 Timeline of globally reported shark strandings over a period of 141 years

meningoencephalitis. Salmon sharks were the most affected by this
pathology, followed by leopard sharks. Capture stress also appears
to be a common cause provided in the reports, along with lethal inju-
ries caused by fishing apparatus; especially noted for scalloped ham-
merheads and shortfin makos (Isurus oxyrinchus, Lamnidae). Other
traumas were also seen in stranded sharks, among them lethal inju-
ries caused by collisions with motorized boats and predator injuries
(e.g. bite marks). Injuries of unknown aetiology and causes are also
being observed.
Small and medium-­sized coastal species, such as leopard and
port Jackson sharks (Heterodontus portusjacksoni, Heterodontidae),
appear to be particularly vulnerable to strandings due to low
tides, leaving them trapped in sandbanks and tide pools. Sharks
may be trapped in shallow waters during foraging, courtship, or
while seeking shallow habitats for parturition. Marine debris and
pollution have also been reported as an associated cause, mainly
for whale and brown smooth-­h ound sharks. Lastly, environmental
disturbances appear to contribute significantly to mass strandings
(e.g. blue sharks and Chilean catsharks) especially when an abrupt
contemporaneously decrease in temperature and dissolved oxy-
gen occurs.
4 | DISCUSSION
4.1 | Shark species
This is the first review on global shark strandings, with focus on
more affected species, locations with the highest occurrences
and overall patterns. Reports of stranding events were recovered
for 22 of 37 shark families and a total of 89 species, indicating
that this phenomenon affects sharks with distinct evolutionary
WOSNICK et al.
F I G U R E 5 Temporal stranding patterns
of the top 10 species identified in the
present study. (a) Graphical representation
for the top 2 species. (b) Graphical
representation for the remaining top
10 species
patterns, ecological specializations, distributions and life histo-
ries. Although there is a predominance of small-­sized species,
medium and large-­sized sharks are also affected by this phenom-
enon. However, mass strandings (apart from singular events such
as the case of the blue sharks in South Africa) appear to affect
small sharks more significantly. It is possible that this pattern is
more related to species-­
s pecific behaviours rather than shark
size, including propensity to travel together in schools. Although
coastal species also showed a predominance in the number of
reports, oceanic and deep-­s ea sharks were also reported in the
stranding records. Moreover, both benthic and pelagic species
were reported. It is worth mentioning that the predominance of
coastal species might be due to higher population density along
shorelines when compared to oceanic species. Also, it is possi-
ble that oceanic sharks that die offshore sink or are consumed
by scavengers before the carcasses reach the shore, resulting in
potential underrepresentation.
Detailed data are available for only a few species, mostly for
which strandings occur most often. Several mass stranding events
| 7
have affected leopard sharks over the years, but most of them
were not systematically documented (Retallack et al., 2019). It is
worth noting that about 1/3 of the reports are from a single mas-
sive leopard shark stranding event off the coast of California, con-
figuring the most significant source of reports from open-­a ccess
databases. The biggest stranding events recorded for the species
were in 1967, 2006, 2011 and 2017, all from California. The sharks
found alive in these strandings were often described as disori-
ented and confused, exhibiting unusual swimming patterns and
behaviours (Retallack et al., 2019). Necropsies performed on the
animals that died in 2017 were diagnostic of necrotizing meningo-
encephalitis, which could explain the disorientation. The specific
cause of meningoencephalitis (e.g. bacterial, viral or other cause)
was not determined. The contributing effects of factors such as
water temperature, salinity and pollution all deserve consideration
and greater study in future stranding events. The repetitive nature
of strandings, as well as occasional mass mortalities in this spe-
cies, may impact population dynamics and local ecology. Improved
monitoring and diagnostics could better determine the cause and
8 | WOSNICK et al.

F I G U R E 6 Survival rates of sharks

How many Sharks stranded
survived to strand?

Did not survive Found alive

x 90.1% 9.9%

Of these,how
many survived?

Returned to
Did not survive the sea

1% x
x 94%
Returned to Died soon after
the sea

5%
No further reports of
new strandings

Causes of shark stranding events F I G U R E 7 Identified causes related

to shark stranding events, with their
SUDDEN respective percentages of occurrence for
ENVIRONMENT ALTERATIONS reports that information is available
65%
COLISION WITH BOATS 0,3% 14% MENINGOENCEPHALITIS

LOW / HIGH TIDE, FORAGING AND

SANDBANKS TIDE POOLS 0,6% 9% REPRODUCTION

CAPTURES STRESS 0,6% 3% FISHERY

EXTERNAL INJURIES 0,8% 2% PREDATION

OTHER PATHOLOGIES 0,9% 1,2% MARINE LITTER

INTERNAL INJURIES 1% 1,2% PREGNANCY AND ABORTIONS

High Low

the influence of stranding events and inform the health of the
environment.
The biggest stranding event for brown smooth-­
hounds hap-
pened in 1967 (Russo & Herald, 1968). A total of 492 sharks stranded
in a period of 33 days, also exhibiting abnormal behaviour and er-
ratic swimming. Although the cause of the mass stranding was
not determined, it is believed that it was caused by a pollutant.
Interestingly, the mass events and individual strandings for this spe-
cies in California occurred concurrently with the strandings of the
leopard sharks. Such a pattern indicates that the same causes may
be affecting both species. In fact, it is possible that pathologies with
a high dispersion potential are the main cause; however, until now,
strandings for this species have not been investigated in depth. The
brown smooth-­hound is not at imminent risk of extinction. However,
mass mortalities can significantly alter the dynamics of the affected
populations, requiring constant monitoring.
Records shows that salmon sharks frequently strand along the
West coast of North America (Carlisle et al., 2015), mainly along
British Columbia, Canada and northern Baja California, Mexico
(Goldman & Musick, 2006). There are also several records for both
WOSNICK et al.
California and Oregon (the United States), and most strandings oc-
curred during fall and summer (Schaffer et al., 2013). Generalized
additive models showed that the probability of stranding was higher
when mean water temperature was high or when individuals were
exposed to cold socks during coastal upwellings (Carlisle et al.,
2015). Since most of the salmon sharks that strand are young of the
year (YOY), it is possible that the species occupies a narrow thermal
niche at this life stage or that cold stress predisposes to bacterial in-
fections such as meningitis or meningoencephalitis (Martinez-­Steele
et al., 2018; Schaffer et al., 2013) when water temperature drops
below 12°C (Carlisle et al., 2015). Sudden changes in temperature do
not seem to be a limiting factor for adults and it is worth mentioning
that they also strand in the region, with some of them presenting
extensive hematomas in the ventral region, a strong indication of
interaction with fisheries. Additionally, some adults were found with
large bites, indicating that prey–­predator interactions might also be
a source of strandings and for carcasses to wash up. Although the
species is listed as Least Concern on a global level, these stranding
events are of particular concern to the North America population,
as YOYs are the most affected and this may reflect an unfavourable
population trajectory, leading to declines in near future that must
be monitored.
Among the large-­sized species, the whale shark is the species
with the most information available. Strandings occur relatively
frequently on the coast of South Africa (Speed et al., 2009) and
Mexico. A reduction in metabolic rate due to sudden changes in
water temperature and/or steeply-­sloping continental shelf com-
bined with heavy wave action have been pointed out as possible
causes (Beckley et al., 1997). As filter feeders, whale sharks are par-
ticularly vulnerable to marine litter and a necropsy performed on a
shark recently stranded in the Philippines indicated the presence of
plastic debris in the gills and stomach (Abreo et al., 2019) suggesting
that it might have contributed to the event. Other potential causes
for whale shark strandings are mentioned in the literature, among
them fisheries pressure, illness, pollution or several environmental
and bathymetric factors, such as sudden changes in surface currents
and tidal differences (Beckley et al., 1997; Pierce & Norman, 2016).
Because they are commonly mistaken for real whales, whale sharks
stranding generates media attention with many people trying to res-
cue the animal. Unfortunately, success rates are low, and many of
them die before anything can be done. Thus, identifying locations
where stranding is recurrent and overall characteristics related to
it informs future efforts, to establish cooperative monitoring (espe-
cially between South Africa and Mexico, even if there is no popula-
tion connectivity) and surveillance programmes.
4.2 | Sex and maturity
There is a clear lack of data regarding the sex of stranded sharks.
The problem is also faced for other marine animals, such as ceta-
ceans, phocids and sea turtles (Chaloupka et al., 2008; Gallini et al.,
2021; Norman et al., 2004). In the case of sharks, sex identification
| 9
is facilitated by the presence of external copulatory organs on male
sharks (i.e. claspers). However, as most reports are made by citizen
scientists, a precise identification by specialists can only be per-
formed when photographs of the ventral region are available. As
individual or small-­scale strandings are rarely reported by special-
ists and monitoring programs, it is necessary that training be made
available to citizen scientists and enthusiasts, so that more accurate
data on sex can be collected. Interestingly, even in scientific reports,
sexual identification is rare, which may be a reflection of personal
communication and instruction for data collection.
According to our results, female sharks are more vulnerable to
stranding, representing 74% of reports (i.e. for which it was possible
to identify the sex). Considering the knowledge available on shark
reproductive behaviour, pregnant females of several species tend to
approach the coast to give birth (Elisio et al., 2016; Heithaus, 2007).
Moreover, the energetic demands during pregnancy can decrease
female's immune response (Guida et al., 2017), which could favour
pathogen's spread responsible for strandings. In addition, stranding
can also induce parturition. This phenomenon is believed to be an
adaptive strategy to increase the chances of litter survival (i.e. self-­
sacrifice hypothesis) (Adams et al., 2018), with one case documented
in the literature for the bluntnose sixgill shark (Hexanchus griseus,
Hexanchidae) (Williams et al., 2010) and in several online videos. In
the present study, five developing embryos of shortspine spurdog
shark were found in a tide pool, suggesting that the mother may
have stranded and expelled them prematurely. Additional factors
related to the greater vulnerability of females cannot be ruled out
as many did not show signs of pregnancy or were juveniles. Another
point to be considered is that this overall pattern may not be repre-
sentative of shark strandings. For example, in the Gulf of California,
male whale sharks are more vulnerable to stranding (Ketchum et al.,
2013).
4.3 | Geographic distribution
To optimize resources and increase the chances of rescue, it is ex-
tremely important to identify places where marine animals strand
the most. Through spatial analysis, it is possible to map locations
with higher carcass deposition, signalling hotspots and directing
surveillance and monitoring efforts (Olson et al., 2020). While shark
stranding is a global phenomenon, stranding reports are likely biased
towards areas where there is public access and high public traffic
(such as piers). The United States, New Zealand, the United Kingdom
and South Africa were hotspots in our study. Interestingly, these
countries are also global hotspots for mammal strandings (Best &
Reeb, 2010; MacLeod et al., 2004; McGovern et al., 2016; Pyenson,
2011), emphasizing their relevance in the global stranding scenario
of marine animals.
The oceanographic aspects, topology and surface currents seem
to be the main conditions involved with strandings in New Zealand
(Brabyn & McLean, 1992). In the United Kingdom, events seem to
be related to the passive loading of carcasses by the North Atlantic
10 | WOSNICK et al.
drift (McGovern et al., 2016). Although the causes identified for
both New Zealand and the United Kingdom are mostly of environ-
mental origin, other variables, including anthropogenic impacts and
recurrent disease outbreaks, cannot be ruled out. In South Africa, a
mass event was recorded for blue sharks due to sudden temperature
changes, with no further information on survival rates. Whale shark
strandings are also relatively common along the South African coast,
with a peak in 1991 (n = 14), including a specific situation where
3 whale sharks stranded in the same location, at the same time
(Beckley et al., 1997). Reasons are uncertain although the narrow,
steeply-­sloping continental shelf and beaches with heavy wave ac-
tion may be contributing factors (Beckley et al., 1997).
The United States is undoubtedly the country with the highest
reported stranding rates for marine animals. Our results indicate the
state of California had the highest rates of shark strandings, corrob-
orating patterns described for mammals. Moreover, California has
one of the longest coast lines of the country, with a high number of
people living in coastal areas, enhancing opportunities for strand-
ings to be recognized. Apart from causes linked to anthropogenic ac-
tivities, the coastal geography of California is also believed to favour
carcass deposition (Danil et al., 2010). Moreover, mass stranding
events were prevalent in the state, and further studies are needed
to understand the causes involved in these events. In addition, the
coast of California is a promising place to monitor shark strandings
more thoroughly, which could allow the creation of rescue and reha-
bilitation protocols to be disseminated.
4.4 | Historical trends on shark strandings
Apart from specific mass strandings, an increase in the number of
reports over the decades has been noted. Although anthropogenic
activities that directly affect marine animals has intensified, it is
more likely that this increase is related to a greater number of peo-
ple (outside the academia, NGOs and monitoring programs) involved
in species identification activities and informal stranding reports.
Between 1880 and 1990, only a portion of the global population
had access to photographic cameras and the internet. Even though
the internet was launched in 1969, dial-­up connections and the high
cost of computers made online access very restricted until the early
2000s. While the collection of scientific data by non-­scientists dates
back a few millennia (Irwin, 2018), it was only in the mid-­1990s that
the concept of citizen science began to be popularized. The emer-
gence of broadband internet and the first cell phones with cameras
in the early 2000s followed by access to wireless internet in the late
2000s made reports of stranding events much easier and more com-
mon in developed countries. Increased photograph documentation
improved the value of reports by allowing identification of species
and estimation of life stage and physical condition.
In addition, the creation of websites aimed at sharing information
generated by citizen scientists (e.g. iNaturalist), further facilitated
the dissemination of stranding reports on a global level. The low
quality of collected data, the lack of standardized methodology and
scientific rigour are pointed out as the main challenges to using cit-
izen science (Gibson et al., 2019). However, a significant part of the
available data recovered for this study were from online platforms,
emphasizing the potential as a global data resource. Measures to en-
sure the usability and standardization of collected data would help
greatly. An overview of social media reporting trends is presented in
Table S5. It is important to mention that most of the data available
are from developed countries (i.e. Australia and the United States)
probably due Internet access. Not searching in foreign languages
may also have affected availability bias as can the socio-­economic
status of the countries. That said, documentation efforts would ben-
efit from the participation of citizens from underdeveloped and de-
veloping regions, where feasible, in order to reduce geographic bias
of shark stranding reports.
4.5 | Survival rates
Most stranded sharks were found already in a moribund condition or
dead. In addition, only a small portion that was reported as stranded
alive survived after release. As sharks perform branchial respiration,
even short periods out of water are potentially lethal. Moreover, due
to the difference in atmospheric pressure, the weight exerted by the
animal's body out of the water tends to cause irreversible damage to
the organs, leading to late mortality. Additional causes should also
be considered, including exposure to solar radiation (Wosnick et al.,
2019), thermal stress/shock and physical injury/stress.
Another point of concern is that, in most cases, shark rescue is
not carried out properly or by professionals trained to handle and
give clinical support to these animals. Furthermore, shark rescue and
rehabilitation programs are rare. In less serious cases, such as strand-
ing in tide pools, the creation and dissemination of simple protocols
can ensure survival. However, in more serious cases or for large-­
sized species, euthanasia may be necessary, emphasizing the need
for trained professionals to respond to calls, as well as veterinarians/
veterinary pathologists and other experts to perform diagnostics.
4.6 | Potential causes
4.6.1 | Infectious disease
There are only a few articles about infectious disease associated
with shark strandings, most of them suggesting meningitis or menin-
goencephalitis caused by Carnobacterium spp. (Carnobacteriaceae),
Streptococcus spp. (Streptococcaceae) or by the ciliated protozoan
Miamiensis avidus (Parauronematidae) (Carlisle et al., 2015; Dagleish
et al., 2010; Martinez-­Steele et al., 2018; Morick et al., 2020; Retallack
et al., 2019; Schaffer et al., 2012, 2013). The genus Carnobacterium
is part of the bacterial community in some teleost and sharks, but in
cases of stress-­induced immunosuppression, these bacteria can be-
come pathogenic leading to systemic infection and subsequent mor-
bidity and mortality (Schaffer et al., 2013). As for Streptococcus spp.,
WOSNICK et al.
there is one report of a fatal infection in a sandbar shark (Carcharhinus
plumbeus, Carcharhinidae) stranded in Israel (Morick et al., 2020). The
post-­mortem examination indicated meningoencephalitis caused by S.
agalactiae, which is usually associated with septicemia and meningoen-
cephalitis in teleost fish, but extremely rare in sharks.
Scuticociliates are opportunistic protozoans, also known to
cause disease in teleost fish and elasmobranchs (Retallack et al.,
2019). This pathogen was proved to cause meningoencephalitis in
leopard sharks and was associated with a mass stranding event in
California (Retallack et al., 2019). Sharks can also acquire infections
through their diets, consuming prey that were infected with patho-
gens (Carlisle et al., 2015). As the data on this matter remain scarce,
more studies are needed to fully understand the pathogenicity and
diseases that can cause or increase the chance of strandings, being
the carcasses found ashore a fundamental source of information.
4.6.2 | Abrupt environmental alterations
As described for invertebrates and other vertebrates, abrupt envi-
ronmental changes (i.e. hypoxia and sudden temperature changes)
can lead to strandings. To date, confirmed cases of abrupt environ-
mental alterations leading to shark strandings are scarce. In 2008, a
massive stranding of 26 species was recorded in Coliumo bay (Chile),
including the Chilean catshark (Hernández-­Miranda et al., 2010).
According to physiological and pathological analyses, the animals
were healthy, and the cause of death was severe hypoxia due to
an abrupt resurgence event through the entry of south-­west winds
(ESSW) into the bay. These events deserve special attention as in a
climate change scenario it can be intensified and increase mortality.
Since most sharks are ectothermic, sudden changes in tempera-
ture can be particularly harmful due to altered thermal windows and
impairment of vital functions (Bernal et al., 2018; Pegado et al., 2020).
However, even endothermic species (as is the case of some species
reported in the present study) can be harmed by abrupt alterations
as relocating metabolic substrates to maintain body temperature can
lead to immunosuppression and subsequent vulnerability to pathogens
(Carlisle et al., 2015). However, as many sharks perform horizontal
migrations on a daily basis and are therefore exposed to abrupt tem-
perature variations, further investigations into the real role of sudden
temperature changes in strandings should be carried out.
4.6.3 | Negative interaction with fisheries
Currently, fishing reports only consider what has been caught and
whether it has been returned/discarded. However, the number of
animals that were released and stranded soon after is not yet con-
sidered. Retained fishing hooks were first reported to cause major
lacerations and pathologies in blue sharks (Borucinska et al., 2002).
Mortality due to fishing gear was also reported for the longfin mako
shark (Isurus paucus, Lamnidae) (Adams et al., 2015). In the present
| 11
study, evidence of capture stress and/or interaction with fishing
gear was noted for other species, including the bluntnose sixgill
shark (Hexanchus griseus, Hexanchidae), the broadnose sevengill
shark (Notorynchus cepedianus, Hexanchidae), the common thresher
shark (Alopias vulpinus, Alopiidae) and the porbeagle shark (Lamna
nasus, Lamnidae). These results indicate that fishing is an important
source of stranding, which could significantly increase the currently
reported post-­release mortality rates and assist in the estimation of
bycatch impacts (Adimey et al., 2014; Peltier et al., 2016).
4.6.4 | Other trauma
Boat collisions are a well-­documented threat to marine life, and
cetaceans and surface-­feeding sharks seem to be more vulner-
able to these encounters (Korman, 2015; Speedie et al., 2009;
Waereebeek et al., 2007). In some cases, collisions are fatal (bask-
ing shark, Cetorhinus maximus, Cetorhinidae -­Speedie et al., 2009);
however, in most cases, sharks are not fatally injured (whale shark
-­Speed et al., 2008; great white shark, Carcharodon carcharias,
Lamnidae -­Domeier, 2012). This does not mean that late mor-
tality does not occur, since the resulting lacerations can be the
gateway to infections, or affect the animal's swimming capacity,
reducing their foraging success and increasing their susceptibility
to predators.
Bites have also been observed in some species, indicating that
prey–­
predator interactions are a significant source of stranding,
mainly for juveniles or small-­sized sharks. However, in the majority
of reports, sharks were found already dead, making it impossible to
confirm whether the bites were lethal (wash-­up carcass) or if the
sharks stranded still alive. It is known that juvenile and small-­sized
sharks can be predated by larger individuals/species and shark bites
are also a significant cause of marine mammal strandings (great
white sharks and sea otters; Moxley et al., 2019).
4.6.5 | Foraging, sandbanks and tide pools
Strand feeding is characterized by an animal coming out of the
water, no longer floating to trap their preys and optimize capture
(McGovern et al., 2019). There are many reports in the media of
sharks performing strand feeding, and one particular case involv-
ing a basking shark chasing a school of salmon has been reported
in the literature (Whitley, 1965 -­cited by Speed et al., 2009).
Sharks can also be trapped in sandbanks and tide pools, and al-
though small-­sized species are more prone to these situations,
according to our results, even reef blacktip sharks (Carcharhinus
melanopterus, Carcharhinidae) have been found trapped. In most
cases, the rescue is successful as sharks are not directly exposed
to air. However, considering that sandbanks and tide pools are
subject to abrupt abiotic fluctuations, post-­rescue mortality might
also occur.
12 | WOSNICK et al.
5 | CO N C LU S I O N S A N D FU T U R E
D I R EC TI O N S
As only reports available in open-­access databases were consid-
ered in this review, it is likely that the number of strandings is much
higher, pointing to the urgent need to establish worldwide rigorous
shark stranding monitoring programs that will record, preserve and
share comprehensive morphometric data and coordinate complete
diagnostic investigations of shark stranding events to permit for
improved study and understanding. Investigations would include
consideration of water quality parameters (e.g. temperature, salinity,
pollution and dissolved oxygen) and post-­mortem examination with
ancillary diagnostics as necessary for infectious and other disease
processes where possible.
Although sharks are much less affected by strandings than
marine mammals, this phenomenon has been systematically ne-
glected resulting in significant knowledge gaps and decades of
poor-­quality data. As a result of such neglect, responses to shark
strandings are inefficient with little chance of success in rescue
and rehabilitation. In fact, even occurring to a lesser extent, shark
mortality is much higher when compared to other taxonomic
groups that undergo stranding. Therefore, it is imperative to have
shark specialists leading rescue and rehabilitation initiatives, as
well as developing specific protocols for shark euthanasia when-
ever necessary. Unfortunately, there are great chances that post-­
stranding survival rates cannot be improved for sharks. However,
understanding the causes might help to mitigate or at least reduce
some of them, especially those of anthropogenic origin. Besides,
in a climate change scenario, shark strandings should also be bet-
ter discussed, since this phenomenon can be further aggravated,
not only by the intensification of weather and climate events on
coastal zones but also by the increase in the dispersing potential
of many pathologies.
In summary, many shark species are subject to stranding, and this
is a global phenomenon. Future studies should focus on evaluating
the patterns presented here in more detail, as well as more efforts
should be directed to the global hotspots identified in the present
study. Also, more attention must be given to sharks, through the
establishment of specific monitoring programs and the creation of
a free-­access global database. A database of professionals and ex-
perts willing to help it is also necessary, aiming to build connections,
assist rescue groups and share resources. Moreover, it is necessary
to establish shark stranding networks aiming to provide critical data
necessary for conservation of imperiled species and inform policy
on a local or regional scale, and to provide online training for citi-
zen scientists and enthusiasts. Lastly, elevating interest of society is
also crucial, as funding opportunities to improve responses to shark
strandings may arise from tax payers, officials and conservationists.
AC K N OW L E D G E M E N T S
Financial support to NW was provided by the Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (CAPES). The authors
thank Erica Padilha and Aline Cristina Prado for their help in data
collection, figure creation and data discussion. We also thank the ref-
erees for their careful and insightful reviews of our manuscript, and
Otto Bismarck Fazzano Gadig and Clinton Duffy for their expertise
in shark identification. Lastly, the authors thank designers Scarlet23,
T. Michael Keesey, rawpixel, pch.vector, Blue Vectors, macrovector,
photo3idea-­studio, Nature Vectors for the icons used in the figures
available through the Freepik, PhyloPic, Flaticon, PhyloPic and vect-
eezy websites. The authors have no conflict of interest to declare.
DATA AVA I L A B I L I T Y S TAT E M E N T
The datasets generated during and/or analysed during the current
study are available from the corresponding author on reasonable
request.
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S U P P O R T I N G I N FO R M AT I O N
Additional supporting information may be found in the online
version of the article at the publisher’s website.
How to cite this article: Wosnick, N., Leite, R. D., Giareta, E.
P., Morick, D., & Musyl, M. (2022). Global assessment of
shark strandings. Fish and Fisheries, 00, 1–­14. https://doi.
org/10.1111/faf.12648