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Wild animals are captured or taken opportunistically, and the meat, body parts, and/or eggs are consumed for local subsistence or used for traditional purposes to some extent across most of the world, particularly in the tropics and subtropics. The consumption of aquatic animals is widespread, in some places has been sustained for millennia, and can be an important source of nutrition, income, and cultural identity to communities. Yet, economic opportunities to exploit wildlife at higher levels have led to unsustainable exploitation of some species. In the literature, there has been limited focus on the exploitation of aquatic non-fish animals for food and other purposes. Understanding the scope and potential threat of aquatic wild meat exploitation is an important first step toward appropriate inclusion on the international policy and conservation management agenda. Here, we conduct a review of the literature, and present an overview of the contemporary use of aquatic megafauna (cetaceans, sirenians, chelonians, and crocodylians) in the global tropics and subtropics, for species listed on the Appendices of the Convention on the Conservation of Migratory Species of Wild Animals (CMS). We find that consumption of aquatic megafauna is widespread in coastal regions, although to varying degrees, and that some species are likely to be at risk from overexploitation, particularly riverine megafauna. Finally, we provide recommendations for CMS in the context of the mandate of the Aquatic Wild Meat Working Group.
fmars-09-837447 March 15, 2022 Time: 18:16 # 1
published: 21 March 2022
doi: 10.3389/fmars.2022.837447
Edited by:
Maria Carolina Viana,
State University of Campinas, Brazil
Reviewed by:
Hector Barrios-Garrido,
University of Zulia, Venezuela
Cristina Brito,
CHAM – Center for the Humanities
NOVA FCSH, Portugal
Daniel J. Ingram;
Daniel J. Ingram
Margi Prideaux
Nicola K. Hodgins
Isabel C. Avila
Tim Collins
Mel Cosentino
Lucy W. Keith-Diagne
Helene Marsh
Matthew H. Shirley
Yusuke Fukuda
Kerstin B. J. Glaus
Rima W. Jabado
Jeffrey W. Lang
Lindsay Porter
Specialty section:
This article was submitted to
Marine Conservation
and Sustainability,
a section of the journal
Frontiers in Marine Science
Received: 16 December 2021
Accepted: 21 January 2022
Published: 21 March 2022
Widespread Use of Migratory
Megafauna for Aquatic Wild Meat in
the Tropics and Subtropics
Daniel J. Ingram1,2*, Margi Prideaux3, Nicola K. Hodgins4*, Heidrun Frisch-Nwakanma5,
Isabel C. Avila6,7, Tim Collins8*, Mel Cosentino9, Lucy W. Keith-Diagne2,
Helene Marsh10, Matthew H. Shirley11, Koen Van Waerebeek12,13 , Maximin K. Djondo14 ,
Yusuke Fukuda15, Kerstin B. J. Glaus16, Rima W. Jabado10,17, Jeffrey W. Lang18,
Sigrid Lüber16 , Charlie Manolis19, Grahame J. W. Webb19 and Lindsay Porter20,21
1Faculty of Natural Sciences, University of Stirling, Stirling, United Kingdom, 2African Aquatic Conservation Fund, Joal,
Senegal, 3Convention on the Conservation of Migratory Species of Wild Animals Aquatic Wild Meat Working Group, Bonn,
Germany, 4Whale and Dolphin Conservation, Chippenham, United Kingdom, 5Convention on the Conservation of Migratory
Species of Wild Animals, Bonn, Germany, 6Grupo de Ecología Animal, Universidad del Valle, Cali, Colombia, 7Institute for
Terrestrial and Aquatic Wildlife Research, University of Veterinary Medicine Hannover, Foundation, Hannover, Germany,
8Global Conservation Program, Wildlife Conservation Society, Bronx, NY, United States, 9Wild Earth Foundation, Puerto
Pirámides, Argentina, 10 College of Science and Engineering, James Cook University, Townsville, QLD, Australia, 11 Institute
of Environment, Florida International University, North Miami, FL, United States, 12 Pro Delphinus, Lima, Peru, 13 Centro
Peruano de Estudios Cetológicos, Lima, Peru, 14 Benin Environment & Education Society, Cotonou, Benin, 15 Department
of Environment, Parks and Water Security, Palmerston, NT, Australia, 16 OceanCare, Wädenswil, Switzerland, 17 Elasmo
Project, Dubai, United Arab Emirates, 18 Gharial Ecology Project, Madras Crocodile Bank Trust, Mahabalipuram, India,
19 Crocodile Specialist Group, Wildlife Management International Pty. Limited, Karama, NT, Australia, 20 Southeast Asia
Marine Mammal Research, Hong Kong, Hong Kong SAR, China, 21 Scientific Committee (Small Cetaceans) of the
International Whaling Commission, Cambridge, United Kingdom
Wild animals are captured or taken opportunistically, and the meat, body parts, and/or
eggs are consumed for local subsistence or used for traditional purposes to some extent
across most of the world, particularly in the tropics and subtropics. The consumption
of aquatic animals is widespread, in some places has been sustained for millennia, and
can be an important source of nutrition, income, and cultural identity to communities.
Yet, economic opportunities to exploit wildlife at higher levels have led to unsustainable
exploitation of some species. In the literature, there has been limited focus on the
exploitation of aquatic non-fish animals for food and other purposes. Understanding
the scope and potential threat of aquatic wild meat exploitation is an important first step
toward appropriate inclusion on the international policy and conservation management
agenda. Here, we conduct a review of the literature, and present an overview of
the contemporary use of aquatic megafauna (cetaceans, sirenians, chelonians, and
crocodylians) in the global tropics and subtropics, for species listed on the Appendices
of the Convention on the Conservation of Migratory Species of Wild Animals (CMS). We
find that consumption of aquatic megafauna is widespread in coastal regions, although
to varying degrees, and that some species are likely to be at risk from overexploitation,
particularly riverine megafauna. Finally, we provide recommendations for CMS in the
context of the mandate of the Aquatic Wild Meat Working Group.
Keywords: bushmeat, cetaceans, health implications, hunting, crocodiles, sirenians, turtles, threatened species
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Ingram et al. Aquatic Wild Meat
Globally, especially in the tropics and subtropics, wild animals
are captured or taken opportunistically and the meat, body
parts, and/or eggs are consumed as food or used for traditional
remedies, cultural purposes, and religious ceremonies (Alves and
Albuquerque, 2017;Ingram et al., 2021). Increasingly, products
are sold locally, nationally, regionally, and internationally,
providing income to many communities (Coad et al., 2019;
Ingram, 2020). When harvests of wildlife are unsustainable, they
cause population declines and pose a threat to species’ survival
(McCauley et al., 2015;Benítez-López et al., 2017;He et al., 2017;
Ripple et al., 2019). Over the past few decades, a vast literature
has amassed on the harvest of wild-caught fish and terrestrial
animals, which includes quantitative studies of the magnitude
and spatial extent of harvests in some regions, sustainability
assessments, the human dimensions of wildlife use, and the
impact of overexploitation on ecosystems (Worm et al., 2009;
Lynch et al., 2016;Coad et al., 2019;Dobson et al., 2019;Ingram
et al., 2021). While the commercial targeting of some taxa has
raised serious international concerns (e.g., whaling), there has
been limited focus on the exploitation of most freshwater and
marine (hereafter “aquatic”) non-fish1animals used for food
and other purposes.
For both terrestrial and aquatic wildlife, the terminology used
to discuss harvest (also called “takes” in the wildlife management
literature, and “catches” in the fisheries literature), consumption,
and trade has changed over the past few decades. In 2000, and
largely focussing on terrestrial species, the International Union
for Conservation of Nature (IUCN) adopted the term “wild
meat” to describe the “meat and other products derived from
wild animals for human consumption,” whether legal or illegal,
to harmonise terminology across regions (Mainka and Trivedi,
2002). In 2017, both the International Whaling Commission
(IWC) and the Convention on the Conservation of Migratory
Species of Wild Animals (CMS) adopted the term “aquatic
wild meat,” defined as “products obtained through all forms of
take, including unregulated, legal, or illegal hunts as well as
deliberate or opportunistic catches from stranded (dead or alive)
and/or bycaught (also known as incidental catch) individuals” to
similarly harmonise this issue for aquatic non-fish species (CMS,
2017a,b;IWC, 2018).
The consumption of aquatic animals is widespread, and is
an important source of nutrition, income, and cultural identity
for many communities (Robards and Reeves, 2011). Considering
aquatic wild meat as all aquatic non-fish vertebrates that are
used by humans for food, medicine, or handicrafts, the IUCN
Red List of Threatened Species identifies 672 species of birds,
322 amphibians, 137 reptiles, and 112 mammals as aquatic
species that are used or traded (IUCN, 2021). Harvest levels
may be high or low, but many are unregulated and unmanaged.
Growing human populations, improved animal hunting and
capture methods, expanded market access, and an escalating
1We use ‘fish’ to denote traditional fisheries products, including fishes and
commercially exploited crustaceans, squids, bivalves, sea cucumbers, and
gastropods, but excluding chondrichthyans (sharks, rays, and chimaeras) for policy
reasons as directed by CMS.
demand for animal-sourced proteins, has increased exploitation
levels and, in some areas for at least some species, unsustainable
harvests are now evident (Milner-Gulland et al., 2003;Cawthorn
and Hoffman, 2015;Coad et al., 2019). Overarching guidance
regarding aquatic wild meat in international policy remains
limited for most species. Understanding the scope and potential
threat of overexploitation of aquatic wild meat species is an
important first step toward developing effective international and
domestic policy.
The CMS has provided some guidance on the use of migratory
species, defined in the Convention text as “the entire population
or any geographically separate part of the population of any
species or lower taxon of wild animals, a significant proportion
of whose members cyclically and predictably cross one or more
national jurisdictional boundaries.” CMS provides a way for
countries to cooperate to ensure species’ migrations can still
continue without barriers, and to address issues such as habitat
destruction and exploitation. As of 2020, 132 countries have
become Parties to the CMS Convention and an additional 65
countries (non-Parties) are signatories to CMS Agreements and
Memoranda of Understanding (Supplementary Material 1). In
2017, the CMS Scientific Council established an Aquatic Wild
Meat Working Group to focus on the exploitation of aquatic
animals (CMS, 2017b). We build here on information that
was prepared for the establishment of the Working Group,
and recognising the growing breadth and depth of this issue,
present an overview of the use of aquatic mammals (specifically
cetaceans and sirenians), turtles, and crocodylians in the tropics
and subtropics (Figure 1).
Many cultures have used aquatic megafauna throughout
history (Robards and Reeves, 2011;Cosentino and Fisher,
2016;Porter and Lai, 2017). Here we focus on contemporary
practises and the degree of conservation concern associated with
them. Specifically, we examine 37 species listed on the CMS
Appendices as endangered (CMS Appendix I) and/or requiring
international cooperation for their conservation (CMS Appendix
II and Table 1) that have been identified as being used for
aquatic wild meat. This review is a first step toward increasing
our understanding of aquatic wild meat use and its impacts,
particularly of aquatic megafauna, and we emphasise the need
for similar efforts for the hundreds of other species harvested for
aquatic wild meat.
The Americas and the Caribbean
The capture, killing, and trade of cetaceans and their products are
illegal in most tropical American countries. Despite this, use for
food and other purposes occurs in many countries, although the
drivers and magnitude of the exploitation vary markedly between
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Ingram et al. Aquatic Wild Meat
FIGURE 1 | Migratory megafauna used for aquatic wild meat, including: (A) Atlantic humpback dolphin (Sousa teuszii), Conkouati-Douli National Park, Republic of
the Congo; (B) African Manatee (Trichechus senegalensis), Lagos Lagoon, Nigeria; (C) Green turtle (Chelonia mydas), Joal, Senegal; and (D) Saltwater crocodile
(Crocodylus porosus) egg collection, Northern Territory, Australia. Photo credits: Tim Collins (A), Christogonus Uzoma Ejimadu (B), Pearson McGovern, African
Aquatic Conservation Fund (C), and Yusuke Fukuda (D).
regions. In some countries (e.g., Argentina), uses are rare events
and are almost exclusively opportunistic, while in others (e.g.,
Peru and Ecuador), there is continued use of cetaceans in spite of
existing legislation, especially as bait in a number of fisheries (Van
Waerebeek and Reyes, 1994;Vidal et al., 1994;Van Waerebeek
et al., 1999, 2018;Avila et al., 2008;Crespo, 2009;Dawson, 2009;
Flores and Da Silva, 2009;Goodall, 2009;Cosentino and Fisher,
2016;Campbell et al., 2020a,b;Castro et al., 2020). There are
marked differences in the target species and methods used to
capture them, both between and within countries. For example,
botos (Inia geoffrensis) and tucuxis (Sotalia fluviatilis) are illegally
harvested for use as bait for catfish (Calophysus macropterus)
in Brazil, Peru, Colombia, and Venezuela (Flores et al., 2008;
Trujillo et al., 2010;Mintzer et al., 2012;Campbell et al., 2020b)
resulting in serious population declines (e.g., even in the more
protected part of their range in Brazil; Martin and Da Silva, 2021),
while dolphins off the coasts of Chile (Van Waerebeek et al.,
1999) and Argentina are rarely recorded as taken incidentally
or directly for either bait or human consumption. Common
bottlenose dolphins (Tursiops truncatus) and the pantropical
spotted dolphin (Stenella attenuata) have been hunted (at least 13
dolphins/year) for bait by local long-line fishers in the northern
Colombian Pacific (Avila et al., 2008). In Chilean Patagonia,
Peale’s dolphins (Lagenorhynchus australis) used to be hunted for
crab bait, although changes in the dynamic of this fishery may
have reduced their direct harvest for this purpose (Lescrauwaet
and Gibbons, 1994;Goodall, 2009).
The largest exploitation of small cetaceans traditionally
occurred in Peru, where they have been caught (intentionally
and otherwise) in artisanal gillnets and by harpooning for
decades, and sold in local markets or transported to the capital
Lima (Read et al., 1988;Van Waerebeek and Reyes, 1994;Van
Waerebeek et al., 1999). The use of dolphins as bait has spread
to all coastal areas, with annual catches increasing since the
early 2000s (Mangel et al., 2010;Tzika et al., 2010;Campbell
et al., 2020a). The use of small cetaceans as bait, including
dusky (Lagenorhynchus obscurus), common (Delphinus delphis),
common bottlenose dolphins, as well as Burmeister’s porpoises
(Phocoena spinipinnis) (e.g., in longline and gillnet shark fisheries
in coastal Peru) appears prolific and has now largely replaced use
for human consumption (Van Waerebeek et al., 1999, 2018;Van
Waerebeek and Würsig, 2009;Mangel et al., 2010;Campbell et al.,
2020a). Nonetheless, dolphin meat was still widely available in
Peruvian fish markets until at least 2010, but increasingly covertly
(Mangel et al., 2010;Tzika et al., 2010). A study of 942 beach-
cast small cetaceans between 2000 and 2017 showed that use still
continues in central and southern Peru, although the high level of
stranded bycaught animals that do not show signs of butchering
suggests it may occur on a significantly lower scale compared to
what was observed in the 1980s and 1990s (Van Waerebeek et al.,
1999, 2018).
The use of small cetaceans in other tropical American
countries is less widespread. Cetacean harvests were reported for
Venezuela 20 years ago (Romero et al., 2001), and exploitation
for meat and bait has been more widespread than commonly
thought (affecting 11 of 20 species), though it is poorly
documented (Romero et al., 1997). Recent data are scarce,
but Sotalia guianensis has been confirmed as a target species
(Barrios-Garrido et al., 2021;Briceño et al., 2021). Recent
studies in Ecuador have revealed the use of the CMS-listed
Eastern Tropical Pacific (ETP) population of pantropical spotted
dolphins as bait (Castro et al., 2020). In Ecuador, fishers use
body parts of aquatic mammals, including dolphins, as bait in
improvised Fish Aggregating Devices (FADs; Castro et al., 2020).
Several species are involved in this illegal practice, caught both
accidentally and via direct targeting, but the extent of harvest is
unknown. CMS-listed species of concern include botos, Guiana
dolphins, dusky dolphins, Burmeister’s porpoise, and common
bottlenose dolphins.
Captures of several species of cetaceans have been
documented throughout the Caribbean including in the
Dominican Republic, St Vincent and the Grenadines (SVG),
Trinidad and Tobago, St Lucia, and Dominica (Vidal et al.,
1994;Bolaños-Jiménez et al., 2014). CMS-listed species
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TABLE 1 | CMS Species of Concern by CMS Appendices and Region.
Species Africa Asia The Americas and
the Caribbean
Oceania CMS Appendix IUCN Category
Atlantic humpback dolphin (Sousa teuszii) X I and II CR
Australian snub-fin dolphin (Orcaella heinsohni) X II VU
Amazon river dolphin/Boto (Inia geoffrensis) X II EN
Burmeister’s porpoise (Phocoena spinipinnis) X II NT
Commerson’s dolphin (Cephalorhynchus commersonii) X II (South American population) LC
Clymene dolphin (Stenella clymene) X X II (West African population) LC
Dall’s porpoise (Phocoenoides dalli) X II LC
Dusky dolphin (Lagenorhynchus obscurus) X II LC
Franciscana (Pontoporia blainvillei) X I and II VU
Fraser’s dolphin (Lagenodelphis hosei) X II (Southeast Asian populations) LC
Ganges River dolphin (Platanista gangetica) X I and II EN
Guiana dolphin (Sotalia guianensis) X II NT
Heaviside’s dolphin (Cephalorhynchus heavisidii) X II NT
Humpback whale (Megaptera novaeangliae) X X X X I LC (Oceania
sub-population EN)
Indo-Pacific bottlenose dolphin (Tursiops aduncus) X II (Arafura/Timor Sea
Indo-Pacific finless porpoise (Neophocaena phocaenoides) X II VU
Indo-Pacific humpback dolphin (Sousa chinensis) X II VU
Irrawaddy dolphin (Orcaella brevirostris) X I and II EN
Orca (Orcinus orca) X II DD
Pantropical spotted dolphin (Stenella attenuata) X X II (eastern tropical Pacific and
Southeast Asian populations)
Peale’s dolphin (Lagenorhynchus australis) X II LC
Spectacled porpoise (Phocoena dioptrica) X II LC
Spinner dolphin (Stenella longirostris) X X X II (eastern tropical Pacific and
Southeast Asian populations)
Tucuxi (Sotalia fluviatilis) X II EN
African manatee (Trichechus senegalensis) X I and II VU
Amazonian manatee (Trichechus inunguis) X II VU
Dugong (Dugong dugon) X X II VU
West Indian manatee (Trichechus manatus) X I and II (populations between
Honduras and Panama)
Green turtle (Chelonia mydas) X X X X I and II EN (North Indian Ocean
sub-pop VU)
Hawksbill turtle (Eretmochelys imbricata) X X X X I and II CR
Kemp’s Ridley turtle (Lepidochelys kempii) X X I and II CR
Leatherback turtle (Dermochelys coriacea) X X X X I and II VU
Loggerhead turtle (Caretta caretta) X X X X I and II VU
Olive Ridley turtle (Lepidochelys olivacea) X X X X I and II VU
South American river turtle (Podocnemis expansa) X I (Upper Amazon populations)
and II
Gharial (Gavialis gangeticus) X I CR
Salt-water crocodile (Crocodylus porosus) X X II LC
IUCN categories listed include: Not Evaluated (NE), Data Deficient (DD), Least Concern (LC), Near Threatened (NT), Vulnerable (VU), Endangered (EN), and Critically
Endangered (CR).
harvested include Fraser’s (Lagenodelphis hosei), spinner
(Stenella longirostris), common (Delphinus delphis), and clymene
dolphins (S. clymene), orcas (Orcinus orca), and humpback
whales (Megaptera novaeangliae). Except for SVG, direct
harvesting for large whales, such as humpback whales, is scarce
in tropical America (Williams-Grey et al., 2014). However, there
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Ingram et al. Aquatic Wild Meat
are some recent records of opportunistic hunting, for example in
2004, in the Colombian Pacific, a humpback whale calf, which
was alone and apparently lost, was killed by artisanal fishermen
for consumption purposes (Avila et al., 2013). In Barrouallie in
SVG, cetaceans are regularly hunted, with records of between
203 and 927 individuals harvested each year by a single whaler
(Fielding and Kiszka, 2021). In 2019, reports presented to the
Scientific Committee of the IWC raised concerns as it was
estimated that the current levels of hunting of short-finned pilot
whales (Globicephala macrorhynchus) and orcas in the waters of
SVG were unsustainable (Fielding, 2018;IWC, 2019).
Manatees have been historically hunted throughout their range
in the Americas and the Caribbean. Direct harvest of the
Amazonian (Trichechus inunguis) and the West Indian manatees
(T. manatus) have been recorded in at least 14 countries.
Both species are mainly used for food, but also for other
purposes, including remedies and traditional ceremonies (Braga
Ferreira et al., 2011;Cosentino and Fisher, 2016). Although
most manatees are harvested opportunistically (e.g., as bycatch),
manatee hunters may still be active in Brazil (Souza et al.,
2014), Colombia (Arévalo-González et al., 2010;Cruz-Antía
and Gómez, 2011;Kiszka, 2014), Ecuador (Denkinger, 2010;
Tirira, 2011), and Peru, where Amazonian manatee calves
are also captured alive to be “grown” in ponds for food
(Perea-Sicchar et al., 2011).
In the past, Amazonian manatees have experienced high
hunting pressure. In the central-western Brazilian Amazon,
trade in manatee hides led to the hunting of 113,000
animals between 1904 and 1969 (Antunes et al., 2016). This
practice no longer occurs (Antunes et al., 2016), but illegal
hunting for meat, largely conducted with traditional harpoons, is
considered among the main contemporary threats to the species
(Marmontel et al., 2016), and occurs throughout the species’
range: Brazil (Calvimontes, 2009;Franzini et al., 2013;Souza
et al., 2014;Pantoja, 2015;Crema et al., 2020); Colombia (Kendall,
2013), Peru (Soto, 2007;Kendall, 2013;Kirkland et al., 2020); and
Ecuador (Denkinger, 2010). Meat is consumed or sold locally
(Marmontel et al., 2016), but can also be marketed in larger
towns and cities (e.g., Franzini et al., 2013;Souza, 2015;El Bizri
et al., 2020). Although not considered a substantial threat to
the species, recent uses of Amazonian manatee for traditional
remedies have been reported in the Caatinga region of Brazil
by Barbosa et al. (2018). Although Amazonian manatee is not
reported to occur in this region, it is plausible that these reports
refer to the West Indian manatee. Changes to water levels as a
result of dam construction is amplifying hunting impacts on the
Amazonian manatee because migrating animals become more
available to hunters in narrow passages along migration routes
(Arraut et al., 2010, 2017).
West Indian manatees are also hunted opportunistically (e.g.,
as bycatch in fishing nets) and directly targeted throughout the
Caribbean, in the Magdalena and Orinoco River systems, and the
Maracaibo Lake system (Montiel-Villalobos and Barrios-Garrido,
2005;Romero and Creswell, 2005;Vail, 2005;Castelblanco-
Martínez et al., 2009, 2016). They were hunted historically for oil,
bones, and meat (e.g., O’Shea et al., 1988), but today hunting is
exclusively for meat, a practice considered a significant threat in
Brazil, Colombia, Costa Rica, Cuba, Dominican Republic, French
Guiana, Guatemala, Honduras, Mexico, Suriname, Trinidad,
and Venezuela (Self-Sullivan and Mignucci-Giannoni, 2012;
Alvarez-Alemán et al., 2021). Killing of bycaught animals for
consumption occurs in Colombia, Dominican Republic and Haiti
(Kiszka, 2014). Although numbers of manatees killed by hunting
in Panama are low, it still occurs (Muschett, 2008;Quintana-
Rizzo and Reynolds, 2010). West Indian manatees in Colombia
have been killed and sold locally for food or bait, mainly via direct
hunting, for decades (Montoya-Ospina et al., 2001;Castelblanco-
Martínez et al., 2009).
All six marine turtle species in the Americas and the Caribbean
are considered threatened according to the IUCN Red List of
Threatened Species (green turtle, Chelonia mydas, hawksbill
turtle, Eretmochelys imbricata, Kemp ridley turtle, Lepidochelys
kempii, leatherback turtle, Dermochelys coriacea, loggerhead
turtle, Caretta caretta, and olive ridley turtle, Lepidochelys
olivacea;IUCN, 2021;Table 1). While the threats vary by species,
exploitation for meat, eggs, and carapaces (e.g., hawksbill turtles)
at varying levels is ubiquitous and considered a significant threat
(Amorocho et al., 2016;Nahill et al., 2020).
In the Caribbean, marine turtles have been harvested by
people continuously, with commercial exploitation and trade
from Cuba for 500 years. Marine turtle harvests were managed
as a commercial fisheries resource in Cuba (unique for the
region) until recently (Carrillo et al., 1999;Fleming, 2001),
and some level of regulation of harvests occurred in other
countries: Cayman Islands (Aiken et al., 2001;Bell et al., 2006),
Turks and Caicos (Richardson et al., 2009), the British Virgin
Islands, Anguilla and Montserrat (Richardson et al., 2006), and
Nicaragua (Garland and Carthy, 2010). Contemporarily, marine
turtle bycatch and direct harvest for subsistence (mainly meat
and medicine) and domestic commercial purposes (mainly eggs,
medicine, and hawksbill shell products) still occurs in parts
of Brazil (Guebert et al., 2013), Costa Rica (Chacón-Chaverri
and Eckert, 2007;Mejías-Balsalobre et al., 2021), Colombia
and Venezuela (Barrios-Garrido et al., 2018), Mexico (Mancini
and Koch, 2009;Senko et al., 2014), Nicaragua (Garland and
Carthy, 2010;Lagueux et al., 2014, 2017;Ankersen et al.,
2015), and several Caribbean islands (Richardson et al., 2006,
2009;Amorocho et al., 2016). Together, the Caribbean has one
of the highest global legal marine turtle harvests. Nicaragua
accounted for 22.3% of the reported global legal harvest of
marine turtles in 2013, with an estimated 9,413 marine turtles
harvested per year, mostly green turtles (Humber et al., 2014).
However, the full extent of the global harvest is difficult
to estimate because many uses by Indigenous Peoples and
Local Communities (IPLCs) are not reported (but see Barrios-
Garrido H.A. et al., 2020). The Turks and Caicos Islands
has the largest documented legal hawksbill turtle fishery in
the western Atlantic, with approximately 114–277 hawksbill
turtles harvested annually, as well as 176–324 green turtles
(Stringell et al., 2013).
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In Colombia, Nicaragua, and Panama, use and trade in
turtle meat, eggs, and handicrafts (from hawksbill shell plates)
is illegal, but reasonably common. It mainly satisfies domestic
consumption demand (e.g., Chacón, 2002), but with some
international flow between countries in the region (e.g.,
Costa Rica, Ecuador, El Salvador, Guyana, Honduras, Mexico,
Suriname, and Venezuela), especially for artefacts made from
hawksbill shell plates (Pilcher and Williams, 2018). In Mexico,
marine turtles have been exploited as a traditional activity for
centuries in Baja California (Caldwell, 1963;Nabhan, 2003) and
along the eastern Pacific coast (Webb and Carrillo, 2000), despite
their fully protected status. Here, turtle hides are still locally cured
for leather, for use as cowboy boots (Kemf et al., 2000). During the
commercial meat fisheries of olive ridleys in Mexico, in the 1970s
and 1980s, turtle skin for leather was a significant byproduct
traded internationally (WWF, 2004). Throughout the Caribbean,
current harvests on nesting beaches, for meat, oil, and eggs,
are reported as contributing to declines in nesting populations
or constraining recovery: Grenada and Guyana (Eckert, 2001),
Dominican Republic (WWF, 2004), Belize (Meerman, 2005), and
Cuba (Amorocho et al., 2016). Tortuguero, Costa Rica, hosts
the largest recovered green turtle rookery in the Atlantic basin
(Troëng and Rankin, 2005). In the Guianas, green turtle status is
adversely affected by illegal harvest of eggs and nesting females,
and by bycatch from near and off-shore fisheries (Amorocho
et al., 2016). Relatively high rates of marine turtle bycatch also
occur in Suriname fisheries (Amorocho et al., 2016). Prolific
historical commercial exploitation devastated the abundance of
nesting turtles in the British Virgin Islands (Richardson et al.,
2006), Jamaica (Amorocho et al., 2016), and the Cayman Islands
(Fleming, 2001). While population recoveries have occurred
in recent years, the abundance of nesting turtles remains a
small fraction of what existed historically (Blumenthal et al.,
2021). That the unregulated, historical, large-scale commercial
harvest was demonstrably unsustainable is clear. The impact
on recovering populations of the widespread low level of use
occurring now, and whether they constitute serious threats to
turtle populations, remains largely unquantified.
In spite of protective legislation, marine turtle eggs are still
collected for subsistence and commercial use (Eckert, 2001). For
example, signs of illegal egg harvesting were observed on 51.1%
of surveys in Tortuguero, Costa Rica (González and Harrison,
2012). However, nesting at the green turtle rookery increased
by 417% between 1971 and 2003 (Troëng and Rankin, 2005).
Caribbean Nicaraguans have legally harvested green turtles for
more than 400 years, and continue to rely on the species as an
available and inexpensive source of protein and income. Over
8,000 green turtles are harvested annually along the Miskito coast
of Nicaragua (Lagueux et al., 2014, 2017), and current rates of
harvest may be unsustainable (Campbell and Lagueux, 2005;
Lagueux et al., 2005). How the increases in green turtle nesting
at Tortuguero in Costa Rica are related to the turtle harvest
in adjoining Nicaragua is largely unknown. Indeed, research is
needed on harvested and source populations of green turtles in
the Caribbean (Garland and Carthy, 2010).
In the Guajira Peninsula (between Colombia and Venezuela),
the Wayuu indigenous people consider marine turtles the most
important animal sustaining their culture, and today, there is
a significant subsistence and commercial harvest of green and
hawksbill turtles by the Wayuu community (Barrios-Garrido,
2012;Barrios-Garrido et al., 2017, 2018;Barrios-Garrido H.A.
et al., 2020). Barrios-Garrido H.A. et al. (2020) estimated
3,402 ±247 turtles/year were extracted in the study area (10
ports). Harvests occur in Colombia and Venezuela, where the
legal framework is unclear, for both domestic and international
trade (Bräutigam and Eckert, 2006;Barrios-Garrido et al., 2017).
In Peru, turtle consumption has been historically important in
the Pisco-San Andres area, for example in 1987, up to 110 fishing
boats were recorded landing turtles in San Andrés, the highest
level of marine turtle exploitation recorded in Peru (Quiñones
et al., 2010). Marine turtle harvest was a legal activity until 1995,
when it was completely banned, for all species, by Peruvian
legislation (Morales and Vargas, 1996). However, the exploitation
continues in Peru’s coastal waters, as bycaught and targeted
harvesting (Quiñones et al., 2017;Alfaro-Shigueto et al., 2018).
In Brazil, marine turtles were exploited historically, until it was
banned in 1986 (Marcovaldi and dei Marcovaldi, 1999). Illegal
harvesting still occurs in some coastal areas (Guebert et al., 2013).
Inland, the South American giant river turtle (Podocnemis
expansa) is harvested for meat, eggs, and oil in the Amazon,
Orinoco and Essequibo basins (Tortoise and Freshwater Turtle
Specialist Group, 1996). The species has been commercially
traded since at least the 1800s (Forero-Medina et al., 2021).
Current harvests occur in Brazil (e.g., Lopes et al., 2012),
Bolivia (Conway-Gómez, 2007), Colombia (Gómez et al., 2016),
Ecuador (De la Montaña, 2013), Peru (Pineda-Catalan et al.,
2012), and Venezuela (Peñaloza et al., 2013). The harvests are
mainly for meat, an important source of protein for remote
communities, but also for traditional remedies (Gómez et al.,
2016). Trade is largely domestic, from rural to urban areas (e.g.,
in Peru; Pineda-Catalan et al., 2012). Despite the harvest being
sustained over time, it is considered likely that wild populations
have declined relative to historical abundance, but quantifying
status – current abundance relative to historical abundance –
is logistically challenging (Mogollones et al., 2010;Forero-
Medina et al., 2021). The current Red List assessment (Least
Concern/Conservation Dependent), may need to be reviewed
if additional data on declines become available (Tortoise and
Freshwater Turtle Specialist Group, 1996).
Although demand for chelonian meat and eggs remains
strong throughout the tropical Americas and the Caribbean,
conservation successes based on the implementation of national
management programmes are evident. For example, the Kemp’s
ridley turtle was seriously depleted and near extinction in
the second half of the 20th century. After more than two
decades of protection, and some management interventions,
an exponential increase of adult females on nesting beaches in
Mexico occurred (Webb and Carrillo, 2000;WWF, 2004;Shaver
and Rubio, 2008), however, this has declined in recent years
(Bevan et al., 2016). In Costa Rica, a managed olive ridley egg
harvest programme was introduced at Ostional in 1987. It is
a legal, commercial harvest, run by the local people through a
community cooperative, which has been sustained and benefits
both turtles (through increased stewardship) and local people
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(Campbell et al., 2007). After 35 years of sustainable operation,
it provides a unique opportunity to re-examine sustainable
harvest rates. There is no evidence that it has stimulated a
nationwide market for eggs illegally extracted outside Ostional
(Sardeshpande and MacMillan, 2019), although such possibilities
are difficult to test (but see Pheasey et al., 2020;Mejías-
Balsalobre et al., 2021). In Brazil, Projeto TAMAR-IBAMA
(Tartarugas Marinhas) involves a marine turtle conservation
programme initiated in 1980, which reduced the harvest of
gravid nesting females and their eggs in all major nesting areas
(Marcovaldi and dei Marcovaldi, 1999). Local participation of
the fishing villages, including the employment of former egg
harvesters to patrol the beaches and protect the nests, along with
education programmes and ecotourism, have all contributed to
the programme’s success.
Commitments to chelonian conservation generally in the
Americas and Caribbean have been strengthened under several
international policy instruments (although challenges still exist,
e.g., Barrios-Garrido et al., 2019), for example: the banning
of international trade under the Convention on International
Trade in Endangered Species of Wild Fauna and Flora (CITES);
bycatch reduction under the Inter-American Tropical Tuna
Commission (IATTC); adaptation to climate change under
the Inter-American Sea Turtle Convention (IAC); and the
activities of the Permanent Commission for the South Pacific
and the Protocol Concerning Specially Protected Areas and
Wildlife (SPAW).
There is evidence of the use of cetaceans in most countries
in tropical Africa, with meat and other body parts used for
human consumption, shark bait, traditional medicine, and other
purposes (Van Waerebeek et al., 2003;Clapham and Van
Waerebeek, 2007;Weir et al., 2010;Robards and Reeves, 2011;
Weir and Pierce, 2012;Cosentino and Fisher, 2016). Dolphins
are both intentionally hunted and landed as bycatch in artisanal
gillnets, drift gillnets, beach seines, and other fishing gear.
The available data from West and Central Africa are limited,
but recent records indicate small cetaceans are consumed
as food in Benin (Sohou et al., 2013), Cameroon (Ayissi
et al., 2011, 2014), Guinea (especially Atlantic humpback
dolphin, Sousa teuszii, and common bottlenose dolphin; Van
Waerebeek et al., 2003, 2017;Bamy et al., 2021), Guinea-Bissau
(Leeney et al., 2015), Nigeria (Uwagbae and Van Waerebeek,
2010;Van Waerebeek et al., 2017), Republic of the Congo
(Collins et al., 2010, 2019), Democratic Republic of the Congo
(Collins et al., 2019), Senegal and The Gambia (Maigret, 1994;
Murphy et al., 1997;Van Waerebeek et al., 2000, 2003;Leeney
et al., 2015;Keith-Diagne et al., 2017), and Togo (Segniagbeto
et al., 2014). Historically whaling was important on the Cape
Verde Islands (Brito et al., 2016), and some consumption
of cetacean meat remains. However, while a recent review
(Segniagbeto et al., 2019) indicates instances of consumption
of pilot whales and stranded melon-headed whales, and use
of teeth for manufacture of local jewellery, there are no signs
of any systematic utilisation on the Cape Verde Islands. This
conclusion coincides with Hazevoet et al. (2010) who stated
that only few bycatches or purposeful catches had come to
their attention. Recent evidence also suggests that opportunistic
harvest of whales occurs on Annobón (Equatorial Guinea),
although rarely, and includes large cetaceans (preferentially
calves), such as humpback whales (Fielding and Barrientos,
2021), and possibly São Tomé and Príncipe (Brito et al.,
2010). In Ghana at least 16 cetacean species are used as
aquatic wild meat and, in some years, more than one
thousand individuals are landed (Ofori-Danson et al., 2003;
Van Waerebeek et al., 2009, 2014;Debrah et al., 2010). In
some countries, including Ghana, as demand increased for
dolphin meat, for human consumption or shark bait, bycatch
gradually transformed into targeted harvesting (Ofori-Danson
et al., 2003). Ghanaian artisanal fishers, operating in Togolese
coastal waters, are thought to promote trade and consumption
of cetacean meat (Segniagbeto et al., 2014). Smoked cetacean
meat from coastal Togo is traded far from the coast as
wild meat in northern Togo, Burkina Faso, Niger, and Mali
(Segniagbeto et al., 2014). In the western most countries, The
Gambia and Senegal, dolphin meat and oil is also used in
traditional remedies (Madge, 1998;Leeney et al., 2015). It is
likely that cetaceans are consumed throughout the Gulf of
Guinea, despite the lack of specific records (Clapham and Van
Waerebeek, 2007;Collins et al., 2010;Robards and Reeves,
2011;Van Waerebeek et al., 2017). The Atlantic humpback
dolphin is considered disproportionately impacted by wild
meat harvest, because of its small population size (IWC,
2019), inshore habitat use, and high vulnerability to capture
in small-scale coastal fisheries (Van Waerebeek et al., 2017;
Bamy et al., 2021).
In Eastern Africa, there are a number of records of
opportunistic use, bycatch, and intentional harvests of dolphins.
In Mozambique, there is concern that increasing intentional
harvest has evolved from commercial use of bycaught animals
(Guissamulo and Cockcroft, 1997). Stranded cetaceans here are
also used for meat (Reeve-Arnold et al., 2020). Bycatch and
direct harvest, possibly for shark bait, have been reported on
the Kenyan coastline (Mwango’mbe et al., 2021), and for human
consumption and bait in Tanzania (Braulik et al., 2018). In
Zanzibar, stranded large cetaceans are utilised as food when
fresh, and when decomposed, for the extraction of oil and for
use in traditional remedies. Small cetaceans are largely used
as bait, rather than for human consumption or other uses
(Amir et al., 2012). Direct harvests of dolphins have been
reported historically around the Seychelles islands group (De
Lestang, 1993), although no recent data confirm this is ongoing.
Direct exploitation of small cetaceans is regularly reported
from the western coast of Madagascar, mostly Indian Ocean
humpback dolphin (Sousa plumbea), bottlenose dolphins (likely
Tursiops truncatus), and spinner dolphins (Andrianarivelo, 2001;
Razafindrakoto et al., 2004;Cerchio et al., 2009a,b;Temple et al.,
2019). Between 2000 and 2018, more than 2,750 small cetaceans,
comprising at least seven species, were intentionally caught or
bycaught by 10 fishing communities in southern Madagascar
(Andrianarivelo et al., 2019).
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African manatees (Trichechus senegalensis), distributed
exclusively in West and Central Africa, are legally protected in
all 21 countries in which they occur. Domestic trade for food,
traditional medicine, and other purposes has been recorded
historically and continues to some degree in all countries (Reeves
et al., 1988;Powell, 1996;Akoi, 2004;Dodman et al., 2008;Keith
Diagne, 2014, 2015;Bachand et al., 2015;Mayaka et al., 2015,
2019;Kamla, 2019). In the past, manatee hunting was conducted
by specialised hunters, but more recently also by fishers and
generalist hunters. Consumption of bycaught animals also occurs
widely in both coastal and inland regions, largely driven by
poverty (Dodman et al., 2008;Bachand et al., 2015;Keith Diagne,
2015;Kamla, 2019). Manatee meat is also sold in markets and
restaurants in urban centres, which has increased its commercial
value (Thibault and Blaney, 2003;Fa et al., 2006;Mvele and
Arrowood, 2013).
The eastern coast of the African mainland and nearby islands
north of 25oS are the south-western extreme of the distribution
range of the dugong (Dugong dugon). Historical dugong hunting
with spears and nets has been recorded in Kenya and Somalia
(Husar, 1975), and probably occurred throughout East Africa
(Muir and Kiszka, 2012). Dugong hunting is banned in most
African countries, and dugongs are now rarely hunted due to
population declines. Poor enforcement enables meat to be sold
from dugongs that are caught incidentally (Marsh et al., 2011) and
recent evidence indicates occasional dugong hunting still occurs
in Kenya, Tanzania, and Mozambique (Muir and Kiszka, 2012).
Some Kenyan and Tanzanian coastal communities use dugong
meat, bones, and oil for food, medicine, and ornamental purposes
(Wamukoya et al., 1995;Marsh et al., 2002, 2011;Muir and
Kiszka, 2012;Said et al., 2020). In Bazaruto Bay, Mozambique,
which supports the most important dugong population in the
region (Marsh et al., 2011), the illegal directed harvest of dugongs
is estimated at 4–6 dugongs per year (Findlay et al., 2011).
Bycaught animals are sold, albeit illegally, because they represent
a windfall for impoverished fishers (Marsh et al., 2011). In
northern Madagascar, hunting for meat is thought to have been
intense historically (Durbin and Ralambo, 1994), although is
considered to have declined in recent decades due to dugong
population decline (Humber et al., 2015). Hunting probably still
occurs along the west coast of Madagascar (Cerchio et al., 2012),
Comoros, and the Department of Mayotte (French territory)
(Muir and Kiszka, 2012).
Marine turtles are also harvested to different degrees for
food and traditional remedies along the West and Central
African coast, including The Gambia, Gabon, Guinea-Bissau,
Sierra Leone, Ghana, Togo, Mauritania, Benin, Cape Verde,
Senegal, Côte d’Ivoire, Guinea, and the Republic of Congo
(CMS, 2000;Thibault and Blaney, 2003;Bal et al., 2007;Fretey
et al., 2007;Catry et al., 2009;Hancock et al., 2017). Marine
turtle exploitation (both illegal, and legal for Urekan people
permitted with a quota) is particularly intensive on Bioko
Island, Equatorial Guinea (Lepidochelys olivacea,Chelonia mydas,
Dermochelys coriacea, Eretmochelys imbricata), with products
sold at local and urban markets (Tomás et al., 2010), São
Tomé and Príncipe (C. mydas, L. olivacea;Veríssimo et al.,
2020), and Senegal (McGovern et al., 2021). For example, in
São Tomé and Príncipe, 25% of rural and 32% of urban
respondents in a survey had consumed marine turtle meat
in the past year (Veríssimo et al., 2020). Catch of marine
turtles in Nigeria is estimated to be in the thousands of
individuals annually, and high numbers of turtle eggs are
harvested (Lewison and Moore, 2012). Reductions in turtle
exploitation in Bioko Island occurred in years when beach
patrols were implemented, but this requires consistent funding
(Tomás et al., 2010).
Harvest of marine turtles and their eggs is also prevalent in
coastal East Africa. In Kenya, it has been estimated that 10–
50 percent of nesting turtles and their eggs are harvested to
supply underground markets (Nzuki, 2004). About 10 percent
of turtle products available in the Tana Delta and Malindi
are thought to have been supplied by foreign fishers, mostly
from Somalia and Tanzania (Nzuki, 2005). In Tanzania, turtle
products were being sold both openly and covertly at landing
sites in Dar es Salaam (West, 2008), and two assessments (2015
and 2016) conducted in the Lindi Region confirmed a directed
fishery was capturing green turtles for both local consumption
and commercial trade (IOSEA, 2019). Egg collection persists
in Tanzania, although on Mafia Island, nest monitoring, nest
protection payments, and education programs about sea turtle
conservation resulted in egg harvest declining from 100% of
eggs (prior to programme initiation in 2001) to 1% (2004) and
4% (2005) (Ferraro, 2007). In Mozambique, marine turtles are
caught by specialist turtle hunters and by some artisanal fishers
(Williams, 2017, 2021;Pilcher and Williams, 2018). Retained
turtles are most commonly consumed domestically for meat,
occasionally for local remedies/cultural practises, or as bait for
fishing. Trade in turtle meat is often clandestine, but turtle
shells are readily available in craft markets (>200 products
observed between 2017 and 2020 [exact survey period unknown];
Nahill et al., 2020). Surveys across three coastal provinces in
Mozambique, although crude, suggested the directed and bycatch
harvest of turtles could be as high as 800,000 turtles per year,
of which ~200,000 may be consumed or traded and many are
reportedly released alive (Pilcher and Williams, 2018).
Illegal domestic use and trade of marine turtles is prevalent
in Madagascar, particularly along the west coast, where hawksbill
and green turtles (live, meat, eggs, and plastron/ligaments)
can be commercialised in other parts of the island including
the capital city, Antananarivo (Muttenzer, 2007;Gough et al.,
2009;Rakotondrazafy and Andrianasolo, 2012;Pilcher and
Williams, 2018). Approximately 52% of fishers stated that they
targeted marine turtles (Pilcher and Williams, 2018), in spite of
national decrees prohibiting all exploitation (IOSEA, 2014, 2019).
A new illegal trade network was uncovered in north-western
Madagascar in 2012, supplying traders in Mahajanga with turtles,
although the final destination was not established (IOSEA, 2014).
In total, it is estimated that up to 100,000 turtles/year (mostly
green turtles) are taken by Madagascan fishers in two areas
alone, Toliara and Antsiranana (Pilcher and Williams, 2018). In
Comoros, turtle hunting is reportedly widespread in the Moheli
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Marine Park (Moheli Marine Park, 2009), Grande Comore, and
Anjouan (IOSEA, 2019).
The town of ¯
Assab, Eritrea, has a well-known market for
turtle products which it trades locally as well as with Yemen
(IOSEA, 2014). Use of marine turtle meat and harvest of eggs
is also reported from Mauritius, Seychelles, South Africa, and
Sudan (IOSEA, 2019).
Robards and Reeves (2011) provided the first systematic
documentation of global marine mammal consumption, collating
data from 1970 to 2009. It demonstrated that for some Asian
cultural groups, the use of marine mammals for food, and likely
for other purposes, is higher than ever before. Asias extensive
coastline and large riverine systems are densely populated and
communities rely heavily on aquatic resources. The review
highlighted an increase in bycatch and showed that in some areas
in Asia, bycatch has developed into deliberate catches and the
harvest of small cetaceans has become commercialised. Although
harming, harassing, killing (when bycaught alive), and hunting
of marine mammals is illegal in most Asian countries, indigenous
and subsistence use still occurs in many areas, at varying levels
(Porter and Lai, 2017;Lee et al., 2019).
Both legal and illegal use and sale of CMS-listed cetaceans
occurs in East Asia (here including China and Taiwan, but
not Japan and South Korea). Most species are used for human
consumption, but some for bait in fisheries (Mintzer et al., 2018).
In Taiwan, despite the sale of bycaught cetaceans being illegal,
the meat of hundreds of bycaught individuals was once sold in
local markets (Sui, 2014), however, recent advances in forensic
surveillance have since curtailed most trade (Lo et al., 2013).
Information for China is anecdotal and limited, e.g., Jefferson
and Van Waerebeek (2002) presented photographic evidence of
bycaught common dolphins landed at fish markets of Putian,
Fujian Province. Wang et al. (2015) presented records of 193
incidents of bycatch, stranding and injuries (mostly fisheries
inflicted) of aquatic mammals nationwide in 2000 to 2006, but
did not report on subsequent use of carcasses. A short study
on the use of Chinese social media and national online news
services to monitor cetacean trade in fish markets in China
indicated that the sale of cetaceans for food, particularly finless
porpoise (Neophocaena spp.), is not uncommon and that some
trade occurs far inland (IWC, 2016). As elsewhere, the lack of
a nationwide reporting scheme and the poorly known status of
most populations of aquatic mammals in China, hampers our
understanding of the impact of both targeted and incidental takes
(Wang et al., 2015).
In Southeast Asia, cetaceans have been used both historically,
and still today, for food and other uses (e.g., Tun, 2006;Porter and
Lai, 2017;Altherr and Hodgins, 2018). In Indonesia, traditional
sperm whale (Physeter macrocephalus) harvests have a long
history, and are restricted to two specific villages, Lamalera
and Lamakera (Leatherwood et al., 1991). These hunts have,
at times, also targeted blackfish (Orcinus orca, Globicephala
macrorhynchus). The hunting of small cetaceans by these
villages is a relatively new practice, but it is believed to be
rapidly increasing, possibly because of declining fish stocks
(Cerchio et al., 2019). Data on catches from non-discriminant
fisheries, such as “experimental nets” or “tiger nets,” indicate that
single net sets may have bycaught thousands of cetaceans, the
meat of which is subsequently used for human consumption and
the pet food industry (CMS, 2015). In the Philippines, many
villages have a long history of hunting cetaceans for bait but
also, within the last two decades, some local communities have
developed a taste for dolphin meat, leading to deliberate harvests
to meet this new demand (Dolar et al., 1994;Dolar, 1999). In the
southern parts of the Philippines, at least one indigenous group
is known to consume dolphin meat as part of their traditional
diet and to use other cetacean products for ceremonies, including
wedding dowries. In Cambodia and Viet Nam, it is commonly
believed that catching a dolphin brings bad luck, thus in general,
dolphin exploitation is rare or non-existent (Perrin et al., 2002;
Beasley and Davidson, 2007). Historically, coastal Irrawaddy
dolphins (Orcaella brevirostris) were exploited for their oil in both
countries (Perrin et al., 2002).
Throughout many parts of South Asia, recent data
suggest cetacean meat is exploited commercially for human
consumption, particularly west India, especially the Indo-Pacific
finless porpoise (N. phocaenoides) and Indo-Pacific bottlenose
(Tursiops aduncus;Kumarran, 2012). A few species, notably
the Indo-Pacific finless porpoise, are hunted for a niche market
along the west coast of India and there is an open market for
cetacean meat in several locations throughout the states of
Goa, Karnataka, and Kerala (Kumarran, 2012). In the town
of Malpe alone, an estimated 2,000 cetaceans are caught for
human consumption every year (Kumarran, 2012). In India
and Bangladesh, Ganges river dolphins (Platanista gangetica)
that become entangled in nylon gill-nets were sometimes killed
by fishers to extract oil to be used as a fish attractant (Smith
and Smith, 1998;Sinha, 2002). In Pakistan, the meat of Indian
Ocean humpback, Indo-Pacific bottlenose and spinner dolphins,
and finless porpoise is used for shark bait, human food, and
traditional remedies (Gore et al., 2012;Kiani and Van Waerebeek,
2015). In the south coast of Sri Lanka, small cetaceans are hunted
using hand-held harpoons and this practice has spread to western
areas driven by the increasing demand for dolphin meat from
inland and urban people (Ilangakoon, 2012).
For western Asia (Arabian region), Robards and Reeves
(2011) note that cetaceans were caught and used for human
consumption in Oman (dwarf sperm whale Kogia sima, common
dolphin, Indo-Pacific humpback dolphin, spinner dolphin,
bottlenose dolphin) and the United Arab Emirates (common
dolphin) between 1970 and 2009. A recent review of finless
porpoise stranding in Kuwait did not indicate the use of this
species as wild meat (Nithyanandan and Bohadi, 2021). Little
is known from this region on the extent of cetacean use or
its sustainability.
Declines in dugong numbers have been linked to overexploitation
both in Southeast Asia and elsewhere in Asia (Marsh et al., 1997,
2002, 2011;Perrin et al., 2002;Mustika, 2006). Although direct
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harvests of dugongs are now banned in most Asian countries
(Marsh et al., 2011), Pilcher et al. (2017) estimated that about
16% of dugongs caught in fishing gears are consumed. These
data come from a questionnaire administered in 20 countries,
including 14 Asian countries; country-specific data have not been
published. In some countries of Southeast Asia, dugong meat
is both locally consumed and commercially traded domestically.
For example, in Cambodia, dugongs are considered to bring good
luck if caught, and the meat and other body parts are sold at
high prices (Beasley et al., 2001). In Malaysias most eastern state,
Sabah, community interviews indicate that substantial numbers
of dugongs are bycaught and, when not discarded, may be utilised
for different purposes (Jaaman et al., 2008). Dugong meat is also
eaten in various parts of Indonesia, where body parts are also
used for traditional remedies and the making of religious artefacts
(Marsh et al., 2002, 2011). Although more recent studies are
limited, dugong parts can still be openly purchased in markets
in some islands (Lee and Nijman, 2015). Dugong hunting has
also been documented throughout the species range in the
Philippines. The meat is valued as an additional source of protein
in remote areas with limited resources, and while it was once
sold openly in markets, it is now sold covertly due to its current
protected status (Kataoka et al., 1995;Marsh et al., 2002, 2011).
In South Asia, in west India (Gulf of Kutch), dugong oil is
valued as a preservative and conditioner for wooden boats, and
the meat is believed to have medicinal value (Jones, 1967;Frazier
and Mundkur, 1990;Marsh et al., 2002). Historically, dugong
meat was highly prised in the Gulf of Mannar (India/Sri Lanka)
(Nair et al., 1975) and, in the Andaman and Nicobar Islands,
dugongs are occasionally hunted, although the frequency has
reduced in recent decades (Das, 1996;Marsh et al., 2002;Hines
et al., 2012;D’Souza et al., 2013).
In western Asia (Arabian region), there is archaeological
evidence of a 6,000-year-old dugong-hunting culture in the
Arabian/Persian Gulf (Méry et al., 2009). Although officially
banned by most range states in the region, some illegal hunting
still occurs, at least in the Red Sea (Nasr et al., 2019). Dugongs
are still caught incidentally in gillnets in both the Red Sea (Nasr
et al., 2019) and the Arabian/Persian Gulf. These animals are
variously released if still alive, given to researchers to study, or
consumed (Nasr et al., 2019). The major drivers for deliberate
captures in fishing gears or illegal hunting are poverty and
declining fish stocks (Nasr et al., 2019), enabled by weak or non-
existent enforcement. Although little is published on the current
status of the dugong population(s) in this region, it appears to
support the largest dugong population outside Australia (Marsh
et al., 2011), mainly in the waters of the United Arab Emirates
(UAE) and Qatar.
Like marine mammals, marine turtles have been used for
centuries within many South and Southeast Asian countries,
as food (both meat and eggs), medicine and ornamentation.
Consumption of marine turtle eggs has been reported across most
of South and Southeast Asia, with turtle meat also consumed
in Sri Lanka, Maldives, Myanmar, Thailand, China, Philippines,
and Indonesia (Yifan, 2018). Marine turtles are also used as
ornamentation in Indonesia, China, Philippines, and Japan,
with Indonesia supplying much of the trade with raw materials
(Yifan, 2018).
In East Asia, consumption of marine turtle meat and eggs is
reported, as well as a market for turtle-derived ornamentation
(Yifan, 2018). In China, most marine turtle nesting beaches have
been lost as a consequence of excessive historic exploitation of
turtles and their eggs, as well as habitat degradation (Lin et al.,
2021). China’s ongoing demand for turtle products is considered
one driver encouraging turtle trade from Southeast Asia, with
at-sea trade reported between Chinese vessels and fisherman in
Southeast Asia, e.g., in the Philippines, Indonesia, and Malaysia
(Lam et al., 2011).
Southeast Asia is a hotspot for marine turtle consumption
and trade. Historically, the island of Bali, Indonesia, was one
of the world’s largest markets for green turtles, and remains an
important hub in Indonesia for both domestic trade and some
continuing international trade (Troëng and Drews, 2004;IOSEA,
2014;Pilcher and Williams, 2018). There is evidence of ongoing
high levels of harvest of turtles and their eggs in other parts of
Indonesia (Dethmers and Baxter, 2011;IOSEA, 2014;Pilcher and
Williams, 2018). Eight seizures occurred in Kalimantan between
2015 and 2018, with 13,200 eggs seised. The sale of eggs, turtle
meat, and live turtles is common and occurs along the coasts
of West Sumatra, Java, Bali, Kalimantan, Sulawesi, Maluku, and
Papua (Pilcher and Williams, 2018). Open trade in markets has
reportedly declined, but shifted to underground markets in some
places (Gomez and Krishnasamy, 2019).
In Malaysia, consumption of turtle eggs is the main form
of use, particularly in Sabah and Terengganu (Gomez and
Krishnasamy, 2019;Joseph et al., 2019;Mohd Jani et al., 2020),
which involves both legal and illegal trade. In most of Peninsular
Malaysia, all marine turtle eggs (except leatherback turtle eggs in
Terengganu and Pahang) are freely and legally traded in local
markets, except in Perak and Melaka. In some places, such as
Perlis and Selangor, trade in unregulated (Mohd Jani et al., 2020).
In Sabah and Sarawak, it is illegal to harvest and consume sea
turtle eggs, but harvest and trade, particularly with Terengganu,
is reportedly common (Troëng and Drews, 2004;Chan, 2006;
IOSEA, 2014;Mohd Jani et al., 2020). Increased regulation of
harvests generally was implemented after a dramatic decline in
leatherback turtle nests at Rantau Abang in Terengganu – from
some 10,000 nests per year in 1956 to just three in 2002 (Troëng
and Drews, 2004), attributed to both continuous exploitation
of eggs over decades, and fisheries bycatch of adults (Chan
and Liew, 1996;Eckert et al., 2012;NMFS and USFWS, 2020).
However, at least on Redang Island, Terengganu, legal protection
of turtle nesting beaches as well as changing attitudes toward egg
consumption, are thought to have led to a decline in demand for
green turtle eggs, although consumption continues (Poti et al.,
2021). Population declines of leatherback turtles in Terengganu
contrast with recoveries of green turtles in Sabah, following the
strong enforcement of the egg protection programme in the early
1970s (Chan, 2006).
In Viet Nam, there is a commercial demand for marine
turtle meat and products, observed for sale in 39 of 436
outlets surveyed (Gomez and Krishnasamy, 2019). The most
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widespread use involves hawksbill turtles, with shells used
to make decorative items including bekko ornaments, and
taxidermied whole animals. These are sold domestically, but are
also seised in international trade mostly with China (IOSEA,
2014;Pilcher and Williams, 2018). In 2002, a domestic ban
on turtle trade in Viet Nam markets diverted trade to at-sea
exchanges (Chan et al., 2009), although they can still be found
in some markets, restaurants, and shops (Hamann et al., 2006).
Numerous seizures in Viet Nam, mostly hawksbill turtles, suggest
that Indonesia and Malaysia remain a source of raw shell plates
for tortoise shell bekko manufacture (IOSEA, 2014). Online sale
of turtle products is increasing, especially in Indonesia and Viet
Nam (Pilcher and Williams, 2018).
There is limited information for other countries in Southeast
Asia. The Philippines report an unquantified increase in illegal
harvest of marine turtles in several regions and continued high
levels of egg harvesting (IOSEA, 2019), including in the Tawi-
Tawi Islands (IOSEA, 2014), where historically 70% of eggs were
harvested (Chan and Shepherd, 2002). Harvest of turtles and eggs
is common along the coast of Myanmar (Win and Lwin, 2012),
especially in remote areas (IOSEA, 2019).
In South Asia, there is limited information on the extent
of turtle harvests in the northern Indian Ocean, although it
does occur, as evidenced by expert elicitation (Riskas et al.,
2018) and media reports of fishing vessels that have been
impounded for operating illegally in other countries’ waters,
which have then been found to have turtles onboard (IOSEA,
2014). In India, reports of fishermen targeting turtles for their
meat are largely anecdotal (IOSEA, 2014). While some areas
report extremely high egg harvests, for example nearly all
Olive Ridley (Lepidochelys olivacea) nests in Tamil Nadu were
reportedly harvested illegally in 2011 (IOSEA, 2014). Others
report egg collection is decreasing, but the reasons why are
unclear (Hasan, 2009). Anecdotal reports suggest a turtle fishery
in Pakistan territorial waters may be supplying neighbouring
countries with turtle meat (IOSEA, 2014). In the Maldives, turtle
hunting is increasingly uncommon whereas the harvesting and
consumption of eggs remains common (IOSEA, 2019). In the last
half of the twentieth century, the communities at Rekawa beach,
Sri Lanka, collected most of the turtle eggs laid on the beach,
for sale and local consumption and the harvesting of turtles for
meat was common in other parts of Sri Lanka (Ekanayake et al.,
2002). A survey conducted in 2007 found consumptive use of
turtle meat and eggs had significantly reduced, and noted that
nesting rates were high in areas, such as Rekawa, that now had
community-based protection of nests (Rajakaruna et al., 2009).
In western Asia (Arabian region), there are reports of high
levels of egg collection in Saudi Arabia, and high rates of illegal
harvest of animals in Iranian waters of the Oman Sea (IOSEA,
2014). Low incidence of turtle use is reported for Oman, whereas
anecdotal reports suggest some level of egg collection in the UAE,
but no hunting or larger scale harvesting (IOSEA, 2019).
Only two crocodylian species are currently included in the
Appendices of CMS, both in Asia. Today, limited gharial
(Gavialis gangeticus) populations are restricted to a few large river
systems in north India and lowland Nepal. The species is now
extinct in Bhutan, Myanmar, Pakistan, and possibly, Bangladesh
(Lang et al., 2019). Historically, gharial were hunted as sport, and
occasionally taken opportunistically for skins and meat. Seasonal
movements related to nesting and feeding, up and down large
open rivers and their tributaries likely regularly resulted in trans-
boundary movements between Pakistan, Bangladesh, Bhutan,
Burma, and India by gharials resident in boundary waterways.
Illegal consumption of gharial eggs sometimes occurs, but is a
minor concern relative to the widespread and frequent habitat
destruction/alteration (sand mining and water extraction) and
mortality from net entanglement due to illegal fishing (Stevenson,
2015;Lang et al., 2019). Eggs were used in some communities
along the Karnali-Girwa Rivers (Nepal–India) and Narayani-
Gandak Rivers (Nepal–India) for food and use in traditional
remedies (Lang et al., 2019).
Like the gharial, the population status of saltwater crocodiles
(Crocodylus porosus) varies greatly between range states in
Asia, effectively extinct in Thailand, Cambodia, Viet Nam, and
the Seychelles, with the largest and most secure Asian wild
population in West Papua and Papua Provinces in Indonesia.
Subsistence use of eggs and crocodiles for food by indigenous
peoples and local communities’ dates back thousands of years
(Webb et al., 2010). Currently, the species is subject to legal
ranching (eggs, juveniles) and wild harvest (direct hunting;
Webb et al., 2010). In the state of Sarawak (Malaysia), a limited
wild harvest of crocodiles was implemented recently, to address
increasing levels of human-crocodile conflict, and the meat is
used for domestic consumption. Cultural beliefs of indigenous
people of Sarawak do not allow crocodiles to be killed and/or
eaten (Tisen et al., 2013). In Myanmar, wild populations were
seriously depleted by hide hunting (1950s and 1960s) and the
meat was consumed by people (Platt et al., 2012). Severe depletion
in Myanmar, as in Thailand, Cambodia, Vietnam and Philippines,
has allowed local people to use waterways freely, which they
could not do historically, complicating efforts to rebuild wild
populations (Platt et al., 2012). In wildlife sanctuaries in the
Ayeyarwady Delta, saltwater crocodiles are recovering, despite
incidental catch in fishing nets (Than et al., 2020). A survey in
Thailand (Webb and Jenkins, 1991) revealed some live saltwater
crocodiles imported legally from a small government operated
farm in Myanmar. Some trade may still exist at low levels (Platt
et al., 2012) given the porous border.
Elsewhere in their Asian range, saltwater crocodile
populations are nominally protected, with varying levels of
“problem” animal control (Webb et al., 2010). Recovery of wild
populations increases human-crocodile conflict on people and
livestock (Brackhane et al., 2018), with many rural communities
not inclined to tolerate these large predators. “Problem”
crocodiles removed for public safety, legally or illegally, may be
used as food, but cultural taboos may dictate whether this occurs
(e.g., Corvera et al., 2017;Brackhane et al., 2018;Webb, 2020).
Contemporary use of cetaceans is reported across Polynesia,
Melanesia, and Micronesia, however, often the species could
not be identified (Robards and Reeves, 2011). Since 1990, the
use of stranded or intentionally caught humpback whales has
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been reported as rare in Tonga, as has the use of bottlenose
dolphins in New Caledonia. Species identification is available for
the Solomon Islands, where Risso’s (Grampus griseus), Fraser’s
(Lagenodelphis hosei), pantropical spotted, spinner dolphins, and
Cuvier’s beaked whale (Ziphius cavirostris) are utilised. In villages
on Malaita, Solomon Islands, a traditional dolphin hunt for meat
and teeth occurs (100,000 dolphin teeth annually; Takekawa,
2000). Teeth are used as a traditional wedding dowry, and are
also used for personal adornment (Takekawa, 2000). Attempts
to stop traditional hunts in 2010, using financial compensation,
were unsuccessful (Oremus et al., 2015).
The Gilbert Islands of Kiribati have a long history of
marine mammal hunting (Di Piazza and Pearthree, 2001),
and data suggest that some species of mesoplodont and
Cuvier’s beaked whales are still targeted for human consumption
(Baker et al., 2013).
Dugongs have high cultural value in many parts of Oceania, and
traditional legal hunting is widespread in Pacific Islands range
states (Marsh et al., 2011). The dugong is a cultural keystone
species in the Torres Strait (Butler et al., 2012) where it has
been harvested for at least 4,000 years (Crouch et al., 2007).
The harvest still has significant provisioning value, although the
value of dugong meat as food is less than its cultural services
(Delisle et al., 2018). The Torres Strait Treaty between Australia
and Papua New Guinea guarantees the rights of Islanders
to their traditional way of life, including hunting dugongs
(Havemann and Smith, 2007).
There is limited effective regulation of the Papua New Guinea
harvest, apart from a national ban on the sale of meat, which
means that some villagers still catch dugongs illegally, especially
using mesh nets (Marsh et al., 2011). Throughout their range
in northern Australia, native title holders are permitted to hunt
dugongs in their own traditional sea country. In the Australian
Torres Strait, governance mechanisms include a ban on selling
dugong meat, traditional owner-only hunting rights, spatial
and gear controls, and community management through 15
nonstatutory, communitybased turtle and dugong hunting
plans (Marsh et al., 2015). These restrictions are bolstered by
socioeconomic realities, for example access to regions where
dugongs are abundant is financially unviable due to the high price
of fuel (Marsh et al., 2015). In the Western Islands of Torres
Strait, the dugong harvest in the 1990’s was estimated to be on the
order of 1,000 individuals per year (Marsh et al., 1997, 2002) and
concerns were expressed about its sustainability (Heinsohn et al.,
2004;Marsh et al., 2004). Multiple lines of evidence, including
revised population size estimates, high calving rates, and the high
genetic diversity of the population indicate that the harvest in
the Western and Central Torres Strait is likely sustainable at a
regional scale (Marsh et al., 2015;Hagihara et al., 2018).
Outside of the Torres Strait, there is also evidence of dugong
hunting. Dugong meat is highly prised in the Solomon Islands
and Vanuatu (Chambers et al., 1989;Kile et al., 2000;Marsh
et al., 2002, 2011). However, at least in some areas, such as
the autonomous region of Bougainville, the largest island in
the Solomon Archipelago, directed hunting of dugongs is now
extremely rare (Kinch, 2008). In New Caledonia, dugong hunting
is now prohibited in some provinces, but allowed in others
with a permit, although some illegal harvests still occur (Cleguer
et al., 2017). In Micronesia, dugong hunting in Palau was largely
practised to obtain meat for special occasions, and had high
traditional value (Marsh et al., 1995). Whilst dugong hunting is
now illegal in Palau, evidence from 2003 suggests that dugong
hunting may still be widespread (Matthews, 2003).
Marine turtles and their eggs are harvested for consumption
across several regions of Oceania, except Hawaii, threatening
all five species (Work et al., 2020). It is estimated that
Papua New Guinea harvests 15,217 turtles per year, which is
equivalent to 36.1% of the estimated global legal harvest of 2013
(Humber et al., 2014), and it is unknown if this level of take
is sustainable (Work et al., 2020). In the Torres Strait, where
large juvenile and adult green turtles are abundant (Fuentes
et al., 2015), turtle harvests have been sustained over time,
although how this is achieved biologically is unclear, hence
concerns remain about the sustainability of current harvest
levels (Hagihara et al., 2016). Turtles play a significant role
in the customary economy of many Aboriginal and Torres
Strait Islander communities in Australia, and use for personal,
domestic, and non-commercial purposes is legal, although the
level and impact of the harvest is unknown (IOSEA, 2019). Illegal
egg harvest on the Solomon Islands has been identified as a threat
to olive ridley, leatherback, and hawksbill sea turtles (Work et al.,
2020), as is the estimated annual harvest of 9,473 turtles (species
unknown) by spear fishers (Vuto et al., 2019).
While universally valued, there is significant variability
between communities and regions about the rules and social
practises surrounding turtle consumption in the Pacific Islands
region (Allen, 2007). In most Polynesian islands, turtle
consumption is typically restricted to high-ranking individuals
(e.g., chiefs), while in Niue, meat is only consumed by males
(Rudrud, 2010). On Rapa Nui (Easter Island), consumption of
turtles was once common, but is now no longer practised by the
islanders (Álvarez-Varas et al., 2020). In parts of Melanesia, turtle
shell trade is an important part of society (Balazs, 1983;Allen,
2007;Rudrud, 2010).
The largest and most secure wild populations of saltwater
crocodiles in the Oceania region occur in Australia and
Papua New Guinea. Within these countries, the species is subject
to legal commercial ranching (eggs and juveniles), and wild
harvest for commercial exports in compliance with CITES.
Crocodiles are also hunted legally for subsistence by indigenous
peoples (e.g., Webb, 2020). Hunting crocodiles for meat also
occurs in some parts of the Solomon Islands, but this tends to
involve crocodiles that have attacked people or livestock (Van
der Ploeg et al., 2019). Fear of being attacked by crocodiles
results in crocodiles and their nests or eggs being destroyed
opportunistically by local people (Van der Ploeg et al., 2019).
Studies on saltwater crocodile movement patterns have
largely been carried out in Australia (Campbell et al., 2010;
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Fukuda et al., 2019;Fukuda et al., 2022). Although long-distance
movements of individuals out of their typical distribution range
have been recorded (e.g., Marshall Islands; Manolis, 2005;
Spennemann, 2020), these appear to be the exception rather than
the rule. Movements of Australian saltwater crocodiles across
international borders (e.g., possibly between Australia, Timor-
Leste, Papua New Guinea, and Indonesia) may be linked to
territorial exclusion as populations reach carrying capacity within
Australia (Fukuda et al., 2011;Brackhane et al., 2018), and are not
likely representative of migratory patterns.
The harvesting of CMS-listed, non-fish migratory megafauna,
for aquatic wild meat for human consumption and other
purposes, is clearly widespread in the tropics and subtropics,
with the extent of harvest highly variable between species and
countries, and the impact on populations – whether sustainable
or not– rarely known with scientific confidence. The legality of
national harvests is equally variable across species and countries.
The purposes of use range from subsistence by Indigenous
Peoples and Local Communities (IPLCs), often in poverty, but
respected under the UN Declaration of the Rights of Indigenous
People (UNDRIP), to purely commercial purposes at the other
extreme, and all manner of combinations within. Aquatic wild
meat is clearly a significant issue that needs to be considered
more seriously at national and international levels, taking into
account diverse value orientations (van Vliet, 2018), including
within conventions like the CMS, and some key areas are
discussed below.
Conservation Concerns
Given that the extent of use relative to the size and trends
(increasing, decreasing or stable) in the wild source population
and rates of immigration and emigration are seldom known
accurately, the impact of harvests for aquatic wild meat on
the conservation status of source populations can seldom be
quantified with any scientific certainty. For example, in mainland
South America, it is unclear whether the harvest and trade of
small cetaceans specifically for human consumption in Peru was
being sustained by the wild population, but from a precautionary
stance, increasing rates of harvest for bait or other purposes
in Peru, Bolivia, Ecuador, Colombia, Venezuela, and Brazil
(Barbosa-Filho et al., 2018;Mintzer et al., 2018;Campbell
et al., 2020a;Castro et al., 2020;Briceño et al., 2021) constitute
a potential threat. Similarly, the harvest of migratory small
cetaceans for aquatic wild meat in the Gulf of Guinea, in
particular Atlantic humpback dolphin, which meet IUCN Red
List criteria for Critically Endangered (Collins et al., 2017),
are assumed to be threatening despite limited information on
harvest levels, population size, trends and distribution. In the
south western coast of India and throughout Southeast Asia,
killing, rather than releasing cetaceans by-caught in fishing gear,
for aquatic wild meat is assumed to be a threat to migratory
cetaceans. Indeed, high human population densities and the
prevalence of fishing activities in Southeast Asia generally make
it a hotspot for aquatic wild meat use, even if the harvest
is opportunistic rather than targeted. Against this, traditional
harvests of dolphins in the Solomon Islands appear to have been
occurring for millennia.
For depleted sirenian populations, even a modest harvest
for human consumption may constrain the rate of recovery
of populations which are highly sensitive to changes in adult
survival (Marsh et al., 2011). Most local populations cannot
withstand human-induced mortality. In Senegal, The Gambia,
Nigeria, Cameroon, and Democratic Republic of the Congo,
more than 1,000 manatees have been killed by illegal hunting,
bycatch, dams and watercraft between 2017 and 2019 (Keith-
Diagne et al., 2019). Aquatic wild meat is a major motivation
for this loss, impacting negatively on most populations. Against
these examples, the harvest of dugong for aquatic wild meat
by indigenous people across northern Australia, which contains
the most abundant sirenian populations (Marsh et al., 2011),
have been sustained for more than 4000 years. This highlights
clearly that differences in local circumstances between areas, such
as drivers of hunting and consumption, hunting technologies
used, human density and other threats to animals and their
habitats, and how they change over time, can influence
harvest sustainability.
Capture and consumption of marine turtle adults and
harvest of their eggs is ubiquitous across much of the species’
ranges, particularly in the Caribbean, Southeast Asia, off
Papua New Guinea, and by Indigenous people across northern
Australia. Whilst there are management programs in some places
(see section “Discussion” below), and evidence of significant
declines in population and nesting due to past and present
exploitation in many countries (Troëng and Drews, 2004;
Amorocho et al., 2016;Blumenthal et al., 2021;Lin et al., 2021),
low-level harvests have been sustained for millennia in some
countries, and harvests can be managed sustainably for aquatic
wild meat and other products (Carrillo et al., 1999;Fleming,
Risks to riverine megafauna from harvest may be particularly
high, even if opportunistic, because the risks are compounded
by unique threats to riverine species, such as dams, intensive
fishing, and pollution where human population density is high
(He et al., 2017). In the Amazon, the tucuxi and Amazon
River Dolphin are considered threatened by the IUCN Red List,
are nominally protected, but are illegally harvested as bait for
fishing enterprises. In India and Bangladesh, the use of Ganges
River dolphins for meat and bait is less common today than
in the recent past, but even small harvests may be problematic
given their small population sizes (Braulik and Smith, 2019).
For the sirenians, both the Amazonian and African manatees
are considered sensitive to any harvest given low reproductive
output and generally small local population sizes, making their
use for aquatic wild meat potentially problematic. In South
America, the giant South American river turtle Podocnemis
expansa has been harvested for food for millennia, despite
harvest levels and population responses being poorly known.
Management interventions are enhancing conservation in some
places (Forero-Medina et al., 2021). Riverine megafauna may
suffer from a lack of management and research as they are seen
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as neither terrestrial species nor fish, and the case for increased
research and management of riverine and freshwater megafauna
is a strong one.
Finally, the use of aquatic megafauna for aquatic wild meat
is likely to be far more widespread in terms of frequency and
species than reported here, especially amongst IPLCs. Monitoring
and reporting is limited, and because many of the species
are protected by national law, or are charismatic, their use is
secretive. The trans-boundary nature of harvests and associated
trade of these oceanic, coastal, and riverine species requires
increased international attention and cooperation. Despite the
wisdom of adopting a precautionary approach in the absence
of quantitative data on harvest levels, population sizes, and
trends for most species used for aquatic wild meat, the case for
researching in more depth the harvests that do appear sustainable
is a strong one. There is a fundamental difference between
assembling information on isolated population dynamics, and
studying “dynamic populations” – how populations compensate
to sustain uses (Webb, 2015). CMS has taken an early step,
agreeing to progress the development of a sub-regional Aquatic
Wild Meat Action Plan for the Gulf of Guinea for consideration
by CMS COP14 (Decision 13.64, COP13; CMS, 2020). Similar
urgent attention will be needed to either work through existing
regional processes or developing targeted action plans for the
remaining highlighted species.
Human Dimensions and Drivers of
Aquatic Wild Meat Use
The drivers of aquatic wild meat consumption and trade are
varied and changing. For some IPLCs, consumption of aquatic
wild meat may have high socio-cultural significance (for example
dugongs in the Torres Strait; Delisle et al., 2018), and is a right
bestowed under UNDRIP. Food is an important driver of low-
level aquatic wild meat harvests in many places (e.g., turtles for
Miskito and Creole Caribbean Nicaraguans; Garland and Carthy,
2010), yet its nutritional contribution relative to alternatives is
rarely quantified (Olmedo and Farnés, 2004). Some freshwater
turtles eaten by indigenous people in northern Australia are
exceptionally rich in iron, which is otherwise deficient in their
diet (Eisemberg, 2020). Increasing market access and integration
into cash-based economies changes the dynamics of aquatic
wild meat consumption and trade in some areas, and as such
the gradient between subsistence use and commercial trade
becomes hazy (Frazier, 1980;Ingram et al., 2021). Subsistence
and commercial uses merge insensibly in many of the examples
of human consumption reviewed here, although some uses, for
example botos and tucuxis in Amazonia, involving deliberate and
illegal harvesting for bait to catch fish of higher commercial value,
adds new complications (Mintzer et al., 2018).
The dynamics of aquatic wild meat harvests, use, and
trade may change in response to climate change influences
on food security and habitats. Declines of fish stocks and
terrestrial wild meat may increase dependence on aquatic
megafauna in coastal communities with few alternatives, for
example cetaceans in West Africa (Leeney et al., 2015;Van
Waerebeek et al., 2017). Fisheries catch potential is predicted
to decline over the 21st century under all emissions scenarios,
especially in the tropics (Bindoff et al., 2019), and livelihoods
and food security of communities currently dependent on
marine resources are predicted to be adversely affected. An
interaction between declining fish stocks and use of terrestrial
wild meat has occurred in West Africa (Brashares et al.,
2004;Rowcliffe et al., 2005), hence increased monitoring
of the extent of aquatic wild meat use in local contexts
could be important.
Potential Implications for Human Health
Recognising the importance CMS Parties place on the human
health consequences of species management actions, it is
considered briefly here. Most research investigating the human
health impacts of consuming wildlife deal with terrestrial
wild meat, but some apply to aquatic wild meat. Food and
nutrition, often provided to people in relative poverty by
wild meat, is fundamental to their health and well-being,
and a basic right (Article 25) within the UN Universal
Declaration of Human Rights. However, there are clearly health
risks associated with wild meat that are context-specific, and
include zoonotic pathogens (viruses, bacteria, parasites) and
excessive human consumption of heavy metals and pollutants.
Recorded Emerging Infectious Disease (EIDs) events, which
can influence economies and public health, nationally and
internationally, were 60.3% zoonoses (Jones et al., 2008),
with 71.8% originating in terrestrial wildlife, and increasing
over time (Jones et al., 2008). Ebola virus disease (Greatorex
et al., 2016), Avian Influenza A (Poovorawan et al., 2013),
and Middle East Respiratory Syndrome (MERS) (Cauchemez
et al., 2014) are all examples. None of these have been
attributed to aquatic wild meat, but they exist side-by-side
in many cultures.
Fundamental hygiene is a risk factor with all meat
consumption, wild or domestic. The consumption of raw or
undercooked pinniped or cetacean meat has resulted in bacterial
(e.g., salmonellosis and botulism) and parasite (trichinellosis and
toxoplasmosis) infections in people (Bender et al., 1972;Tryland,
2000;McLaughlin et al., 2004;Van Bressem et al., 2009;Tryland
et al., 2014). Reptile meat and eggs, if not subject to inspection
and hygienic treatment, can cause bacterial (Salmonella spp.,
Vibrio spp.) and potentially parasite infections (Spirometra,
Trichinella, Gnathostoma, pentastomids) and biotoxin problems
(Magnino et al., 2009;Cantlay et al., 2017). Heavy metals and
pollutants in some chelonian life stages (Frías-Espericueta et al.,
2006), cetaceans (Fielding and Evans, 2014), and sirenians
(Marsh et al., 2002, 2011) exceed international food safety
standards, and a growing list of bacterial, viral, and fungal agents
in marine mammals (Waltzek et al., 2012) could be problematic.
Viable substitutes for wild meat, if they can be afforded, also
have a multitude of health risk implications (Tomley and Shirley,
2009), and clearly further research is needed.
Where the use of aquatic wild meat is not sustainable, clearly
compromises the conservation of the species involved, and has
definitive health risks that exceed those of substitute meat,
strategies for reducing aquatic wild meat use and consumption
may be possible through partnerships between local people,
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governments, public health professions, veterinarians, and both
wildlife managers and ecologists.
Focused Research and Action
For most species used for aquatic wild meat reviewed here,
there are vast gaps in knowledge about the extent of harvest,
the source population size and trends, and the degree to which
it is influenced by immigration and emigration. Added to this,
there are relatively few insights into the population processes
involved in aquatic wild meat harvests that have been sustained,
sometimes over millennia. Hence conservation action has been
precautionary, in many cases wisely implementing species
protection, but tending to ignore ongoing uses or treat them as
unavoidable collateral damage. This is especially so where the
rights of IPLCs to use aquatic wild meat is engrained in national
legislation and international agreements, such as UNDRIP.
Given many of these uses will continue, and be vulnerable to
expansion for economic reasons, a strong case exists to better
understand the sustainability of current uses, and be better
able to apply management interventions when needed to ensure
sustainability is achieved, and that unsustainable uses can be
avoided. Harvests of cetaceans and sirenians are considered
compromised by low reproductive rates. Crocodylians and
chelonians share many life history traits, but whereas managed,
sustainable uses of crocodylians are common (e.g., egg harvests),
they are reasonably rare with marine turtles. The total annual
legal take of wild marine turtles was thought to have decreased
from 116,420 in the 1980s to 45,387 in the 2000s (Humber
et al., 2014), but real levels of harvest, including illegal harvest,
may be much higher (Pilcher and Williams, 2018). At some
sites, successful recovery of depleted marine turtle nesting
populations has been attributed to protecting nest sites from
egg collection (Chan, 2006;Hamilton et al., 2015;Sardeshpande
and MacMillan, 2019). At other sites, managed harvest of
eggs has been sustainable and created increased stewardship
of nesting animals (Campbell et al., 2007;Sardeshpande and
MacMillan, 2019). Bycatch and subsequent use of turtles through
industrial and small-scale fisheries also need to be addressed
simultaneously. Life history parameters can be used to make
crude predictions about sustainability, but factors like density-
dependent adjustments are difficult to account for. It is clear that
much more research is needed on the dynamics of populations
where harvests have been sustained over time to understand the
circumstances under which it is biologically possible.
Active management interventions, supported by local
communities (community-based management), are critical
to ensuring the sustainability of uses of aquatic wild meat
for sustenance, maintenance of culture and tradition, and
improved livelihoods. It is also important for identifying
levels of use likely to be unsustainable. When a species is not
used for aquatic wild meat, management action tends to be
ignored. Investment in management is more likely when aquatic
megafauna contributes to improved livelihoods of local people
(Campbell et al., 2007;Sardeshpande and MacMillan, 2019);
for example, the community turtle egg harvest at Ostional,
Costa Rica (Sardeshpande and MacMillan, 2019). Public support
for rebuilding wild populations of predators, such as saltwater
crocodiles, has been achieved by generating economic benefits
through sustainable use (Webb, 2020), and by targeting problem
animals for removal and use (Fukuda et al., 2014). In northern
Australia, a conservation goal of recovering a severely depleted
wild population of saltwater crocodiles back to carrying capacity
was achieved whilst being used sustainably to generate economic
benefits in compliance with CITES.
Such case histories are unlikely to be directly transferable
to most cetaceans and sirenians, given different population
parameters and market values. However, traditional uses will
continue and management may be needed if sustainability
is compromised. Increased uses for human consumption
and bait, driven by commercial incentives, are likely to
compromise sustainability and need appropriate management
interventions (public education, alternative food/bait substitutes,
altered fishing methods and bycatch mitigation measures,
increased inspections/enforcement, etc.). The drivers of
consumptive use are often relative poverty, and legal
protection, sustainability for the future, risk of punishment,
and accessing substitutes are simply overpowered by immediate
needs. Top–down legal restrictions alone are unlikely to be
sufficient in many socioeconomic contexts where poverty is
rife, cultural values are not understood, and management
capacity is low (Barrios-Garrido H. et al., 2020;Ingram,
For both crocodylian species, it is questionable whether today,
they meet the criteria for inclusion as migratory species on the
CMS appendices. Historically, riverine species such as the gharial
made seasonal movements up and down large, wide-open rivers,
prior to the advent of numerous man-made obstructions. Now,
the many barrages and dams constructed along these waterways
have resulted in predominantly unidirectional movements, e.g.,
downstream, but not upstream, for one or more life history
stages, typically the younger, smaller age cohorts. Depending
on river stretches traversed, cyclical seasonal movements may
cross national boundaries, e.g., seasonal movements on rivers
flowing between Nepal and India. Similarly, transboundary
movements of saltwater crocodiles likely occur in certain
estuarine/riverine settings, but are poorly documented, even
though long distance movements at sea occur (Webb and Messel,
1978). Other freshwater species, living in wetlands, riverine,
and/or coastal areas at confluences of national boundaries may
regularly or occasionally make daily or seasonal movements
across international borders. Technically, these species pose a
special challenge to the existing definitions of transboundary
migrations. The result is their inclusion in the CMS listing, albeit
exceptional relative to other listings.
Whilst this review focuses on megafauna that are considered
migratory and threatened and are listed in CMS Appendices,
there are hundreds if not thousands of other vertebrate species
(including fish) used for consumption of aquatic wild meat. Many
of these species occur across the tropics and subtropics: for most,
sustainable uses are most likely in question. Many of these are
included on the IUCN Red List, and are covered by CITES,
but some are not because trans-border movements are not
evident. Examples include: crocodylians (Genera: Osteolaemus,
Mecistops, and Crocodylus) and otters (Genera: Aonyx, Hydrictis)
Frontiers in Marine Science | 15 March 2022 | Volume 9 | Article 837447