RES EARCH

CONSERVATION ened with extinction (7), a key question for

coral reef ecosystems lies in understanding
Widespread diversity deficits of coral reef sharks the global extent of species loss in elasmo-
branch assemblages. We characterized elas-
and rays mobranch assemblage structure on coral reefs
across a gradient of human pressures to esti-
mate the local depletion and global extinction
Colin A. Simpfendorfer1,2*, Michael R. Heithaus3, Michelle R. Heupel2,4, M. Aaron MacNeil5, Mark Meekan6, risk of the most common reef species, reveal-
Euan Harvey7, C. Samantha Sherman1,8, Leanne M. Currey-Randall4, Jordan S. Goetze9,10, Jeremy J. Kiszka3, ing the human and environmental factors that
Matthew J. Rees6,11, Conrad W. Speed6, Vinay Udyawer12, Mark E. Bond3, Kathryn I. Flowers3,13, influence assemblage structure and that lead
Gina M. Clementi3, Jasmine Valentin-Albanese14, M. Shiham Adam15, Khadeeja Ali3,16, Jacob Asher17, to a deficit in predator diversity that could
Eva Aylagas17, Océane Beaufort18, Cecilie Benjamin19, Anthony T. F. Bernard20,21, Michael L. Berumen22, affect reef ecological functioning.
Stacy Bierwagen4, Chico Birrell23, Erika Bonnema3, Rosalind M. K. Bown24, Edward J. Brooks25, To understand the extent of the reef elasmo-
J. Jed Brown26, Dayne Buddo27, Patrick J. Burke28,29, Camila Cáceres3, Marta Cambra30,31, branch diversity deficit, we surveyed 391 coral reefs
Diego Cardeñosa3, Jeffrey C. Carrier32, Sara Casareto3, Jennifer E. Caselle33, Venkatesh Charloo34,
in 67 nations and territories using 22,756 baited
Joshua E. Cinner35, Thomas Claverie36, Eric E. G. Clua37,38, Jesse E. M. Cochran22, Neil Cook39,40, remote underwater video stations (BRUVS).
Jessica E. Cramp41,42, Brooke M. D’Alberto1,43, Martin de Graaf44, Mareike C. Dornhege45,
We examined reef-level species richness, spe-
Mario Espinoza30,31, Andy Estep46, Lanya Fanovich40, Naomi F. Farabaugh3, Daniel Fernando24,

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cies composition of elasmobranch assem-
Carlos E. L. Ferreira47, Candace Y. A. Fields3,25, Anna L. Flam48, Camilla Floros49,50,
blages, and species relative abundance (MaxN;
Virginia Fourqurean51,52, Laura Gajdzik22,53, Laura García Barcia3, Ricardo Garla54,55, Kirk Gastrich3, the maximum number of each species ob-
Lachlan George2, Tommaso Giarrizzo56,57, Rory Graham5, Tristan L. Guttridge59,60, Valerie Hagan13,
served in a single frame of each 60-min de-
Royale S. Hardenstine16,22, Stephen M. Heck13, Aaron C. Henderson61, Patricia Heithaus3, ployment then averaged across all deployments
Heidi Hertler61, Mauricio Hoyos Padilla62,63, Robert E. Hueter64,65, Rima W. Jabado1,66,
on one reef) (8). We examined how elasmo-
Jean-Christophe Joyeux67, Vanessa Jaiteh68,69, Mohini Johnson70, Stacy D. Jupiter71, branch species assemblages changed in re-
Muslimin Kaimuddin72,70, Devanshi Kasana3, Megan Kelley3, Steven T. Kessel73, Benedict Kiilu74,
sponse to human pressures, using unweighted
Taratau Kirata75†, Baraka Kuguru76, Fabian Kyne77, Tim Langlois78,79, Frida Lara80,81, Jaedon Lawe82, pair group with arithmetic mean (UPGMA)
Elodie J. I. Lédée1, Steve Lindfield83, Andrea Luna-Acosta84, Jade Q. Maggs85,
clustering to identify reefs with the most simi-
B. Mabel Manjaji-Matsumoto86, Andrea Marshall87, Lucy Martin88, Daniel Mateos-Molina89,90, lar assemblages (8). We then compared these
Philip Matich60, Erin McCombs91, Ashlie McIvor22,92, Dianne McLean6,93, Llewelyn Meggs82,
clusters with estimated depletion of key resi-
Stephen Moore1, Sushmita Mukherji1,2, Ryan Murray94, Stephen J. Newman95, Josep Nogués88, dent elasmobranch species at the reef level and
Clay Obota96,97, Domingo Ochavillo98, Owen O'Shea99,100, Kennedy E. Osuka96,101,
examined whether socioeconomic, management,
Yannis P. Papastamatiou3, Nishan Perera23, Bradley Peterson14, Caio R. Pimentel67,102, or environmental factors could predict cluster
Fabián Pina-Amargós103,104, Hudson T. Pinheiro105, Alessandro Ponzo106, Andhika Prasetyo107, membership, using linear discriminant analy-
L. M. Sjamsul Quamar108, Jessica R. Quinlan3, José Amorim Reis-Filho109, Hector Ruiz110, sis. Reef-level depletion was estimated by divid-
Alexei Ruiz-Abierno104, Enric Sala111, Pelayo Salinas-de-León112,113, Melita A. Samoilys96,114, ing the observed mean MaxN of a species at
William R. Sample3, Michelle Schärer-Umpierre110, Audrey M. Schlaff1, Kurt Schmid55,115, individual reefs by a model-estimated baseline
Sara N. Schoen3, Nikola Simpson116, Adam N. H. Smith117, Julia L. Y. Spaet118, abundance (without human pressures) for each
Lauren Sparks119, Twan Stoffers120, Akshay Tanna24, Rubén Torres121, Michael J. Travers95, sampling site (a small group of closely asso-
Maurits van Zinnicq Bergmann3,58, Laurent Vigliola122, Juney Ward123, Joseph D. Warren14, ciated reefs) and subtracting this value from 1.
Alexandra M. Watts47,124, Colin K. Wen125, Elizabeth R. Whitman3, Aaron J. Wirsing126, Aljoscha Wothke40, Baseline abundance (also expressed as MaxN)
Esteban Zarza-González127,128, Demian D. Chapman3,60 was estimated from a general linear model
relating observed MaxN to sampling site, hu-
A global survey of coral reefs reveals that overfishing is driving resident shark species toward man pressure [represented by total market
extinction, causing diversity deficits in reef elasmobranch (shark and ray) assemblages. Our species- gravity, the size and travel time to human mar-
level analysis revealed global declines of 60 to 73% for five common resident reef shark species kets (2)], and marine protected area (MPA)
and that individual shark species were not detected at 34 to 47% of surveyed reefs. As reefs status [closed to all fishing, open to fishing,
become more shark-depleted, rays begin to dominate assemblages. Shark-dominated assemblages or restricted (some fishing but with restric-
persist in wealthy nations with strong governance and in highly protected areas, whereas tions)]. The baseline was estimated by setting
poverty, weak governance, and a lack of shark management are associated with depauperate all parameters to those expected at a site with
assemblages mainly composed of rays. Without action to address these diversity deficits, loss of no human pressure (gravity to the minimum
ecological function and ecosystem services will increasingly affect human communities. for an ocean basin and protection status to
closed) (8).

C
Sampling identified 104 distinct elasmobranch
oral reef ecosystems are under increas- on coral reefs as predators and prey across species or species complexes (table S1), repre-
ing pressure from human activities— multiple trophic levels and in the cycling and senting more than 77% of elasmobranch spe-
including intense fishing, degraded water movement of nutrients (3–5). Recent evidence cies known to occur on coral reefs at some
quality, and climate change (1, 2)—that indicates that overfishing has driven sharks point during their lives (9). More than half
threaten species supporting a wide range toward functional extinction on many reefs. In (n = 53) of the species were rarely observed,
of ecosystem functions (3). Sharks and rays a global survey, sharks were not observed on with 10 or fewer sightings. We estimated reef-
(hereafter “elasmobranchs”) have diverse roles nearly 20% of reefs surveyed (6). Yet until re- level depletion for the nine most commonly
cently, reef shark species were listed in lower occurring species of shark [n = 5; Caribbean
risk extinction categories by the International reef sharks (Carcharhinus perezi) and nurse
Affiliations are listed at the end of this paper.
*Corresponding author. Email: [email protected] Union for the Conservation of Nature (IUCN). sharks (Ginglymostoma cirratum) in the Atlantic;
†Deceased. With ~37% of all elasmobranch species threat- grey reef sharks (Carcharhinus amblyrhynchos),

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RES EARCH | R E S E A R C H A R T I C L E

blacktip reef sharks (Carcharhinus melanopterus), A

and whitetip reef sharks (Triaenodon obesus) 30N
in the Indo-Pacific] and rays [n = 4; yellow
stingrays (Urobatis jamaicensis) and southern
stingrays (Hypanus americanus) in the Atlan- 15N
tic; blue spotted mask rays (Neotrygon spp.)
and blue spotted ribbontail rays (Taeniura

Latitude
lymma and Taeniura lessoni) in the Indo-
0
Pacific]. The Galapagos shark was excluded Depletion
from estimates of global depletion because 1.00
sampling only covered a relatively small pro- 0.75
portion of its range, but the results for this 15S 0.50
species were broadly similar. The nine key 0.25
resident species represented 77.7% of all elas- 0.00
mobranchs observed in the study and are 30S
those that serve important ecological roles
(10) and contribute the most to, and under- 30E 60E 90E 120E 150E 180E 150W 120W 90W 60W 30W
Longitude

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pin, livelihoods through fishing (11) and dive
tourism (12). B C D
We found that mean depletion of five key 4 1.00 25
Basin

Prop. elasmobranch MaxN comprised shark

resident reef sharks on individual reefs ranged
Abundance (elasmobranch MaxN per drop)

from 100% depletion (none observed) to 0% Atlantic

Indo−Pacific
(no depletion), averaging 62.8% (Fig. 1A). Mean 20
depletion of key resident reef sharks followed 3 0.75
the overall decline in elasmobranch abun-

Species richness
dance as measured with MaxN (Fig. 1B), de-
15
creased as the fraction of the elasmobranch
assemblage comprised of sharks decreased 2 0.50
(Fig. 1C), and showed little change across a
range of elasmobranch species richness (Fig. 10
1D); these patterns were generally consistent
between ocean basins. Across the range of
1 0.25
depletion, five main clusters of reefs were iden- 5
tified in the Atlantic, and eight were identified
in the Indo-Pacific (Figs. 2 and 3), including at
least one cluster in each ocean basin (cluster
1 in the Atlantic and cluster 2 in the Indo- 0 0.00 0
Pacific) having shark populations in a rela- 1.00 0.75 0.50 0.25 0.00 1.00 0.75 0.50 0.25 0.00 1.00 0.75 0.50 0.25 0.00
tively intact state, with low levels of depletion Shark depletion Shark depletion Shark depletion
of the five main resident reef shark species
(Caribbean reef and nurse sharks in the At- Fig. 1. The global decline of coral reef elasmobranchs. (A) Reef-scale estimates of depletion of
lantic; grey reef, blacktip reef, and whitetip resident coral reef shark species. Depletion is proportion of unfished population lost, represented as
reef sharks in the Indo-Pacific) (8). Remain- the measured MaxN as a proportion of MaxN in an unfished state (gravity, lowest in basin; MPA status,
ing clusters represented assemblages with closed) (8). Open circles indicate no sharks or rays were observed; gray circles indicate none of the
increasing depletion of resident shark spe- resident shark species used to calculate mean depletion were present. (B) Relationship between
cies and greater proportions of the overall elas- depletion of resident shark species and MaxN by ocean basin. (C) Relationship between depletion of
mobranch assemblage represented by rays resident shark species and the proportion of elasmobranch MaxN that comprised shark, demonstrating
(Figs. 2C and 3B). Both ocean basins show a the transition from shark- to ray-dominated assemblages. (D) Relationship between depletion of
similar transition through these assemblages resident shark species and species richness.
as key resident shark species became depleted.
The four key ray species (yellow and south-
ern stingrays in the Atlantic; blue spotted
mask and blue spotted ribbontail rays in the ment factors, with linear discriminant analysis tected area (MPA) or whether a reef was within
Indo-Pacific) increased only with depletion of (LDA) accounting for ~85% of variance be- a nation where all targeted shark fishing and
one or more resident reef shark species, with tween clusters (tables S2 and S3). Important trade is prohibited, known as a “shark sanc-
rays dominating in the most shark-depleted socioeconomic factors included the Human tuary.” Given that shark sanctuaries have
areas. These predictable changes in assem- Development Index (an index of a nation’s largely been implemented in nations in which
blage provide the ability to infer the status of level of education, life expectancy and stan- fishing for sharks was limited for economic
reef shark populations, and the level of hu- dard of living) and Voice and Accountability or cultural reasons (6), their effectiveness as
man pressure they are experiencing, in future Index (an index of the extent to which people tools for recovering reef shark populations
surveys. in each nation can participate in governance, remains an open question. Total market grav-
Elasmobranch species assemblage clusters free expression, free media, and free associ- ity was more important in the Indo-Pacific
on reefs in both basins were significantly relat- ation). Important management factors were than the Atlantic, possibly because remote reefs
ed to certain socioeconomic and manage- whether the reef occurred in a marine pro- (>4 hours travel time from human settlements)

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RES EARCH | R E S E A R C H A R T I C L E

Fig. 2. Structure of shark and ray assemblages A 90W 80W 70W 60W B 45W 30W
on Atlantic coral reefs. (A and B) Clusters of 0 0
reefs with similar species composition from 30N Cluster 30N
1 6
UPGMA clustering of 106 reefs in the Atlantic 2 10
Brazil

Latitude
basin based on a global set of 31 coral reef– 3 10S 10S
associated species. Five main clusters, 20N 20N
representing 87.0% of reefs, were identified.
Their locations are indicated with colored triangles. 20S 20S
Reefs with minor clusters are indicated with gray 10N 10N
dots (n = 7). Reefs where no elasmobranchs were
90W 80W 70W 60W 45W 30W
observed are indicated with black dots (n = 5). Longitude Longitude
(C) Regime plot showing all species assemblage
C 1.00 0.75 0.50 0.25 0.00 D Caribbean reef shark
clusters as a function of the mean depletion of the 1.0
1.00 1 1.00
resident reef shark species (Caribbean reef and 0.5
nurse sharks) and the proportion of all observed 1 0.0
elasmobranchs that were sharks. Size of points 25 1 3 2 6 10
12 Nurse shark
(and numbers) indicate the number of reefs in 0.75 0.75
1.0
Proportion of shark

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each cluster, and colors indicate cluster identity as

Proportion original
0.5
per (A). (D) Population level relative to original levels
0.0
of four resident reef species in each of the five main 2
1 3 2 6 10
0.50 0.50
clusters. Proportion of original level = 1 – depletion.
Southern stingray
Horizontal lines indicate mean, boxes indicates
4
25 to 75 percentile, and whiskers indicate 95% 14 6 2
confidence interval. 0.25 6 0.25 0
1 3 2 6 10
2
Yellow stingray

0.00 0.00 4
2
1.00 0.75 0.50 0.25 0.00 0
Shark depletion 1 3 2 6 10

are relatively rare in the Atlantic compared (fig. S2). Among rays, deficits were even more tional depletion levels. To estimate an overall
with the Indo-Pacific (fig. S1) (13). Environ- stark: 78.9% (75 of 95) for yellow stingray, global depletion level by species, we weighted
mental factors (coral cover and relief) had lit- 62.8% (81 of 129) for blue spotted ribbontail the jurisdictional depletion by the percent-
tle influence in predicting cluster membership. rays, and 55.6% (79 of 142) for blue spotted age of the world’s coral reefs in their waters
Elasmobranch assemblage structure on coral maskrays. An exception was the southern sting- and produced a weighted global mean de-
reefs in both the Atlantic and Indo-Pacific are ray, which was not detected at only 19.8% (n = pletion (8). Extinction risk was estimated
therefore mainly driven by management and 20 of 101) of expected reefs in the Atlantic. A by comparing proportional global depletion
socioeconomic factors, with shark-dominated failure to detect rays may not always indicate to the criteria for the IUCN Red List A2 (pop-
assemblages more likely to occur in wealthy, absence because they are often cryptic and ulation decline) category (17), assuming that
well-governed nations and in highly protected therefore missed on BRUVS, especially when the decline had occurred in the past three
areas or shark sanctuaries, whereas poverty, sharks are present (14). Collectively, these di- generations (29 to 90 years). In IUCN assess-
limited governance, and a lack of shark pro- versity deficits show that elasmobranch loss ments before the availability of this global
tection are associated with assemblages mainly on coral reefs is more extensive than previ- survey, all reef-resident shark species were
composed of rays. ously demonstrated, with widespread losses considered at lower risk of extinction (Near
To further characterize the diversity deficits of key species across many of the world’s coral Threatened) (18). Grey reef shark had the
that underpin these assemblage differences, we reefs, especially in Asia, eastern Africa, conti- highest level of global decline [69.8% ± 1 stan-
compared species observations in our BRUVS nental South America, and the central-eastern dard error (SE) 62.6 to 77.1], followed by
with their historical ranges drawn from pub- Caribbean. nurse shark (68.6% ± 49.7 to 87.4), Caribbean
lished literature, including historical accounts, Previous estimates of the status of reef shark reef shark (64.8% ± 42.0 to 87.5), blacktip
and found that sharks were not detected at and ray species have been geographically reef shark (64.5% ± 58.7 to 70.4), and white-
13.6% of reefs (19 Atlantic and 34 Indo-Pacific), limited, varying among surveyed reefs from tip reef shark (60.4% ± 51.2 to 70.2) (Fig. 4).
whereas rays were not detected at 21.5% of very high abundances (15) to local extinction The estimated declines of resident species of
reefs (10 Atlantic and 74 Indo-Pacific); both (16). This disparity has made it difficult to reef sharks met the IUCN Red List criteria for
groups were not detected at 6.6% of reefs sur- assess the global status of individual species. Endangered. Population changes of rays were
veyed (5 Atlantic and 19 Indo-Pacific). At the Therefore, we used our estimates of reef-level more variable, with increasing populations
species level, absences were severe. On the depletion to estimate the global depletion in some nations and declines in others (fig.
basis of their known historic distribution, def- and extinction risk of the most common res- S3), reflecting the compositional changes seen
icits were 46.9% of reefs (112 of 246) for black- ident reef sharks (five species) and rays (four across our gradient of human pressures. When
tip reef sharks, 41.3% (31 of 75) for Caribbean species). Mean and standard error reef-level examined at the global level, no ray species ex-
reef sharks, 40.8% (102 of 250) for grey reef depletion was calculated within jurisdic- amined met criteria for elevated extinction risk,
sharks, 36.2% (89 of 246) for whitetip reef tions (nations or remote territories) and used which is consistent with current nonthreatened
sharks, and 34.7% (n = 26 of 75) for nurse sharks to produce confidence intervals for jurisdic- status of these species on the Red List.

Simpfendorfer et al., Science 380, 1155–1160 (2023) 16 June 2023 3 of 6

RES EARCH | R E S E A R C H A R T I C L E

A 30E 60E 90E 120E 150E 180E 150W 120W 90W

30N 30N
Cluster
1
15N 15N
2

Latitude
5
0 0 7
10
15S 15S 13
14
18
30S 30S
30E 60E 90E 120E 150E 180E 150W 120W 90W
Longitude
B 1.00 0.75 0.50 0.25 0.00 C Grey reef shark
1.0
0.5
1.00 1.00 0.0
7 2
85 2 5 1 10 13 14 18 7
1

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57 Blacktip reef shark
6 1.0
27 0.5
0.75 1 0.75 0.0
3 2 5 1 10 13 14 18 7
Proportion of shark

Proportion original Whitetip reef shark

1.0
0.50 2 0.50 0.5
0.0
2 5 1 10 13 14 18 7

4 12 Blue spotted maskray

0.25 3 0.25 2
22
0
2 5 1 10 13 14 18 7
1
Blue spotted ribbontail ray
0.00 1
6 0.00 3
2
1
1.00 0.75 0.50 0.25 0.00 0
Shark depletion 2 5 1 10 13 14 18 7

Fig. 3. Structure of shark and ray assemblages on Indo-Pacific coral reefs. resident species of reef shark (grey reef, blacktip reef, whitetip reef, and
(A) Clusters of reefs with similar species composition from UPGMA clustering Galapagos sharks) and the proportion of all observed elasmobranchs that were
of 285 reefs in the Indo-Pacific basin based on a global set of 31 coral reef- sharks. Size of points (and numbers) indicate the number of reefs in each cluster,
associated species. Eight main clusters, representing 82.1% of reefs, were and colors indicate cluster identity as per (A); minor clusters are indicated
identified. Their locations are indicated with colored triangles. Reefs with minor in gray. (C) Population level relative to original levels of five core shark and
clusters are indicated with gray dots (n = 30). Reefs where no elasmobranchs ray species in each of the eight main species assemblage clusters. Proportion
were observed are indicated with black dots (n = 21). (B) Regime plot showing of original level = 1 – depletion. Horizontal lines indicate mean, boxes indicate
all species assemblage clusters as a function of the mean depletion of the 25 to 75 percentile, and whiskers indicate 95% confidence interval.

Our study of nations hosting ~90% of global These changes wrought on coral reef elasmo- tant. Thus, despite populations being func-
reefs reveals that resident reef shark species branch assemblages demonstrate the per- tionally extinct at the reef level, the potential
are at much higher risk of extinction than vasiveness of fishing on coral reefs (19) and to rebuild abundances remains relatively high
previously thought. Local declines, shaped the substantial risks to reef-dependent hu- if there are protected areas or strong fisheries
by human pressures that vary across ocean man communities of continued overfishing. management within a region (6). These source
basins, have led to consistent changes in the Elasmobranch species vary widely in their eco- populations are present among many small
structure of coral reef elasmobranch assem- nomic value, with some fished for subsistence, oceanic islands where low human populations
blages that may have profound effects on others fished for local or export markets, and and the high cultural value of sharks has re-
the broader ecosystem. The direct and indi- others valued alive as tourism resources (12, 20). sulted in fishing levels that are below those
rect effects of fishing have driven shifts in Thus, understanding threats and conservation seen elsewhere (21). MPAs also provide the
species composition from shark-dominated options for rebuilding populations at a species opportunity to act as source populations; how-
to ray-dominated assemblages and ultimate- level will assist in developing effective man- ever, their designation alone is insufficient to
ly the complete loss of sharks and rays at a agement of coral reef elasmobranchs as part deliver benefits. As others have observed (22),
small proportion (~ 7%) of reefs surveyed. In of a sustainable social-ecological system. high compliance is required. We show that
addition to changes in the structure of as- Although reef sharks are at considerable there are reefs in regions with widespread
semblages, all major resident shark species risk over broad spatial scales, our results show depletion of reef shark species that had metrics
have declined to such levels that they qualify that declines at one reef will have little effect indicating that they are in a relatively healthy
as Endangered by the IUCN Red List Criteria. on reefs tens to hundreds of kilometers dis- state compared with those around them. These

Simpfendorfer et al., Science 380, 1155–1160 (2023) 16 June 2023 4 of 6

RES EARCH | R E S E A R C H A R T I C L E

A Sri Lanka
B
Kenya Martinique

Qatar Species
Japan Jamaica Caribbean reef
Indonesia Nurse
Tanzania Montserrat
Taiwan
Saudi Arabia (Red Sea) Dominican Republic
Philippines
Vanuatu Species
Barbados
USA−Pacific Blacktip reef
Madagascar Grey reef
Colombia
South Africa Whitetip reef
Mayotte
USA−Western Atlantic
Mozambique

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Christmas Island
American Samoa Trinidad and Tobago
Malaysia
Nation

Guam Brazil
Samoa
Papua New Guinea Cuba
Seychelles
Cook Islands
Belize
Maldives
French Polynesia
Pedro Bank
Australia−Indian Ocean
Fiji
The Bahamas
Kiribati
Fed. States of Micronesia
Niue Turks and Caicos
Cocos−Keeling
Palau Colombia (Offshore Islands)
Solomon Islands
Australia−Pacific Dutch Caribbean South
New Caledonia
Tonga Antigua and Barbuda
Jarvis Island
1.00 0.75 0.50 0.25 0.00 1.00 0.75 0.50 0.25 0.00
Depletion level Depletion level

Fig. 4. Depletion of core coral reef shark species in the Indo-Pacific and Atlantic basins at national or near-national scale. (A) Indo-Pacific basin.
(B) Atlantic basin. Depletion was calculated by comparing reef-level species MaxN values to unfished, estimated by using a linear model in which market
gravity (a measure of the human pressure from population and access to reefs) was set to the ocean basin minimum and reef protected status was
“closed” (no take MPA) (8). Reef-level depletion scores were modeled by nation and used to estimate a global level of depletion (vertical dashed
lines) ± 1 standard error (shaded area) calculated by weighting national-level depletion by coral reef area (as a percent of global total coral reef area that
occurs within the range of each shark species).

included Tubbataha (Philippines), Sipidan species-specific reef shark management pro- nations to develop, implement, and enforce
(Island Malaysia), Glover’s Reef and Light- vides the best way forward for conservation regulations, and the likelihood that fishers
house Reef (Belize), and Misool (Indonesia); and rebuilding of reef sharks in places where comply with regulations, will be critical to
in all of these locations, there are programs to they have declined, among nations with the maintaining or rebuilding populations and
actively manage and enforce MPA regulations desire and capacity to do so (7, 8). Recent diverse elasmobranch assemblages. If not ad-
that are likely to account for these successes studies show that populations of reef sharks dressed, pressures causing the shark and ray
(23–25). can rebound in under a decade if appropri- diversity deficits we outline will continue to
Multiple nations have strong management ate management strategies that reduce fish- result in a loss of species, ecological func-
measures (such as spatial protections and/or ing pressure are in place (26). Although direct tions, and ecosystem services that support
fishing restrictions) in place that benefit management is critical, local and national sustainable livelihoods for millions of people
reef species. This study builds the case that socioeconomic factors that affect the ability of worldwide.

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72
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 Widespread diversity deficits of coral reef sharks and rays
Colin A. Simpfendorfer, Michael R. Heithaus, Michelle R. Heupel, M. Aaron MacNeil, Mark Meekan, Euan Harvey, C.
Samantha Sherman, Leanne M. Currey-Randall, Jordan S. Goetze, Jeremy J. Kiszka, Matthew J. Rees, Conrad W. Speed,
Vinay Udyawer, Mark E. Bond, Kathryn I. Flowers, Gina M. Clementi, Jasmine Valentin-Albanese, M. Shiham Adam,
Khadeeja Ali, Jacob Asher, Eva Aylagas, Ocane Beaufort, Cecilie Benjamin, Anthony T. F. Bernard, Michael L. Berumen,
Stacy Bierwagen, Chico Birrell, Erika Bonnema, Rosalind M. K. Bown, Edward J. Brooks, J. Jed Brown, Dayne Buddo,
Patrick J. Burke, Camila Cceres, Marta Cambra, Diego Cardeosa, Jeffrey C. Carrier, Sara Casareto, Jennifer E. Caselle,
Venkatesh Charloo, Joshua E. Cinner, Thomas Claverie, Eric E. G. Clua, Jesse E. M. Cochran, Neil Cook, Jessica E.
Cramp, Brooke M. DAlberto, Martin de Graaf, Mareike C. Dornhege, Mario Espinoza, Andy Estep, Lanya Fanovich,
Naomi F. Farabaugh, Daniel Fernando, Carlos E. L. Ferreira, Candace Y. A. Fields, Anna L. Flam, Camilla Floros, Virginia

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Fourqurean, Laura Gajdzik, Laura Garca Barcia, Ricardo Garla, Kirk Gastrich, Lachlan George, Tommaso Giarrizzo, Rory
Graham, Tristan L. Guttridge, Valerie Hagan, Royale S. Hardenstine, Stephen M. Heck, Aaron C. Henderson, Patricia
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Jaiteh, Mohini Johnson, Stacy D. Jupiter, Muslimin Kaimuddin, Devanshi Kasana, Megan Kelley, Steven T. Kessel,
Benedict Kiilu, Taratau Kirata, Baraka Kuguru, Fabian Kyne, Tim Langlois, Frida Lara, Jaedon Lawe, Elodie J. I. Lde,
Steve Lindfield, Andrea Luna-Acosta, Jade Q. Maggs, B. Mabel Manjaji-Matsumoto, Andrea Marshall, Lucy Martin, Daniel
Mateos-Molina, Philip Matich, Erin McCombs, Ashlie McIvor, Dianne McLean, Llewelyn Meggs, Stephen Moore, Sushmita
Mukherji, Ryan Murray, Stephen J. Newman, Josep Nogus, Clay Obota, Domingo Ochavillo, Owen O'Shea, Kennedy E.
Osuka, Yannis P. Papastamatiou, Nishan Perera, Bradley Peterson, Caio R. Pimentel, Fabin Pina-Amargs, Hudson T.
Pinheiro, Alessandro Ponzo, Andhika Prasetyo, L. M. Sjamsul Quamar, Jessica R. Quinlan, Jos Amorim Reis-Filho, Hector
Ruiz, Alexei Ruiz-Abierno, Enric Sala, Pelayo Salinas de-Len, Melita A. Samoilys, William R. Sample, Michelle Schrer-
Umpierre, Audrey M. Schlaff, Kurt Schmid, Sara N. Schoen, Nikola Simpson, Adam N. H. Smith, Julia L. Y. Spaet, Lauren
Sparks, Twan Stoffers, Akshay Tanna, Rubn Torres, Michael J. Travers, Maurits van Zinnicq Bergmann, Laurent Vigliola,
Juney Ward, Joseph D. Warren, Alexandra M. Watts, Colin K. Wen, Elizabeth R. Whitman, Aaron J. Wirsing, Aljoscha
Wothke, Esteban Zarza-Gonzlez, and Demian D. Chapman

Science, 380 (6650), .

DOI: 10.1126/science.ade4884

Editor’s summary
In recent years, much attention has been given to catastrophic declines in sharks. Most of this attention has focused
on large pelagic species that are highly threatened by direct and indirect harvest. Simpfendorfer et al. looked globally
at the smaller, coral reef–associated species of sharks and rays and found steep declines in shark species (see the
Perspective by Shiffman). Five of the most common reef shark species have experienced a decline of up to 73%.
As shark species decline on coral reefs, ray species increase, indicating a community-wide shift. Species are best
protected when active protections are in place, suggesting routes for better conservation. —Sacha Vignieri

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