FISHERIES The smallworld of global marine fisheries:The ... · longest coastlines. In the...
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FISHERIES
The small world of global marinefisheries: The cross-boundaryconsequences of larval dispersalNandini Ramesh1*, James A. Rising2, Kimberly L. Oremus3
Fish stocks are managed within national boundaries and by regional organizations, butthe interdependence of stocks between these jurisdictions, especially as a result oflarval dispersal, remains poorly explored. We examined the international connectivity of747 commercially fished taxonomic groups by building a global network of fish larvaldispersal. We found that the world’s fisheries are highly interconnected, forming asmall-world network, emphasizing the need for international cooperation. Wequantify each country’s dependence on its neighbors in terms of landed value, foodsecurity, and jobs. We estimate that more than $10 billion in annual catch from 2005to 2014 is attributable to these international flows of larvae. The economic risksassociated with these dependencies is greatest in the tropics.
Marine fisheries supply food and liveli-hoods to millions of people around theworld (1). Though fisheries are typicallymanaged at the scale of national exclu-sive economic zones (EEZs), many fish
populations are connected beyond EEZ bound-aries (2–6). Whereas pelagic species can betracked across international borders as adults(7), nonpelagic populations connect primarilyvia the dispersal of fish eggs and larvae, formsthat cannot yet swim by ocean currents (2, 8).
Larval connectivity patterns have been ana-lyzed at both the regional (6, 9–12) and global(4, 13, 14) levels and have been used to sug-gest changes for spatial management andconservation (12, 15). However, the impacton fisheries of larval connectivity across EEZsis not well understood, even though morethan 90% of the world’s fish are caught with-in EEZs (16).On the scale of a single species or region,
this connectivity can be analyzed empirically
through genetic testing (9, 10). For analyses onlarger scales, dispersal patterns can be estimatedusing biophysical models that combine oceano-graphic data with an understanding of the bi-ology of the stocks (4, 14, 17). Such efforts can bechallenging, because species vary widely in larvaltiming and duration and currents vary with theseasons; therefore, generalizations can be mis-leading. More realistic inputs can be achievedby using life history traits for each species, in-cluding time and place of spawning and larvalduration. Sensitivity analyses can help to en-sure that results are robust to changes in keyassumptions (14), while empirical bounding cansafeguard against predicting unrealistic disper-sion outcomes (6).Network analysis has previously been applied
to marine systems to describe the connectivity ofplankton communities (18), local fishing com-munities (19, 20), and marine reserves (14).Networks of larval flows have been used toidentify “hub” subpopulations for protectionon a regional scale (12).In this study, we combined oceanographic and
life history data for 706 species and 434 generaof commercially harvested fish to estimate theirconnectivity across 249 EEZs and constructeda network representing the larval flows between
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1Department of Earth and Planetary Science, Universityof California, Berkeley, Berkeley, CA 94720, USA. 2GranthamResearch Institute, London School of Economics, London,UK. 3School of Marine Science and Policy, Universityof Delaware, Newark, DE 19716, USA.*Corresponding author. Email: [email protected]
Mexico
Bermuda
Cuba
Nicaragua
Honduras
BahamasTurks and Caicos
Cayman Islands
Venezuela
Colombia - Jamaica
Colombia
Panama
Puerto Rico
British Virgin Islands
Anguilla
Saint Kitts and NevisAntigua and Barbuda
Barbados
Curacao
Saba
Northern Saint-Martin
Aruba
Sint-Eustasius
Martinique
US Virgin Islands
Saint Lucia
Grenada
Saint Vincent and the Grenadines
Trinidad and Tobago
Guyana
United States
Dominican Republic
Jamaica
Haiti
Costa Rica
Bonaire
Saint Pierre and MiquelonCanada
IcelandGreenland
Svalbard
Alaska
Guatemala
SpainPortugal
El Salvador
Suriname
Ecuador
Peru Brazil
French Guiana
Montserrat
Dominica
Guadeloupe
Argentina
Liberia
Equatorial GuineaSao Tome and Principe
Falkland Islands
Uruguay
Chile
Micronesia
Japan
Palau
Indonesia
Russia
South Korea
Philippines
Paracel Islands
Japan - South Korea Conflict Zone
Vietnam
Southern Kuriles
North Korea
Spratly Islands
Conflict ZoneChina
Taiwan
Japan - Korea
Brunei
Italy
Croatia
Turkey
Germany
Denmark
Poland
Sweden
LatviaLithuania
Norway Estonia
Bulgaria
RomaniaGeorgia
Ukraine
Iran
Tunisia
France
Algeria Libya
Malta GreeceAlbania
Cyprus
EgyptIsrael
Montenegro
Morocco
United Kingdom BelgiumNetherlandsIreland
JerseyGuernsey
Madeira
Faeroe IslandsJan Mayen
Canary IslandsWestern Sahara
SyriaLebanon
Saudi Arabia
SudanYemenEritreaUnited Arab Emirates
KuwaitBahrain
Qatar
Somalia
Djibouti
Mozambique
South Africa
Bassas da IndiaIle Europa
Comoro Islands
Mauritius
Kenya
Tanzania
Mayotte
Glorioso Islands
Ile Tromelin
British Indian Ocean Territory
Maldives
Namibia
Seychelles
Juan de Nova IslandMadagascar
Oman
Pakistan
India
Papua New Guinea
Malaysia
CambodiaThailand
Singapore
Myanmar
Kiribati
American Samoa
SamoaNiue
Tonga
Wallis and Futuna
Tuvalu
Fiji
Howland Isl. and Baker Isl.
Solomon Islands
New Caledonia
Nauru
Australia - Papua New Guinea
Australia
Sri Lanka
East Timor
Andaman and Nicobar
Australia - East Timor
Australia/Indonesia Vanuatu
Marshall Islands
Bangladesh
New Zealand
Macquarie Island
Finland
Angola
GabonRepublique du Congo
Sierra LeoneTogoBenin
Ghana
Ivory CoastNigeria
Cameroon
DR Congo
Cape Verde
Western Sahara/MauritaniaMauritania
GambiaSenegal
Guinea BissauGuinea
Reunion
AntarcticaAsia PacificCaribbeanEast AfricaMediterraneanMiddle East
North AmericaNorthern EuropeSouth AmericaSouth AsiaWest AfricaWest Pacific
source clockwise sink
Fig. 1. The network of spawn-attributed catch flows between EEZs. Each EEZ is a node (circle) of the network and its color represents its networkcommunity. The connectors or edges in this network flow clockwise from source to sink, with their thicknesses representing the magnitude of the net flowof caught biomass between the EEZs. Only the edges in the upper tercile of edge weights are shown, for clarity (see SM 3.2 for the full network). The sizeof each node represents its out-degree, i.e., the number of other EEZs for which it acts as a source of fish larvae, including connections not shown in this image.
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nations. Nations that depend heavily upon theirneighbors for recruitment risk losing part oftheir catch if the fisheries in the source EEZsoutside their jurisdiction are poorly managed.We quantified these risks in economic termsand identified regional “hotspots” of risk forcatch, fishery employment, and food security.We used a particle-tracking system (21) with
time-varying ocean currents (22) and species-specific life histories (23) to simulate the dis-persal of eggs and larvae through a dynamicocean. We placed multiple simulated particlesfor each species based on the timing and loca-tion of that species’ spawning and let them driftfor their larval duration to obtain a probabilisticestimate of species-specific larval trajectories.We used a random-walk parameterization (21)that adds a small velocity at every time step toaccount for turbulent motion at small scales[see section 3.1.2 in the supplementary mate-rials (SM 3.1.2)].We empirically bounded our results by dis-
carding particles that arrived in regions wherethe species is not present in observed catchdata (16). For a given EEZ, catch is attributedbased on the proportion of particles arrivingthere from each spawning country (see SM 1.1).This proportionality forms the core assumptionof our model. We tested our main results with aseries of analyses of sensitivity to this assumption.These included reducing spawn floating dura-tion to account for uncertainties in spawningmortality (2, 24), introducing return adult spawn-ing migration (25) (see SM 3.6), and distinguish-ing different levels of recruitment limitation.We estimated how much of each country’s ob-
served catch comes from its neighbors by con-structing for each species a transitionmatrix thatdescribes the probability of its offspring dispers-ing from one EEZ to another. This transfer ofbiomass between nations’ EEZs is representedas a network in Fig. 1.Each connector of the network represents
net flows of fish from one country to another.Countries that depend on inflows of juvenilefish to maintain their local populations requireinternational cooperation to ensure sustainablefisheries. Our analysis of these flows revealedthat a large proportion ofmarine fisherieswithinEEZs form a single, global network (Fig. 1).We found that the global network of marine
fisheries is a scale-free, small-world network.The scale-free network property, common innatural systems (26), is characterized by an ex-ponential distribution of the number of con-nections from each node (see SM 3.2). Thisexponential degree distribution results in a“hub-and-spoke” structure that is resilient torandom disturbances because of the large num-ber of less-connected countries from which dis-turbances do not easily propagate to other partsof the network. However, a disturbance to anyof the highly connected hubs in a scale-free net-work can affect numerous surrounding nodes.In this context, habitat destruction, overfishing,or environmental change in a hub EEZ couldhave impacts that spread beyond its own bound-
aries. Conversely, targeted efforts to managefisheries within these hub EEZs could benefitmany nations.To demonstrate the relationship between cur-
rents and the network of larval dispersal, wefocused more closely on four regions (Fig. 2).The differences between the regional networksand average current speed arise from the details
of current speeds during spawning, larval du-ration, and empirical observations of speciespresence or catch. The influence of the GuineaCurrent on the connectivity of West Africa’s fish-eries can be seen in the large number of EEZsthat act as sources to their eastward neighbors,especially between Guinea-Bissau and Nigeria.While the strongest connections are typically
Ramesh et al., Science 364, 1192–1196 (2019) 21 June 2019 2 of 4
A EWest Africa
Baltic Sea
Caribbean
B F
C G
DH
Western Pacific
Asia PacificCaribbeanNorth AmericaNorthern EuropeSouth AmericaWest AfricaWest Pacific
source clockwise sink
Communities
Liberia
Canary Islands
Western Sahara
Sierra Leone Togo
BeninGhana
Ivory Coast
Nigeria
Cape Verde
Western Sahara/Mauritania
Mauritania
Gambia
Senegal
Guinea Bissau
Guinea
Russia
Germany
Denmark
Poland
Sweden
Latvia
Lithuania
Estonia
Finland
Norway
MicronesiaPalau
Indonesia
Philippines
Northern Mariana Islands and Guam
Paracel Islands
Spratly Islands
ChinaTaiwan
Brunei
Papua New Guinea
Malaysia
Kiribati
American Samoa
Line Group
Samoa Cook Islands
NiueTonga
Phoenix Group
Wallis and Futuna
Tuvalu
Jarvis Island
Tokelau
Fiji
Solomon Islands
New Caledonia
Nauru
Australia - Papua New Guinea
Australia
Australia - East Timor
Australia/Indonesia
Vanuatu
Norfolk Island
Marshall Islands
Christmas Island
New Zealand
Palmyra Atoll
Wake Island
Pitcairn
French Polynesia
Howland Island and Baker Island
Cuba
Nicaragua
Honduras
Bahamas
Cayman Islands
Venezuela
Colombia - Jamaica
Colombia
Panama
Puerto Rico
British Virgin Islands
Anguilla
Saint Kitts and NevisAntigua and Barbuda
Barbados
Curacao
SabaNorthern Saint-Martin
Aruba
Martinique
U.S. Virgin Islands
Saint Lucia
Grenada
Saint Vincent and the Grenadines
Trinidad and Tobago
Guyana
Dominican Republic
Jamaica
Haiti
Suriname
Brazil
French Guiana
Montserrat
DominicaGuadeloupe
Costa Rica
Fig. 2. Regional currents and community networks. (A to D) The speed (shown in colors, incentimeters per second) and direction (arrows) of ocean surface currents in four regions withinterconnected fisheries (West Africa, Baltic Sea, the Caribbean, and Western Pacific) duringthe month of maximum spawning activity in each (August, May, June, and May, respectively).(E to H) The corresponding subset of the global network encompassed by the four regions.Colors, node sizing, and connector directions are as for Fig. 1. Nodes are arranged toapproximately correspond to geographic locations of the EEZs.
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between adjacent EEZs, many connections alsoextend over longer distances. In contrast, theBaltic Sea has substantially weaker currents.There, the largest outward flows originate fromSweden and Norway, which have the region’slongest coastlines. In the Caribbean, the NorthBrazil Current flows northwestward along theSouth American coast, and consequently manyof the EEZs lying along this current act assources for the Lesser Antilles. Within the LesserAntilles, the density of small EEZs gives rise to ahighly interconnected, complex network struc-
ture. The effect of the northward flow along thisisland chain can be inferred from the larger nodesizes among the EEZs lying in its southern por-tion. In the Western Pacific, strong currentsdominate in the equatorial ocean, with weakercurrents at higher latitudes. The large areas en-compassed by this region’s EEZs mean that, un-like the other regions, most connections arebetween immediate neighbors.The small-world property implies that it is
possible to traverse the global network in a smallnumber of steps, on average. Within this net-
work, there exist smaller clusters or communi-ties that are tightly connected. Most of theseclusters internally exhibit the small-world prop-erty. In theory, this property of the global fish-eries network suggests that disturbances to alarge hub could propagate via cascading effectson the surrounding spokes.A key question is whether disruptions to a
given EEZ actually propagate in this manner. Astock’s response to external shocks depends onboth its population dynamics and mortality fromfishing, which can be affected by management
Ramesh et al., Science 364, 1192–1196 (2019) 21 June 2019 3 of 4
Catch outflows (1000 tons) Value outflows (USD)
0 500 1000 0 300M 600M 900M
IcelandPeru
DenmarkSweden
IndonesiaArgentina
BrazilTaiwan
AustraliaSouth Korea
NorwayPapua New Guinea
RussiaUnited States
United KingdomIndia
ChinaAlaskaJapan
Outflows to other EEZsCatch inflows (1000 tons) Value inflows (USD)
0 500 1000 0 300M 600M 900M
North KoreaSwedenUruguay
DenmarkChile
TaiwanKiribatiTurkey
MyanmarPapua New Guinea
United KingdomPakistanVietnam
JapanNorwayMexico
IndonesiaChina
South KoreaRussia
Inflows from other EEZs
Resilience:
High (> 99%)
Medium (> 95%)
Low (< 95%)
Resilience:
High (> 99%)
Medium (> 95%)
Low (< 95%)
Fig. 3. Countries with highest outflowing and inflowing catches.(Left) Top 20 countries sorted by total outflowing catch (in thousands oftons) and value [in U.S. dollars (USD)] at risk. (Right) Top 20 countriessorted by total inflow of catch (thousands of tons) and value (USD) at risk.
For both catch and landed values, data from 2005 to 2014 were usedand attributed to larvae, by species. Resilience levels represent theestimated decline a population can endure without being consideredvulnerable to local extinction.
Poland
under 129129 848848 53925392 1879718797 66848over 66848
Sweden
Belgium
Netherlands
Poland
Latvia
Estonia
Lithuania
Romania
Finland
North Korea
Bahamas
Sao Tomeand Principe
St. Lucia
Uruguay
Dominica
GuyanaSt. Vincent andthe Grenadines
Barbados
Suriname
GambiaMauritania
Cameroon
Guinea-Bissau
Comoros
Maldives
Pakistan
Bangladesh
Indonesia
Philippines MarshallIslands
Kiribati
NauruPapua New Guinea
TuvaluSolomonIslands
Palau
Catch Value
Food Security
GDP
Labor
< 129129-848848-5,3925,392-18,79718,797-66,848> 66,848N/A
Catch (1000 tons)
Fed. Statesof Micronesia
Fig. 4. Hotspot map showing fishing dependency on spawninggrounds in neighboring waters, by country. Countries are shadedby catch (in thousands of metric tons) at risk, with darker shadesrepresenting higher catches. Icons depict EEZs that are the mostdependent on their neighbors. The catch icon indicates that more than
30% of a country’s catch value is dependent on neighboring spawninggrounds, the GDP icon represents a risk to more than 0.8% of itsGDP, the labor icon represents that more than 1.5% of its jobs arevulnerable, and the food security icon represents a food securitydependence index >1.1%.
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(27). Some fish stocks are biologically capableof replenishing themselves when their numbersdwindle, provided that fishing pressure is re-lieved, reducing the likelihood that disturbanceswill propagate. However, “recruitment-limited”stocks are vulnerable to a decline in spawningpopulation, making it more likely that distur-bances will spread across the network even ifthe receiving fisheries are managed. We adoptedFishBase’s classification of stock resilienceas a proxy for this type of density dependence.For high-resilience stocks, which are generallynot recruitment limited, our measure of stockdependence overestimates the extent to whichstocks will be reduced if recruitment inflows fail.For those stocks classified as having medium orlow resilience, however, we found a strong cor-relation between our simulation’s predictionsand observed variances in stock levels (see SM3.5). Even for countries whose fisheries mostlycomprise non–density-dependent stocks, theselarval inflows serve as a buffer against fisherycollapse within their waters.To contextualize our results, we estimated the
economic significance of the network’s interna-tional connections. First, we considered theamount and value of catch that flows in and outof each EEZ (Fig. 3). Japan, China, and Alaskaare responsible for the greatest outflows, reflect-ing their productive waters. However, havingfewer neighbors makes them smaller hubs(Fig. 1). Indonesia has the most landed value at-tributable to other countries, due to its high-value catch andmany neighbors. The countrieswith the greatest catch inflows are generallythose with the largest fisheries.Next, we identified nations that are potentially
most vulnerable, in socioeconomic terms, tothe management of neighboring waters (seetable S5). In Fig. 4, we highlight countries thatdepend the most on the spawning grounds ofneighbors in terms of their total catch, grossdomestic product (GDP), number of jobs in thefishery industry, and a fishery food security de-pendence index (28). Themost vulnerable nationsare concentrated in the hotspot regions of theCaribbean, West Africa, Northern Europe, andOceania. The risks to national GDP and laborforce are generally highest in the tropics. How-ever, our measure of food security risk alsoidentified a few European nations.
Our analysis showed that about $10 billionworth of annual marine catch may rely ontransnational exchanges of fish offspring. Thesedependencies form a single global network, in-dicating that marine fisheries, even within na-tional boundaries, constitute an interconnected,globally shared resource.This network’s scale-free and small-world prop-
erties imply that fish stocks from a small num-ber of EEZs provide benefits to a large number of“downstream” countries. The most vulnerablenations are clustered in a few hotspot regions(Fig. 4). This pattern lends further support to theuse of international frameworks, such as largemarine ecosystems and marine protected areanetworks (29, 30).Further research is needed to understand how
small-scale coastal processes, larval behavior, andfisheries management affect this connectivity.Beyond the spawning connections studiedhere, national fisheries are interdependent throughthe movement of adult fish, population shiftsunder climate change, and international fishingtreaties. In particular, the role of adult fish mi-gration in driving international connectivity re-mains an important question. While a moredetailed analysis is required to accurately de-scribe dispersal pathways of individual species,this study highlights the role of larval connec-tivity across international boundaries and theneed for multilateral cooperation for sustain-able management of these shared resources.
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ACKNOWLEDGMENTS
The authors thank D. Dookie, M. Burgess, M. A. Cane,A. Chaintreau, A. Carlisle, J. Cohen, C. Costello, R. Defries,S. Gaines, S. Hsiang, C. Moffat, and C. Szuwalski for comments,suggestions, and references. This work was initiated and partlyconducted at Columbia University in the City of New York, whichprovided computing resources. Funding: N.R. was partiallysupported by the National Aeronautics and Space Administration(NASA) Headquarters under the NASA Earth and Space ScienceFellowship Program, grant NNX-14AK96H. Author contributions:N.R. performed the network analysis and Lagrangian modeling.J.A.R. performed the country-level risk analysis. N.R., J.A.R., andK.L.O. designed the study, collected data, and wrote the paper.Competing interests: The authors declare no competing interests.Data and materials availability: All newly organized data usedin this study and the intermediate and final results data arepublicly available at Zenodo (31). Analysis reproduction code isavailable at https://github.com/openmodels/small-world-fisheries.
SUPPLEMENTARY MATERIALS
science.sciencemag.org/content/364/6446/1192/suppl/DC1Materials and MethodsFigs. S1 to S13Tables S1 to S8References (32–44)
6 September 2018; accepted 23 May 201910.1126/science.aav3409
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dispersalThe small world of global marine fisheries: The cross-boundary consequences of larval
Nandini Ramesh, James A. Rising and Kimberly L. Oremus
DOI: 10.1126/science.aav3409 (6446), 1192-1196.364Science
, this issue p. 1192Sciencemanagement, and food supplies globally.particular hubs of productivity are widely important. Such connectivity has wide-ranging implications for conservation,that global fish populations represent a small-world network where connections across populations are tight and
seemsanalysis to assess the degree to which populations found in one part of the world may have come from another. It model how these currents distribute the fish larvae of more than 700 species. They used networket al.currents. Ramesh
marine fish, perhaps more than any other vertebrate group, are connected across large distances through ocean Countries manage their fisheries as if they were a local resource. To some degree, this may reflect reality, but
A small, interconnected world
ARTICLE TOOLS http://science.sciencemag.org/content/364/6446/1192
MATERIALSSUPPLEMENTARY http://science.sciencemag.org/content/suppl/2019/06/19/364.6446.1192.DC1
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