Aquaculture 576 (2023) 739837

Contents lists available at ScienceDirect

Aquaculture
journal homepage: www.elsevier.com/locate/aquaculture

A global assessment of species diversification in aquaculture

Junning Cai a, *, Hing Ling Chan b, Xue Yan c, PingSun Leung d
a
Fisheries and Aquaculture Division, Food and Agriculture Organization of the United Nations (FAO), Viale delle Terme di Caracalla, 00153 Rome, Italy
b
Pacific Islands Fisheries Science Center, NOAA - National Oceanic & Atmospheric Administration, Honolulu, United States of America
c
Chinese Academy of Fishery Sciences (CAFS), Beijing, China
d
University of Hawai'i at Manoa, Honolulu, United States of America

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

Keywords: Aquaculture is a diverse food production system, and a high species diversity in aquaculture can make the global
Aquaculture food system more resilient. Species diversification could facilitate aquaculture growth through multiple mech­
Diversity anisms and increase the sector's resilience and long-term sustainability. Facing increasing challenges from
Species diversification
climate change, disease outbreaks, market fluctuations, and other disturbances, species diversification has
Effective number of species
Within-group diversity
become a widely recognized and endorsed development strategy in the policy and scientific communities for the
Between-group diversity growth and resilience of the aquaculture sector. However, many attempts to establish new species have yielded
little long-term success, and the private sector often concentrates efforts on the most advantageous species for
rapid growth. This paper presents a comprehensive assessment of aquaculture species diversification at the
global, regional (more than 30 country groups), and national (nearly 200 countries) levels, covering the period
1950 to 2020 with a focus on 1990 to 2020. The assessment employs the concept of “effective number of species”
as a measure of diversity and uses two sub-indicators to assess within-group versus between-group diversity. The
indicator system reveals detailed patterns of species diversity and uncovers different driving forces of species
diversification. Additionally, the study maps the status and trends of species diversity to offer a refined
perspective on species diversity profiles and diversification patterns. The findings reveal that beneath high
species diversity in world aquaculture lies generally low diversity at the national level; nearly half of national
aquaculture has no within-group or between-group diversity. Furthermore, species diversification has been
losing momentum in recent decades, and concentration has become a more prominent development pathway,
with a tendency to drive aquaculture towards a low-diversity system similar to terrestrial farming. Public in­
terventions are crucial to promote species diversification in aquaculture for the sector's resilience and long-term
sustainability. It is important to not only reduce the cost of species diversification for the private sector but also
to dedicate more public efforts towards increasing its benefits and viability. While diverse diversification patterns
among national aquaculture indicate no one-style-fits-all species diversification pathway, a country may draw
guidance from others' experiences, especially since countries with similar species diversity profiles or diversi­
fication patterns tend to cluster geographically. Evidence-based policymaking and sector management regarding
species diversification entail collaborative efforts among policymakers, scientists, and the aquaculture commu­
nity to expand and refine assessment frameworks, improve data availability and quality, and efficiently utilize
information, knowledge, and insights generated by these assessments to inform decision making at various levels.

1. Introduction
As a rapidly expanding sector in the food industry, aquaculture
stands out for its diverse range of species compared to terrestrial
farming, which could enhance the resilience of the global food system
(Troell et al., 2014; FAO, 2019a). Since 1950, more than 600 aquatic
species have been farmed in global aquaculture, with more than 400
species still being actively cultivated. Among these, 10 species account
for approximately half of the total production (FAO, 2019b). In contrast,
two thirds of global crop production come from 10 species (out of <200
plant species with significant production globally), while only ~40
terrestrial animal species are cultivated for food such as meat, milk, and
eggs, and production is concentrated in only a handful, although di­
versity comes from numerous breeds and varieties (FAO, 2019c).

* Corresponding author.
E-mail addresses: junning.cai@fao.org (J. Cai), hingling.chan@noaa.gov (H.L. Chan), yanx@cafs.ac.cn (X. Yan), psleung@hawaii.edu (P. Leung).

https://doi.org/10.1016/j.aquaculture.2023.739837
Received 17 February 2023; Received in revised form 21 May 2023; Accepted 25 June 2023
Available online 26 June 2023
0044-8486/© 2023 Food and Agriculture Organization of the United Nations. Published by Elsevier B.V
J. Cai et al.
High species diversity could also be advantageous for the aquacul­
ture sector per se. Technically, species diversification can increase
farming efficiency through polyculturing multiple species in the same
farming system, cultivating the most suitable species in different
farming environments, or rotating different species according to sea­
sonal variations (Ravisankar et al., 2005; Thomas et al., 2021; Newton
et al., 2021). Economically, species diversification can help the sector
overcome market satiation and broaden its market base (Abellán and
Basurco, 1999; Liao, 2000). Finally yet perhaps most importantly, spe­
cies diversification can enhance aquaculture's resilience against climate
change, disease outbreaks, market fluctuations, and other shocks that
pose challenges to the sector's long-term sustainability (Wilson and
Archer, 2010; Gephart et al., 2017; Metian et al., 2020). Therefore,
species diversification is widely endorsed as a prominent strategy for
sustainable aquaculture development by both policy and scientific
communities (Abellán and Basurco, 1999; CIHEAM, 2000; Naylor and
Burke, 2005; Brummett, 2007; Martínez-Cordero, 2007; Megahed and
Aly, 2009; Le Francois et al., 2010; FAO, 2011; Schmidt et al., 2011; Guy
et al., 2014; Troell et al., 2014; Harvey et al., 2017; Roy, 2019; Boyd
et al., 2020; Metian et al., 2020; García-Márquez et al., 2021; Thomas
et al., 2021; Oboh, 2022).
However, although aquaculture is one of the most diverse food
production systems at the global level, species diversity is generally low
in national aquaculture (Metian et al., 2020; Cai et al., 2022). Many
attempts to promote new species have yielded little long-term success
(Metian et al., 2020; Harvey et al., 2017; Teletchea and Fontaine, 2014;
Muñoz-Lechuga et al., 2018). Failure to establish a novel species could
be attributed to various hindrances, such as technical difficulties
(Fernández-Polanco and Bjorndal, 2017), limited markets (Basurco and
Abellán, 1999), or institutional constraints (Barrington et al., 2009). Yet
fundamentally, when aquaculture species that appear promising indi­
vidually compete for limited resources and markets, market mecha­
nisms, such as economies of scale and maximization of risk-adjusted
return, tend to make aquaculture production concentrated towards a
few winner species (Cai et al., 2022).
Should a country pursue species diversification or adopt a “concen­
tration” pathway to focus on a few most productive species for high
growth and exploit their intra-specific potential for diversity? The
answer tends to vary for different countries or for the same country at
different stages of aquaculture development, as both pathways have
pros and cons (Bilio, 2008; Teletchea and Fontaine, 2014). Information,
knowledge, and understanding of past experiences of species diversifi­
cation is crucial for guiding aquaculture development, especially for
countries with a less developed aquaculture sector.
Here we conduct a comprehensive, in-depth assessment of aquacul­
ture species diversification at the global, regional (30+ country groups),
and national (~200 countries) levels. Our analysis covers seven decades
(1950–2020), with an emphasis on the past three decades (1990–2020).
Similar to Cai et al. (2022), we use the “effective number of species” as a
measure of “true” diversity (Hill, 1973; Jost, 2006), which is more
intuitive than the Shannon (entropy) index used by Metian et al. (2020).
While Metian et al. (2020) quantified the diversity of aquaculture spe­
cies at the national level, we measure it at the national, regional, and
global levels. This helps identify inconsistent species diversification
patterns at different geographic scales. We further decompose the
effective number of species into two sub-indicators: one captures di­
versity among species groups (i.e. between-group diversity) and the
other measures species diversity within species groups (i.e. within-group
diversity). The decomposition reveals detailed patterns of species di­
versity and uncovers different driving forces of species diversification.
Similar to Metian et al. (2020), we map species diversity to examine its
geographic patterns. Yet the scope of our mapping is broader, including
both a comparison of species diversity across countries and a categori­
zation of species diversification trends.
We highlight and discuss key findings in the main text. Some results
may be part of conventional wisdom, yet most are less apparent or
2
Aquaculture 576 (2023) 739837
underappreciated. More comprehensive results are documented in
supplementary materials, which can be used to facilitate evidence-based
policymaking and sector management in aquaculture at the global,
regional, and national levels. We conclude the paper with discussion of
lessons learned, future prospects, and way forward, including potential
areas for further study.
2. Data and methods
2.1. Data
We use aquaculture production statistics from the Food and Agri­
culture Organization of the United Nations (FAO) Global Aquaculture
Production Statistics 1950–2020 (FAO, 2022). Reporting entities in the
database are classified as countries, including non-sovereign territories.
In order to facilitate analysis, we combine the statistics of mainland
Tanzania and Zanzibar into the United Republic of Tanzania. For all
other reporting entities, we use the classifications adopted in the FAO
database. In addition to assessments at the country and global levels, we
also examine species diversity and diversification patterns in 32 coun­
tries groups. For narrative convenience, we refer to aquaculture in these
32 groups as “regional aquaculture” (Table 1).
All ASFIS (Aquatic Sciences and Fisheries Information System) spe­
cies items recorded in the database are included in our analysis. These
species items could refer to individual species, hybrids or “nei” (not
elsewhere included) species items that are groups of related species (e.g.
genera) when identification to species was not recorded in the database
(FAO, 2019b, 2020; Metian et al., 2020). According to the International
Standard Statistical Classification of Aquatic Animals and Plants (ISS­
CAAP), the ASFIS species items fall into eight divisions: (i) marine fishes,
(ii) freshwater fishes, (iii) diadromous fishes, (iv) crustaceans, (v) mol­
luscs, (vi) aquatic plants, (vii) miscellaneous aquatic animals, and (viii)
miscellaneous aquatic animal products. We group the first three di­
visions into “finfish” and the last two into “miscellaneous aquatic ani­
mals and animal products” (abbreviated as “MAA”), and we rename
“aquatic plants” into “algae” to more accurately reflect its species
composition. As a result, we have five species groups (finfish, crusta­
ceans, molluscs, MAA, and algae) for the examination of diversity within
each group and between them. For narrative convenience, we will refer
to the ASFIS species items simply as “species” throughout the rest of this
article.
2.2. Measuring species diversity
As a unifying notation of commonly used measures of diversity (Hill,
1973), a general formula for the effective number of species is
( )1−1 q
∑n
Dq = sq (1)
i=1 i
where n represents the total number of species; si denotes the share of
species i in the production of all species; and q indicates the order of
diversity. When q = 0, D0 = n is equal to the total number of species,
which measures solely richness and gives zero weight to evenness. As q
increases, more weight is given to evenness in the measure of diversity.
While efforts in biodiversity conservation may focus on species richness,
evenness is a crucial dimension of species diversity for sustainable
aquaculture development (Harvey et al., 2017).
Like Cai et al. (2022), we measure species diversity in aquaculture
using the effective number of species at q = 1, i.e.
∑n
ENS ≡ D1 = e− i=1 si ln(si ) , (2)
where the summation term is the Shannon index:
∑n
H≡ − si ln(si ) = ln(ENS). (3)
i=1
 J. Cai et al.
Table 1
Status and trends of species diversity in world and regional aquaculture, 1990–2020.
I II III IV V VI VII VIII IX X XI XII

World and 32 regional aquaculture1 Aquaculture production in Species diversity measures: ENS (WGENS; ENSG) Scenarios or patterns of species diversification2
2020

000t Share of world (%) Year 1990 Year 2000 Year 2010 Year 2020 Overall: 1990–2020 1990–2000 2000–2010 2010–2020

World 122580 100.00 35.61 (11.06; 3.22) 34.88 (10.71; 3.26) 50.59 (14.6; 3.47) 47.46 (13.56; 3.50) Inverted N U1 D2 U1 D2
Row 52097 42.50 38.46 (11.26; 3.42) 45.87 (14.72; 3.12) 39.84 (13.07; 3.05) 33.18 (11.4; 2.91) Inverted U D3 U2 D1 D1
Developed regions 5147 4.20 19.68 (6.60; 2.98) 20.15 (7.29; 2.76) 18.47 (6.98; 2.65) 15.68 (6.33; 2.48) Inverted U D1 U2 D1 D1
Developing regions 117433 95.80 28.11 (8.77; 3.21) 29.03 (8.85; 3.28) 45.66 (13.08; 3.49) 44.30 (12.58; 3.52) Inverted U U1 U1 U1 D2
SIDS 57 0.05 7.68 (2.56; 3.00) 10.71 (3.23; 3.31) 12.73 (3.97; 3.20) 13.75 (4.66; 2.95) Straight up U2 U1 U2 U2
LLDCs 590 0.48 4.89 (4.88; 1.00) 9.45 (9.43; 1.00) 11.19 (11.13; 1.01) 14.07 (14.02; 1.00) Straight up U1 U1 U1 U2
Asia 112301 91.61 29.70 (9.23; 3.22) 28.70 (8.71; 3.30) 44.06 (12.56; 3.51) 42.28 (11.96; 3.53) Inverted N U1 D2 U1 D2
Eastern Asia 74770 61.00 20.45 (6.41; 3.19) 20.48 (6.28; 3.26) 31.58 (8.67; 3.64) 30.35 (8.08; 3.76) Inverted U U1 U3 U1 D2
South-eastern Asia 24830 20.26 15.84 (4.81; 3.3) 17.51 (5.23; 3.35) 17.54 (5.74; 3.06) 14.39 (5.06; 2.85) Inverted U D3 U1 U2 D1
Western Asia 598 0.49 4.57 (4.52; 1.01) 6.26 (5.63; 1.11) 7.52 (5.60; 1.34) 7.52 (5.48; 1.37) Inverted U U1 U1 U3 D2
Southern Asia 11995 9.79 6.44 (5.39; 1.20) 8.20 (6.34; 1.29) 7.79 (6.36; 1.23) 10.49 (7.56; 1.39) N-shape U1 U1 D3 U1
Central Asia 108 0.09 2.07 (2.07; 1.00) 3.05 (3.05; 1.00) 3.47 (3.47; 1.00) 5.71 (5.71; 1.00) Straight up U2 U2 U2 U2
Americas 4401 3.59 13.06 (4.17; 3.13) 16.04 (6.84; 2.34) 15.93 (5.78; 2.76) 11.46 (4.18; 2.74) Inverted U D3 U2 D2 D1
LAC 3781 3.08 9.05 (2.93; 3.09) 13.70 (5.65; 2.43) 11.13 (4.02; 2.77) 8.70 (3.24; 2.68) Inverted U D3 U2 D2 D1
South America 3307 2.70 6.20 (2.01; 3.08) 12.40 (5.28; 2.35) 11.23 (4.13; 2.72) 8.47 (3.16; 2.68) Inverted U U2 U2 D2 D1
Central America 442 0.36 6.59 (2.67; 2.47) 4.25 (2.02; 2.10) 2.75 (1.41; 1.96) 3.29 (1.59; 2.07) U-shape D1 D1 D1 U1
3

Caribbean 32 0.03 5.95 (3.91; 1.52) 5.06 (3.25; 1.56) 5.82 (3.49; 1.67) 6.29 (3.40; 1.85) U-shape U3 D2 U1 U3
Northern America 620 0.51 6.63 (2.87; 2.31) 6.66 (3.52; 1.89) 7.71 (3.20; 2.41) 8.06 (3.09; 2.61) Straight up U1 U2 U3 U3
Europe 3284 2.68 9.78 (4.76; 2.06) 10.22 (5.12; 2.00) 8.19 (4.64; 1.76) 7.29 (4.30; 1.69) Inverted U D1 U2 D1 D1
Southern Europe 610 0.50 5.49 (2.89; 1.90) 6.79 (3.46; 1.96) 8.05 (3.92; 2.05) 8.05 (3.90; 2.06) Straight up U1 U1 U1 U3
Eastern Europe 448 0.37 2.37 (2.22; 1.07) 4.52 (4.1; 1.10) 5.39 (5.07; 1.06) 10.44 (6.45; 1.62) Straight up U1 U1 U2 U1
Northern Europe 1956 1.60 2.72 (2.09; 1.30) 2.11 (1.7; 1.24) 1.97 (1.61; 1.22) 1.67 (1.48; 1.13) Straight down D1 D1 D1 D1
Western Europe 270 0.22 4.57 (2.66; 1.72) 5.32 (2.95; 1.80) 6.35 (3.37; 1.89) 5.93 (3.20; 1.85) Inverted U U1 U1 U1 D1
Africa 2354 1.92 7.92 (4.95; 1.60) 7.65 (4.93; 1.55) 7.88 (5.50; 1.43) 7.00 (5.61; 1.25) Inverted N D3 D1 U2 D3
Sub-Saharan Africa 732 0.60 9.77 (3.81; 2.56) 8.17 (3.3; 2.47) 7.63 (3.88; 1.96) 8.60 (5.25; 1.64) U-shape D3 D1 D3 U2
Eastern Africa 353 0.29 3.61 (1.75; 2.06) 2.92 (1.44; 2.03) 4.15 (1.92; 2.16) 5.35 (2.73; 1.96) U-shape U2 D1 U1 U2
Middle Africa 11 0.01 2.19 (2.19; 1.00) 1.92 (1.92; 1.00) 2.47 (2.37; 1.04) 3.12 (2.99; 1.04) U-shape U1 D2 U1 U1
Western Africa 346 0.28 5.45 (5.21; 1.05) 8.76 (8.58; 1.02) 6.14 (6.11; 1.01) 6.71 (6.61; 1.01) N-shape U2 U2 D1 U1
Southern Africa 13 0.01 4.67 (2.27; 2.06) 5.22 (2.23; 2.34) 7.56 (2.55; 2.97) 5.32 (1.78; 2.98) Inverted U U3 U3 U1 D2
Northern Africa 1632 1.33 3.80 (3.65; 1.04) 4.13 (4.09; 1.01) 4.04 (4.00; 1.01) 4.00 (3.94; 1.02) Inverted U U2 U2 D2 D2
Oceania 240 0.20 5.87 (2.23; 2.64) 5.79 (2.16; 2.68) 7.56 (2.65; 2.85) 6.26 (2.22; 2.82) Inverted N U3 D2 U1 D1
AU & NZ 225 0.18 4.43 (2.65; 1.67) 4.06 (2.16; 1.88) 5.66 (2.52; 2.25) 5.06 (2.13; 2.37) Inverted N U3 D2 U1 D2
Pacific islands 15 0.01 1.51 (1.04; 1.45) 2.59 (1.12; 2.31) 5.09 (1.96; 2.59) 3.54 (1.30; 2.72) Inverted U U1 U1 U1 D2

Notes: 1. Regional aquaculture (32) = 5 geographic regions (bold font) + 20 non-overlapping geographic sub-regions +7 custom country groups (italic font). The classification of geographic regions and sub-regions follows
the United Nations (UN) M49 standard. Rest of the world (ROW) = World aquaculture, excluding China. According to the UN M49 standard (the 2018 definition), Developed Regions include Europe, Northern America,

Aquaculture 576 (2023) 739837

Japan, Australia, New Zealand, Cyprus, and Israel, while other countries belong to Developing Regions. Small Island Developing States (SIDS) and Landlocked Developing Countries (LLDCs) follow the classifications adopted by
the UN Office of the High Representative for the Least Developed Countries, Landlocked Development countries and Small Island Developing States (UN-OHRLLS). Two customary groups are ad hoc geographic groups:
Latin America and the Caribbean (LAC) = Central America + South America + the Caribbean; sub-Saharan Africa = Eastern Africa + Middle Africa + Western Africa + Southern Africa + Sudan in Northern Africa. AU & NZ
= Australia and New Zealand. 2. Six scenarios (U1, U2, U3, D1, D2, and D3) and six patterns (e.g. inverted U-shape) are categorized in Table 2.
J. Cai et al.
The Shannon index is a well-known diversity index, and it was used
by Metian et al. (2020) and Sicuro (2021) to examine species diversity in
global aquaculture. ENS is less widely used, but it is considered a more
intuitive measure of true diversity (Hill, 1973; Jost, 2006). The scale of
ENS ranges from 1 to the total number of species (n). When production is
evenly distributed among all species, the ENS value is equal to n. The
more uneven the distribution of production is, the closer the ENS value
will be to 1.
ENS in eq. (2) can be partitioned into two components:
ENS = WGENS × ENSG. (4)
The first component (WGENS), which is a measure of within-group
diversity, is calculated as a weighted geometric mean of ENS within
different species groups (denoted as ENSj). As discussed in Section 2.1,
here we consider five species groups (finfish, crustaceans, molluscs,
MAA, and algae). The weight assigned to each group is based on its share
of total production (denoted as sj), i.e.
∏( )s
WGENS = ENSj j . (5)
j
The second component (ENSG) represents the effective number of
species groups, i.e.
∑ molluscs (14.5%) contributed 90% of the production. The rest primarily
ENSG = e j j ( j ) ,
− s ln s
(6)
which measures the evenness of production distribution among the five
species groups (i.e. between-group diversity). WGENS, ENSG, and ENS
are equivalent to three diversity measures (alpha, beta, and gamma di­
versity, respectively) that have been used to assess biological diversity in
an ecosystem (Jost, 2007; Tuomisto, 2010). While ENS has been used in
Cai et al. (2022) to facilitate benchmarking species diversification in
aquaculture, this is the first study, to the best of our knowledge, to apply
within-group diversity (WGENS) and between-group diversity (ENSG)
as sub-indicators of species diversity in aquaculture.
According to eqs. (3)–(6), the Shannon index can also be partitioned
into
H = Hwithin + Hbetween , (7)
∑ ∑
where Hwithin = jsjHj and Hbetween = j − sj ln (sj) are alternative
measures of within-group and between-group diversity.
While ENS and its two components (WGENS and ENSG) are more
intuitive diversity measures – see numerical examples at the beginning
of Section 3.1, the multiplicative partition of ENS in eq. (4) makes it
inconvenient to calculate the shares of WGENS and ENSG in ENS.
Therefore, we used the partition of the Shannon index defined in eq. (7)
to calculate the shares of within-group diversity and between-group
diversity in overall species diversity: Hwithin/H measures the within-
group share; Hbetween/H measures the between-group share; and the
two shares sum up to 100%. The within-group share can be further
decomposed into the share of each species group in overall diversity (i.e.
sjHj/H).
2.3. Categorizing changes in species diversity
Table 2 presents six scenarios of ENS changes and six patterns of ENS
trends. The six scenarios links a change in ENS to changes in its within-
group and between-group components, while the six patterns categorize
the trends of species diversification during three decades (1990–2020)
or two decades (2000− 2020).
While originally used to measure biological diversity, the concept of
“effective number” has been applied to measure diversity in other areas,
such as the effective number of parties in a political system (Laakso and
Taagepera, 1979) and the effective number of firms in an economy
(Ordover et al., 1982). Similarly, we use the “effective number of
diversification patterns” (ENDP) to measure the diversity of the six
4
Aquaculture 576 (2023) 739837
patterns among countries (i.e. the evenness of the distribution of these
six patterns among all countries worldwide or in a country group). The
formula is
∑6
ENDP = e− i=1 pi ln(pi ) , (8)
∑
where pi = ni/ ni measures the prevalence of diversification pattern i
among countries, as ni is the number of countries with pattern i. Simi­
larly, eq. (8) can be used to calculate the ENDP among regional
aquaculture.
3. Results
3.1. Species diversification in world aquaculture
In 2020, a total of 448 species contributed the world aquaculture
production of 123 million tonnes. Yet the production was concentrated
on a much lower number of species – 46 species contributed 90% of the
production. The diversity of the species composition can be measured by
the effective number of species (ENS) of 47.5 (Fig. 1a).
While the 123 million tonnes of world aquaculture production
spread across five species groups, finfish (46.9%), algae (28.6%), and
came from crustaceans (9.2%), with a small amount of miscellaneous
aquatic animals and animal products (MAA; 0.9%). As a measure of
between-group diversity, the 3.5 effective number of species groups
(ENSG) reflects this imbalanced taxonomic composition (Fig. 1b).
In 2020, finfish accounted for nearly half of world aquaculture
production, its species diversity (30.5 ENS) was also much higher than
the other four species groups (Fig. 1c–g). The production of algae was
nearly twice of molluscs, yet its species diversity (6.1 ENS) was lower
than that of molluscs (9.4 ENS). The species diversity in crustaceans (4.9
ENS) was the lowest, whereas the 6.3 ENS of MAA, the smallest species
group notwithstanding, was higher than algae and crustaceans. As a
measure of within-group diversity, the 13.56 within-group ENS
(WGENS) measures the average species diversity within these five spe­
cies groups (Fig. 1b).
In summary, world aquaculture production in 2020 (123 million
tonnes) comprised 448 species, with a species diversity of 47.5 ENS.
Between-group diversity (3.5 ENSG) accounted for 32% of the overall
species diversity, whereas the rest 68% came from within-group di­
versity (13.56 WGENS), including 41.5% from (species diversity within)
finfish, 13.4% from algae, 8.4% from molluscs, 3.8% from crustaceans,
and 0.4% from MAA.
Table 2
Categorizing changes in species diversity.
Three scenarios of ENS increases (¼ Up)
• U1 = ENS increase (both components up)
• U2 = ENS increase (only WGENS up)
• U3 = ENS increase (only ENSG up)
Three scenarios of ENS decreases (¼ Down)
• D1 = ENS decrease (both components down)
• D2 = ENS decrease (only WGENS down)
• D3 = ENS decrease (only ENSG down)
Six patterns of ENS trends during three decades (1990–2020) or two decades
(2000–2020)
• Straight-down = ENS decreased in all three decades (or the second and third decades
in case of no aquaculture production in 1990)
• Inverted U-shape = ENS increased in the first decade (or the second decade if no
production in 1990) yet declined in the last decade
• Inverted N-shape (И) = ENS decreased during first and third decades whereas
increased during the second decade
• Straight-up = ENS increased in all three decades (or the second and third decades in
case of no aquaculture production in 1990)
• U-shape = ENS declined in the first decade (or the second decade if no production in
1990) yet increased in the last decade
• N-shape = ENS increased during first and third decades whereas declined during the
second decade.
J. Cai et al.
Historically, species diversity in world aquaculture more than
doubled in four decades between 1950 and 1990 through a “straight-up”
diversification trend, with increased ENS in all eight mid-term (5-year)
intervals during the period (Fig. 1a). Species diversity increased further
during the recent three decades (1990–2020), yet the diversification
trend was less monotonic. Two ENS declines have resulted in an
“inverted N-shape (И)” ENS trend that declined in the first decade
(1990–2000) and the last decade (2010− 2020), whereas increased in
the middle decade (2000− 2010). The first decline occurred in the first
half of the 1990s, with a partial recovery in the second half. A similar
pattern occurred in the 2010s (Fig. 1a).
The future will tell whether the second decline (during 2010–2020)
was a temporary dip similar to the first one (during 1990–2000); or it
may be the beginning of a longer term downward ENS trend in world
aquaculture similar to the pattern for the rest of the world, excluding
China (i.e. the ROW aquaculture) since 2000 (Supplementary Fig. 1a).
However, the downward ENS trend in world aquaculture in the third
decade (2010–2020) differed from the downward trend in the first
decade (1990–2000) in that the former was consistent with regional and
national situations, whereas the latter was not.
3.2. Inconsistent species diversification at different geographic scales
In the first decade (1990–2000), species diversity declined in world
aquaculture yet increased in most (22 out of 32) regional aquaculture
(Table 1; column X) as well as nearly two thirds of national aquaculture
that accounted for nearly 90% of world production (Fig. 2b). The
inconsistency could seemingly be explained by the fact that species
diversification in world aquaculture mimicked that of Asia, the largest
aquaculture region that contributed more than 90% of world production
(Table 1, column III). A deeper look, however, reveals another
Fig. 1. Species diversification in world aquaculture, 1950–2020.
Notes: There was a major disaggregation in the reporting of China's data in the early 2000s, with the number of species increased from 30 in 2002 to 84 in 2003. This
improvement of data quality nevertheless makes the ENS in world aquaculture after 2003 (inclusive) not directly comparable with those in past years, as China
accounted nearly 70% of world production at the time. We aggregated China's aquaculture production in 2005 based on the species composition in 2000. Based on
the adjusted data, the comparable ENS in world aquaculture 2005 was 41.4, which is still higher than the 34.9 ENS in 2000.
5
Aquaculture 576 (2023) 739837
inconsistency: declined species diversity in Asia versus increased di­
versity in all five Asian sub-regions (Table 1; column X). A similar
inconsistency occurred between 1990 and 2020 when species diversity
declined in Africa (as well as sub-Saharan Africa) while increased in all
five African sub-regions (Table 1; column IX).
Inconsistencies could occur not only on diversification trends but
also on the status of species diversity. As China has been the largest
national aquaculture with the most diverse species composition, one
may expect that species diversity in world aquaculture should always be
higher than diversity in the ROW aquaculture. The opposite occurred,
nevertheless, in 1990 and 2000 (Table 1; columns IV and V). Such in­
consistencies indicate that species diversification in world or regional
aquaculture may not necessarily reflect situations in national aquacul­
ture. After all, if every country cultivated a unique species at the same
amount, world aquaculture would have a high species diversity even
with no species diversity at the national level.
3.3. Decelerating species diversification
During the three decades between 1990 and 2020, species diversity
increased in most (22 out of 32) regional aquaculture. This, however,
reflects the situation in the first and second decades (Table 1; Supple­
mentary Table 1a), whereas species diversity declined in most (19)
regional aquaculture in the third decade (Table 1; Supplementary
Table 1b). Such decelerating diversification process also occurred in
national aquaculture.
Between 1990 and 2020, species diversity increased in nearly three
quarters of national aquaculture that accounted for more than 80% of
world production (Fig. 2a). The situations in the first decade (Fig. 2b)
and the second decade (Fig. 2c) were consistent with the overall trend.
Yet species diversity declined in a higher proportion of national
J. Cai et al.
aquaculture in the second decade compared with the first one (37%
versus 34.6%), and the production share of countries with a declined
species diversity in the second decade (23.7%; Fig. 2c) nearly doubled
the level in the first decade (11.9%; Fig. 2b). Indeed, the 23.7% would be
higher than the remaining 11.1% when China's production share (65%)
is excluded from the 76.1% production share of national aquaculture
with an increased diversity in the second decade (Fig. 2c).
The diversification trend appeared to reach a turning point in the
third decade when the prevalence of national aquaculture with an
increased diversity declined to exactly 50%, and they contributed <20%
of world production (Fig. 2d). The large decline in the production share
of more diversified aquaculture was primarily due to the shift of China's
aquaculture from diversification trends in the first and second decades
to a concentration trend in the third decade. However, even when
China’ production share (58%) is excluded from the 81% production
share of national aquaculture with a declined diversity in the third
decade, the remaining 23% would still be higher than the 18.9% pro­
duction share of those with an increased diversity (Fig. 2d).
3.4. Diverse species diversification patterns
Among 196 national aquaculture in 2020, we identify six patterns of
ENS trends (Table 2) for 161 countries (Fig. 2e). Most of the 161 ENS
trends were 30-year trends between 1990 and 2020, with a few 20-year
trends between 2000 and 2020 in case of no aquaculture production in
1990 (Fig. 2e). Thirty-five relatively young national aquaculture, which
accounted for <0.1% of world production in 2020, had no such 30- or
20-year trends, resulting in their exclusion from the six diversification
patterns.
As species diversity increased in 89 countries (nearly two thirds of
national aquaculture) in the first decade (Fig. 2b) while declined in 91
countries (exactly half) in the third decade (Fig. 2d), an inverted U-
shape trend emerged as the prevailing pattern in national aquaculture
during 1990–2020 (Fig. 2e). This most common pattern nevertheless
occurred in only 59 countries (around one third). These 59 countries
spread across Africa (17), Asia (17), Europe (12), the Americas (10), and
Oceania (3), and they contributed nearly 80% of world production in
2020, including six of the top 10 largest national aquaculture (Fig. 2e;
Supplementary Table 2a).
The inverted N-shape pattern is similar to the inverted U-shape
pattern in that both are associated with a decline in diversity in the third
decade. While characterizing the ENS trend in world aquaculture, the
inverted N-shape pattern occurred in only 18 countries that contributed
to just half a percent of world production (Fig. 2e; Supplementary
Table 2b). This further exemplifies the inconsistency discussed in Sec­
tion 3.2. Straight-down, which is another pattern with a declined di­
versity in the third decade, was the least common pattern that occurred
in only eight national aquaculture (0.2% of world production) (Fig. 2e;
Supplementary Table 2c). In contrast, straight-up ENS trends occurred in
31 countries that accounted for 6.7% of world production; most of which
are in Europe (14) or Asia (10) (Fig. 2e; Supplementary Table 2d). U-
shape and N-shape are another two patterns with increased diversity in
the third decade, which occurred in 28 countries (4.8% of world pro­
duction; Supplementary Table 2e) and 17 countries (8.5%; Supple­
mentary Table 2f), respectively.
Based on eq. (8), we calculate the effective number of diversification
patterns (ENDP) to measure the diversity of diversification patterns, i.e.
the evenness of the distribution of the six patterns among countries. The
results indicate diverse diversification patterns among national aqua­
culture: 5.06 ENDP for the 161 national aquaculture worldwide, ~4–5
ENDP for national aquaculture in each of the five regions (from 4.34
ENDP for national aquaculture in Oceania to 5.03 ENDP for national
aquaculture in the Americas), 5.21 ENDP for landlocked national
aquaculture, and 4.51 ENDP for island national aquaculture (Supple­
mentary Fig. 2).
However, there were clusters of national aquaculture with the same
6
Aquaculture 576 (2023) 739837
or similar patterns (Fig. 2e), such as (i) a cluster of (national aquaculture
with) straight-up ENS trends in East-Central Europe; (ii) two clusters of
U-shape or N-shape trends: one in South-Central Asia; the other
stretching from the United States of America southward through Central
America (except Nicaragua) to three of the four members of the Andean
Community (Bolivia, Colombia and Peru, except Ecuador) as well as two
associated members (Paraguay and Uruguay); and (iii) several clusters
of inverted U-shape or inverted N-shape trends in (a) the Mediterranean
Basin, (b) Western and Southern Africa, (c) the south and east parts of
South America, and (d) three countries in Maritime Southeast Asia
(Indonesia, Malaysia and Brunei Darussalam) plus Australia and New
Zealand plus four nearby Pacific SIDS (Papua New Guinea, New Cale­
donia, Guam, and Palau).
The distribution of the six diversification patterns among the 32
regional aquaculture was similar to the situation in the 161 national
aquaculture, i.e. inverted U-shape being the most common pattern
(occurred in 14 regional aquaculture), followed by straight-up (6), U-
shape (5), inverted N-shape (4), N-shape (2), and straight-down in only
one regional aquaculture (Supplementary Table 3). The diversity of the
six patterns among the 32 regional aquaculture was 4.5 ENDP.
3.5. Within-group versus between-group diversity
Within-group diversity contributed approximately 60 to 70% of
species diversity in world aquaculture during 1950–2020. This share
was generally on an upward trend, increasing from 63.1% in 1960 to
67.5% in 2020 (Supplementary Fig. 3). For the 32 regional aquaculture,
the average within-group share in species diversity during 1990–2020
varied from 23% in Pacific Islands to 100% in Central Asia, yet the share
was above 50% for most regional aquaculture with a few exceptions
(Supplementary Table 4).
The pattern of within-group share in national aquaculture has
remained relatively unchanged over 1990, 2000, 2010 and 2020 (Sup­
plementary Fig. 4). In 2020, the within-group share in national aqua­
culture varied from 0 to 100%, yet it was more than 50% in 126 of all
196 national aquaculture (Supplementary Fig. 4d). These 126 countries
(illustrated with different shades of blue in Fig. 3d) accounted for more
than 90% of world production.
As highlighted in Section 3.1, finfish diversity was the primary
source that contributed more than 40% of species diversity in world
aquaculture in 2020. For national aquaculture, finfish was the primary
source of species diversity in 114 countries. Among all finfish farming
countries, the median finfish share in species diversity was nearly 80%
(Supplementary Fig. 5b).
Apparently from Fig. 1a and b, species diversification in world
aquaculture mimicked the trend of within-group diversity. Similarly,
within-group diversity has been the main driving force of species
diversification in regional aquaculture (Table 1; Supplementary
Tables 1a and 1b) and national aquaculture (Fig. 2; Supplementary
Fig. 6). During 1990–2020, ENS increased in 109 national aquaculture;
93 of which had a higher WGENS (Fig. 2a: U1 and U2), whereas 33 of 40
countries with a declined ENS had a lower WGENS (Fig. 2a: D1 and D2).
Similarly, within-group diversity was the primary driving force of spe­
cies diversification in all three sub-decades (Fig. 2b–d).
In 2020, within-group diversity was the sole source of species di­
versity (ENSG =1; the within-group share = 100%) in 67 national
aquaculture; nearly all (65) of them farmed only finfish (Supplementary
Table 5). Representing more than one third of all 196 national aqua­
culture notwithstanding, these 67 countries contributed only half a
percent of world production. They were mostly clustered in Africa,
Central Asia, and Eastern and Central Europe (Fig. 3d, highlighted in
dark blue). Nearly half (29) of them are landlocked countries, including
24 Landlocked Developing Countries (LLDCs). Among 37 landlocked
countries (Supplementary Table 6), the within-group share in 2020 was
equal to 100% in 29 countries, above 90% in 34 countries, and above
60% in nearly all of them (except one country with no species diversity,
J. Cai et al. Aquaculture 576 (2023) 739837

Fig. 2. Categorizing species diversification in national aquaculture.

Notes: Table 2 categorizes the six scenarios of ENS changes (U1, U2, U3, D1, D2, and D3) illustrated in Fig. 2a–d. Countries with no changes in ENS are not reflected in
these maps but illustrated in Supplementary Fig. 6. Table 2 also categorizes the six diversification patterns illustrated in Fig. 2e. The first three patterns (illustrated
with different shades of pink) are associated with a decline in ENS in the third decade, whereas the last three patterns (illustrated with different shades of green) are
associated with an increase in ENS in the third decade. In a bracket, the first number represents the number of national aquaculture in the category; the second
number (in percentage) represents the prevalence of the category, calculated as the first number divided by the total number of countries in all categories. Lastly, the
third number (in percentage) represents the category's share in world production (during the respective period for Fig. 2a–d and in 2020 for Fig. 2e). The production
share of all categories may not sum up exactly to 100% because the categorization does not cover all national aquaculture. These absent national aquaculture, along
with countries with no aquaculture, are illustrated in grey. The maps were created using Datawrapper; the country/territory boundaries depicted therein do not
represent authors' opinions. Countries/territories with a small land area may not be discernible on the maps. (For interpretation of the references to colour in this
figure legend, the reader is referred to the web version of this article.)

7
J. Cai et al.
i.e. ENS = 1).
As highlighted in Section 3.1, between-group diversity accounted for
around one third of species diversity in world aquaculture in 2020,
which was lower than the contribution of finfish yet higher than each of
the other four species groups. In 2020, between-group diversity was the
primary source (compared with each of the five species groups) of
species diversity in 61 countries (~30% of all 196 national aquaculture).
Among them, between-group diversity was greater than within-group
diversity (ENSG > WGENS; the within-group share <50%) in 55 coun­
tries that contributed 7.8% of world production (Supplementary
Table 7). Most of these 55 countries, highlighted in warm colors in
Fig. 3d, were located in the Americas (24 countries; 57% of national
aquaculture in the region) and Oceania (10 countries; 53%). The
remaining 21 countries were spread across the other three regions, with
eight in Asia, seven in Europe, and six in Africa. More than half (28) of
the 55 countries were island economies, including 22 Small Island
Developing States (SIDS). These 28 island economies represented nearly
half of 60 island aquaculture in 2020 (Supplementary Table 8).
During 1990–2020 and the three sub-decades, between-group di­
versity remained unchanged in ~30–40% of national aquaculture where
only one species group was farmed, whereas it increased in most of the
other countries (Supplementary Fig. 7). During 1990–2020, between-
group and within-group diversity moved in different directions in
nearly 100 national aquaculture that accounted for nearly 20% of world
production (Fig. 2a: U2, U3, D2, and D3). Among them, ENSG dictated
the direction of ENS trends in only 23 countries (7.7% of world pro­
duction; Fig. 2a: U3 and D3), including (i) 16 countries where increased
between-group diversity drove up the overall species diversity in spite of
declined or unchanged within-group diversity (6%; Fig. 2a: U3; Sup­
plementary Table 9a) and (ii) seven countries where overall species
diversity declined because of between-group diversity despite increased
or unchanged within-group diversity (1.7%; Fig. 2a: D3; Supplementary
Table 9b). The pattern of between-group diversity dictating the direc­
tion of species diversification in a small portion of national aquaculture
occurred in all three sub-decades (Fig. 2b–d).
3.6. Generally low (often no) diversity in national aquaculture
In 2020, ENS varied from 1 to 28 among 196 national aquaculture,
yet half of them were <2.5 and three fourths <4 (Fig. 3a; Supplementary
Fig. 8). The median WGENS was below 2 (Fig. 3b), whereas the median
ENSG was slightly above 1 (Fig. 3c). Nearly half (94) of the 196 coun­
tries had no within-group or between-group diversity, including 82
countries (42% of all 196 national aquaculture) with no between-group
diversity (ENSG = 1), 27 countries (14%) with no within-group diversity
(WGENS = 1), and among them, 15 countries (8%) with neither (i.e. no
species diversity; WGENS = ENSG = ENS = 1). No species diversity was
a more common phenomenon within the five species groups (Supple­
mentary Table 10). In 2020, finfish was the only species group with
median ENS (among all countries that farmed finfish) above 2; molluscs
was another species group (besides finfish) with median ENS >1; and
median ENS was equal to 1 for all the other three species groups (Sup­
plementary Fig. 9).
In 2020, countries with relatively low (below-median) ENS
contributed <5% of world production, whereas a quarter of countries
with the highest ENS contributed more than 90% (Fig. 3a). Thirty-three
countries had extraordinarily high diversity in terms of at least one of
the eight diversity measures (ENS, WGENS, ENSG and ENS in each of the
five species groups). Most (21) of these countries belonged to the top 50
national aquaculture (Supplementary Table 11), and China (the largest
national aquaculture with extraordinarily high diversity in terms of all
eight measures) contributed nearly 60% of world production.
There were large national aquaculture with low species diversity,
such as export-oriented aquaculture in Norway (1.5 million tonnes of
production with 1.31 ENS) and Ecuador (0.8 million tonnes with 1.11
ENS). A small national aquaculture could nevertheless have a high
8
Aquaculture 576 (2023) 739837
species diversity. For instance, Singapore, a high-income city state with
strong consumer preferences for a variety of aquatic foods, had a small
aquaculture sector (<5000 t of production in 2020) with the third
highest species diversity among all national aquaculture (Supplemen­
tary Table 11).
3.7. Correlations between production and diversity
It has been recognized that larger aquaculture production tend to be
associated with higher species diversity (Metian et al., 2020). Cai et al.
(2022) found a positive correlation (r = ~0.6) between aquaculture
production and species diversity based on data from over 200 countries
for three decades (over 5000 observations). They used this relationship
to set benchmarks for species diversification.
The positive correlation between aquaculture production and species
diversity raises the question of whether species composition would tend
to become more diverse as aquaculture production grows bigger (i.e.
species diversification as a natural course of aquaculture expansion). A
closer examination that distinguishes between cross-sectional correla­
tion (i.e., correlation between production and diversity for a number of
countries in a specific time) and temporal correlation (i.e., correlation
between production and diversity for a specific country over time) will
shed light on this question.
In 2020, a strong, positive correlation between aquaculture pro­
duction and ENS was found among all 196 countries (r = 0.65). This
positive cross-sectional correlation was consistent across the years
1990–2020, with a median value of r = 0.64 (Supplementary Table 12).
In addition to the positive correlation between aquaculture production
and overall species diversity, cross-sectional correlations between pro­
duction and the two components of ENS (WGENS and ENSG) were also
found to be positive, albeit weaker (median r = 0.24 and 0.34, respec­
tively). So were the correlations between production and ENS in each of
the five species groups, with median r varied from 0.38 for finfish to 0.84
for MAA (Supplementary Table 12). These positive cross-sectional cor­
relations indicate that countries with a relatively large aquaculture
production generally have a higher species diversity than those with a
relatively small production.
Temporal correlation between a country's production and diversity
over time is less straightforward. During 1990–2020, temporal corre­
lation between production and ENS varied between − 1 and 0.99 in all
countries, and it was positive in 55% of the countries (108 out of 196)
yet negative in 45% (88 out of 196), with the median r = 0.1. In 73
countries (37%), the temporal correlation was significantly positive (p-
value <0.05), which indicates that these countries' aquaculture pro­
duction and ENS generally moved in the same direction during the
period. The temporal correlation was significantly negative in 49
countries (25%) where aquaculture production and ENS generally
moved in different directions. In the remaining 74 countries (38%), the
temporal correlation was not statistically significant (Supplementary
Table 13). While these findings may be deemed evidence of generally
positive, albeit weak, temporal correlation between production and ENS
in national aquaculture during 1990–2020, such a generally positive
relationship was less obvious in the first two decades (median r around
-0.03 in both periods) and became non-existent in the last decade
(Supplementary Table 13). During the third decade (2010–2020), the
temporal correlation between production and ENS was negative in more
than half (101) of all 189 national aquaculture with median r = − 0.08.
The correlation was significantly positive in 46 countries (<5% of world
production) while significantly negative in 49 countries (over 20%)
(Supplementary Fig. 10; Supplementary Table 14). Similarly, there was
a lack of generally positive temporal correlations between production
and the other seven diversity measures (Supplementary Table 13). These
findings indicate that while countries with a larger aquaculture pro­
duction tend to have a higher species diversity than those with a lower
production, there is no obvious tendency that a country's species di­
versity would move in the same direction with its production. In other
J. Cai et al. Aquaculture 576 (2023) 739837

Fig. 3. Comparisons of species diversity among countries, 2020.

Notes: In parentheses, the first number is the number of countries in the category; the second number (in percentage) represents the category's share in world
production. In map (a), (b), and (c), the boundaries of the ranges are determined by quartile distributions presented in Supplementary Fig. 8. The lower bound of a
range is exclusive, whereas the upper bound is inclusive. In map (d), both the lower and upper bounds of a range are exclusive. In map (a) or (b), “Min” + “Q1”
include a quarter of all 196 national aquaculture with the lowest ENS or WGENS; “Q2” includes the quarter between the first quartile and the second quartile
(median); Q3 includes the quarter between the median and the third quartile; and “Q4” + “XL” include the quarter with the highest ENS or WGENS. “XL” includes
countries with extraordinarily large diversity – see Supplementary Table 11. In map (c), “Min/Q1” + “Q2” indicate that among the 98 countries with relatively low
(below-median) ENSG in 2020, ENSG = 1 in 82 countries, and the ENSG of the rest 16 countries in “Q2” was above 1 yet no >1.02. The definitions of “Q3”, “Q4” and
“XL” in map (c) are similar to those in maps (a) and (b). All maps were created using Datawrapper; the country/territory boundaries depicted therein do not represent
authors' opinions. Countries/territories with a small land area may not be discernible on the maps.

9
J. Cai et al.
words, species diversification has no tendency to be a natural course of
aquaculture expansion.
3.8. Geo-clustering of countries with similar species diversity
National aquaculture within close geographic proximity tend to have
similar species diversity profiles. We have highlighted clusters of
countries with similar diversification patterns in Section 3.4 and those
with similar diversity profiles in Section 3.5. Here we systematically
examine geo-clustering patterns by grouping countries into four cate­
gories according to their within-group and between-group diversity
(Fig. 3e).
In 2020, 48 countries (illustrated in blue in Fig. 3e) had a relatively
high (above-median) within-group diversity as well as a relatively high
between-group diversity; all of them also had relatively high overall
diversity (Supplementary Table 15a). This “both > median” category
primarily includes 19 countries in Asia (primarily Eastern and South-
eastern Asia) and 11 countries in Europe (primarily Western and
Southern Europe). The rest spread in Africa (8 countries), the Americas
(7), and Oceania (3). Representing less than a quarter of all 196 national
aquaculture notwithstanding, “both > median” appears to be the most
prominent pattern in the map (Fig. 3e) because it includes nine of the top
10 countries with the largest land area, except Kazakhstan (Supple­
mentary Table 16). This may reflect that abundant, diverse natural re­
sources, such as indigenous species and suitable farm sites, tend to be
conducive to high species diversity. Indeed, while Kazakhstan, like most
landlocked countries, farmed only finfish hence had no between-group
diversity, its overall species diversity was higher than three quarters
of ~200 national aquaculture because of its extraordinarily high species
diversity in finfish aquaculture (Supplementary Table 11). The “both >
median” category accounted for more than 90% of world production,
including six of the top 10 national aquaculture, 20 of the top 30, and 25
of the top 50 (Supplementary Table 15a).
In contrast, another 48 countries (illustrated in red in Fig. 3e) in the
category of “both < median” contributed only 1.5% of world production
(Supplementary Table 15b). The contribution primarily came from
Norway, which was the only top 10 largest national aquaculture with
relatively low within-group diversity and between-group diversity; and
Ghana was the only other top-50 national aquaculture in this category. A
large proportion of the countries in the “both < median” category are
located in Africa (21 countries), and the rest spread across the other four
regions: six in Asia (primarily Western Asia) and seven each in the
Americas (primarily the Caribbean), Europe (primarily Northern
Europe) and Oceania (entirely Pacific SIDS).
Fifty countries (illustrated in green in Fig. 3e) had a relatively high
within-group diversity yet a relatively low between-group diversity;
most (36) of them had a relatively high overall diversity (Supplementary
Table 15c). This category of “WGENS > median; ENSG ≤ median” pri­
marily includes 16 countries in Asia (mostly landlocked countries in
Central, Southern, or Western Asia), 15 countries in Europe (mostly
East-Central Europe), and 13 countries in Africa (nearly half of them
clustered in Western Africa). Only five countries in the Americas
belonged to this category, which nevertheless included the only two
landlocked countries in the region (Fig. 3e). Indeed, most (24) of the 37
landlocked national aquaculture worldwide, including 19 of the 31
LLDCs, belonged to this category. Landlocked aquaculture's relatively
low between-group diversity may primarily reflect their lack of coastal
resources (needed for the farming of most algae and molluscs species) as
well as their relatively low preferences for aquatic foods (Cai and Leung,
2022). The “WGENS > median; ENSG ≤ median” category accounted for
2.2% of world production, including one top-10 national aquaculture
(Egypt), two top-30, and nine top-50 (Supplementary Table 15c).
Another 50 countries (illustrated in purple in Fig. 3e) had a relatively
high between-group diversity yet a relatively low within-group di­
versity, with a relatively low overall diversity in most (36) of them
(Supplementary Table 15d). This is the exact opposite of the previous
10
Aquaculture 576 (2023) 739837
category. Nearly half (23) of the 50 countries in this category of
“WGENS ≤ median; ENSG > median” are in the Americas. This is
nevertheless not obvious in Fig. 3e, because these 23 countries include a
cluster of 13 Caribbean SIDS indiscernible in the map. Indeed, more than
half (28) of the 50 countries in this category are island economies,
including 22 SIDS. While island states tend to have a relatively high
between-group diversity thanks to long coastlines – algae and molluscs
are primarily cultivated in marine areas, a small island aquaculture
usually has a low within-group diversity because of small market (con­
strained by its small population and high transportation costs) and low
production scale. Other clusters in the “WGENS ≤ median; ENSG >
median” category include (i) 10 countries in Central America, and the
east and north coasts of South America; (ii) seven Pacific SIDS in Oce­
ania, (iii) five countries in Northern Europe and (iv) five countries in
South and South-eastern Africa (Supplementary 16d; Fig. 3e). This
category accounted for 5.5% of world production, including two top-10
national aquaculture (the Philippines and Chile), seven top-30, and 13
top-50 (Supplementary Table 15d).
4. Discussion
4.1. Lessons learned
Species diversification may not be consistent at different geographic
scales (Section 3.2). Hypothetically, if there were 200 countries each
cultivating a unique species in equal amounts, the species composition
of global aquaculture would be highly diverse (ENS = 200), even with
no species diversity (ENS = 1) at the national level. In reality, beneath
high species diversity in world aquaculture lies low species diversity in
most national aquaculture (Section 3.6). Therefore, species diversifica­
tion should be assessed at various levels (world, regional, sub-regional,
national, and sub-national) for evidence-based policymaking and sector
management in aquaculture.
Policy and planning for species diversification should take a holistic
approach recognizing that species diversity in aquaculture is shaped by
many factors, such as resource endowments, climate conditions, farming
systems and technologies, consumer preferences, market structure, and
institutional arrangements, among others (Guy et al., 2014; Metian
et al., 2020; Cai et al., 2022). Species diversification in aquaculture
tends to be a long-term, fluctuating process with diverse diversification
patterns varying across countries (Section 3.4).
A young aquaculture sector may start with a relatively high species
diversity when farmers attempt to cultivate a variety of species in the
exploratory stage (Brummett, 2007). Yet technical, market, or other
breakthroughs would tend to first occur in the most promising species
and result in production concentrated towards a few major species. The
development of minor species, together with diminishing growth po­
tential in major species because of various constraints, such as depletion
of suitable farm sites, productivity limit, and market satiation, could
turn the concentration process into a diversification trend with pro­
duction expansion primarily driven by more advantageous minor spe­
cies. These minor species may eventually become major species; then
their further expansion could turn the diversification trend back into a
concentration process.
Technical, economic, or institutional constraints may hinder the
establishment of minor species, whereas the advantages of major species
could be sustained or amplified by constant innovations, economies of
scales, insatiable global market, or other facilitating factors. Under this
situation, a concentration trend may go on for a long period. On the
other hand, certain factors, such as high consumer preferences for new
varieties of aquatic foods or aquaculture entrepreneurs' passion and
perseverance in cultivating new species, may constantly bring about
breakthroughs in new species to sustain a straight-up species diversifi­
cation trend. Therefore, there is no one-style-fits-all species diversifi­
cation pathway, and species diversification may not be a suitable
development strategy for all countries, or for a particular country at
J. Cai et al.
different stages of aquaculture development.
Species diversification could stimulate aquaculture growth under
certain situations, such as (i) increasing market demand in places where
consumers have high preferences over a variety of aquatic foods, (ii)
improving farming efficiency through polyculture, or (iii) utilizing
diverse farming environments and resources. However, instead of
spreading efforts across a range of species, concentrating on the most
advantageous ones tends to be a more effective way to promote aqua­
culture growth, especially in a young aquaculture sector that lacks ca­
pacities (e.g. entrepreneurship, technical expertise, and supply chain) to
expand aquaculture through species diversification. For an aquaculture
sector at its infancy, focusing on the most promising species would tend
to result in production concentration towards a few major species. Not
only can such species concentration facilitate rapid growth, it could also
be a process of building capacities for the development of novel or minor
species through spillover effects to lay a foundation for potential species
diversification in the future.
Species diversification is often considered a panacea to addressing
challenges faced by existing species, such as market satiation (Abellán
and Basurco, 1999), low productivity (Carrera-Quintana et al., 2022), or
disease outbreaks (Guy et al., 2014). Yet focusing on improving existing
species could sometimes be a more effective way to address such chal­
lenges. For instance, after a shrimp disease outbreak in 1999 devastated
Ecuador's aquaculture sector that was dominated by whiteleg shrimp
(Litopenaeus vannamei), the recovery was not via species diversification
but through improving shrimp farming regulations and management
(Marcillo, 2017). As a result, the share of whiteleg shrimp in the coun­
try's aquaculture production increased from 82% in 2000 to 98% in
2020, and the country experienced rapid aquaculture growth at a rate of
13.5% a year, compared to 7.7% in South America, 5.7% in the Amer­
icas, and 5.4% in world aquaculture.
In contrast with enthusiasm in the public sector, the private sector is
often more cautious about species diversification because establishing
novel species tends to be a technically challenging, financially
demanding, time-consuming, and risky process that may deter even an
advanced aquaculture sector (Harvey et al., 2017). For instance, Nor­
way's attempt to diversify its salmon-dominated aquaculture industry
through the farming of Atlantic cod (Gadus morhua) was hindered by
technical difficulties (e.g. knowledge gaps in intensive cod juvenile
production, inadequate broodstock development, and high mortality)
and foiled by competition from increased wild cod production
(Fernández-Polanco and Bjorndal, 2017; Puvanendran et al., 2021).
Institutional arrangements may be a subtle yet important factor
affecting species diversification. While measures for biodiversity con­
servation may constrain diversification via non-native species, regula­
tions that force farmers to internalize the impacts of their operations on
the ecosystem could motivate them to adopt a more diversified farming
system, such as Integrated Multi-trophic Aquaculture (IMTA) of finfish,
bivalves, and seaweeds (Cai et al., 2021). An aquaculture sector
comprised by numerous small or medium-scale farmers could be more
conducive to species diversification than one dominated by a few large
operators. Small or medium entrepreneurs tend to have more incentives
to pursue species diversification as a means to gain competitive ad­
vantages because it is easier for them reap the full benefit of a successful
diversification by shifting their entire operations to the new species. In
contrast, large operators usually lack such flexibilities, and new species
may encroach the market of their existing species. Conversely, large
operators tend to be better rewarded by concentrating efforts on
strengthening their dominance in established species to benefit from
economies of scale along the supply chain.
4.2. Future prospects
Accumulated efforts (e.g. domesticated seed, tailor-made feed, and
dedicated marketing) in improving the performance of established
species, which are usually the most advantageous species in the first
11
Aquaculture 576 (2023) 739837
place, would tend to further strengthen their dominance and make
aquaculture production concentrated towards a few winner species.
Indeed, species concentration has become an increasingly prominent
aquaculture development pathway in recent decades (Sections 3.3 and
3.7).
The characteristics of modern aquaculture (e.g. monoculture,
formulated feed, specialized seed production, global markets, and
industrialized operations), which tend to facilitate the accumulation of
economies of scale along the supply chain, are conducive to fostering
winner species (Cai et al., 2022). Increasingly stringent restrictions over
the use of non-native species in aquaculture for biodiversity conserva­
tion could facilitate initial species diversification among indigenous
species, such as the case of tilapia farming in Malawi (Cai et al., 2022). In
the long run, however, the production may have a tendency to
concentrate towards a few local winner species, which could result in
high species diversity in world aquaculture cum low diversity in national
aquaculture.
On the demand side, consumers' willingness to pay for novelty and
variety is a crucial factor to motivate and sustain species diversification,
yet such willingness tends to be weak in countries with low consumer
preferences for aquatic foods. Unfortunately, most countries have rela­
tively low preferences for aquatic foods, and these countries are ex­
pected to be the primary contributors to world population growth (Cai
and Leung, 2022).
In light of these supply- and demand-side factors that are unfav­
ourable to species diversification, we envision that the decelerating
trend of species diversification in recent decades (Section 3.3) would
tend to continue and eventually drive aquaculture towards a low-
diversity system similar to terrestrial farming, unless there are effec­
tive public interventions to turn the tide.
4.3. Way forward
Diversity is one of the 10 elements of agroecology that have been
synthesized to guide the transition to sustainable food and agricultural
systems (FAO, 2018). Public interventions to facilitate species diversi­
fication in aquaculture are beneficial for the sector's resilience and long-
term sustainability. Public support has largely focused on reducing the
cost of species diversification for the private sector. This includes
research in areas such as species selection methods and criteria (Abellán
and Basurco, 1999; Leung et al., 2007; Le Francois et al., 2010; Alvarez-
Lajonchère and Ibarra-Castro, 2013) as well as development projects
such as breeding programmes (Harvey et al., 2017). There has also been
a strong emphasis on building close partnership between research and
the industry (Metian et al., 2020).
More public efforts should be dedicated to increasing the benefits
and viability of species diversification, such as (i) promoting farming
systems and technologies that are conducive to species diversification
(e.g. polyculture); (ii) market development to increase consumer pref­
erences for aquatic foods and foster diverse dietary habits and culinary
cultures; and (iii) creating policy and business environments that
encourage long-term investments and award entrepreneurship.
While funding and other supports to develop potential species for
aquaculture would always be worthwhile in the long run, limited ca­
pacities (e.g. natural resources, human resources, and markets) do not
allow all promising species to establish at the same time. Therefore,
prioritization is necessary for efficient allocation of limited public re­
sources, and a country may seek guidance from species diversification
pathways in other countries, especially with geo-clustering of countries
with similar species diversity profiles or diversification patterns (Section
3.4 and Section 3.8).
For instance, while efforts in promoting species diversification have
been mostly focused on finfish diversity (Le Francois et al., 2010),
between-group diversity deserves more attention in light of the finding
of no between-group diversity in more than 40% of national aquaculture
(Section 3.6). Nearly all landlocked national aquaculture had virtually
J. Cai et al.
no between-group diversity, and the experiences of two exceptions (i.e.
a relatively high between-group diversity in Central African Republic
and Chad because of the farming of spirulina) may not offer much
guidance because of their low production scale (Supplementary
Table 6). China's experiences in inland aquaculture could provide useful
guidance in this regard (Newton et al., 2021). Better yet, landlocked
countries may learn from the global experiences of relatively high
between-group diversity in inland aquaculture of different scales. For
example, in 2020, Singapore had a nearly 700-t inland aquaculture
production, with a between-group diversity of 2.04 ENSG. Other inland
aquaculture sectors with a relatively high between-group diversity
include Chile (nearly 2000 t with 1.86 ENSG), Spain (nearly 20,000 t
with 1.54 ENSG), and Taiwan Province of China (over 100,000 t with
1.72 ENSG), among others (Supplementary Table 17).
Based on ENS as an intuitive diversity measure, Cai et al. (2022)
established a benchmarking system to synthesize global experiences to
guide policy and planning for species diversification in national aqua­
culture. The in-depth assessment conducted here could expand the
benchmarking system to incorporate the measures of within-group di­
versity (WGENS) and between-group diversity (ENSG) (Section 2.2). The
two measures provide important insights about the sources of species
diversity in aquaculture (Section 3.5), such as relatively low between-
group diversity in landlocked countries versus relatively low within-
group diversity in small island states (Section 3.8). It has been recog­
nized that the impact of biological diversity on the resilience of a dy­
namic ecosystem depends on the diversity of functional groups and the
species diversity within these groups (Elmqvist et al., 2003). Similarly,
different sources of species diversity in aquaculture would tend to in­
fluence its contribution to the sector's growth and resilience. Further
study in this area is worthwhile.
ENS and its within- and between-group components, which corre­
spond to gamma, alpha and beta diversity in ecology, should become a
standard indicator system widely used in the study of species diversity in
aquaculture and mainstreamed in policy dialogues. The within- and
between-group partition here is based on the categorization of five
customary taxonomic groups; other partitions can be adopted according
to different planning or research purposes.
4.4. Concluding remarks
The results of our systematic assessment have uncovered a wealth of
information that sheds light on species diversification in aquaculture.
We document the comprehensive results in the supplementary materials
(including a summary in Supplementary Table 18) to facilitate further
analysis. Policymakers and practitioners can use this information to
inform their decision-making at the national, regional, and global levels.
However, the usefulness of these results depends on the quality of the
underlying data. The existence of nei species items in the FAO statistics,
i.e. inadequate data disaggregation at the species level (Section 2.1)
affects accuracies in the measure of species diversity (FAO, 2020; Metian
et al., 2020). Cai et al. (2022) used an example to demonstrate that this
thorny issue tends to have a less impact on ENS as a diversity measure
that captures both richness and evenness, because nei species items are
usually associated with relatively small production. However, the data
imperfection tends to hinder more in-depth analyses, such as examining
species richness and evenness separately. Joint efforts from all stake­
holders (governments, international organizations, research commu­
nities, the private sector, and others) are needed to narrow the data gap.
Authors' statements
During the preparation of this work, the authors utilized GPT-3.5 for
language editing assistance. Subsequently, the authors thoroughly
reviewed and edited the content as necessary, assuming full re­
sponsibility for the publication's final content.
12
Aquaculture 576 (2023) 739837
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
Data will be made available on request.
Acknowledgements
We would like to acknowledge the valuable support provided by the
FAO Fisheries and Aquaculture Division (NFI) through the World
Aquaculture Performance Indicators (WAPI) initiative. This work aimed
to provide technical assistance in the development of information and
knowledge products, including factsheets, policy briefs, country/case
studies, and project reports, to facilitate evidence-based policymaking
and sector management in aquaculture. We extend our sincere gratitude
to Michael Parke at NOAA Fisheries and several anonymous reviewers
for their thorough review of this paper. Their insightful feedback greatly
contributed to the quality of this work. We would also like to express our
appreciation to the following individuals for their valuable comments,
suggestions, and support: Manuel Barange, Arnljotur Bergsson, Ian
Cowx, Lionel Dabbadie, Kim Friedman, Carlos Fuentevilla, Nathanael
Hishamunda, Audun Lem, Graham Mair, Felix Marttin, Ana Menezes,
KwangSuk Oh, Fernanda Sampaio, Doris Soto, Austin Stankus, Xinhua
Yuan, Wenbo Zhang, Xiaowei Zhou, and all participants of the NFI
seminar held on 1 March 2023, where this work was presented. Finally,
we want to emphasize that the opinions expressed in this paper are
solely those of the authors and do not necessarily reflect the positions of
their respective affiliations.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.aquaculture.2023.739837.
References
Abellán, E., Basurco, B. (Eds.), 1999. Marine Finfish Species Diversification: Current
Situation and Prospects in Mediterranean Aquaculture. CIHEAM, Zaragoza.
Alvarez-Lajonchère, L., Ibarra-Castro, L., 2013. Aquaculture species selection method
applied to marine fish in the Caribbean. Aquaculture. 408–409, 20–29. https://doi.
org/10.1016/j. aquaculture.2013.05.020.
Barrington, K., Chopin, T., Robinson, S., 2009. Integrated multi-trophic aquaculture
(IMTA) in marine temperate waters. In: Soto, D. (Ed.), Integrated Mariculture: A
Global Review, FAO Fisheries and Aquaculture Technical Paper No. 529, pp. 7–46.
Rome.
Basurco, B., Abellán, E., 1999. Finfish species diversification in the context of
Mediterranean marine fish farming development. In: Abellán, E., Basurco, B. (Eds.),
Marine Finfish Species Diversification: Current Situation and Prospects in
Mediterranean Aquaculture. Zaragoza, CIHEAM, pp. 9–25.
Bilio, M., 2008. Controlled reproduction and domestication in aquaculture – the current
state of the art. Part IV. Aquacult. Eur. 33 (2), 12–24.
Boyd, C.E., D'Abramo, L.R., Glencross, B.D., et al., 2020. Achieving sustainable
aquaculture: historical and current perspectives and future needs and challenges.
J. World Aquac. Soc. 51, 578–633.
Brummett, R.E., 2007. Indigenous species for African aquaculture development. In:
Bert, T.M. (Ed.), Ecological and Genetic Implications of Aquaculture Activities,
Methods and Technologies in Fish Biology and Fisheries, vol. 6. Springer, Dordrecht,
pp. 229–245. https://doi.org/10.1007/978-1-4020-6148-6_13.
Cai, J., Leung, P.S., 2022. Unlocking the potential of aquatic foods in global food security
and nutrition: a missing piece under the lens of seafood liking index. Global Food
Secur. 33, 100641 https://doi.org/10.1016/j.gfs.2022.100641.
Cai, J., Lovatelli, A., Aguilar-Manjarrez, J., Cornish, L., Dabbadie, L., Desrochers, A.,
Diffey, S., Garrido Gamarro, E., Geehan, J., Hurtado, A., Lucente, D., Mair, G.,
Miao, W., Potin, P., Przybyla, C., Reantaso, M., Roubach, R., Tauati, M., Yuan, X.,
2021. Seaweeds and microalgae: an overview for unlocking their potential in global
aquaculture development. In: FAO Fisheries and Aquaculture Circular No. 1229.
https://doi.org/10.4060/cb5670en. Rome.
Cai, J., Yan, X., Leung, P.S., 2022. Benchmarking species diversification in global
aquaculture. In: FAO Fisheries and Aquaculture Technical Paper No. 605. https://
doi.org/10.4060/cb8335en. Rome.
J. Cai et al.
Carrera-Quintana, S.C., Gentile, P., Girón-Hernández, J., 2022. An overview on the
aquaculture development in Colombia: current status, opportunities and challenges.
Aquaculture. 561, 738583 https://doi.org/10.1016/j.aquaculture.2022.738583.
CIHEAM, 2000. Recent advances in Mediterranean aquaculture finfish species
diversification. CIHEAM. (Cahiers Options Méditerranéennes; n. 47), Zaragoza.
Seminar of the CIHEAM Network on Technology of Aquaculture in the
Mediterranean on “Recent advances in Mediterranean aquaculture finfish species
diversification”, 1999/05/24-28, Zaragoza (Spain). https://om.ciheam.org/option.
php?IDOM=760#prettyPhoto. Seminar of the CIHEAM Network on Technology of
Aquaculture in the Mediterranean on “Recent advances in Mediterranean
aquaculture finfish species diversification”, 1999/05/24-28, Zaragoza (Spain).
Elmqvist, T., Folke, C., Nystrӧm, M., Peterson, G., Bengtsson, J., Walker, B., et al., 2003.
Response diversity, ecosystem change, and resilience. Front. Ecol. Environ. 1,
488–494.
FAO (Food and Agriculture Organization of the United Nations), 2011. World
aquaculture 2010. In: FAO Fisheries and Aquaculture Department. Technical Paper.
No. 500/1. Rome.
FAO (Food and Agriculture Organization of the United Nations), 2018. The 10 elements
of agroecology. In: Guiding the Transition to Sustainable Food and Agricultural
Systems. Rome.
FAO (Food and Agriculture Organization of the United Nations), 2019a. The State of the
World’s Aquatic Genetic Resources for Food and Agriculture. FAO Commission on
Genetic Resources for Food and Agriculture Assessments, Rome.
FAO (Food and Agriculture Organization of the United Nations), 2019b. Top 10 species
groups in global aquaculture 2017. In: World Aquaculture Performance Indicators
(WAPI) Factsheet. Rome. www.fao.org/3/ca5224en/ca5224en.pdf. Rome.
FAO (Food and Agriculture Organization of the United Nations), 2019c. The state of the
world’s biodiversity for food and agriculture. In: Bélanger, J., Pilling, D. (Eds.), FAO
Commission on Genetic Resources for Food and Agriculture Assessments. Rome. htt
p://www.fao.org/3/CA3129EN/CA3129EN.pdf.
FAO (Food and Agriculture Organization of the United Nations), 2020. The State of
World Fisheries and Aquaculture 2020. Sustainability in Action. Rome. https://doi.
org/10.4060/ca9229en.
FAO (Food and Agriculture Organization of the United Nations), 2022. Fishery and
Aquaculture Statistics. Global Production by Production Source 1950–2020
(FishStatJ). Rome. www.fao.org/fishery/en/statistics/software/fishstatj.
Fernández-Polanco, J., Bjorndal, T., 2017. Aquaculture diversification in Europe: the
Kingdom of Spain and the Kingdom of Norway. In: Harvey, B., Soto, D.,
Carolsfeld, J., Beveridge, M., Bartley, D.M. (Eds.), Planning for aquaculture
diversification: the importance of climate change and other drivers. FAO Technical
Workshop, 23–25 June 2016, FAO, Rome. FAO Fisheries and Aquaculture
Proceedings No. 47. Rome, pp. 37–49.
García-Márquez, J., Galafat, A., Alarcón, F.J., Figueroa, F.L., Martínez-Manzanares, E.,
Arijo, S., Abdala-Díaz, R.T., 2021. Cultivated and wild juvenile thick-lipped grey
mullet, Chelon labrosus: a comparison from a nutritional point of view. Animals
(Basel). 11 (7), 2112. https://doi.org/10.3390/ani11072112. PMID: 34359240;
PMCID: PMC8300304.
Gephart, J.A., Deutsch, L., Pace, M.L., Troell, M., Seekell, D.A., 2017. Shocks to fish
production: identification, trends, and consequences. Glob. Environ. Change. 42,
24–32. https://doi.org/10.1016/J.GLOENVCHA.2016.11.003.
Guy, J.A., McIlgorm, A., Waterman, P., 2014. Aquaculture in regional Australia:
responding to trade externalities. A northern NSW case study. J. Econ. Soc. Policy.
16 (1), 1–29.
Harvey, B., Soto, D., Carolsfeld, J., Beveridge, M., Bartley, D.M., 2017. Planning for
aquaculture diversification: the importance of climate change and other drivers. In:
FAO Technical Workshop, 23–25 June 2016, FAO, Rome. FAO Fisheries and
Aquaculture Proceedings No. 47. Rome.
Hill, M.O., 1973. Diversity and evenness: a unifying notation and its consequences.
Ecology. 54 (2), 427–432.
Jost, L., 2006. Entropy and diversity. Oikos. 113, 363–375. https://doi.org/10.1111/
j.2006.0030-1299.14714.x.
Jost, L., 2007. Partitioning diversity into independent alpha and beta components.
Ecology 88 (10), 2427–2439. https://doi.org/10.1890/06-1736.1. Erratum in:
Ecology. 2009. 90 (12), 3593. PMID: 18027744.
Laakso, M., Taagepera, R., 1979. "Effective" number of parties: a measure with
application to West Europe. Comp. Polit. Stud. 12 (1), 3–27. https://doi.org/
10.1177/001041407901200101.
13
Aquaculture 576 (2023) 739837
Le Francois, N.R., Jobling, M., Carter, C., Blier, P.U., Savoie, A. (Eds.), 2010. Finfish
Aquaculture Diversification. CAB International, Oxford.
Leung, P.S., Lee, C.S., O’Bryen, P.J. (Eds.), 2007. Species and System Selection for
Sustainable Aquaculture. Blackwell Publishing, Ames.
Liao, I.C., 2000. The state of finfish diversification in Asian aquaculture. In: Recent
Advances in Mediterranean Aquaculture Finfish Species Diversification. CIHEAM,
Zaragoza, pp. 109–125. Cahiers Options Méditerranéennes, No. 47.) Seminar of the
CIHEAM Network on Technology of Aquaculture in the Mediterranean on “Recent
advances in Mediterranean aquaculture finfish species diversification”, 1999/05/
24–28, Zaragoza (Spain). http://om.ciheam.org/om/pdf/c47/00600610.pdf.
Marcillo, F., 2017. Shrimp farming and the environment in Ecuador: past and present.
World Aquacult. 48 (3), 39–42.
Martínez-Cordero, F.J., 2007. Socioeconomic aspects of species and system selection for
sustainable aquaculture development in Mexico: Historic overview and current
general trends. In: Leung, P.S., Lee, C.S., O’Bryen, P.J. (Eds.), Species and System
Selection for Sustainable Aquaculture. Blackwell Publishing, Ames, pp. 225–239.
Megahed, M.E., Aly, S.M., 2009. Domestication and species diversification to improve
marine aquaculture in Egypt (a prospective view). Abbassa Int. J. Aquacult.
529–546. Special issue for Global Fisheries and Aquaculture Research Conference,
Cairo International Convention Center, 24–26 October 2009.
Metian, M., Troell, M., Christensen, V., Steenbeek, J., Pouil, S., 2020. Mapping diversity
of species in global aquaculture. Rev. Aquac. 12, 1090–1100. https://doi.org/
10.1111/raq.12374.
Muñoz-Lechuga, R., Sanz-Fernández, V., Cabrera-Castro, R., 2018. An overview of
freshwater and marine finfish aquaculture in Spain: emphasis on regions. Rev. Fish.
Sci. Aquacult. 26 (2), 195–213. https://doi.org/10.1080/23308249.2017.1381832.
Naylor, R.L., Burke, M., 2005. Aquaculture and ocean resources: raising tigers of the sea.
Annu. Rev. Environ. Resour. 30, 185–218.
Newton, R., Zhang, W., Xian, Z., McAdam, B., Little, D.C., 2021. Intensification,
regulation and diversification: the changing face of inland aquaculture in China.
Ambio. 65, 1–18.
Oboh, A., 2022. Diversification of farmed fish species: a means to increase aquaculture
production in Nigeria. Rev. Aquacult. 14 (4), 2089–2098. https://doi.org/10.1111/
raq.12690.
Ordover, J.A., Sykes, A.O., Willig, R.D., 1982. Herfindahl concentration, rivalry, and
mergers. Harvard Law Rev. 95 (8), 1857–1874. https://doi.org/10.2307/1340652.
Puvanendran, V., Mortensen, A., Johansen, L.-H., et al., 2021. Development of cod
farming in Norway: past and current biological and market status and future
prospects and directions. Rev. Aquacult. 14 (1), 308–342. https://doi.org/10.1111/
raq.12599.
Ravisankar, T., Sarada, C., Krishnan, M., 2005. Diversification of fish culture and exports
among major shrimp-producing countries of Asia: a spatial and temporal analysis.
Agricult. Econ. Res. Rev. 18, 187–195.
Roy, K., 2019. Technicalities to be considered for culture fisheries development in Indian
inland waters: seed and feed policy review. Environ. Dev. Sustain. 21, 281–302.
https://doi.org/10.1007/s10668-017-0037-3.
Schmidt, G., Espinos, F., Ruiz, F., Segarrax, M., Mañanos, E., Muñoz, J.L., et al., 2011.
Diversification in Aquaculture: A Tool for Sustainability. Spanish Ministry of
Environmental, Rural and Marine Affairs, Publications Centre, Madrid.
Sicuro, B., 2021. World aquaculture diversity: Origins and perspectives. Rev. Aquac. 13
(3), 1619–1634. https://doi.org/10.1111/raq.12537.
Teletchea, F., Fontaine, P., 2014. Levels of domestication in fish: implications for the
sustainable future of aquaculture. Fish Fish. 15, 181–195.
Thomas, M., Pasquet, A., Aubin, J., Nahon, S., Lecocq, T., 2021. When more is more:
taking advantage of species diversity to move towards sustainable aquaculture. Biol.
Rev. 96, 767–784. https://doi.org/10.1111/brv.12677.
Troell, M., Naylor, R.L., Metian, M., Beveridge, M., Tyedmers, P.H., Folke, C., et al.,
2014. Does aquaculture add resilience to the global food system? Proc. Natl. Acad.
Sci. U. S. A. 111, 13257–13263.
Tuomisto, H., 2010. A diversity of beta diversities: straightening up a concept gone awry.
Part 1. Defining beta diversity as a function of alpha and gamma diversity.
Ecography. 33 (1), 2–22. http://www.jstor.org/stable/20696326.
Wilson, J.R., Archer, B., 2010. Diversification pays: Economic perspectives on investment
in diversified aquaculture. In: Le Francois, N.R., Jobling, M., Carter, C., Blier, P.U.,
Savoie, A. (Eds.), Finfish Aquaculture Diversification. CAB International, Oxford,
pp. 514–530.
Supplementary Figures

Supplementary Figure 1: Species diversification in the ROW aquaculture (i.e. world aquaculture, excluding China), 1950–2020 1
Supplementary Figure 2: Effective number of diversification patterns (ENDP) 2
Supplementary Figure 3: Decomposition of species diversity in world aquaculture, 1950–2020 3
Supplementary Figure 4: Within-group share in species diversity: situations in four years 4
Supplementary Figure 5: Six components of species diversity, 2020 5
Supplementary Figure 6: Comparison of ENS trends and WGENS trends in national aquaculture 6
Supplementary Figure 7: ENSG trends in national aquaculture, 1990–2020 7
Supplementary Figure 8: Box-and-whisker plots of species diversity in national aquaculture, 2020 8
Supplementary Figure 9: Quartile distribution of measures of species diversity, 2020 9
Supplementary Figure 10: Temporal correlations between production and ENS, 2010–2020 10
Supplementary Tables
Supplementary Table 1a: Regional aquaculture with INCREASED species diversity, 1990–2020 and three sub-decades 11
Supplementary Table 1b: Regional aquaculture with DECREASED species diversity, 1990–2020 and three sub-decades 12
Supplementary Table 2a: Inverted U-shape ENS trends, 1990–2020 13
Supplementary Table 2b: Inverted N-shape ENS trends, 1990–2020 15
Supplementary Table 2c: Straight-down ENS trends, 1990–2020 16
Supplementary Table 2d: Straight-up ENS trends, 1990–2020 17
Supplementary Table 2e: U-shape ENS trends, 1990–2020 18
Supplementary Table 2f: N-shape ENS trends, 1990–2020 19
Supplementary Table 3: Categorizing ENS trends in regional aquaculture, 1990–2020 20
Supplementary Table 4: Within-group share in species diversity in regional aquaculture 21
Supplementary Table 5: National aquaculture in 2020 where within-group species diversity was the sole source of species diversity 22
Supplementary Table 6: Species diversity in landlocked aquaculture, 2020 24
Supplementary Table 7: National aquaculture in 2020 where between-group diversity was greater than within-group diversity 25
Supplementary Table 8: Species diversity in island aquaculture, 2020 27
Supplementary Table 9a: National aquaculture whose ENS trend during 1990–2020 belonged to the category of U3 29
Supplementary Table 9b: National aquaculture whose ENS trend during 1990–2020 belonged to the category of D3 29
Supplementary Table 10: National aquaculture with no species diversity (ENS = 1) 30
Supplementary Table 11: National aquaculture with extraordinarily high diversity (aka outliers) 31
Supplementary Table 12: Cross-sectional correlations between aquaculture production and species diversity in national aquaculture 32
Supplementary Table 13: Temporal correlation between a country’s production and diversity during 2010–2020 33
Supplementary Table 14: National aquaculture with statistically significant temporal correlations between production and ENS in the most recent decade (2010–
2020) 34
Supplementary Table 15a: Countries in the “Both > median” category (i.e. relatively high within-group diversity and between-group diversity) 36
Supplementary Table 15b: Countries in the “Both < median” category (i.e. relatively low within-group diversity and between-group diversity) 37
Supplementary Table 15c: Countries in the “WGENS > median; ENSG ≤ median” category (i.e. relatively high within-group diversity yet relatively low
between-group diversity) 38
Supplementary Table 15d: Countries in the “WGENS ≤ median; ENSG > median” category (i.e. relatively low within-group diversity yet relatively high
between-group diversity) 49
Supplementary Table 16: Species diversity in countries with the largest land area 40
Supplementary Table 17: Between-group diversity in inland aquaculture, 2020 41
Supplementary Table 18: A summary of key results 42
Supplementary Figure 1: Species diversification in the ROW aquaculture (i.e. world aquaculture, excluding China), 1950–2020

1
Supplementary Figure 2: Effective number of diversification patterns (ENDP)

World (161 national aquaculture) 5.06

Africa (38 national aquaculture) 4.74

Americas (33 national aquaculture) 5.03

Asia (42 national aquaculture) 4.57

Europe (37 national aquaculture) 4.39

Oceania (11 national aquaculture) 4.34

Landlocked countries (32 national aquaculture) 5.21

LLDCs (27 national aquaculture) 5.12

Island economies (38 national aquaculture) 4.51

SIDS (25 national aquaculture) 5.21

Notes: The “effective number of diversification patterns” (ENDP) measures the evenness of the distribution of six diversification
patterns among countries – see more details in section 2.3. LLDCs = Landlocked Developing Countries. SIDS = Small Island
Developing States.

2
Supplementary Figure 3: Decomposition of species diversity in world aquaculture, 1950–2020

69.6

68.2 68.3
67.7 67.7 67.5
67.3
66.8
66.1 66.2

64.4 64.5
63.8
62.9 14.60
63.1 13.48 13.12 13.56
11.06 10.71
9.43 10.04
7.43 7.15 7.67 7.87
6.62 6.16 6.34

2.90 2.97 3.05 3.08 3.11 3.16 3.22 3.25 3.26 3.45 3.47 3.41 3.50
2.41 2.40

1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 2010 2015 2020

Within-group diversity (WGENS) Between-group diversity (ENSG) Share of within-group diversity (%)

3
Supplementary Figure 4: Within-group share in species diversity: situations in four years

(a) Year 1990 (b) Year 2000

100 100
90 90

Within-group share (%)

Within-group share (%)

80 80
70 70
60 60
50 50
40 40
30 30
20 20
10 10
0 0
0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 100 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 100
Percentile of all countries (%) Percentile of all countries (%)

(c) Year 2010 (d) Year 2020

100 100
90 90
Within-group share (%)

Within-group share (%)

80 80
70 70
60 60
50 50
40 40
30 30
20 20
10 10
0 0
0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 100 0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 100
Percentile of all countries (%) Percentile of all countries (%)

4
Supplementary Figure 5: Six components of species diversity, 2020

Notes: The boundaries of the ranges are determined by quartile distributions presented in the table below. In parentheses, the first number is the
number of countries in the category; the second number (in percentage) represents the category’s share in world production.
 Chart (a): Among 181 countries with ENS > 1 in 2020, the contribution of between-group diversity to species diversity was zero (i.e.
no between-group diversity) in 67 countries (0.43 percent of world production).
 Chart (b): Finfish diversity accounted for 100 percent of species diversity in 65 countries (more than one third of 179 finfish farming
countries).
 Chart (c): Nine countries in “XL” had an extraordinarily large molluscs share in species diversity, which is greater than the third
quartile plus 1.5 times of the difference between the third quartile and the first quartile, yet smaller than the maximum.
 Charts (d), (e) and (f): Crustaceans had no contribution to species diversity in more than half of crustacean farming countries. Same for
MAA and algae.
Contribution to species diversity (%)
Quartiles Between-group share Finfish share Molluscs share Crustaceans share MAA share Algae share
(181 countries) (179) (63) (77) (26) (48)
Minimum 0 0 0 0 0 0
1st quartile 0 26.9 0 0 0 0
2nd quartile (median) 17.28 79.82 0.67 0 0 0
3rd quartile 54.67 100 8.99 0.37 0.09 0.09
Maximum, excluding outliers n.a. n.a. 21.31 0.51 0.1 0.19
Maximum 100 100 100 34.57 9.79 100
All maps were created using Datawrapper; the country/territory boundaries depicted therein do not represent authors’ opinions.
Countries/territories with a small land area may not be discernible on the maps.

5
Supplementary Figure 6: Comparison of ENS trends and WGENS trends in national aquaculture

Notes: Up = ENS or WGENS increase; Down = ENS or WGENS decrease; and Flat = ENS or WGENS unchanged. In
parentheses, the first number represents the number of national aquaculture in the category; the second number (in percentage)
represents the prevalence of the category (i.e. the first number divided by the total number of countries in all categories); and the
third number (in percentage) represents the category’s share in world production during the period. National aquaculture that are
not covered by the categorization, along with countries with no aquaculture, are illustrated in grey. All maps were created using
Datawrapper; the country/territory boundaries depicted therein do not represent authors’ opinions. Countries/territories with a
small land area may not be discernible on the maps.

6
Supplementary Figure 7: ENSG trends in national aquaculture, 1990–2020

Notes: All maps were created using Datawrapper; the country/territory boundaries depicted therein do not represent authors’ opinions. Countries/territories with a small land area
may not be discernible on the maps. Down = ENSG decrease; Up = ENSG increase; and Flat = ENSG unchanged. In parentheses, the first number represents the number of
national aquaculture in the category; the second number (in percentage) represents the prevalence of the category (i.e. the first number divided by the total number of countries in
all categories); and the third number (in percentage) represents the category’s share in world production during the period. National aquaculture that are not covered by the
categorization, together with countries with no aquaculture, are illustrated in the grey colour.

7
Supplementary Figure 8: Box-and-whisker plots of species diversity in national aquaculture, 2020

Notes: In the first bar (on ENS), the five numeric labels from bottom to top represent, respectively, the minimum ENS, the first
quartile (25 percent), the median (50 percent, in bold and italic font), the third quartile (75 percent), and the “maximum” ENS
excluding “outliers” that are extraordinarily large ENS with distance from the third quartile greater than 1.5 times of the height of
the box (i.e. distance between the third quartile and the first quartile). When overlapped, only the label of a higher quartile is
shown. In the legends, a number in parentheses represent the number of countries (including outliers) that farmed the species
group. The outliers are not shown in the chart for clarity but presented in Supplementary Table 11. The quartile distributions are
mapped in Supplementary Figure 9.

8
Supplementary Figure 9: Quartile distribution of measures of species diversity, 2020

Notes: The boundaries of the ranges are based on the quartile distributions illustrated in Supplementary Figure 8. The first three
maps here are identical to map (a), (b), and (c) in Figure 3 in the main text. Readers may refer to the notes accompanying these
two figures to better understand the maps presented here. All maps were created using Datawrapper; the country/territory
boundaries depicted therein do not represent authors’ opinions. Countries/territories with a small land area may not be discernible
on the maps.

9
Supplementary Figure 10:

Notes: In parentheses, the first number represents the number of national aquaculture in the category; the second
number (in percentage) represents represents the category’s share in world production during the period. For each
country, there were up to 11 observations during 2010–2020. Thus, the temporal correlation between its production
and ENS was statistically significant (p-value < 0.05) when r > 0.6 (dark blue) or r < 0.6 (dark red). Countries or
territories that belong to these two categories are presented in Supplementary Table 14. All maps were created using
Datawrapper; the country/territory boundaries depicted therein do not represent authors’ opinions.
Countries/territories with a small land area may not be discernible on the maps.

10
Supplementary Table 1a: Regional aquaculture with INCREASED species diversity, 1990–2020 and three sub-decades
Ranked by
Overall three decades (1990–2020): First decade (1990–2000): Second decade (2000–2010): Third decade (2010–2020):
production
22 regional aquaculture with increased ENS 22 regional aquaculture with increased ENS 20 regional aquaculture with increased ENS 13 regional aquaculture with increased ENS
share
U1: ENS increased with both WGENS and ENSG up (share of world production during the period)
1 Developing regions (93.39 %) Developing regions (88.24 %) Developing regions (92.7 %) Southern Asia (8.09 %)
2 Asia (91.11 %) South-eastern Asia (8.69 %) Asia (90.83 %) Central America (0.34 %)
3 Eastern Asia (66.19 %) Southern Asia (6.69 %) Eastern Asia (70.28 %) Western Africa (0.32 %)
4 Southern Asia (7.52 %) Southern Europe (1.54 %) Southern Europe (0.98 %) Eastern Europe (0.32 %)
5 Northern America (0.91 %) Western Europe (1.41 %) Western Europe (0.6 %) Middle Africa (0.01 %)
6 Southern Europe (0.84 %) Eastern Europe (0.7 %) Eastern Africa (0.4 %)
7 Western Europe (0.55 %) Western Asia (0.21 %) Oceania (0.27 %)
8 Eastern Europe (0.38 %) Landlocked developing countries (0.19 %) Australia and New Zealand (0.25 %)
9 Western Asia (0.32 %) Small Island Developing States (0.15 %) Landlocked developing countries (0.24 %)
10 Landlocked developing countries (0.31 %) Pacific islands (0.04 %) Caribbean (0.06 %)
11 Pacific islands (0.02 %) Pacific islands (0.02 %)
12 Middle Africa (0.01 %) Southern Africa (0.01 %)
13 Middle Africa (0.01 %)
U2: ENS increased with only WGENS up (share of world production during the period)
1 South America (2.15 %) World excluding China (36.16%) South-eastern Asia (13.42 %) Sub-Saharan Africa (0.78 %)
2 Northern Africa (1.01 %) Developed regions (11.66 %) Africa (1.5 %) Eastern Africa (0.44 %)
3 Eastern Africa (0.4 %) Europe (5.34 %) Eastern Europe (0.39 %) Landlocked developing countries (0.39 %)
4 Western Africa (0.23 %) Americas (3.36 %) Small Island Developing States (0.11 %) Small Island Developing States (0.06 %)
5 Small Island Developing States (0.09 %) Latin America and the Caribbean (1.78 %) Central Asia (0.01 %) Central Asia (0.04 %)
6 Central Asia (0.04 %) Northern America (1.57 %)
7 South America (1.5 %)
8 Northern Africa (0.38 %)
9 Western Africa (0.07 %)
10 Central Asia (0.06 %)
U3: ENS increased with only ENSG up (share of world production during the period)
1 Oceania (0.25 %) Eastern Asia (74.5 %) Northern America (1.12 %) Northern America (0.61 %)
2 Australia and New Zealand (0.22 %) Southern Africa (0.01 %) Western Asia (0.3 %) Southern Europe (0.6 %)
3 Caribbean (0.05 %) Caribbean (0.03 %)
4 Southern Africa (0.01 %)
Notes: This table is a rearrangement of relevant contents in Table 1. The share of each regional aquaculture in world production is presented in parentheses.

11
Supplementary Table 1b: Regional aquaculture with DECREASED species diversity, 1990–2020 and three sub-decades
Ranked by
Overall three decades (1990–2020): First decade (1990–2000): Second decade (2000–2010): Third decade (2010–2020):
production
10 regional aquaculture with decreased ENS 10 regional aquaculture with decreased ENS 12 regional aquaculture with decreased ENS 19 regional aquaculture with decreased ENS
share
D1: ENS declined with both WGENS and ENSG down (share of world production during the period)
1 Developed regions (6.53 %) Northern Europe (1.68 %) World excluding China (34.91%) World excluding China (42.01%)
2 Europe (3.48 %) Africa (0.72 %) Developed regions (7.27 %) South-eastern Asia (21.44 %)
3 Northern Europe (1.71 %) Sub-Saharan Africa (0.34 %) Europe (3.74 %) Developed regions (4.71 %)
4 Central America (0.32 %) Eastern Africa (0.25 %) Northern Europe (1.77 %) Americas (3.32 %)
5 Central America (0.19 %) Central America (0.34 %) Europe (2.88 %)
6 Western Africa (0.15 %) Latin America and the Caribbean (2.71 %)
7 South America (2.34 %)
8 Northern Europe (1.69 %)
9 Western Europe (0.28 %)
10 Oceania (0.21 %)
D2: ENS declined with only WGENS down (share of world production during the period)
1 Asia (90.16 %) Americas (3.63 %) Developing regions (95.29 %)
2 Oceania (0.33 %) Latin America and the Caribbean (2.51 %) Asia (91.55 %)
3 Australia and New Zealand (0.29 %) South America (2.1 %) Eastern Asia (61.61 %)
4 Caribbean (0.09 %) Northern Africa (0.94 %) Northern Africa (1.26 %)
5 Middle Africa (0.01 %) Western Asia (0.37 %)
6 Australia and New Zealand (0.19 %)
7 Pacific islands (0.02 %)
8 Southern Africa (0.01 %)
D3: ENS declined with only ENSG down (share of world production during the period)
1 World excluding China (39.6%) Southern Asia (6.83 %) Africa (2.04 %)
2 South-eastern Asia (17.05 %) Sub-Saharan Africa (0.57 %)
3 Americas (3.43 %)
4 Latin America and the Caribbean (2.51 %)
5 Africa (1.66 %)
6 Sub-Saharan Africa (0.64 %)
Notes: This table is a rearrangement of relevant contents in Table 1. The share of each regional aquaculture in world production is presented in parentheses.

12
Supplementary Table 2a: Inverted U-shape ENS trends, 1990–2020
Inverted U-shape: ENS increased in the first decade (or the second decade if no Aquaculture production in 2020 Effective number of species (ENS)
production in 1990) yet declined in the last decade.
 Upward overall trends in blue: ENS2020 > ENS1990 (or ENS2020 in case of no ENS1990)

 Downward overall trends in red: ENS2020 < ENS1990 (or ENS2000 in case of no ENS1990) Share of world
Countries/territories indexed by their ranking in world aquaculture 2020 Tonnes 1990 2000 2010 2020
production (%)
LLDCs = Landlocked Developing Countries
SIDS = Small Island Developing States
Total (59) 97 046 818 79.1701 n.a. n.a. n.a. n.a.
Africa (17) 1 840 400 1.5014 n.a. n.a. n.a. n.a.
8. Egypt 1 591 896 1.2987 3.27 3.95 3.90 3.77
Landlocked (LLDCs) 35. Uganda 123 897 0.1011 2.75 3.43 1.96 1.89
45. Ghana 64 010 0.0522 1.70 1.90 1.39 1.36
58. Tunisia 23 486 0.0192 6.92 7.34 5.45 2.95
Island 74. Madagascar 13 550 0.0111 1.75 2.42 3.10 2.11
79. South Africa 9 753 0.0080 4.39 4.47 5.86 4.66
Landlocked (LLDCs) 85. Mali 7 686 0.0063 1.72 1.93 2.13 1.78
114. Angola 2 062 0.0017 1.00 1.15 1.08
Landlocked (LLDCs) 123. Burundi 1 450 0.0012 1.00 1.18 1.43 1.30
131. Congo 900 0.0007 1.00 1.00 2.49 1.60
136. Togo 730 0.0006 1.00 1.00 2.13 1.31
Landlocked (LLDCs) 142. Ethiopia 534 0.0004 1.69 3.75 2.96 2.17
146. Namibia 321 0.0003 1.00 3.59 4.15 1.61
160. Sierra Leone 85 0.0001 1.00 1.00 1.28 1.25
Island 173. Mayotte 20 0.0000 1.00 1.64 1.00
180. Libya 10 0.0000 1.00 2.95 2.55 1.00
Island 181. Réunion 10 0.0000 1.26 2.43 2.96 2.00
Americas (10) 2 912 662 2.3761 n.a. n.a. n.a. n.a.
9. Chile 1 505 486 1.2282 3.92 5.05 4.48 3.39
14. Ecuador 774 569 0.6319 1.53 1.77 1.66 1.11
16. Brazil 630 200 0.5141 1.60 8.19 7.93 5.13
113. Argentina 2 085 0.0017 1.00 2.71 3.20 3.12
Island (SIDS) 153. Guyana 138 0.0001 1.96 3.84 4.95 2.00
Island (SIDS) 159. Saint Lucia 92 0.0001 1.00 1.00 2.11 1.44
Island (SIDS) 162. Guadeloupe 45 0.0000 1.00 1.00 2.36 1.53
Island (SIDS) 163. Martinique 42 0.0000 2.17 2.75 2.47 1.68
Island (SIDS) 187. Trinidad and Tobago 4.2 0.0000 1.00 1.20 3.11 1.73
190. French Guiana 2.0 0.0000 1.00 1.63 2.83 2.00
Asia (17) 89 940 649 73.3729 n.a. n.a. n.a. n.a.
1. China 70 483 538 57.4999 13.57 16.61 28.18 28.00
Island 2. Indonesia 14 845 014 12.1105 10.26 10.30 6.40 5.63
6. Republic of Korea 2 327 903 1.8991 6.90 7.14 7.74 6.95
11. Myanmar 1 145 018 0.9341 1.11 1.22 4.70 3.98
20. Cambodia 400 400 0.3266 5.52 6.03 8.20 7.70
21. Malaysia 400 017 0.3263 4.26 9.47 9.12 8.47

13
Inverted U-shape: ENS increased in the first decade (or the second decade if no Aquaculture production in 2020 Effective number of species (ENS)
production in 1990) yet declined in the last decade.
 Upward overall trends in blue: ENS2020 > ENS1990 (or ENS2020 in case of no ENS1990)

 Downward overall trends in red: ENS2020 < ENS1990 (or ENS2000 in case of no ENS1990) Share of world
Countries/territories indexed by their ranking in world aquaculture 2020 Tonnes 1990 2000 2010 2020
production (%)
LLDCs = Landlocked Developing Countries
SIDS = Small Island Developing States
Island 24. Taiwan Province of China 278 503 0.2272 11.40 15.66 11.21 11.04
59. Iraq 22 704 0.0185 1.00 1.00 1.91 1.71
Landlocked (LLDCs) 66. Armenia 18 350 0.0150 1.93 2.70 3.51 2.77
Landlocked (LLDCs) 84. Afghanistan 8 050 0.0066 1.00 1.00 1.28 1.04
98. China, Hong Kong Special Administrative Region 3 835 0.0031 12.05 12.27 9.52 7.25
Island 103. Brunei Darussalam 3 500 0.0029 3.46 4.86 3.82 2.47
117. Georgia 2 027 0.0017 1.18 3.11 4.65 3.34
134. Lebanon 828 0.0007 1.00 1.00 1.40 1.20
Landlocked (LLDCs) 135. Tajikistan 775 0.0006 2.06 2.41 3.82 2.94
Landlocked (LLDCs) 150. Bhutan 181 0.0001 1.00 1.00 7.15 6.31
Island (SIDS) 186. Bahrain 5.5 0.0000 1.00 2.48 1.36
Europe (12) 2 246 879 1.8330 n.a. n.a. n.a. n.a.
10. Norway 1 490 412 1.2159 1.17 1.43 1.41 1.31
25. Spain 276 571 0.2256 1.91 2.41 2.97 2.97
28. France 191 350 0.1561 4.03 4.59 5.36 4.62
36. Italy 122 778 0.1002 3.96 4.43 4.80 4.51
Island 42. Faeroe Islands 89 055 0.0727 1.03 1.17 1.20 1.01
Island 51. Iceland 40 595 0.0331 1.23 2.07 3.45 1.65
Landlocked (LLDCs) 75. Republic of Moldova 12 600 0.0103 1.99 5.27 4.57 4.17
76. Romania 12 200 0.0100 4.49 5.34 7.01 5.93
81. Albania 9 284 0.0076 1.58 2.14 3.72 3.36
127. Channel Islands 1 107 0.0009 1.30 1.81 1.65 1.35
137. Latvia 717 0.0006 1.00 1.80 2.51 2.51
149. Belgium 209 0.0002 2.50 5.15 2.42 1.00
Oceania (3) 106 228 0.0867 n.a. n.a. n.a. n.a.
39. Australia 106 088 0.0865 6.24 7.61 7.19 4.43
Island (SIDS) 154. Guam 118 0.0001 2.50 3.01 2.85 2.79
Island (SIDS) 172. Palau 22 0.0000 1.00 1.00 2.71 2.53

14
Supplementary Table 2b: Inverted N-shape ENS trends, 1990–2020
Inverted N-shape ENS trend: ENS decreased during first and third decades whereas Aquaculture production in 2020 Effective number of species (ENS)
increased during the second decade.
 Upward overall trends in blue: ENS2020 > ENS1990

 Downward overall trends in red: ENS2020 < ENS1990 Share of world

Countries/territories indexed by their ranking in world aquaculture 2020 Tonnes 1990 2000 2010 2020
production (%)
LLDCs = Landlocked Developing Countries
SIDS = Small Island Developing States
Total (18) 673 246 0.5492 n.a. n.a. n.a. n.a.
Africa (6) 62 533 0.0510 n.a. n.a. n.a. n.a.
Landlocked (LLDCs) 48. Zambia 45 670 0.0373 2.93 2.88 3.54 3.21
92. Algeria 5 436 0.0044 3.34 2.76 5.51 4.48
94. Côte d'Ivoire 4 620 0.0038 3.43 1.86 2.95 2.48
102. Cameroon 3 556 0.0029 1.99 1.65 3.96 2.12
Landlocked (LLDCs) 109. Lesotho 2 600 0.0021 2.08 1.00 1.01 1.00
Landlocked (LLDCs) 138. Burkina Faso 651 0.0005 2.00 1.00 2.56 2.33
Americas (3) 56 528 0.0461 n.a. n.a. n.a. n.a.
56. Nicaragua 29 410 0.0240 1.27 1.09 1.11 1.00
Island (SIDS) 57. Cuba 26 200 0.0214 3.98 3.25 4.00 3.97
Island (SIDS) 130. Jamaica 918 0.0007 1.08 1.02 1.19 1.05
Asia (2) 172 139 0.1404 n.a. n.a. n.a. n.a.
Landlocked (LLDCs) 34. Lao People's Democratic Republic 130 020 0.1061 8.00 5.12 10.39 10.33
Island 50. Sri Lanka 42 119 0.0344 1.14 1.07 5.88 5.17
Europe (3) 255 882 0.2087 n.a. n.a. n.a. n.a.
Island 27. United Kingdom 221 000 0.1803 2.44 1.79 2.13 1.69
Island 63. Malta 19 829 0.0162 1.89 1.48 2.00 1.71
69. Finland 15 053 0.0123 1.08 1.07 1.31 1.26
Oceania (4) 126 164 0.1029 n.a. n.a. n.a. n.a.
Island 37. New Zealand 118 582 0.0967 1.74 1.53 1.60 1.57
Island (SIDS) 89. Papua New Guinea 6 102 0.0050 1.82 1.21 2.86 2.30
Island (SIDS) 122. New Caledonia 1 478 0.0012 1.51 1.11 1.25 1.03
Island (SIDS) 192. Kiribati 2.0 0.0000 1.04 1.01 1.02 1.00

15
Supplementary Table 2c: Straight-down ENS trends, 1990–2020
Straight down: ENS decreased in all three decades or the second and third decades in Aquaculture production in 2020 Effective number of species (ENS)
case of no aquaculture production in 1990
 Upward overall trends in blue: ENS2020 > ENS1990 (or ENS2020 in case of no ENS1990)

 Downward overall trends in red: ENS2020 < ENS1990 (or ENS2000 in case of no ENS1990) Share of world
Countries/territories indexed by their ranking in world aquaculture 2020 Tonnes 1990 2000 2010 2020
production (%)
LLDCs = Landlocked Developing Countries
SIDS = Small Island Developing States
Total (8) 273 105 0.2228 n.a. n.a. n.a. n.a.
Africa (3) 18 978 0.0155 n.a. n.a. n.a. n.a.
Landlocked (LLDCs) 68. Zimbabwe 15 425 0.0126 2.20 1.22 1.15 1.03
Island (SIDS) 104. Mauritius 3 298 0.0027 3.64 3.07 1.53 1.15
147. Liberia 255 0.0002 3.35 1.69 1.48
Americas (2) 224 616 0.1832 n.a. n.a. n.a. n.a.
30. Canada 171 007 0.1395 6.75 4.40 3.60 3.15
46. Venezuela (Bolivarian Republic of) 53 609 0.0437 4.26 3.03 2.34 1.85
Asia (2) 7 793 0.0064 n.a. n.a. n.a. n.a.
Island 86. Cyprus 7 343 0.0060 2.53 2.38 2.20 2.05
144. Kuwait 450 0.0004 3.76 1.15 1.00
Europe (1) 21 718 0.0177 n.a. n.a. n.a. n.a.
60. Croatia 21 718 0.0177 7.30 7.14 5.12

16
Supplementary Table 2d: Straight-up ENS trends, 1990–2020
Straight-up: ENS increased in all three decades or the second and third decades in Aquaculture production in 2020 Effective number of species (ENS)
case of no aquaculture production in 1990
 Upward overall trends in blue: ENS2020 > ENS1990 (or ENS2020 in case of no ENS1990)

 Downward overall trends in red: ENS2020 < ENS1990 (or ENS2000 in case of no ENS1990) Share of world
Countries/territories indexed by their ranking in world aquaculture 2020 Tonnes 1990 2000 2010 2020
production (%)
LLDCs = Landlocked Developing Countries
SIDS = Small Island Developing States
Total (31) 8 205 452 6.6939 n.a. n.a. n.a. n.a.
Africa (3) 4 299 0.0035 n.a. n.a. n.a. n.a.
101. Democratic Republic of the Congo 3 590 0.0029 1.00 1.01 1.13 1.15
Landlocked (LLDCs) 139. Niger 649 0.0005 1.00 1.00 1.82 1.92
161. Gabon 60 0.0000 1.00 1.24 1.31 1.33
Americas (3) 71 734 0.0585 n.a. n.a. n.a. n.a.
44. Honduras 71 151 0.0580 1.20 1.62 1.96 1.99
Island (SIDS) 141. Belize 565 0.0005 1.00 1.00 1.43 1.47
Island (SIDS) 176. Puerto Rico 18 0.0000 1.13 1.79 1.97 2.65
Asia (10) 7 562 660 6.1696 n.a. n.a. n.a. n.a.
4. Viet Nam 4 614 692 3.7646 2.91 4.22 6.43 8.24
5. Bangladesh 2 583 866 2.1079 1.65 7.69 12.01 14.30
31. Pakistan 162 462 0.1325 1.03 1.04 5.56 5.67
40. Saudi Arabia 99 907 0.0815 1.21 2.16 2.19 3.95
Landlocked (LLDCs) 43. Nepal 76 822 0.0627 1.75 4.40 5.87 6.98
71. Israel 14 700 0.0120 3.09 5.35 5.77 5.80
Island (SIDS) 93. Singapore 4 829 0.0039 4.38 4.79 8.07 11.67
Landlocked (LLDCs) 110. Kyrgyzstan 2 550 0.0021 1.80 2.09 2.95 3.98
111. Syrian Arab Republic 2 300 0.0019 1.69 2.35 3.42 3.67
Landlocked (LLDCs) 143. Azerbaijan 533 0.0004 1.57 1.68 2.68 4.74
Europe (14) 561 255 0.4579 n.a. n.a. n.a. n.a.
22. Russian Federation 291 194 0.2376 2.02 4.67 5.45 10.03
47. Poland 47 700 0.0389 1.67 2.28 2.85 3.69
49. Denmark 42 629 0.0348 1.13 1.31 1.62 2.24
52. Netherlands 39 940 0.0326 1.14 1.61 1.95 2.18
Island 53. Ireland 37 751 0.0308 2.41 3.13 3.18 3.29
55. Germany 32 258 0.0263 3.51 3.56 4.15 5.42
Landlocked 62. Czechia 20 401 0.0166 1.87 1.96 2.09
70. Bulgaria 15 052 0.0123 1.93 5.37 6.02 6.24
77. Sweden 12 090 0.0099 2.14 2.17 2.20 2.24
Landlocked 82. Belarus 9 266 0.0076 1.03 1.68 2.20 2.48
95. Austria 4 527 0.0037 1.89 2.60 4.18 7.02
96. Lithuania 4 477 0.0037 1.07 1.22 1.52 2.85
Landlocked 112. Slovakia 2 296 0.0019 1.85 2.11 3.82
118. Slovenia 1 673 0.0014 2.78 3.64 4.38
Oceania (1) 5 504 0.0045 n.a. n.a. n.a. n.a.
Island (SIDS) 91. Solomon Islands 5 504 0.0045 1.00 1.00 1.00 1.01

17
Supplementary Table 2e: U-shape ENS trends, 1990–2020
U-shape: ENS declined in the first decade (or the second decade if no production in Aquaculture production in 2020 Effective number of species (ENS)
1990) yet increased in the last decade
 Upward overall trends in blue: ENS2020 > ENS1990 (or ENS2020 in case of no ENS1990)

 Downward overall trends in red: ENS2020 < ENS1990 (or ENS2000 in case of no ENS1990) Share of world
Countries/territories indexed by their ranking in world aquaculture 2020 Tonnes 1990 2000 2010 2020
production (%)
LLDCs = Landlocked Developing Countries
SIDS = Small Island Developing States
Total (28) 5 889 312 4.8045 n.a. n.a. n.a. n.a.
Africa (5) 140 111 0.1143 n.a. n.a. n.a. n.a.
38. United Republic of Tanzania 108 568 0.0886 1.18 1.03 1.03 1.70
61. Kenya 20 831 0.0170 4.61 3.54 2.14 2.33
Landlocked (LLDCs) 80. Malawi 9 393 0.0077 2.52 1.32 3.10 3.56
128. Senegal 1 100 0.0009 1.82 1.41 3.30 3.67
Landlocked (LLDCs) 148. Central African Republic 219 0.0002 1.21 1.00 3.33 4.86
Americas (10) 899 209 0.7336 n.a. n.a. n.a. n.a.
18. United States of America 448 535 0.3659 5.08 4.82 5.76 6.60
23. Mexico 278 694 0.2274 5.80 3.55 2.14 2.78
32. Peru 143 830 0.1173 3.17 2.75 2.75 3.62
67. Costa Rica 16 269 0.0133 2.65 1.99 2.02 2.12
97. Panama 3 879 0.0032 2.07 2.02 2.01 3.04
Landlocked (LLDCs) 100. Bolivia (Plurinational State of) 3 720 0.0030 2.93 1.78 3.88 5.09
Island (SIDS) 108. Dominican Republic 2 680 0.0022 5.01 3.20 3.96 7.67
Island (SIDS) 121. Haiti 1 560 0.0013 1.76 1.38 1.52
Island (SIDS) 165. Suriname 37 0.0000 1.94 1.18 2.54
Island (SIDS) 185. Bahamas 6.0 0.0000 1.58 1.00 1.00 1.57
Asia (7) 4 712 007 3.8440 n.a. n.a. n.a. n.a.
Island 7. Philippines 2 322 831 1.8949 5.45 4.12 3.56 3.93
Island 12. Japan 996 297 0.8128 8.98 8.53 8.08 8.68
13. Thailand 962 467 0.7852 8.28 7.48 6.70 7.24
19. Türkiye 421 411 0.3438 3.47 2.86 3.03 3.61
Landlocked (LLDCs) 88. Kazakhstan 6 795 0.0055 1.89 1.31 3.73 7.06
115. Jordan 2 055 0.0017 1.89 1.06 2.00 2.00
Landlocked (LLDCs) 151. Turkmenistan 150 0.0001 1.97 1.14 2.46 4.45
Europe (4) 136 509 0.1114 n.a. n.a. n.a. n.a.
33. Greece 131 737 0.1075 4.41 3.80 3.45 3.69
Landlocked 116. Switzerland 2 048 0.0017 1.55 1.15 1.73 3.58
Landlocked (LLDCs) 119. North Macedonia 1 634 0.0013 2.64 1.72 2.13
129. Estonia 1 090 0.0009 1.97 1.67 1.82 1.85
Oceania (2) 1 476 0.0012 n.a. n.a. n.a. n.a.
Island (SIDS) 124. French Polynesia 1 371 0.0011 2.43 1.48 1.14 1.54
Island (SIDS) 155. Tonga 105 0.0001 3.13 1.01 1.01

18
Supplementary Table 2f: N-shape ENS trends, 1990–2020
N-shape: ENS increased during first and third decades whereas declined during the Aquaculture production in 2020 Effective number of species (ENS)
second one
 Upward overall trends in blue: ENS2020 > ENS1990

 Downward overall trends in red: ENS2020 < ENS1990 Share of world

Countries/territories indexed by their ranking in world aquaculture 2020 Tonnes 1990 2000 2010 2020
production (%)
LLDCs = Landlocked Developing Countries
SIDS = Small Island Developing States
Total (17) 10 458 029 8.5316 n.a. n.a. n.a. n.a.
Africa (4) 270 420 0.2206 n.a. n.a. n.a. n.a.
26. Nigeria 261 711 0.2135 4.39 6.84 5.39 5.66
Landlocked (LLDCs) 87. Rwanda 7 059 0.0058 1.31 1.32 1.13 1.40
120. Morocco 1 618 0.0013 4.01 6.85 5.68 6.51
166. Gambia 33 0.0000 1.00 2.87 2.04 2.56
Americas (5) 235 705 0.1923 n.a. n.a. n.a. n.a.
29. Colombia 179 351 0.1463 3.26 5.52 4.51 5.25
54. Guatemala 33 651 0.0275 1.86 5.12 1.18 1.98
Landlocked (LLDCs) 73. Paraguay 14 100 0.0115 2.79 4.85 2.09 2.83
83. El Salvador 8 500 0.0069 1.83 2.12 1.38 1.51
157. Uruguay 103 0.0001 1.00 1.98 1.59 3.27
Asia (4) 9 900 089 8.0764 n.a. n.a. n.a. n.a.
3. India 8 641 286 7.0495 5.51 6.15 4.55 6.58
15. Democratic People's Republic of Korea 680 300 0.5550 1.91 2.20 1.55 1.59
17. Iran (Islamic Republic of) 480 500 0.3920 3.56 4.49 4.16 4.90
Landlocked (LLDCs) 41. Uzbekistan 98 003 0.0800 2.01 3.16 2.82 4.97
Europe (3) 51 493 0.0420 n.a. n.a. n.a. n.a.
64. Ukraine 18 568 0.0151 1.64 2.57 1.89 3.08
Landlocked 65. Hungary 18 373 0.0150 2.36 3.04 2.97 3.05
72. Portugal 14 552 0.0119 3.79 6.15 5.94 6.49
Oceania (1) 322 0.0003 n.a. n.a. n.a. n.a.
Island (SIDS) 145. Fiji 322 0.0003 1.06 2.09 1.94 3.28

19
Supplementary Table 3: Categorizing ENS trends in regional aquaculture, 1990–2020
Inverted U-shape (14) Straight-up (6) U-shape (5) Inverted N-shape (4) N-shape (2) Straight-down (1)

ENS increased between 1990 and 2020

Developing regions (95.8%; U1, U1, D2) Northern America (0.51%; U2, U3, U3) Eastern Africa (0.29%; D1, U1, U2) Asia (91.61%; D2, U1, D2) Southern Asia (9.79%; U1, D3, U1)

Western Africa (0.28%; U2, D1,

Eastern Asia (61%; U3, U1, D2) Southern Europe (0.5%; U1, U1, U3) Caribbean (0.03%; D2, U1, U3) Oceania (0.2%; D2, U1, D1)
U1)

South America (2.7%; U2, D2, D1) LLDCs (0.48%; U1, U1, U2) Middle Africa (0.01%; D2, U1, U1) AU & NZ (0.18; D2, U1, D2)

Northern Africa (1.33%; U2, D2, D2) Eastern Europe (0.37%; U1, U2, U1)

Western Asia (0.49%; U1, U3, D2) Central Asia (0.09%; U2, U2, U2)

Western Europe (0.22%; U1, U1, D1) SIDS (0.05%; U1, U2, U2)

Pacific islands (0.01%; U1, U1, D2)

Southern Africa (0.01%; U3, U1, D2)

ENS declined between 1990 and 2020

Sub-Saharan Africa (0.6%; D1, D3, Africa (1.92%; D1, U2, D3) Northern Europe (1.6%; D1, D1, D1)
World excluding China (42.5%; U2, D1, D1)
U2)
South-eastern Asia (20.26%; U1, U2, D1) Central America (0.36%; D1, D1, U1)

Developed regions (4.2%; U2, D1, D1)

Americas (3.59%; U2, D2, D1)

LAC (3.08%; U2, D2, D1)

Europe (2.68%; U2, D1, D1)

Notes: This table is a rearrangement of relevant contents in Table 1. The share of each regional aquaculture in world production is presented in parentheses. The categorization of the six scenarios
(U1, U2, U3, D1, D2, and D3) is presented in Table 2.

20
Supplementary Table 4: Within-group share in species diversity in regional aquaculture
Average within-group share in overall species diversity (%)
World and 32 regional aquaculture Third decade
Overall 1990– First decade Second decade Third decade
minus
2020 1990–2000 2000–2010 2010–2020
first decade
World 67.33 66.46 67.62 67.94 1.49
World excluding China 68.79 67.70 69.95 68.93 1.23
Developed regions 65.83 64.72 66.23 66.65 1.93
Developing regions 65.84 64.22 66.22 67.12 2.90
Small Island Developing States 53.92 53.09 54.55 53.74 0.65
Landlocked developing countries 99.85 99.95 99.82 99.78 -0.17
Africa 78.22 72.95 77.41 84.69 11.74
Sub-Saharan Africa 63.44 56.23 60.94 72.85 16.62
Western Africa 99.38 99.39 99.47 99.27 -0.12
Northern Africa 98.76 98.33 99.15 98.91 0.58
Middle Africa 98.53 100.00 99.60 95.85 -4.15
Eastern Africa 44.61 37.34 43.13 52.49 15.15
Southern Africa 43.39 49.03 41.69 40.17 -8.87
Americas 62.10 59.90 64.79 62.39 2.49
Latin America and the Caribbean 57.59 54.92 60.47 58.18 3.26
Caribbean 72.93 79.49 73.60 65.52 -13.97
Northern America 58.82 59.66 60.46 56.86 -2.80
South America 55.85 49.85 60.82 58.07 8.23
Central America 44.99 49.85 43.48 40.95 -8.91
Asia 65.65 64.48 65.86 66.60 2.11
Central Asia 100.00 100.00 100.00 100.00 0.00
Western Asia 90.40 95.28 87.86 87.96 -7.32
Southern Asia 88.39 89.49 88.23 87.56 -1.93
Eastern Asia 61.85 61.08 62.25 62.20 1.12
South-eastern Asia 58.35 56.30 58.66 60.28 3.98
Europe 71.47 69.60 72.12 72.72 3.13
Eastern Europe 93.12 91.46 97.40 90.84 -0.62
Northern Europe 71.76 73.73 69.12 72.25 -1.49
Western Europe 65.85 64.66 66.03 66.73 2.07
Southern Europe 64.75 64.18 65.18 64.96 0.78
Oceania 46.77 47.97 47.66 44.55 -3.42
Australia and New Zealand 54.62 60.63 53.71 49.42 -11.21
Pacific islands 23.09 12.87 24.67 32.37 19.50
Notes: See Table 1 for the definitions of the 32 regional aquaculture.

21
Supplementary Table 5: National aquaculture in 2020 where within-group species diversity was the
sole source of species diversity
Countries or territories with within-group share = 100 in 2020
Aquaculture production Single species
(indexed by ranking in world production 2020) ENS (=
group farmed
AF = Africa; AM = Americas; AS = Asia; ER = Europe; ON = Oceania Share of WGENS)
Tonnes
LLDCs = Landlocked Developing Countries world (%)
Overall 67 countries/territories 653 103 0.5328 n.a. n.a.
AF Landlocked LLDCs 35. Uganda 123 897 0.1011 1.888 Finfish
AS Landlocked LLDCs 41. Uzbekistan 98 003 0.0800 4.967 Finfish
AS Landlocked LLDCs 43. Nepal 76 822 0.0627 6.982 Finfish
AF Landlocked LLDCs 48. Zambia 45 670 0.0373 3.212 Finfish
AS 59. Iraq 22 704 0.0185 1.708 Finfish
ER Landlocked 62. Czechia 20 401 0.0166 2.085 Finfish
ER 63. Malta 19 829 0.0162 1.713 Finfish
ER 64. Ukraine 18 568 0.0151 3.078 Finfish
ER Landlocked 65. Hungary 18 373 0.0150 3.053 Finfish
AS Landlocked LLDCs 66. Armenia 18 350 0.0150 2.770 Finfish
AF Landlocked LLDCs 68. Zimbabwe 15 425 0.0126 1.027 Finfish
ER 69. Finland 15 053 0.0123 1.264 Finfish
AS 71. Israel 14 700 0.0120 5.804 Finfish
AM Landlocked LLDCs 73. Paraguay 14 100 0.0115 2.829 Finfish
ER 76. Romania 12 200 0.0100 5.934 Finfish
AF 78. Sudan 9 850 0.0080 1.656 Finfish
AF Landlocked LLDCs 80. Malawi 9 393 0.0077 3.560 Finfish
ER Landlocked 82. Belarus 9 266 0.0076 2.479 Finfish
AS Landlocked LLDCs 84. Afghanistan 8 050 0.0066 1.038 Finfish
AF Landlocked LLDCs 85. Mali 7 686 0.0063 1.778 Finfish
AS Landlocked LLDCs 88. Kazakhstan 6 795 0.0055 7.064 Finfish
ER Landlocked 90. Serbia 6 010 0.0049 2.301 Finfish
AF 94. Côte d'Ivoire 4 620 0.0038 2.485 Finfish
ER 95. Austria 4 527 0.0037 7.018 Finfish
ER 96. Lithuania 4 477 0.0037 2.847 Finfish
ER 99. Bosnia and Herzegovina 3 776 0.0031 1.893 Finfish
AM Landlocked LLDCs 100. Bolivia (Plurinational State of) 3 720 0.0030 5.088 Finfish
AF 101. Democratic Republic of the Congo 3 590 0.0029 1.153 Finfish
AF 102. Cameroon 3 556 0.0029 2.123 Finfish
AF 107. Benin 2 987 0.0024 1.998 Finfish
AF Landlocked LLDCs 109. Lesotho 2 600 0.0021 1.003 Finfish
AS Landlocked LLDCs 110. Kyrgyzstan 2 550 0.0021 3.978 Finfish
AS 111. Syrian Arab Republic 2 300 0.0019 3.672 Finfish
ER Landlocked 112. Slovakia 2 296 0.0019 3.821 Finfish
AF 114. Angola 2 062 0.0017 1.078 Finfish
AS 115. Jordan 2 055 0.0017 2.000 Finfish
AS 117. Georgia 2 027 0.0017 3.337 Finfish
ER Landlocked LLDCs 119. North Macedonia 1 634 0.0013 2.132 Finfish
AM 121. Haiti 1 560 0.0013 1.522 Finfish
AF Landlocked LLDCs 123. Burundi 1 450 0.0012 1.298 Finfish
AS 125. Oman 1 307 0.0011 1.624 Finfish
AF 126. Guinea 1 135 0.0009 3.711 Finfish
ER 127. Channel Islands 1 107 0.0009 1.346 Molluscs
AF 131. Congo 900 0.0007 1.600 Finfish
AS Landlocked LLDCs 135. Tajikistan 775 0.0006 2.943 Finfish
AF 136. Togo 730 0.0006 1.305 Finfish
ER 137. Latvia 717 0.0006 2.511 Finfish
AF Landlocked LLDCs 139. Niger 649 0.0005 1.924 Finfish

22
 Countries or territories with within-group share = 100 in 2020
Aquaculture production Single species
(indexed by ranking in world production 2020) ENS (=
group farmed
AF = Africa; AM = Americas; AS = Asia; ER = Europe; ON = Oceania Share of WGENS)
Tonnes
LLDCs = Landlocked Developing Countries world (%)
AS 140. State of Palestine 600 0.0005 1.697 Finfish
AF Landlocked LLDCs 142. Ethiopia 534 0.0004 2.172 Finfish
AS Landlocked LLDCs 143. Azerbaijan 533 0.0004 4.739 Finfish
AF 147. Liberia 255 0.0002 1.485 Finfish
AS Landlocked LLDCs 150. Bhutan 181 0.0001 6.315 Finfish
AS Landlocked LLDCs 151. Turkmenistan 150 0.0001 4.452 Finfish
AF Landlocked LLDCs 152. Botswana 146 0.0001 1.760 Finfish
ON 155. Tonga 105 0.0001 1.014 Algae
AM 157. Uruguay 103 0.0001 3.275 Finfish
AF 160. Sierra Leone 85 0.0001 1.251 Finfish
AF 161. Gabon 60 0.0000 1.332 Finfish
AF Landlocked LLDCs 167. South Sudan 30 0.0000 1.278 Finfish
ON 172. Palau 22 0.0000 2.533 Finfish
AF 177. Equatorial Guinea 15 0.0000 2.693 Finfish
AF 181. Réunion 10 0.0000 2.000 Finfish
AM 184. United States Virgin Islands 8 0.0000 2.828 Finfish
AS 186. Bahrain 6 0.0000 1.356 Finfish
AM 187. Trinidad and Tobago 4 0.0000 1.731 Finfish
AM 191. French Guiana 2 0.0000 2.000 Finfish

23
Supplementary Table 6: Species diversity in landlocked aquaculture, 2020
Landlocked aquaculture in 2020 Aquaculture
Diversity measures
(indexed by their ranking in world production 2020) production, 2020 Within-
AF = Africa; AM = Americas; AS = Asia; ER = Europe; group
Share of
ON = Oceania Tonnes ENS WGENS ENSG share (%)
world (%)
LLDCs = Landlocked Developing Countries
Overall 37 landlocked aquaculture 648 292 0.5289 n.a. n.a. n.a n.a.
AS LLDCs 34. Lao People's Democratic Republic 130 020 0.1061 10.33 10.26 1.01 99.7
AF LLDCs 35. Uganda 123 897 0.1011 1.89 1.89 1.00 100.0
AS LLDCs 41. Uzbekistan 98 003 0.0800 4.97 4.97 1.00 100.0
AS LLDCs 43. Nepal 76 822 0.0627 6.98 6.98 1.00 100.0
AF LLDCs 48. Zambia 45 670 0.0373 3.21 3.21 1.00 100.0
ER 62. Czechia 20 401 0.0166 2.09 2.09 1.00 100.0
ER 65. Hungary 18 373 0.0150 3.05 3.05 1.00 100.0
AS LLDCs 66. Armenia 18 350 0.0150 2.77 2.77 1.00 100.0
AF LLDCs 68. Zimbabwe 15 425 0.0126 1.03 1.03 1.00 100.0
AM LLDCs 73. Paraguay 14 100 0.0115 2.83 2.83 1.00 100.0
ER LLDCs 75. Republic of Moldova 12 600 0.0103 4.17 4.16 1.00 99.8
AF LLDCs 80. Malawi 9 393 0.0077 3.56 3.56 1.00 100.0
ER 82. Belarus 9 266 0.0076 2.48 2.48 1.00 100.0
AS LLDCs 84. Afghanistan 8 050 0.0066 1.04 1.04 1.00 100.0
AF LLDCs 85. Mali 7 686 0.0063 1.78 1.78 1.00 100.0
AF LLDCs 87. Rwanda 7 059 0.0058 1.40 1.39 1.00 98.8
AS LLDCs 88. Kazakhstan 6 795 0.0055 7.06 7.06 1.00 100.0
ER 90. Serbia 6 010 0.0049 2.30 2.30 1.00 100.0
AM LLDCs 100. Bolivia (Plurinational State of) 3 720 0.0030 5.09 5.09 1.00 100.0
AF LLDCs 109. Lesotho 2 600 0.0021 1.003 1.003 1.00 100.0
AS LLDCs 110. Kyrgyzstan 2 550 0.0021 3.98 3.98 1.00 100.0
ER 112. Slovakia 2 296 0.0019 3.82 3.82 1.00 100.0
ER 116. Switzerland 2 048 0.0017 3.58 3.32 1.08 94.0
ER LLDCs 119. North Macedonia 1 634 0.0013 2.13 2.13 1.00 100.0
AF LLDCs 123. Burundi 1 450 0.0012 1.30 1.30 1.00 100.0
AS LLDCs 135. Tajikistan 775 0.0006 2.94 2.94 1.00 100.0
AF LLDCs 138. Burkina Faso 651 0.0005 2.33 2.22 1.05 93.8
AF LLDCs 139. Niger 649 0.0005 1.92 1.92 1.00 100.0
AF LLDCs 142. Ethiopia 534 0.0004 2.17 2.17 1.00 100.0
AS LLDCs 143. Azerbaijan 533 0.0004 4.74 4.74 1.00 100.0
AF LLDCs 148. Central African Republic 219 0.0002 4.86 2.84 1.71 66.0
AS LLDCs 150. Bhutan 181 0.0001 6.31 6.31 1.00 100.0
AS LLDCs 151. Turkmenistan 150 0.0001 4.45 4.45 1.00 100.0
AF LLDCs 152. Botswana 146 0.0001 1.76 1.76 1.00 100.0
AF LLDCs 156. Chad 105 0.0001 3.42 2.10 1.63 60.4
AF LLDCs 158. Eswatini 100 0.0001 1.00 1.00 1.00 n.a.
AF LLDCs 167. South Sudan 30 0.00002 1.28 1.28 1.00 100.0

24
Supplementary Table 7: National aquaculture in 2020 with between-group diversity greater than within-group diversity (ENSG > WGNES)
55 countries/territories with ENSG > WGENS in 2020 Between-group
Aquaculture production Overall Within-group diversity
(indexed by ranking in world production 2020) diversity
Species
AF = Africa; AM = Americas; AS = Asia; ER = Europe; ON = Oceania Within- Between-
Share of diversity
SIDS = Small Island Developing States Tonnes WGENS group ENSG group
world (%) (ENS)
share (%) share (%)
Overall 55 countries/territories 9 597 813 7.8298 n.a. n.a. n.a. n.a. n.a.
AS Island 7. Philippines 2 322 831 1.8949 3.934 1.632 36 2.411 64
AM 9. Chile 1 505 486 1.2282 3.385 1.770 47 1.913 53
AS Island 12. Japan 996 297 0.8128 8.684 2.776 47 3.128 53
AS 13. Thailand 962 467 0.7852 7.237 2.653 49 2.728 51
AM 14. Ecuador 774 569 0.6319 1.109 1.013 12 1.095 88
AS 15. Democratic People's Republic of Korea 680 300 0.5550 1.593 1.056 12 1.508 88
AM 18. United States of America 448 535 0.3659 6.604 2.353 45 2.807 55
AS 21. Malaysia 400 017 0.3263 8.467 2.719 47 3.114 53
AM 23. Mexico 278 694 0.2274 2.776 1.356 30 2.048 70
ER 25. Spain 276 571 0.2256 2.966 1.607 44 1.846 56
AM 32. Peru 143 830 0.1173 3.615 1.235 16 2.928 84
ON Island 37. New Zealand 118 582 0.0967 1.573 1.067 14 1.474 86
AF 38. United Republic of Tanzania 108 568 0.0886 1.705 1.091 16 1.563 84
AS 40. Saudi Arabia 99 907 0.0815 3.952 1.980 50 1.996 50
ER Island 42. Faeroe Islands 89 055 0.0727 1.010 1.001 9 1.009 91
AM 44. Honduras 71 151 0.0580 1.992 1.000 - 1.992 100
AM 46. Venezuela (Bolivarian Republic of) 53 609 0.0437 1.853 1.047 7 1.770 93
ER 49. Denmark 42 629 0.0348 2.237 1.464 47 1.528 53
ER 52. Netherlands 39 940 0.0326 2.179 1.463 49 1.490 51
ER Island 53. Ireland 37 751 0.0308 3.285 1.688 44 1.947 56
AM 54. Guatemala 33 651 0.0275 1.981 1.016 2 1.951 98
AM 56. Nicaragua 29 410 0.0240 1.003 1.000 - 1.003 100
AM 67. Costa Rica 16 269 0.0133 2.116 1.418 47 1.493 53
AF Island 74. Madagascar 13 550 0.0111 2.114 1.004 1 2.105 99
ER 77. Sweden 12 090 0.0099 2.237 1.375 40 1.627 60
AF 79. South Africa 9 753 0.0080 4.660 1.701 35 2.740 65
AM 83. El Salvador 8 500 0.0069 1.506 1.008 2 1.494 98
ON SIDS Island 89. Papua New Guinea 6 102 0.0050 2.302 1.252 27 1.840 73
ON SIDS Island 91. Solomon Islands 5 504 0.0045 1.006 1.000 - 1.006 100
AM 97. Panama 3 879 0.0032 3.038 1.535 39 1.979 61
ON SIDS Island 122. New Caledonia 1 478 0.0012 1.031 1.004 14 1.026 86
ON SIDS Island 124. French Polynesia 1 371 0.0011 1.539 1.043 10 1.476 90
AF 128. Senegal 1 100 0.0009 3.665 1.837 47 1.995 53

25
 55 countries/territories with ENSG > WGENS in 2020 Between-group
Aquaculture production Overall Within-group diversity
(indexed by ranking in world production 2020) diversity
Species
AF = Africa; AM = Americas; AS = Asia; ER = Europe; ON = Oceania Within- Between-
Share of diversity
SIDS = Small Island Developing States Tonnes WGENS group ENSG group
world (%) (ENS)
share (%) share (%)
AM SIDS Island 130. Jamaica 918 0.0007 1.048 1.018 38 1.029 62
ER 132. Montenegro 885 0.0007 2.869 1.589 44 1.806 56
AS SIDS Island 133. Timor-Leste 835 0.0007 1.732 1.105 18 1.568 82
AM SIDS 141. Belize 565 0.0005 1.474 1.000 - 1.474 100
ON SIDS Island 145. Fiji 322 0.0003 3.279 1.170 13 2.802 87
AF 146. Namibia 321 0.0003 1.609 1.124 25 1.431 75
AM SIDS 153. Guyana 138 0.0001 1.999 1.165 22 1.716 78
AM SIDS Island 159. Saint Lucia 92 0.0001 1.441 1.000 - 1.441 100
AM SIDS Island 163. Martinique 42 0.0000 1.680 1.115 21 1.507 79
ON SIDS Island 164. Northern Mariana Islands 41 0.0000 2.205 1.103 12 1.999 88
AM SIDS 165. Suriname 37 0.0000 2.543 1.419 37 1.792 63
AF 166. Gambia 33 0.0000 2.563 1.311 29 1.955 71
AM SIDS Island 168. Barbados 26 0.0000 1.101 1.000 - 1.101 100
AS 169. Qatar 23 0.0000 1.164 1.000 - 1.164 100
AM SIDS Island 170. Grenada 23 0.0000 1.112 1.000 - 1.112 100
AM SIDS Island 174. Antigua and Barbuda 20 0.0000 2.000 1.000 - 2.000 100
AM SIDS Island 176. Puerto Rico 18 0.0000 2.648 1.401 35 1.890 65
ON SIDS Island 179. Samoa 10 0.0000 1.011 1.001 7 1.011 93
ON SIDS Island 182. Marshall Islands 10 0.0000 1.579 1.000 - 1.579 100
AM SIDS Island 185. Bahamas 6 0.0000 1.569 1.000 - 1.569 100
ON SIDS Island 188. Vanuatu 3 0.0000 1.989 1.226 30 1.622 70
AM SIDS Island 190. Saint Kitts and Nevis 2 0.0000 2.000 1.000 - 2.000 100

26
Supplementary Table 8: Species diversity in island aquaculture, 2020
Island aquaculture in 2020 Aquaculture production, 2020 Diversity measures
Between-
(indexed by their ranking in world production 2020)
Share of group
AF = Africa; AM = Americas; AS = Asia; ER = Europe; ON = Oceania Tonnes ENS WGENS ENSG
world (%) share (%)
SIDS = Small Island Developing States
Overall 60 island aquaculture 19 092 507 15.5755 n.a. n.a. n.a n.a.
28 island aquaculture with ENSG > WGENS (highlighted in red) 3 595 628 2.9333 n.a. n.a. n.a n.a.
22 island aquaculture with ENSG < WGENS 15 496 796 12.6422 n.a. n.a. n.a n.a.
10 island aquaculture with no species diversity (ENS = 1; italic) 83 0.0001 n.a. n.a. n.a n.a.
AS Island 2. Indonesia 14 845 014 12.1105 5.63 2.47 2.28 48
AS Island 7. Philippines 2 322 831 1.8949 3.93 1.63 2.41 64
AS Island 12. Japan 996 297 0.8128 8.68 2.78 3.13 53
AS Island 24. Taiwan Province of China 278 503 0.2272 11.04 4.65 2.38 36
ER Island 27. United Kingdom 221 000 0.1803 1.69 1.34 1.26 45
ON Island 37. New Zealand 118 582 0.0967 1.57 1.07 1.47 86
ER Island 42. Faeroe Islands 89 055 0.0727 1.01 1.00 1.01 91
AS Island 50. Sri Lanka 42 119 0.0344 5.17 2.93 1.77 35
ER Island 51. Iceland 40 595 0.0331 1.65 1.65 1.00 0
ER Island 53. Ireland 37 751 0.0308 3.29 1.69 1.95 56
AM Island SIDS 57. Cuba 26 200 0.0214 3.97 2.09 1.90 47
ER Island 63. Malta 19 829 0.0162 1.71 1.71 1.00 0
AF Island 74. Madagascar 13 550 0.0111 2.11 1.00 2.10 99
AS Island 86. Cyprus 7 343 0.0060 2.05 2.02 1.02 2
ON Island SIDS 89. Papua New Guinea 6 102 0.0050 2.30 1.25 1.84 73
ON Island SIDS 91. Solomon Islands 5 504 0.0045 1.01 1.00 1.01 100
AS Island SIDS 93. Singapore 4 829 0.0039 11.67 6.06 1.93 27
AS Island 103. Brunei Darussalam 3 500 0.0029 2.47 1.68 1.47 43
AF Island SIDS 104. Mauritius 3 298 0.0027 1.15 1.11 1.04 26
AM Island SIDS 108. Dominican Republic 2 680 0.0022 7.67 5.26 1.46 18
AM Island SIDS 121. Haiti 1 560 0.0013 1.52 1.52 1.00 0
ON Island SIDS 122. New Caledonia 1 478 0.0012 1.03 1.00 1.03 86
ON Island SIDS 124. French Polynesia 1 371 0.0011 1.54 1.04 1.48 90
AM Island SIDS 130. Jamaica 918 0.0007 1.05 1.02 1.03 62
AS Island SIDS 133. Timor-Leste 835 0.0007 1.73 1.10 1.57 82
AM SIDS 141. Belize 565 0.0005 1.47 1.00 1.47 100
ON Island SIDS 145. Fiji 322 0.0003 3.28 1.17 2.80 87
AM SIDS 153. Guyana 138 0.0001 2.00 1.17 1.72 78
ON Island SIDS 154. Guam 118 0.0001 2.79 1.94 1.44 36

27
 Island aquaculture in 2020 Aquaculture production, 2020 Diversity measures
Between-
(indexed by their ranking in world production 2020)
Share of group
AF = Africa; AM = Americas; AS = Asia; ER = Europe; ON = Oceania Tonnes ENS WGENS ENSG
world (%) share (%)
SIDS = Small Island Developing States
ON Island SIDS 155. Tonga 105 0.0001 1.01 1.01 1.00 0
AM Island SIDS 159. Saint Lucia 92 0.0001 1.44 1.00 1.44 100
AM Island SIDS 162. Guadeloupe 45 0.0000 1.53 1.27 1.20 43
AM Island SIDS 163. Martinique 42 0.0000 1.68 1.11 1.51 79
ON Island SIDS 164. Northern Mariana Islands 41 0.0000 2.21 1.10 2.00 88
AM SIDS 165. Suriname 37 0.0000 2.54 1.42 1.79 63
AM Island SIDS 168. Barbados 26 0.0000 1.10 1.00 1.10 100
AM Island SIDS 170. Grenada 23 0.0000 1.11 1.00 1.11 100
AF Island SIDS 171. Cabo Verde 22 0.0000 1.00 1.00 1.00
ON Island SIDS 172. Palau 22 0.0000 2.53 2.53 1.00 0
ON Island SIDS 173. American Samoa 20 0.0000 1.00 1.00 1.00
AM Island SIDS 174. Antigua and Barbuda 20 0.0000 2.00 1.00 2.00 100
AF Island 175. Mayotte 20 0.0000 1.00 1.00 1.00
AM Island SIDS 176. Puerto Rico 18 0.0000 2.65 1.40 1.89 65
AM Island SIDS 178. Saint Vincent and the Grenadines 13 0.0000 1.00 1.00 1.00
ON Island SIDS 179. Samoa 10 0.0000 1.01 1.00 1.01 93
AF Island 181. Réunion 10 0.0000 2.00 2.00 1.00 0
ON Island SIDS 182. Marshall Islands 10 0.0000 1.58 1.00 1.58 100
ON Island SIDS 183. Cook Islands 8.5 0.0000 2.92 2.03 1.44 34
AM Island SIDS 184. United States Virgin Islands 8.0 0.0000 2.83 2.83 1.00 0
AM Island SIDS 185. Bahamas 6.0 0.0000 1.57 1.00 1.57 100
AS Island SIDS 186. Bahrain 5.5 0.0000 1.36 1.36 1.00 0
AM Island SIDS 187. Trinidad and Tobago 4.2 0.0000 1.73 1.73 1.00 0
ON Island SIDS 188. Vanuatu 3.1 0.0000 1.99 1.23 1.62 70
ON Island SIDS 189. Tuvalu 2.5 0.0000 1.00 1.00 1.00
ON Island SIDS 191. Kiribati 2.0 0.0000 1.00 1.00 1.00
AM Island SIDS 192. Saint Kitts and Nevis 2.0 0.0000 2.00 1.00 2.00 100
AM Island SIDS 193. Aruba 1.5 0.0000 1.00 1.00 1.00
AM Island SIDS 194. British Virgin Islands 1.0 0.0000 1.00 1.00 1.00
ON Island SIDS 195. Nauru 0.5 0.0000 1.00 1.00 1.00
AM Island 196. Bonaire, Sint Eustatius and Saba 0.2 0.0000 1.00 1.00 1.00

28
Supplementary Table 9a: National aquaculture whose ENS trend during 1990–2020 belonged to the category of U3
16 national aquaculture whose Aquaculture Diversity change between 1990 and
Diversity in 1990 Diversity in 2020
diversity change between 1990 production, 2020 2020
and 2020 belonged to the category Share of
of U3 = ENS increase (only ENSG Tonnes world ENS WGENS ENSG ENS WGENS ENSG ΔENS ΔWGENS ΔENSG
up); see Figure 2a. (%)
AS India 8 641 286 7.0495 5.51 4.74 1.16 6.58 4.59 1.43 1.07 -0.15 0.27
AS Iran (Islamic Republic of) 480 500 0.3920 3.56 3.56 1.00 4.90 3.53 1.39 1.33 -0.04 0.39
AS Türkiye 421 411 0.3438 3.47 3.47 1.00 3.61 3.42 1.06 0.14 -0.05 0.06
AM Peru 143 830 0.1173 3.17 1.46 2.17 3.62 1.23 2.93 0.44 -0.23 0.75
AM Honduras 71 151 0.0580 1.20 1.02 1.18 1.99 1.00 1.99 0.80 -0.02 0.81
AM Guatemala 33 651 0.0275 1.86 1.15 1.61 1.98 1.02 1.95 0.13 -0.13 0.34
AF Madagascar 13 550 0.0111 1.75 1.50 1.17 2.11 1.00 2.10 0.37 -0.49 0.94
ER Sweden 12 090 0.0099 2.14 1.45 1.47 2.24 1.38 1.63 0.09 -0.08 0.15
AF South Africa 9 753 0.0080 4.39 2.13 2.06 4.66 1.70 2.74 0.27 -0.43 0.68
ON Papua New Guinea 6 102 0.0050 1.82 1.82 1.00 2.30 1.25 1.84 0.48 -0.57 0.84
ON Solomon Islands 5 504 0.0045 1.00 1.00 1.00 1.01 1.00 1.01 0.01 0.00 0.01
AM Panama 3 879 0.0032 2.07 1.54 1.35 3.04 1.54 1.98 0.96 -0.01 0.63
AM Belize 565 0.0005 1.00 1.00 1.00 1.47 1.00 1.47 0.47 0.00 0.47
ON Guam 118 0.0001 2.50 2.50 1.00 2.79 1.94 1.44 0.29 -0.56 0.44
AM Saint Lucia 92 0.0001 1.00 1.00 1.00 1.44 1.00 1.44 0.44 0.00 0.44
AM Saint Kitts and Nevis 2 0.0000 1.00 1.00 1.00 2.00 1.00 2.00 1.00 0.00 1.00

Supplementary Table 9b: National aquaculture whose ENS trend during 1990–2020 belonged to the category of D3
7 national aquaculture whose Aquaculture Diversity change between 1990 and
Diversity in 1990 Diversity in 2020
diversity change between 1990 production, 2020 2020
and 2020 belonged to the category Share of
of D3 = ENS decrease (only ENSG Tonnes world ENS WGENS ENSG ENS WGENS ENSG ΔENS ΔWGENS ΔENSG
down); see Figure 2a. (%)
AM Chile 1 505 486 1.2282 3.92 1.66 2.36 3.39 1.77 1.91 -0.53 0.11 -0.44
AS Taiwan Province of China 278 503 0.2272 11.40 4.63 2.46 11.04 4.65 2.38 -0.36 0.02 -0.09
ER Greece 131 737 0.1075 4.41 2.26 1.95 3.69 2.40 1.54 -0.71 0.14 -0.41
AM Costa Rica 16 269 0.0133 2.65 1.27 2.08 2.12 1.42 1.49 -0.53 0.15 -0.59
ON New Caledonia 1 478 0.0012 1.51 1.00 1.51 1.03 1.00 1.03 -0.48 0.00 -0.48
AM Jamaica 918 0.0007 1.08 1.01 1.06 1.05 1.02 1.03 -0.03 0.01 -0.03
AM Martinique 42 0.0000 2.17 1.09 1.98 1.68 1.11 1.51 -0.49 0.02 -0.48

29
Supplementary Table 10: National aquaculture with no species diversity (ENS = 1)
All national aquaculture National aquaculture with no species diversity in the species group (ENS = 1)
Number of
Species groups Prevalence Aquaculture production
countries that Total production
farmed the (tonnes) Number of
Share of total (%) Tonnes Share of total (%)
species group countries
Year 1990
All species 158 17 280 438 32 20.25 6 350 0.04
Finfish 143 8 676 399 31 21.68 148 256 1.71
Algae 17 4 195 304 11 64.71 171 012 4.08
Molluscs 48 3 608 731 19 39.58 67 538 1.87
Crustaceans 61 754 794 30 49.18 25 603 3.39
MAA 11 45 210 8 72.73 4 581 10.13
Year 2000
All species 173 43 016 624 27 15.61 7 879 0.02
Finfish 162 20 813 371 35 21.60 121 122 0.58
Algae 23 10 595 565 14 60.87 123 781 1.17
Molluscs 61 9 757 564 25 40.98 111 067 1.14
Crustaceans 69 1 693 442 37 53.62 53 938 3.19
MAA 12 156 682 6 50.00 1 160 0.74
Year 2010
All species 189 77 994 008 13 6.88 2 560 0.00
Finfish 182 37 752 735 25 13.74 36 450 0.10
Algae 34 20 174 325 20 58.82 27 641 0.14
Molluscs 62 13 728 260 21 33.87 65 359 0.48
Crustaceans 79 5 481 791 45 56.96 316 873 5.78
MAA 26 856 897 14 53.85 30 105 3.51
Year 2020
All species 196 122 580 187 15 7.65 4 014 0.00
Finfish 191 57 461 094 31 16.23 147 412 0.26
Algae 49 35 077 578 30 61.22 233 528 0.67
Molluscs 63 17 740 526 17 26.98 62 478 0.35
Crustaceans 79 11 237 016 45 56.96 956 144 8.51
MAA 26 1 063 972 15 57.69 4 599 0.43

30
Supplementary Table 11: National aquaculture with extraordinarily high diversity (aka outliers)
All species Finfish Crustaceans Molluscs MAA Algae
Country or territory
ENS ENSG WGENS ENS ENS ENS ENS ENS
(11; 5.6 %) (8; 4 %) (11; 5.6 %) (7; 3.6 %) (8; 10 %) (3; 4.8 %) (5; 18.5 %) (8; 15.7 %)
1. China 28.00 3.77 7.43 15.07 5.14 5.82 5.34 3.92
5. Bangladesh 14.30 11.48 12.25 3.97
94. Singapore 11.67 6.06 8.40 4.76
24. Taiwan Province of China 11.04 2.40 1.97
34. Lao People's Democratic
10.33 10.26 10.28
Republic
22. Russian Federation 10.03 2.01
12. Japan 8.68 3.13 2.39
21. Malaysia 8.47 3.11 11.45
4. Viet Nam 8.24 3.34 2.01
20. Cambodia 7.70 6.19
109. Dominican Republic 7.67
32. Peru 2.93
18. United States of America 2.81
146. Fiji 2.80 2.23
80. South Africa 2.74
13. Thailand 2.73
89. Kazakhstan 7.06 7.06
96. Austria 7.02
43. Nepal 6.98
151. Bhutan 6.31
77. Romania 5.93
72. Israel 5.80
6. Republic of Korea 7.40 3.39
7. Philippines 3.05
190. Vanuatu 2.96
38. Australia 3.90
73. Portugal 3.89
15. Democratic People's Republic
2.42
of Korea
28. France 2.50
42. Faeroe Islands 2.18
25. Spain 1.90
10. Norway 1.78
58. Tunisia 1.70
Notes: ENS = effective number of species. ENSG = effective number of species group. WGENS = within-group ENS. MAA = Miscellaneous
aquatic animals and animal products. In parentheses, the first number represents the number of “outliers” (i.e. countries with extraordinarily high
species diversity); and the second number represents the prevalence of outliers (i.e. the percentage of outliers in the total number of countries with
aquaculture production of the species group).

31
Supplementary Table 12: Cross-sectional correlation between aquaculture production and species diversity in national aquaculture
Year ENS WGENS ENSG ENS (finfish) ENS (crustaceans) ENS (molluscs) ENS (MAA) ENS (Algae)
1990 0.4976*** (158) 0.229*** (158) 0.356*** (158) 0.2907*** (143) 0.2763** (61) 0.6674*** (48) 0.7125** (11) 0.4914** (17)
1991 0.4707*** (158) 0.1968** (158) 0.3662*** (158) 0.2933*** (145) 0.2669** (61) 0.6806*** (49) 0.8601*** (10) 0.4346* (16)
1992 0.4438*** (161) 0.1741** (161) 0.3433*** (161) 0.2837*** (150) 0.2348* (67) 0.6698*** (50) 0.5883* (10) 0.4091* (18)
1993 0.4173*** (161) 0.1614** (161) 0.3261*** (161) 0.2614*** (150) 0.2995** (66) 0.6513*** (51) 0.5535* (11) 0.3851 (18)
1994 0.4168*** (162) 0.158** (162) 0.3277*** (162) 0.2529*** (149) 0.3131*** (68) 0.5516*** (50) 0.4875 (11) 0.4557* (18)
1995 0.4344*** (164) 0.1737** (164) 0.3156*** (164) 0.2682*** (152) 0.3863*** (68) 0.582*** (53) 0.5479* (11) 0.4251* (20)
1996 0.4172*** (168) 0.1679** (168) 0.3015*** (168) 0.266*** (157) 0.4942*** (68) 0.5819*** (55) 0.4738 (11) 0.2896 (20)
1997 0.4324*** (167) 0.1737** (167) 0.305*** (167) 0.277*** (156) 0.5219*** (69) 0.6208*** (56) 0.6097** (12) 0.3162 (19)
1998 0.4356*** (169) 0.1754** (169) 0.3113*** (169) 0.2753*** (156) 0.4182*** (69) 0.5828*** (57) 0.6639** (12) 0.445** (20)
1999 0.4489*** (170) 0.1825** (170) 0.3047*** (170) 0.2945*** (157) 0.5082*** (67) 0.58*** (59) 0.6037** (14) 0.3932* (22)
2000 0.4498*** (173) 0.1825** (173) 0.303*** (173) 0.2792*** (162) 0.6278*** (69) 0.5044*** (61) 0.5178* (12) 0.4652** (23)
2001 0.4745*** (173) 0.1898** (173) 0.2994*** (173) 0.2905*** (162) 0.6901*** (66) 0.5182*** (60) 0.4759 (13) 0.4774** (26)
2002 0.4809*** (173) 0.1887** (173) 0.2964*** (173) 0.2877*** (163) 0.7835*** (68) 0.5842*** (58) 0.52* (14) 0.5077*** (26)
2003 0.6162*** (177) 0.2722*** (177) 0.3191*** (177) 0.3801*** (167) 0.8883*** (71) 0.6322*** (58) 0.9403*** (13) 0.7433*** (25)
2004 0.6332*** (180) 0.2847*** (180) 0.3327*** (180) 0.37*** (169) 0.8727*** (71) 0.6178*** (60) 0.9385*** (16) 0.8147*** (25)
2005 0.648*** (183) 0.2748*** (183) 0.3354*** (183) 0.3864*** (171) 0.8262*** (78) 0.5955*** (61) 0.9003*** (16) 0.7543*** (27)
2006 0.6767*** (189) 0.3044*** (189) 0.3404*** (189) 0.4285*** (177) 0.8183*** (80) 0.6024*** (60) 0.9139*** (18) 0.7789*** (26)
2007 0.6904*** (188) 0.303*** (188) 0.3498*** (188) 0.4376*** (180) 0.8393*** (84) 0.6088*** (60) 0.9505*** (17) 0.7908*** (27)
2008 0.6958*** (187) 0.2928*** (187) 0.3417*** (187) 0.4421*** (179) 0.8198*** (84) 0.6378*** (60) 0.9596*** (21) 0.7536*** (30)
2009 0.6758*** (189) 0.277*** (189) 0.3428*** (189) 0.4291*** (181) 0.7527*** (83) 0.6422*** (61) 0.942*** (24) 0.7523*** (30)
2010 0.679*** (189) 0.2637*** (189) 0.3624*** (189) 0.4284*** (182) 0.7559*** (79) 0.6349*** (62) 0.8913*** (26) 0.6908*** (34)
2011 0.6822*** (192) 0.2647*** (192) 0.3452*** (192) 0.4382*** (187) 0.7056*** (81) 0.6196*** (64) 0.826*** (26) 0.7811*** (36)
2012 0.6875*** (192) 0.267*** (192) 0.3515*** (192) 0.4477*** (187) 0.7201*** (80) 0.618*** (63) 0.804*** (25) 0.7587*** (37)
2013 0.665*** (191) 0.2698*** (191) 0.3502*** (191) 0.462*** (186) 0.8148*** (79) 0.6345*** (63) 0.8109*** (26) 0.7261*** (40)
2014 0.641*** (194) 0.2454*** (194) 0.3452*** (194) 0.4572*** (188) 0.7616*** (77) 0.6167*** (63) 0.8794*** (29) 0.5912*** (43)
2015 0.6369*** (198) 0.2436*** (198) 0.346*** (198) 0.4556*** (191) 0.6904*** (77) 0.5659*** (63) 0.8782*** (27) 0.4798*** (51)
2016 0.6459*** (198) 0.2505*** (198) 0.3496*** (198) 0.465*** (193) 0.6608*** (80) 0.6036*** (62) 0.8689*** (28) 0.5318*** (51)
2017 0.6477*** (199) 0.2456*** (199) 0.3653*** (199) 0.4659*** (193) 0.6058*** (79) 0.6309*** (63) 0.8999*** (30) 0.5966*** (52)
2018 0.6454*** (198) 0.2427*** (198) 0.3578*** (198) 0.4643*** (192) 0.5955*** (78) 0.5903*** (65) 0.8942*** (28) 0.5351*** (52)
2019 0.6405*** (196) 0.242*** (196) 0.3693*** (196) 0.4593*** (190) 0.5676*** (83) 0.5847*** (62) 0.8985*** (27) 0.5494*** (48)
2020 0.6454*** (196) 0.2388*** (196) 0.3759*** (196) 0.4677*** (191) 0.5519*** (79) 0.5815*** (63) 0.8984*** (26) 0.6318*** (49)
Minimum 0.4168 0.1580 0.2964 0.2529 0.2348 0.5044 0.4738 0.2896
Median 0.6351 0.2424 0.3422 0.3832 0.6443 0.6062 0.8431 0.5334
Maximum 0.6958 0.3044 0.3759 0.4677 0.8883 0.6806 0.9596 0.8147
Notes: ***p < 0.01; **p < 0.05; *p < 0.1. The number in parentheses represents the number of national aquaculture available to calculate the
cross-sectional correlation in each year.

32
Supplementary Table 13: Temporal correlation between a country’s production and diversity during 2010–2020
Correlation coefficient (Pearson’s r) Share of all national aquaculture (count) or world production (prn)
Temporal correlation between
r not statistically Positive r, statistically Negative r, statistically
aquaculture production and eight Number of
Min r Median r Mean r Max r significant significant significant
diversity measures countries
Count (%) Prn (%) Count (%) Prn (%) Count (%) Prn (%)
Overall three decades: 1990–2020
ENS, all species 196 -1.0000 0.1008 0.0856 0.9910 37.8 15.2 37.2 68.1 25.0 16.5
ENSG 143 -1.0000 -0.0542 -0.0150 0.9727 45.5 6.0 27.3 70.4 27.3 23.3
WGENS 188 -1.0000 0.0648 0.0780 0.9910 39.9 14.3 35.6 72.8 24.5 12.8
ENS, finfish 180 -1.0000 0.1347 0.1245 0.9910 40.0 21.5 41.1 69.9 18.9 8.6
ENS, molluscs 56 -1.0000 -0.0055 0.0764 1.0000 51.8 9.3 28.6 84.0 19.6 6.6
ENS, crustaceans 59 -0.8385 -0.1236 -0.0250 0.9734 50.8 18.4 22.0 62.2 27.1 18.9
ENS, MAA 13 -0.8026 0.4193 0.1996 0.8959 46.2 8.8 38.5 88.0 15.4 2.3
ENS, algae 31 -0.9272 0.1564 0.1598 0.9995 61.3 3.5 25.8 88.9 12.9 7.0
First decade: 1990–2000
ENS, all species 154 -1.0000 -0.0296 0.0191 1.0000 62.3 19.1 20.1 73.0 17.5 7.8
ENSG 101 -1.0000 -0.0839 -0.0503 1.0000 61.4 84.4 17.8 2.1 20.8 12.4
WGENS 147 -1.0000 0.0452 0.0189 0.9714 65.3 20.6 17.7 74.4 17.0 4.7
ENS, finfish 140 -1.0000 0.1004 0.0529 1.0000 62.1 13.6 20.0 81.2 17.9 5.0
ENS, molluscs 38 -1.0000 -0.0287 0.0798 1.0000 63.2 25.2 23.7 1.9 13.2 71.7
ENS, crustaceans 41 -0.8779 -0.2260 -0.0569 0.9325 70.7 44.8 17.1 28.5 12.2 25.5
ENS, MAA 6 -0.4353 0.5582 0.3238 0.8276 66.7 33.3 54.3
ENS, algae 9 -0.7390 -0.1095 0.0260 0.8714 55.6 22.7 33.3 68.8 11.1 6.6
Second decade: 2000–2010
ENS, all species 181 -1.0000 -0.0315 -0.0025 0.9583 61.9 12.9 21.0 72.3 17.1 14.7
ENSG 125 -0.9908 -0.2252 -0.1194 0.9507 56.8 18.5 16.0 65.9 27.2 15.0
WGENS 176 -1.0000 0.0105 0.0216 0.9705 63.6 20.4 21.0 71.9 15.3 7.5
ENS, finfish 167 -1.0000 0.1859 0.1510 1.0000 58.1 24.9 29.9 72.2 12.0 2.8
ENS, molluscs 48 -1.0000 -0.1360 -0.1027 1.0000 62.5 12.2 14.6 81.1 22.9 6.7
ENS, crustaceans 49 -0.9808 -0.0676 -0.0160 1.0000 57.1 82.5 20.4 11.9 22.4 3.8
ENS, MAA 12 -0.9225 0.4497 0.2491 1.0000 50.0 7.8 33.3 87.9 16.7 2.8
ENS, algae 19 -1.0000 0.0018 0.0283 1.0000 57.9 4.8 21.1 82.8 21.1 11.7
Third decade: 2010–2020
ENS, all species 189 -1.0000 -0.0812 -0.0341 1.0000 49.7 74.0 24.3 4.9 25.9 20.9
ENSG 131 -1.0000 -0.0259 0.0032 0.9883 55.0 22.4 23.7 68.2 21.4 8.8
WGENS 176 -1.0000 -0.1118 -0.0427 1.0000 54.0 18.0 21.6 5.2 24.4 76.7
ENS, finfish 168 -1.0000 -0.0133 -0.0240 1.0000 51.8 14.7 23.8 69.0 24.4 16.2
ENS, molluscs 52 -0.9226 -0.0977 -0.0574 0.9783 53.8 90.8 19.2 3.4 26.9 5.7
ENS, crustaceans 41 -0.9323 -0.1882 -0.0838 0.9866 56.1 14.7 14.6 1.0 29.3 77.2
ENS, MAA 13 -0.9481 0.1688 -0.0124 0.9676 53.8 8.4 23.1 89.4 23.1 1.3
ENS, algae 29 -0.9990 0.0681 0.1304 1.0000 62.1 92.3 27.6 7.2 10.3 0.0
Notes: The first row (in italic fonts) contains the following information. Among the estimated temporal correlations during 1990–2020 for 196 national aquaculture (one correlation for each country),
the minimum, median, mean and maximum correlations were, respectively, -1.0000, 0.1008, 0.0856, and 0.991. For 37.8 percent of the 196 national aquaculture (accounting of 15.2 percent of world
production), the correlations were statistically insignificant, whereas the correlation was significantly positive in 37.2 percent of the 196 national aquaculture (68.1 percent of world production), while
significantly negative in 25 percent (16.5 percent of production). Information in other rows can be interpreted in similar ways.

33
Supplementary Table 14: National aquaculture with statistically significant temporal correlations between production and ENS in the most
recent decade (2010–2020)
46 national aquaculture with POSITIVE, statistically significant correlation between production 49 national aquaculture with NEGATIVE, statistically significant correlation
and ENS during 2010–2020 (ranked by production share; 24.3 percent of all national aquaculture; between production and ENS during 2010–2020 (ranked by production share;
4.9 percent of world production) 25.9 percent of all national aquaculture; 20.9 percent of world production)
Share of Share of
world Number of world Number of
National aquaculture Pearson’s r p-value National aquaculture Pearson’s r p-value
production years production years
(%) (%)
Viet Nam 3.4981 11 0.7945 0.0035 Indonesia 12.8975 11 -0.6031 0.0495
Iran (Islamic Republic of) 0.3501 11 0.8630 0.0006 Philippines 2.3237 11 -0.8751 0.0004
Türkiye 0.2590 11 0.7757 0.0050 Norway 1.2764 11 -0.7355 0.0099
Russian Federation 0.1752 11 0.9889 0.0000 Chile 1.1148 11 -0.6985 0.0168
Pakistan 0.1481 11 0.9687 0.0000 Thailand 1.0016 11 -0.6534 0.0292
Greece 0.1164 11 0.6473 0.0313 Brazil 0.5210 11 -0.7841 0.0043
Colombia 0.1100 11 0.6565 0.0282 Ecuador 0.4457 11 -0.9130 0.0001
United States of
Honduras 0.0553 11 0.7355 0.0099 0.4332 11 -0.6946 0.0177
America
Nepal 0.0480 11 0.7708 0.0055 Spain 0.2814 11 -0.7364 0.0098
Poland 0.0370 11 0.7263 0.0114 Cambodia 0.1689 11 -0.9352 0.0000
Denmark 0.0362 11 0.8186 0.0021 Australia 0.0859 11 -0.7274 0.0112
Czechia 0.0202 11 0.6219 0.0411 Faeroe Islands 0.0764 11 -0.7294 0.0109
Paraguay 0.0082 11 0.6347 0.0359 Netherlands 0.0518 11 -0.6701 0.0241
Singapore 0.0052 11 0.6047 0.0488 Ireland 0.0382 11 -0.8586 0.0007
Bosnia and Herzegovina 0.0041 11 0.6770 0.0221 Germany 0.0331 11 -0.9184 0.0001
Lithuania 0.0038 11 0.6465 0.0316 Sri Lanka 0.0263 11 -0.7141 0.0136
Austria 0.0034 11 0.9840 0.0000 Iraq 0.0229 11 -0.6500 0.0304
Democratic Republic of the Congo 0.0031 11 0.7979 0.0033 Croatia 0.0162 11 -0.8934 0.0002
Bolivia (Plurinational State of) 0.0024 11 0.8911 0.0002 Iceland 0.0152 11 -0.9234 0.0001
Dominican Republic 0.0020 11 0.8514 0.0009 Tunisia 0.0150 11 -0.8145 0.0023
United Arab Emirates 0.0017 11 0.7771 0.0049 Armenia 0.0130 11 -0.7212 0.0123
Slovakia 0.0016 11 0.8242 0.0018 Malta 0.0114 11 -0.7782 0.0048
Switzerland 0.0016 11 0.9380 0.0000 Republic of Moldova 0.0107 11 -0.8756 0.0004
Kyrgyzstan 0.0013 11 0.6080 0.0472 Zimbabwe 0.0097 11 -0.9172 0.0001
Lebanon 0.0010 11 0.6986 0.0168 Solomon Islands 0.0079 11 -0.6435 0.0326
Senegal 0.0009 11 0.8079 0.0026 Panama 0.0079 11 -0.7246 0.0117
Latvia 0.0007 11 0.7775 0.0049 El Salvador 0.0061 11 -0.7266 0.0113

34
 46 national aquaculture with POSITIVE, statistically significant correlation between production 49 national aquaculture with NEGATIVE, statistically significant correlation
and ENS during 2010–2020 (ranked by production share; 24.3 percent of all national aquaculture; between production and ENS during 2010–2020 (ranked by production share;
4.9 percent of world production) 25.9 percent of all national aquaculture; 20.9 percent of world production)
Share of Share of
world Number of world Number of
National aquaculture Pearson’s r p-value National aquaculture Pearson’s r p-value
production years production years
(%) (%)
Namibia 0.0005 11 0.8638 0.0006 Afghanistan 0.0053 11 -0.6954 0.0175
Guinea 0.0005 11 0.7362 0.0098 Papua New Guinea 0.0045 11 -0.9358 0.0000
Central African Republic 0.0002 11 0.9640 0.0000 Belize 0.0036 11 -0.6836 0.0204
Guam 0.0001 11 0.8917 0.0002 Mali 0.0035 11 -0.6592 0.0273
Chad 0.0001 11 0.8030 0.0029 French Polynesia 0.0019 11 -0.9402 0.0000
Turkmenistan 0.0001 11 0.8461 0.0010 Cameroon 0.0019 11 -0.7832 0.0044
Réunion 0.0001 11 0.8975 0.0002 Georgia 0.0016 11 -0.9368 0.0000
Northern Mariana Islands 0.0000 11 0.9101 0.0001 Mozambique 0.0015 11 -0.7498 0.0079
Gambia 0.0000 11 0.9638 0.0000 Mauritius 0.0013 11 -0.6405 0.0338
Botswana 0.0000 6 0.9910 0.0001 Channel Islands 0.0012 11 -0.6148 0.0441
Libya 0.0000 11 1.0000 - Lesotho 0.0012 11 -0.8955 0.0002
South Sudan 0.0000 9 0.7091 0.0324 Timor-Leste 0.0008 11 -0.9233 0.0001
Antigua and Barbuda 0.0000 11 0.8536 0.0008 Fiji 0.0006 11 -0.9307 0.0000
Grenada 0.0000 6 0.9363 0.0060 Azerbaijan 0.0005 11 -0.7561 0.0071
Cook Islands 0.0000 11 0.8822 0.0003 Kuwait 0.0003 11 -0.7259 0.0114
Bahamas 0.0000 11 0.7014 0.0162 Liberia 0.0002 11 -0.6024 0.0498
French Guiana 0.0000 11 0.7732 0.0053 Sierra Leone 0.0001 11 -1.0000 0.0000
Marshall Islands 0.0000 7 0.8008 0.0305 Saint Lucia 0.0000 11 -0.8413 0.0012
Saint Kitts and Nevis 0.0000 11 0.9281 0.0000 Guadeloupe 0.0000 11 -0.7131 0.0138
Barbados 0.0000 11 -0.8221 0.0019
United States Virgin
0.0000 11 -0.9930 0.0000
Islands
Dominica 0.0000 7 -1.0000 0.0000

35
Supplementary Table 15a: Countries in the “Both > median” category (i.e. relatively high within-group
diversity and between-group diversity)
Countries (or territories) in this category are (i) indexed
Aquaculture production, 2020 Diversity measures, 2020
according to ranking in world production; (ii) highlighted in
red text when ENS < median and in blue text when
Tonnes Share of world (%) ENS WGENS ENSG
ENS > median; and (iii) mapped in Figure 3e in blue.
World (48 countries) 111 188 012 90.7063 n.a. n.a. n.a.
Africa (8) 53 446 0.0436 n.a. n.a. n.a.
58. Tunisia 23 486 0.0192 2.95 2.82 1.04
61. Kenya 20 831 0.0170 2.33 1.96 1.19
92. Algeria 5 436 0.0044 4.48 3.99 1.12
120. Morocco 1 618 0.0013 6.51 2.66 2.45
128. Senegal 1 100 0.0009 3.67 1.84 2.00
Landlocked (LLDCs) 138. Burkina Faso 651 0.0005 2.33 2.22 1.05
Landlocked (LLDCs) 148. Central African Republic 219 0.0002 4.86 2.84 1.71
Landlocked (LLDCs) 156. Chad 105 0.0001 3.42 2.10 1.63
Americas (7) 1 460 057 1.1911 n.a. n.a. n.a.
16. Brazil 630 200 0.5141 5.13 3.29 1.56
18. United States of America 448 535 0.3659 6.60 2.35 2.81
29. Colombia 179 351 0.1463 5.25 4.60 1.14
30. Canada 171 007 0.1395 3.15 1.98 1.59
Island (SIDS) 57. Cuba 26 200 0.0214 3.97 2.09 1.90
Island (SIDS) 108. Dominican Republic 2 680 0.0022 7.67 5.26 1.46
113. Argentina 2 085 0.0017 3.12 3.00 1.04
Asia (19) 108 734 650 88.7049 n.a. n.a. n.a.
1. China 70 483 538 57.4999 28.00 7.43 3.77
Island 2. Indonesia 14 845 014 12.1105 5.63 2.47 2.28
3. India 8 641 286 7.0495 6.58 4.59 1.43
4. Viet Nam 4 614 692 3.7646 8.24 3.79 2.17
5. Bangladesh 2 583 866 2.1079 14.30 11.48 1.25
6. Republic of Korea 2 327 903 1.8991 6.95 3.30 2.11
11. Myanmar 1 145 018 0.9341 3.98 3.13 1.27
Island 12. Japan 996 297 0.8128 8.68 2.78 3.13
13. Thailand 962 467 0.7852 7.24 2.65 2.73
17. Iran (Islamic Republic of) 480 500 0.3920 4.90 3.53 1.39
19. Türkiye 421 411 0.3438 3.61 3.42 1.06
20. Cambodia 400 400 0.3266 7.70 6.19 1.24
21. Malaysia 400 017 0.3263 8.47 2.72 3.11
Island 24. Taiwan Province of China 278 503 0.2272 11.04 4.65 2.38
40. Saudi Arabia 99 907 0.0815 3.95 1.98 2.00
Island 50. Sri Lanka 42 119 0.0344 5.17 2.93 1.77
Island (SIDS) 93. Singapore 4 829 0.0039 11.67 6.06 1.93
98. China, Hong Kong Special
Administrative Region 3 835 0.0031 7.25 4.64 1.56
106. United Arab Emirates 3 048 0.0025 4.49 3.31 1.36
Europe (11) 833 644 0.6801 n.a. n.a. n.a.
22. Russian Federation 291 194 0.2376 10.03 5.29 1.89
28. France 191 350 0.1561 4.62 2.60 1.77
33. Greece 131 737 0.1075 3.69 2.40 1.54
36. Italy 122 778 0.1002 4.51 2.31 1.95
55. Germany 32 258 0.0263 5.42 2.73 1.99
60. Croatia 21 718 0.0177 5.12 4.58 1.12
70. Bulgaria 15 052 0.0123 6.24 4.11 1.52
72. Portugal 14 552 0.0119 6.49 3.28 1.98
81. Albania 9 284 0.0076 3.36 2.93 1.15
Landlocked 116. Switzerland 2 048 0.0017 3.58 3.32 1.08
118. Slovenia 1 673 0.0014 4.38 2.52 1.74
Oceania (3) 106 215 0.0866 n.a. n.a. n.a.
39. Australia 106 088 0.0865 4.43 2.14 2.07
Island (SIDS) 154. Guam 118 0.0001 2.79 1.94 1.44
Island (SIDS) 183. Cook Islands 9 0.0000 2.92 2.03 1.44

36
Supplementary Table 15b: Countries in the “Both < median” category (i.e. relatively low within-group
diversity and between-group diversity)
Countries (or territories) in this category are (i) indexed
Aquaculture production, 2020 Diversity measures, 2020
according to ranking in world production; (ii) highlighted in red
text when ENS < median and in blue text when ENS > median; Share of world
Tonnes ENS WGENS ENSG
and (iii) mapped in Figure 3e in red. (%)
World (48 countries) 1 845 263 1.5054 n.a. n.a. n.a.
Africa (21) 119 252 0.0973 n.a. n.a. n.a.
45. Ghana 64 010 0.0522 1.36 1.36 1.00
Landlocked (LLDCs) 68. Zimbabwe 15 425 0.0126 1.03 1.03 1.00
78. Sudan 9 850 0.0080 1.66 1.66 1.00
Landlocked (LLDCs) 85. Mali 7 686 0.0063 1.78 1.78 1.00
Landlocked (LLDCs) 87. Rwanda 7 059 0.0058 1.40 1.39 1.00
101. Democratic Republic of the Congo 3 590 0.0029 1.15 1.15 1.00
105. Mozambique 3 162 0.0026 1.00 1.00 1.00
Landlocked (LLDCs) 109. Lesotho 2 600 0.0021 1.00 1.00 1.00
114. Angola 2 062 0.0017 1.08 1.08 1.00
Landlocked (LLDCs) 123. Burundi 1 450 0.0012 1.30 1.30 1.00
131. Congo 900 0.0007 1.60 1.60 1.00
136. Togo 730 0.0006 1.31 1.31 1.00
147. Liberia 255 0.0002 1.48 1.48 1.00
Landlocked (LLDCs) 152. Botswana 146 0.0001 1.76 1.76 1.00
Landlocked (LLDCs) 158. Eswatini 100 0.0001 1.00 1.00 1.00
160. Sierra Leone 85 0.0001 1.25 1.25 1.00
161. Gabon 60 0.0000 1.33 1.33 1.00
Landlocked (LLDCs) 167. South Sudan 30 0.0000 1.28 1.28 1.00
Island (SIDS) 172. Cabo Verde 22 0.0000 1.00 1.00 1.00
Island 173. Mayotte 20 0.0000 1.00 1.00 1.00
181. Libya 10.0 0.0000 1.00 1.00 1.00
America (7) 30 990 0.0253 n.a. n.a. n.a.
56. Nicaragua 29 410 0.0240 1.00 1.00 1.00
Island (SIDS) 121. Haiti 1 560 0.0013 1.52 1.52 1.00
Island (SIDS) 178. Saint Vincent and the Grenadines 13 0.0000 1.00 1.00 1.00
Island (SIDS) 187. Trinidad and Tobago 4 0.0000 1.73 1.73 1.00
Island (SIDS) 193. Aruba 2 0.0000 1.00 1.00 1.00
Island (SIDS) 194. British Virgin Islands 1 0.0000 1.00 1.00 1.00
Island 196. Bonaire, Sint Eustatius and Saba 0.0000 1.00 1.00 1.00
Asia (6) 33 117 0.0270 n.a. n.a. n.a.
59. Iraq 22 704 0.0185 1.71 1.71 1.00
Landlocked (LLDCs) 84. Afghanistan 8 050 0.0066 1.04 1.04 1.00
125. Oman 1 307 0.0011 1.62 1.62 1.00
140. State of Palestine 600 0.0005 1.70 1.70 1.00
144. Kuwait 450 0.0004 1.00 1.00 1.00
Island (SIDS) 186. Bahrain 6 0.0000 1.36 1.36 1.00
Europe (7) 1 656 261 1.3512 n.a. n.a. n.a.
10. Norway 1 490 412 1.2159 1.31 1.29 1.01
Island 42. Faeroe Islands 89 055 0.0727 1.01 1.00 1.01
Island 51. Iceland 40 595 0.0331 1.65 1.65 1.00
Island 63. Malta 19 829 0.0162 1.71 1.71 1.00
69. Finland 15 053 0.0123 1.26 1.26 1.00
127. Channel Islands 1 107 0.0009 1.35 1.35 1.00
149. Belgium 209 0.0002 1.00 1.00 1.00
Oceania (7) 5 644 0.0046 n.a. n.a. n.a.
Island (SIDS) 91. Solomon Islands 5 504 0.0045 1.01 1.00 1.01
Island (SIDS) 155. Tonga 105 0.0001 1.01 1.01 1.00
Island (SIDS) 174. American Samoa 20 0.0000 1.00 1.00 1.00
Island (SIDS) 179. Samoa 10 0.0000 1.01 1.00 1.01
Island (SIDS) 189. Tuvalu 3 0.0000 1.00 1.00 1.00
Island (SIDS) 191. Kiribati 2 0.0000 1.00 1.00 1.00
Island (SIDS) 195. Nauru 1 0.0000 1.00 1.00 1.00

37
Supplementary Table 15c: Countries in the “WGENS > median; ENSG ≤ median” category (i.e.
relatively high within-group diversity yet relatively low between-group diversity)
Countries (or territories) in this category are (i) indexed
according to ranking in world production; (ii) highlighted in Aquaculture production, 2020 Diversity measures, 2020
red text when ENS < median and in blue text when
ENS > median; and (iii) mapped in Figure 3e in green. Tonnes Share of world (%) ENS WGENS ENSG
World (50 countries) 2 752 730 2.2457 n.a. n.a. n.a.
Africa (13) 2 046 073 1.6692 n.a. n.a. n.a.
8. Egypt 1 591 896 1.2987 3.77 3.73 1.01
26. Nigeria 261 711 0.2135 5.66 5.64 1.00
Landlocked (LLDCs) 35. Uganda 123 897 0.1011 1.89 1.89 1.00
Landlocked (LLDCs) 48. Zambia 45 670 0.0373 3.21 3.21 1.00
Landlocked (LLDCs) 80. Malawi 9 393 0.0077 3.56 3.56 1.00
94. Côte d'Ivoire 4 620 0.0038 2.48 2.48 1.00
102. Cameroon 3 556 0.0029 2.12 2.12 1.00
107. Benin 2 987 0.0024 2.00 2.00 1.00
126. Guinea 1 135 0.0009 3.71 3.71 1.00
Landlocked (LLDCs) 139. Niger 649 0.0005 1.92 1.92 1.00
Landlocked (LLDCs) 142. Ethiopia 534 0.0004 2.17 2.17 1.00
177. Equatorial Guinea 15 0.0000 2.69 2.69 1.00
Island 181. Réunion 10 0.0000 2.00 2.00 1.00
Americas (5) 17 933 0.0146 n.a. n.a. n.a.
Landlocked (LLDCs) 73. Paraguay 14 100 0.0115 2.83 2.83 1.00
Landlocked (LLDCs) 100. Bolivia (Plurinational State of) 3 720 0.0030 5.09 5.09 1.00
157. Uruguay 103 0.0001 3.27 3.27 1.00
Island (SIDS) 184. United States Virgin Islands 8 0.0000 2.83 2.83 1.00
190. French Guiana 2 0.0000 2.00 2.00 1.00
Asia (16) 525 066 0.4283 n.a. n.a. n.a.
31. Pakistan 162 462 0.1325 5.67 5.59 1.01
Landlocked (LLDCs) 34. Lao People's Democratic Republic 130 020 0.1061 10.33 10.26 1.01
Landlocked (LLDCs) 41. Uzbekistan 98 003 0.0800 4.97 4.97 1.00
Landlocked (LLDCs) 43. Nepal 76 822 0.0627 6.98 6.98 1.00
Landlocked (LLDCs) 66. Armenia 18 350 0.0150 2.77 2.77 1.00
71. Israel 14 700 0.0120 5.80 5.80 1.00
Island 86. Cyprus 7 343 0.0060 2.05 2.02 1.02
Landlocked (LLDCs) 88. Kazakhstan 6 795 0.0055 7.06 7.06 1.00
Landlocked (LLDCs) 110. Kyrgyzstan 2 550 0.0021 3.98 3.98 1.00
111. Syrian Arab Republic 2 300 0.0019 3.67 3.67 1.00
115. Jordan 2 055 0.0017 2.00 2.00 1.00
117. Georgia 2 027 0.0017 3.34 3.34 1.00
Landlocked (LLDCs) 135. Tajikistan 775 0.0006 2.94 2.94 1.00
Landlocked (LLDCs) 143. Azerbaijan 533 0.0004 4.74 4.74 1.00
Landlocked (LLDCs) 150. Bhutan 181 0.0001 6.31 6.31 1.00
Landlocked (LLDCs) 151. Turkmenistan 150 0.0001 4.45 4.45 1.00
Europe (15) 163 637 0.1335 n.a. n.a. n.a.
47. Poland 47 700 0.0389 3.69 3.69 1.00
Landlocked 62. Czechia 20 401 0.0166 2.09 2.09 1.00
64. Ukraine 18 568 0.0151 3.08 3.08 1.00
Landlocked 65. Hungary 18 373 0.0150 3.05 3.05 1.00
Landlocked (LLDCs) 75. Republic of Moldova 12 600 0.0103 4.17 4.16 1.00
76. Romania 12 200 0.0100 5.93 5.93 1.00
Landlocked 82. Belarus 9 266 0.0076 2.48 2.48 1.00
Landlocked 90. Serbia 6 010 0.0049 2.30 2.30 1.00
95. Austria 4 527 0.0037 7.02 7.02 1.00
96. Lithuania 4 477 0.0037 2.85 2.85 1.00
99. Bosnia and Herzegovina 3 776 0.0031 1.89 1.89 1.00
Landlocked 112. Slovakia 2 296 0.0019 3.82 3.82 1.00
Landlocked (LLDCs) 119. North Macedonia 1 634 0.0013 2.13 2.13 1.00
129. Estonia 1 090 0.0009 1.85 1.84 1.01
137. Latvia 717 0.0006 2.51 2.51 1.00
Oceania (1) 22 0.0000 n.a. n.a. n.a.
Island (SIDS) 172. Palau 22 0.0000 2.53 2.53 1.00

38
Supplementary Table 15d: Countries in the “WGENS ≤ median; ENSG > median” category (i.e.
relatively low within-group diversity yet relatively high between-group diversity)
Countries (or territories) in this category are (i) indexed
according to ranking in world production; (ii) highlighted in
Aquaculture production, 2020 Diversity measures, 2020
red text when ENS < median and in blue text when
ENS > median; and (iii) mapped in Figure 3e in purple. Tonnes Share of world (%) ENS WGENS ENSG
Total (50) 6 794 182 5.5426 n.a. n.a. n.a.
Africa (6) 135 523 0.1106 n.a. n.a. n.a.
38. United Republic of Tanzania 108 568 0.0886 1.70 1.09 1.56
Island 74. Madagascar 13 550 0.0111 2.11 1.00 2.10
79. South Africa 9 753 0.0080 4.66 1.70 2.74
Island (SIDS) 104. Mauritius 3 298 0.0027 1.15 1.11 1.04
146. Namibia 321 0.0003 1.61 1.12 1.43
166. Gambia 33 0.0000 2.56 1.31 1.96
Americas (23) 2 891 567 2.3589 n.a. n.a. n.a.
9. Chile 1 505 486 1.2282 3.39 1.77 1.91
14. Ecuador 774 569 0.6319 1.11 1.01 1.10
23. Mexico 278 694 0.2274 2.78 1.36 2.05
32. Peru 143 830 0.1173 3.62 1.23 2.93
44. Honduras 71 151 0.0580 1.99 1.00 1.99
46. Venezuela (Bolivarian Republic of) 53 609 0.0437 1.85 1.05 1.77
54. Guatemala 33 651 0.0275 1.98 1.02 1.95
67. Costa Rica 16 269 0.0133 2.12 1.42 1.49
83. El Salvador 8 500 0.0069 1.51 1.01 1.49
97. Panama 3 879 0.0032 3.04 1.54 1.98
Island (SIDS) 130. Jamaica 918 0.0007 1.05 1.02 1.03
Island (SIDS) 141. Belize 565 0.0005 1.47 1.00 1.47
Island (SIDS) 153. Guyana 138 0.0001 2.00 1.17 1.72
Island (SIDS) 159. Saint Lucia 92 0.0001 1.44 1.00 1.44
Island (SIDS) 162. Guadeloupe 45 0.0000 1.53 1.27 1.20
Island (SIDS) 163. Martinique 42 0.0000 1.68 1.11 1.51
Island (SIDS) 165. Suriname 37 0.0000 2.54 1.42 1.79
Island (SIDS) 168. Barbados 26 0.0000 1.10 1.00 1.10
Island (SIDS) 170. Grenada 23 0.0000 1.11 1.00 1.11
Island (SIDS) 173. Antigua and Barbuda 20 0.0000 2.00 1.00 2.00
Island (SIDS) 176. Puerto Rico 18 0.0000 2.65 1.40 1.89
Island (SIDS) 185. Bahamas 6 0.0000 1.57 1.00 1.57
Island (SIDS) 192. Saint Kitts and Nevis 2 0.0000 2.00 1.00 2.00
Asia (6) 3 008 317 2.4542 n.a. n.a. n.a.
Island 7. Philippines 2 322 831 1.8949 3.93 1.63 2.41
15. Democratic People's Republic of Korea 680 300 0.5550 1.59 1.06 1.51
Island 103. Brunei Darussalam 3 500 0.0029 2.47 1.68 1.47
Island (SIDS) 133. Timor-Leste 835 0.0007 1.73 1.10 1.57
134. Lebanon 828 0.0007 1.20 1.10 1.09
169. Qatar 23 0.0000 1.16 1.00 1.16
Europe (7) 630 865 0.5147 n.a. n.a. n.a.
25. Spain 276 571 0.2256 2.97 1.61 1.85
Island 27. United Kingdom 221 000 0.1803 1.69 1.34 1.26
49. Denmark 42 629 0.0348 2.24 1.46 1.53
52. Netherlands 39 940 0.0326 2.18 1.46 1.49
Island 53. Ireland 37 751 0.0308 3.29 1.69 1.95
77. Sweden 12 090 0.0099 2.24 1.38 1.63
132. Montenegro 885 0.0007 2.87 1.59 1.81
Oceania (8) 127 909 0.1043 n.a. n.a. n.a.
Island 37. New Zealand 118 582 0.0967 1.57 1.07 1.47
Island (SIDS) 89. Papua New Guinea 6 102 0.0050 2.30 1.25 1.84
Island (SIDS) 122. New Caledonia 1 478 0.0012 1.03 1.00 1.03
Island (SIDS) 124. French Polynesia 1 371 0.0011 1.54 1.04 1.48
Island (SIDS) 145. Fiji 322 0.0003 3.28 1.17 2.80
Island (SIDS) 164. Northern Mariana Islands 41 0.0000 2.21 1.10 2.00
Island (SIDS) 182. Marshall Islands 10 0.0000 1.58 1.00 1.58
Island (SIDS) 188. Vanuatu 3 0.0000 1.99 1.23 1.62

39
Supplementary Table 16: Species diversity in countries with the largest land area
Share of world (%) Diversity measures, 2020
Top 10 countries with the largest land Aquaculture
area Land area production, ENS WGENS ENSG
2020
1. Russian Federation 12.75 0.1752 7.01 5.38 1.31
2. Canada 7.45 0.1734 3.55 2.09 1.70
3. United States of America 7.33 0.4332 6.68 2.47 2.71
4. China 7.16 57.9137 28.10 7.68 3.66
5. Brazil 6.35 0.5210 6.57 3.86 1.69
6. Australia 5.77 0.0859 5.28 2.33 2.26
7. India 2.45 5.5156 6.29 4.59 1.37
8. Argentina 2.07 0.0031 3.35 2.97 1.14
9. Kazakhstan 2.03 0.0025 4.88 4.88 1.00
10. Algeria 1.78 0.0031 5.14 4.52 1.13
Notes: Land area data from FAO. 2016. AQUASTAT Main Database – Food and Agriculture Organization of the United Nations
(FAO).

40
Supplementary Table 17: Examples of relatively large between-group diversity in inland aquaculture, 2020
Inland aquaculture production, 2020

Crustacean share (%) Molluscs share (%) MAA share (%) Algae share (%)
Countries/territories Between-
Finfish
(ranked by production) group
Tonnes share
diversity Largest Largest Largest Largest
(%) Overall Overall Overall Overall
(ENSG) species species species species

China 30 945 335 1.72 83.58 13.76 PC (7.74) 0.60 CC (0.29) 1.86 TS (1.07) 0.20 SP (0.20)
Thailand 405 656 1.39 91.09 8.00 MR (8.00) 0.91
United States of America 243 721 1.84 70.34 29.66 PC (29.66)
Taiwan Province of China 119 496 1.72 85.79 8.45 MR (5.42) 3.42 CF (3.42) 2.34 RC (1.37)
Spain 19 266 1.54 84.41 15.57 PC (15.57) - 0.02 SP (0.02)
Chile 1 928 1.86 31.48 - 68.52 SP (65.35)
Australia 1 460 1.36 90.94 9.06 CQ (4.22)
Tunisia 994 1.29 92.96 - 7.04 SP (7.04)
Singapore 673 2.04 50.14 0.32 MR (0.23) 49.54 RC (49.54)
Central African Republic 219 1.71 77.17 - 22.83 SP (22.83)
Fiji 145 1.86 69.02 30.98 MR (30.98)
Chad 105 1.63 80.95 - 19.05 SP (19.05)
Notes: Including countries or territories with inland aquaculture production in 2020 greater than 100 tonnes and between-group diversity higher than 1.2 ENSG.
PC = Procambarus clarkii; MR = Macrobrachium rosenbergii; CQ = Cherax quadricarinatus; CC = Cipangopaludina chinensis; CF = Corbicula fluminea; TS = Trionyx sinensis;
RC = Rana catesbeiana; SP = Spirulina spp.

41
 Supplementary Table 18: A summary of key results
Country or area: U1: ENS increase (both components up)
 Regional aquaculture in italic Diversity measures U2: ENS increase (only WGENS up)
 National aquaculture indexed by Aquaculture production ENS = Effective number of species (a measure of overall species diversity); WGENS = Within-group ENS (a measure of within-group diversity);
U3: ENS increase (only ENSG up)
production ranking in 2020 ENSG = Effective number of species groups (a measure of between-group diversity); ENS = WGENS × ENSG
D1: ENS decrease (both components down)
 LDLK = Landlocked D2: ENS decrease (only WGENS down)
 LLDCs = Landlocked Developing Year 2020 Year 1990 Year 2000 Year 2010 Year 2020 D3: ENS decrease (only ENSG down)
Countries
 ISLD = Island Thousand % of world Overall: 1990– 2000– 2010–
ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG
 SIDS = Small Island Developing States tonnes production 1990–2020 2000 2010 2020
World 122 580 187 100.0000 35.61 11.06 3.22 34.88 10.71 3.26 50.59 14.60 3.47 47.46 13.56 3.50 Inverted N U1 D2 U1 D2
World excluding China 52 096 649 42.5001 38.46 11.26 3.42 45.87 14.72 3.12 39.84 13.07 3.05 33.18 11.40 2.91 Inverted U D3 U2 D1 D1
Developed regions 5 146 959 4.1989 19.68 6.60 2.98 20.15 7.29 2.76 18.47 6.98 2.65 15.68 6.33 2.48 Inverted U D1 U2 D1 D1
Developing regions 117 433 228 95.8011 28.11 8.77 3.21 29.03 8.85 3.28 45.66 13.08 3.49 44.30 12.58 3.52 Inverted U U1 U1 U1 D2
Small Island Developing
56 507 0.0461 7.68 2.56 3.00 10.71 3.23 3.31 12.73 3.97 3.20 13.75 4.66 2.95 Straight up U2 U1 U2 U2
States
Landlocked developing
589 897 0.4812 4.89 4.88 1.00 9.45 9.43 1.00 11.19 11.13 1.01 14.07 14.02 1.00 Straight up U1 U1 U1 U2
countries
Asia 112 301 149 91.6144 29.70 9.23 3.22 28.70 8.71 3.30 44.06 12.56 3.51 42.28 11.96 3.53 Inverted N U1 D2 U1 D2
Eastern Asia 74 770 377 60.9971 20.45 6.41 3.19 20.48 6.28 3.26 31.58 8.67 3.64 30.35 8.08 3.76 Inverted U U1 U3 U1 D2
South-eastern Asia 24 829 623 20.2558 15.84 4.81 3.30 17.51 5.23 3.35 17.54 5.74 3.06 14.39 5.06 2.85 Inverted U D3 U1 U2 D1
Southern Asia 11 995 286 9.7857 6.44 5.39 1.20 8.20 6.34 1.29 7.79 6.36 1.23 10.49 7.56 1.39 N-shape U1 U1 D3 U1
Western Asia 597 590 0.4875 4.57 4.52 1.01 6.26 5.63 1.11 7.52 5.60 1.34 7.52 5.48 1.37 Inverted U U1 U1 U3 D2
Central Asia 108 273 0.0883 2.07 2.07 1.00 3.05 3.05 1.00 3.47 3.47 1.00 5.71 5.71 1.00 Straight up U2 U2 U2 U2
1. China 70 483 538 57.4999 13.57 4.56 2.98 16.61 5.15 3.23 28.18 7.75 3.64 28.00 7.43 3.77 Inverted U U1 U1 U1 D2
ISLD 2. Indonesia 14 845 014 12.1105 10.26 4.22 2.43 10.30 4.23 2.44 6.40 2.67 2.39 5.63 2.47 2.28 Inverted U D1 U1 D1 D1
3. India 8 641 286 7.0495 5.51 4.74 1.16 6.15 4.89 1.26 4.55 3.76 1.21 6.58 4.59 1.43 N-shape U3 U1 D1 U1
4. Viet Nam 4 614 692 3.7646 2.91 1.33 2.19 4.22 1.80 2.35 6.43 3.55 1.81 8.24 3.79 2.17 Straight up U2 U1 U2 U1
5. Bangladesh 2 583 866 2.1079 1.65 1.20 1.37 7.69 5.58 1.38 12.01 9.39 1.28 14.30 11.48 1.25 Straight up U2 U1 U2 U2
6. Republic of Korea 2 327 903 1.8991 6.90 2.74 2.52 7.14 2.61 2.74 7.74 3.22 2.41 6.95 3.30 2.11 Inverted U U2 U3 U2 D3
ISLD 7. Philippines 2 322 831 1.8949 5.45 1.87 2.91 4.12 1.71 2.41 3.56 1.67 2.14 3.93 1.63 2.41 U-shape D1 D1 D1 U3
11. Myanmar 1 145 018 0.9341 1.11 1.11 1.00 1.22 1.00 1.22 4.70 3.21 1.46 3.98 3.13 1.27 Inverted U U1 U3 U1 D1
ISLD 12. Japan 996 297 0.8128 8.98 2.92 3.07 8.53 2.78 3.07 8.08 2.60 3.11 8.68 2.78 3.13 U-shape D2 D1 D2 U1
13. Thailand 962 467 0.7852 8.28 2.83 2.93 7.48 2.58 2.90 6.70 2.31 2.90 7.24 2.65 2.73 U-shape D1 D1 D2 U2
15. Democratic People's
680 300 0.5550 1.91 1.49 1.28 2.20 1.42 1.55 1.55 1.03 1.51 1.59 1.06 1.51 N-shape D2 U3 D1 U2
Republic of Korea
17. Iran (Islamic Republic
480 500 0.3920 3.56 3.56 1.00 4.49 3.25 1.38 4.16 3.63 1.15 4.90 3.53 1.39 N-shape U3 U3 D3 U3
of)
19. Türkiye 421 411 0.3438 3.47 3.47 1.00 2.86 2.78 1.03 3.03 2.99 1.01 3.61 3.42 1.06 U-shape U3 D2 U2 U1
20. Cambodia 400 400 0.3266 5.52 5.52 1.00 6.03 5.24 1.15 8.20 6.98 1.17 7.70 6.19 1.24 Inverted U U1 U3 U1 D2
21. Malaysia 400 017 0.3263 4.26 2.04 2.09 9.47 2.89 3.28 9.12 2.46 3.71 8.47 2.72 3.11 Inverted U U1 U1 D2 D3
24. Taiwan Province of
ISLD 278 503 0.2272 11.40 4.63 2.46 15.66 5.61 2.79 11.21 4.39 2.55 11.04 4.65 2.38 Inverted U D3 U1 D1 D3
China
31. Pakistan 162 462 0.1325 1.03 1.00 1.03 1.04 1.00 1.04 5.56 5.52 1.01 5.67 5.59 1.01 Straight up U2 U3 U2 U1
34. Lao People's
LLDCs 130 020 0.1061 8.00 8.00 1.00 5.12 5.12 1.00 10.39 10.37 1.00 10.33 10.26 1.01 Inverted N U1 D2 U1 D2
Democratic Republic
40. Saudi Arabia 99 907 0.0815 1.21 1.08 1.13 2.16 1.15 1.88 2.19 1.30 1.69 3.95 1.98 2.00 Straight up U1 U1 U2 U1
LLDCs 41. Uzbekistan 98 003 0.0800 2.01 2.01 1.00 3.16 3.16 1.00 2.82 2.82 1.00 4.97 4.97 1.00 N-shape U2 U2 D2 U2
LLDCs 43. Nepal 76 822 0.0627 1.75 1.75 1.00 4.40 4.40 1.00 5.87 5.87 1.00 6.98 6.98 1.00 Straight up U2 U2 U2 U2
ISLD 50. Sri Lanka 42 119 0.0344 1.14 1.00 1.14 1.07 1.00 1.07 5.88 2.96 1.99 5.17 2.93 1.77 Inverted N U1 D3 U1 D1
59. Iraq 22 704 0.0185 1.00 1.00 1.00 1.00 1.00 1.00 1.91 1.91 1.00 1.71 1.71 1.00 Inverted U U2 U2 D2
LLDCs 66. Armenia 18 350 0.0150 1.93 1.93 1.00 2.70 2.70 1.00 3.51 3.42 1.03 2.77 2.77 1.00 Inverted U U2 U2 U1 D1
71. Israel 14 700 0.0120 3.09 3.08 1.00 5.35 5.35 1.00 5.77 5.77 1.00 5.80 5.80 1.00 Straight up U2 U2 U2 U2
LLDCs 84. Afghanistan 8 050 0.0066 1.00 1.00 1.00 1.00 1.00 1.00 1.28 1.28 1.00 1.04 1.04 1.00 Inverted U U2 U2 D2
ISLD 86. Cyprus 7 343 0.0060 2.53 2.53 1.00 2.38 2.04 1.16 2.20 2.20 1.00 2.05 2.02 1.02 Straight down D2 D2 D3 D2
LLDCs 88. Kazakhstan 6 795 0.0055 1.89 1.89 1.00 1.31 1.31 1.00 3.73 3.73 1.00 7.06 7.06 1.00 U-shape U2 D2 U2 U2
SIDS 93. Singapore 4 829 0.0039 4.38 1.65 2.66 4.79 2.17 2.21 8.07 5.77 1.40 11.67 6.06 1.93 Straight up U2 U2 U2 U1

42
 Country or area: U1: ENS increase (both components up)
 Regional aquaculture in italic Diversity measures U2: ENS increase (only WGENS up)
 National aquaculture indexed by Aquaculture production ENS = Effective number of species (a measure of overall species diversity); WGENS = Within-group ENS (a measure of within-group diversity);
U3: ENS increase (only ENSG up)
production ranking in 2020 ENSG = Effective number of species groups (a measure of between-group diversity); ENS = WGENS × ENSG
D1: ENS decrease (both components down)
 LDLK = Landlocked D2: ENS decrease (only WGENS down)
 LLDCs = Landlocked Developing Year 2020 Year 1990 Year 2000 Year 2010 Year 2020 D3: ENS decrease (only ENSG down)
Countries
 ISLD = Island Thousand % of world Overall: 1990– 2000– 2010–
ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG
 SIDS = Small Island Developing States tonnes production 1990–2020 2000 2010 2020
98. China, Hong Kong
Special Administrative 3 835 0.0031 12.05 9.15 1.32 12.27 9.27 1.32 9.52 6.28 1.52 7.25 4.64 1.56 Inverted U D2 U1 D2 D2
Region
ISLD 103. Brunei Darussalam 3 500 0.0029 3.46 3.46 1.00 4.86 2.62 1.86 3.82 2.08 1.83 2.47 1.68 1.47 Inverted U D2 U3 D1 D1
106. United Arab Emirates 3 048 0.0025 1.32 1.00 1.32 4.49 3.31 1.36 U1
LLDCs 110. Kyrgyzstan 2 550 0.0021 1.80 1.80 1.00 2.09 2.09 1.00 2.95 2.95 1.00 3.98 3.98 1.00 Straight up U2 U2 U2 U2
111. Syrian Arab Republic 2 300 0.0019 1.69 1.69 1.00 2.35 2.35 1.00 3.42 3.42 1.00 3.67 3.67 1.00 Straight up U2 U2 U2 U2
115. Jordan 2 055 0.0017 1.89 1.89 1.00 1.06 1.06 1.00 2.00 2.00 1.00 2.00 2.00 1.00 U-shape U2 D2 U2 U2
117. Georgia 2 027 0.0017 1.18 1.18 1.00 3.11 3.11 1.00 4.65 4.65 1.00 3.34 3.34 1.00 Inverted U U2 U2 U2 D2
125. Oman 1 307 0.0011 1.00 1.00 1.00 1.62 1.62 1.00 U2
SIDS 133. Timor-Leste 835 0.0007 1.20 1.03 1.17 1.73 1.10 1.57 U1
134. Lebanon 828 0.0007 1.00 1.00 1.00 1.00 1.00 1.00 1.40 1.26 1.11 1.20 1.10 1.09 Inverted U U1 U1 D1
LLDCs 135. Tajikistan 775 0.0006 2.06 2.06 1.00 2.41 2.41 1.00 3.82 3.82 1.00 2.94 2.94 1.00 Inverted U U2 U2 U2 D2
140. State of Palestine 600 0.0005 2.81 2.81 1.00 1.70 1.70 1.00 D2
LLDCs 143. Azerbaijan 533 0.0004 1.57 1.57 1.00 1.68 1.68 1.00 2.68 2.68 1.00 4.74 4.74 1.00 Straight up U2 U2 U2 U2
144. Kuwait 450 0.0004 3.76 3.76 1.00 1.15 1.15 1.00 1.00 1.00 1.00 Straight down D2 D2
LLDCs 150. Bhutan 181 0.0001 1.00 1.00 1.00 1.00 1.00 1.00 7.15 7.15 1.00 6.31 6.31 1.00 Inverted U U2 U2 D2
LLDCs 151. Turkmenistan 150 0.0001 1.97 1.97 1.00 1.14 1.14 1.00 2.46 2.46 1.00 4.45 4.45 1.00 U-shape U2 D2 U2 U2
169. Qatar 23 0.0000 1.00 1.00 1.00 1.16 1.00 1.16 U3
SIDS 186. Bahrain 6 0.0000 1.00 1.00 1.00 2.48 2.48 1.00 1.36 1.36 1.00 Inverted U U2 D2
Americas 4 400 548 3.5899 13.06 4.17 3.13 16.04 6.84 2.34 15.93 5.78 2.76 11.46 4.18 2.74 Inverted U D3 U2 D2 D1
Northern America 619 542 0.5054 6.63 2.87 2.31 6.66 3.52 1.89 7.71 3.20 2.41 8.06 3.09 2.61 Straight up U1 U2 U3 U3
Latin America and the
3 781 006 3.0845 9.05 2.93 3.09 13.70 5.65 2.43 11.13 4.02 2.77 8.70 3.24 2.68 Inverted U D3 U2 D2 D1
Caribbean
South America 3 307 229 2.6980 6.20 2.01 3.08 12.40 5.28 2.35 11.23 4.13 2.72 8.47 3.16 2.68 Inverted U U2 U2 D2 D1
Central America 442 119 0.3607 6.59 2.67 2.47 4.25 2.02 2.10 2.75 1.41 1.96 3.29 1.59 2.07 U-shape D1 D1 D1 U1
Caribbean 31 658 0.0258 5.95 3.91 1.52 5.06 3.25 1.56 5.82 3.49 1.67 6.29 3.40 1.85 U-shape U3 D2 U1 U3
9. Chile 1 505 486 1.2282 3.92 1.66 2.36 5.05 2.71 1.86 4.48 2.20 2.04 3.39 1.77 1.91 Inverted U D3 U2 D2 D1
14. Ecuador 774 569 0.6319 1.53 1.47 1.04 1.77 1.10 1.61 1.66 1.03 1.61 1.11 1.01 1.10 Inverted U D2 U3 D2 D1
16. Brazil 630 200 0.5141 1.60 1.07 1.50 8.19 4.07 2.02 7.93 4.22 1.88 5.13 3.29 1.56 Inverted U U1 U1 D3 D1
18. United States of
448 535 0.3659 5.08 2.15 2.36 4.82 2.52 1.91 5.76 2.25 2.57 6.60 2.35 2.81 U-shape U1 D3 U3 U1
America
23. Mexico 278 694 0.2274 5.80 2.53 2.29 3.55 1.64 2.17 2.14 1.24 1.73 2.78 1.36 2.05 U-shape D1 D1 D1 U1
29. Colombia 179 351 0.1463 3.26 1.65 1.97 5.52 3.42 1.61 4.51 2.92 1.54 5.25 4.60 1.14 N-shape U2 U2 D1 U2
30. Canada 171 007 0.1395 6.75 3.82 1.77 4.40 2.50 1.76 3.60 2.07 1.74 3.15 1.98 1.59 Straight down D1 D1 D1 D1
32. Peru 143 830 0.1173 3.17 1.46 2.17 2.75 1.11 2.47 2.75 1.14 2.42 3.62 1.23 2.93 U-shape U3 D2 U2 U1
44. Honduras 71 151 0.0580 1.20 1.02 1.18 1.62 1.00 1.62 1.96 1.00 1.96 1.99 1.00 1.99 Straight up U3 U3 U3 U3
46. Venezuela (Bolivarian
53 609 0.0437 4.26 1.42 3.00 3.03 1.51 2.01 2.34 1.31 1.79 1.85 1.05 1.77 Straight down D1 D3 D1 D1
Republic of)
54. Guatemala 33 651 0.0275 1.86 1.15 1.61 5.12 2.55 2.00 1.18 1.01 1.17 1.98 1.02 1.95 N-shape U3 U1 D1 U1
56. Nicaragua 29 410 0.0240 1.27 1.03 1.23 1.09 1.06 1.03 1.11 1.00 1.11 1.00 1.00 1.00 Inverted N D1 D3 U3 D3
SIDS 57. Cuba 26 200 0.0214 3.98 2.62 1.52 3.25 2.07 1.57 4.00 2.39 1.68 3.97 2.09 1.90 Inverted N D2 D2 U1 D2
67. Costa Rica 16 269 0.0133 2.65 1.27 2.08 1.99 1.33 1.50 2.02 1.40 1.45 2.12 1.42 1.49 U-shape D3 D3 U2 U1
LLDCs 73. Paraguay 14 100 0.0115 2.79 2.79 1.00 4.85 4.85 1.00 2.09 2.09 1.00 2.83 2.83 1.00 N-shape U2 U2 D2 U2
83. El Salvador 8 500 0.0069 1.83 1.07 1.71 2.12 1.23 1.72 1.38 1.01 1.37 1.51 1.01 1.49 N-shape D1 U1 D1 U1
97. Panama 3 879 0.0032 2.07 1.54 1.35 2.02 1.01 2.00 2.01 1.29 1.56 3.04 1.54 1.98 U-shape U3 D2 D3 U1
100. Bolivia (Plurinational
LLDCs 3 720 0.0030 2.93 2.93 1.00 1.78 1.78 1.00 3.88 3.88 1.00 5.09 5.09 1.00 U-shape U2 D2 U2 U2
State of)
SIDS 108. Dominican Republic 2 680 0.0022 5.01 2.51 2.00 3.20 1.84 1.74 3.96 1.98 2.00 7.67 5.26 1.46 U-shape U2 D1 U1 U2
113. Argentina 2 085 0.0017 1.00 1.00 1.00 2.71 1.97 1.38 3.20 2.32 1.38 3.12 3.00 1.04 Inverted U U1 U1 U1 D3
SIDS 121. Haiti 1 560 0.0013 1.76 1.76 1.00 1.38 1.38 1.00 1.52 1.52 1.00 U-shape D2 U2

43
 Country or area: U1: ENS increase (both components up)
 Regional aquaculture in italic Diversity measures U2: ENS increase (only WGENS up)
 National aquaculture indexed by Aquaculture production ENS = Effective number of species (a measure of overall species diversity); WGENS = Within-group ENS (a measure of within-group diversity);
U3: ENS increase (only ENSG up)
production ranking in 2020 ENSG = Effective number of species groups (a measure of between-group diversity); ENS = WGENS × ENSG
D1: ENS decrease (both components down)
 LDLK = Landlocked D2: ENS decrease (only WGENS down)
 LLDCs = Landlocked Developing Year 2020 Year 1990 Year 2000 Year 2010 Year 2020 D3: ENS decrease (only ENSG down)
Countries
 ISLD = Island Thousand % of world Overall: 1990– 2000– 2010–
ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG
 SIDS = Small Island Developing States tonnes production 1990–2020 2000 2010 2020
SIDS 130. Jamaica 918 0.0007 1.08 1.01 1.06 1.02 1.00 1.02 1.19 1.01 1.18 1.05 1.02 1.03 Inverted N D3 D1 U1 D3
SIDS 141. Belize 565 0.0005 1.00 1.00 1.00 1.00 1.00 1.00 1.43 1.04 1.38 1.47 1.00 1.47 Straight up U3 U1 U3
SIDS 153. Guyana 138 0.0001 1.96 1.00 1.96 3.84 2.15 1.79 4.95 3.60 1.37 2.00 1.17 1.72 Inverted U U2 U2 U2 D2
157. Uruguay 103 0.0001 1.00 1.00 1.00 1.98 1.54 1.29 1.59 1.59 1.00 3.27 3.27 1.00 N-shape U2 U1 D3 U2
SIDS 159. Saint Lucia 92 0.0001 1.00 1.00 1.00 1.00 1.00 1.00 2.11 1.00 2.11 1.44 1.00 1.44 Inverted U U3 U3 D3
SIDS 162. Guadeloupe 45 0.0000 1.00 1.00 1.00 1.00 1.00 1.00 2.36 1.23 1.93 1.53 1.27 1.20 Inverted U U1 U1 D3
SIDS 163. Martinique 42 0.0000 2.17 1.09 1.98 2.75 1.42 1.94 2.47 1.96 1.26 1.68 1.11 1.51 Inverted U D3 U2 D3 D2
SIDS 165. Suriname 37 0.0000 1.94 1.00 1.94 1.18 1.00 1.18 2.54 1.42 1.79 U-shape D3 U1
SIDS 168. Barbados 26 0.0000 1.75 1.00 1.75 1.10 1.00 1.10 D3
SIDS 170. Grenada 23 0.0000 1.75 1.00 1.75 1.11 1.00 1.11
SIDS 174. Antigua and Barbuda 20 0.0000 1.00 1.00 1.00 1.00 1.00 1.00 2.00 1.00 2.00 U3
SIDS 176. Puerto Rico 18 0.0000 1.13 1.00 1.13 1.79 1.30 1.38 1.97 1.14 1.73 2.65 1.40 1.89 Straight up U1 U1 U3 U1
178. Saint Vincent and the
SIDS 13 0.0000 1.00 1.00 1.00
Grenadines
184. United States Virgin
SIDS 8 0.0000 2.00 2.00 1.00 2.83 2.83 1.00 U2
Islands
SIDS 185. Bahamas 6 0.0000 1.58 1.25 1.26 1.00 1.00 1.00 1.00 1.00 1.00 1.57 1.00 1.57 U-shape D2 D1 U3
SIDS 187. Trinidad and Tobago 4 0.0000 1.00 1.00 1.00 1.20 1.20 1.00 3.11 2.65 1.17 1.73 1.73 1.00 Inverted U U2 U2 U1 D1
190. French Guiana 2 0.0000 1.00 1.00 1.00 1.63 1.00 1.63 2.83 1.61 1.75 2.00 2.00 1.00 Inverted U U2 U3 U1 D3
SIDS 192. Saint Kitts and Nevis 2 0.0000 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 2.00 1.00 2.00 U3 U3
SIDS 193. Aruba 2 0.0000 1.00 1.00 1.00
SIDS 194. British Virgin Islands 1 0.0000 1.00 1.00 1.00
196. Bonaire, Sint
ISLD 0.2 0.0000 1.00 1.00 1.00
Eustatius and Saba
Europe 3 284 407 2.6794 9.78 4.76 2.06 10.22 5.12 2.00 8.19 4.64 1.76 7.29 4.30 1.69 Inverted U D1 U2 D1 D1
Northern Europe 1 955 977 1.5957 2.72 2.09 1.30 2.11 1.70 1.24 1.97 1.61 1.22 1.67 1.48 1.13 Straight down D1 D1 D1 D1
Southern Europe 610 447 0.4980 5.49 2.89 1.90 6.79 3.46 1.96 8.05 3.92 2.05 8.05 3.90 2.06 Straight up U1 U1 U1 U3
Eastern Europe 447 650 0.3652 2.37 2.22 1.07 4.52 4.10 1.10 5.39 5.07 1.06 10.44 6.45 1.62 Straight up U1 U1 U2 U1
Western Europe 270 332 0.2205 4.57 2.66 1.72 5.32 2.95 1.80 6.35 3.37 1.89 5.93 3.20 1.85 Inverted U U1 U1 U1 D1
10. Norway 1 490 412 1.2159 1.17 1.16 1.00 1.43 1.42 1.01 1.41 1.39 1.01 1.31 1.29 1.01 Inverted U U1 U1 D2 D1
22. Russian Federation 291 194 0.2376 2.02 1.81 1.11 4.67 3.81 1.23 5.45 5.03 1.08 10.03 5.29 1.89 Straight up U1 U1 U2 U1
25. Spain 276 571 0.2256 1.91 1.29 1.49 2.41 1.50 1.61 2.97 1.61 1.84 2.97 1.61 1.85 Inverted U U1 U1 U1 D2
ISLD 27. United Kingdom 221 000 0.1803 2.44 1.96 1.25 1.79 1.35 1.33 2.13 1.38 1.54 1.69 1.34 1.26 Inverted N D2 D2 U1 D1
28. France 191 350 0.1561 4.03 2.49 1.62 4.59 2.69 1.71 5.36 3.02 1.77 4.62 2.60 1.77 Inverted U U1 U1 U1 D2
33. Greece 131 737 0.1075 4.41 2.26 1.95 3.80 2.16 1.76 3.45 2.30 1.50 3.69 2.40 1.54 U-shape D3 D1 D3 U1
36. Italy 122 778 0.1002 3.96 1.87 2.12 4.43 2.23 1.99 4.80 2.52 1.90 4.51 2.31 1.95 Inverted U U2 U2 U2 D2
ISLD 42. Faeroe Islands 89 055 0.0727 1.03 1.03 1.00 1.17 1.17 1.00 1.20 1.20 1.00 1.01 1.00 1.01 Inverted U D2 U2 U2 D2
47. Poland 47 700 0.0389 1.67 1.67 1.00 2.28 2.28 1.00 2.85 2.85 1.00 3.69 3.69 1.00 Straight up U1 U2 U2 U1
49. Denmark 42 629 0.0348 1.13 1.13 1.00 1.31 1.30 1.00 1.62 1.30 1.24 2.24 1.46 1.53 Straight up U1 U1 U3 U1
ISLD 51. Iceland 40 595 0.0331 1.23 1.23 1.00 2.07 2.01 1.03 3.45 3.32 1.04 1.65 1.65 1.00 Inverted U U1 U1 U1 D1
52. Netherlands 39 940 0.0326 1.14 1.07 1.07 1.61 1.21 1.34 1.95 1.39 1.40 2.18 1.46 1.49 Straight up U1 U1 U1 U1
ISLD 53. Ireland 37 751 0.0308 2.41 1.34 1.80 3.13 1.60 1.95 3.18 1.65 1.93 3.29 1.69 1.95 Straight up U1 U1 U2 U1
55. Germany 32 258 0.0263 3.51 1.88 1.87 3.56 1.84 1.93 4.15 2.86 1.45 5.42 2.73 1.99 Straight up U1 U3 U2 U3
60. Croatia 21 718 0.0177 7.30 4.65 1.57 7.14 4.84 1.48 5.12 4.58 1.12 Straight down D3 D1
LDLK 62. Czechia 20 401 0.0166 1.87 1.87 1.00 1.96 1.96 1.00 2.09 2.09 1.00 Straight up U2 U2
ISLD 63. Malta 19 829 0.0162 1.89 1.89 1.00 1.48 1.48 1.00 2.00 2.00 1.00 1.71 1.71 1.00 Inverted N D2 D2 U2 D2
64. Ukraine 18 568 0.0151 1.64 1.64 1.00 2.57 2.56 1.00 1.89 1.85 1.02 3.08 3.08 1.00 N-shape U2 U1 D2 U2
LDLK 65. Hungary 18 373 0.0150 2.36 2.36 1.00 3.04 3.04 1.00 2.97 2.97 1.00 3.05 3.05 1.00 N-shape U2 U2 D2 U2
69. Finland 15 053 0.0123 1.08 1.08 1.00 1.07 1.07 1.00 1.31 1.31 1.00 1.26 1.26 1.00 Inverted N U2 D2 U2 D2
70. Bulgaria 15 052 0.0123 1.93 1.93 1.00 5.37 5.27 1.02 6.02 4.43 1.36 6.24 4.11 1.52 Straight up U1 U1 U3 U3
72. Portugal 14 552 0.0119 3.79 1.90 2.00 6.15 3.09 1.99 5.94 3.01 1.97 6.49 3.28 1.98 N-shape U2 U2 D1 U1

44
 Country or area: U1: ENS increase (both components up)
 Regional aquaculture in italic Diversity measures U2: ENS increase (only WGENS up)
 National aquaculture indexed by Aquaculture production ENS = Effective number of species (a measure of overall species diversity); WGENS = Within-group ENS (a measure of within-group diversity);
U3: ENS increase (only ENSG up)
production ranking in 2020 ENSG = Effective number of species groups (a measure of between-group diversity); ENS = WGENS × ENSG
D1: ENS decrease (both components down)
 LDLK = Landlocked D2: ENS decrease (only WGENS down)
 LLDCs = Landlocked Developing Year 2020 Year 1990 Year 2000 Year 2010 Year 2020 D3: ENS decrease (only ENSG down)
Countries
 ISLD = Island Thousand % of world Overall: 1990– 2000– 2010–
ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG
 SIDS = Small Island Developing States tonnes production 1990–2020 2000 2010 2020
LLDCs 75. Republic of Moldova 12 600 0.0103 1.99 1.99 1.00 5.27 5.08 1.04 4.57 4.49 1.02 4.17 4.16 1.00 Inverted U U1 U1 D1 D1
76. Romania 12 200 0.0100 4.49 4.49 1.00 5.34 5.34 1.00 7.01 7.01 1.00 5.93 5.93 1.00 Inverted U U2 U2 U2 D2
77. Sweden 12 090 0.0099 2.14 1.45 1.47 2.17 1.58 1.37 2.20 1.50 1.47 2.24 1.38 1.63 Straight up U3 U2 U3 U3
81. Albania 9 284 0.0076 1.58 1.13 1.40 2.14 1.09 1.97 3.72 1.83 2.03 3.36 2.93 1.15 Inverted U U2 U3 U1 D3
LDLK 82. Belarus 9 266 0.0076 1.03 1.03 1.00 1.68 1.68 1.00 2.20 2.20 1.00 2.48 2.48 1.00 Straight up U2 U2 U2 U2
LDLK 90. Serbia 6 010 0.0049 2.38 2.38 1.00 2.30 2.30 1.00 D2
95. Austria 4 527 0.0037 1.89 1.89 1.00 2.60 2.59 1.00 4.18 4.18 1.00 7.02 7.02 1.00 Straight up U2 U1 U2 U2
96. Lithuania 4 477 0.0037 1.07 1.07 1.00 1.22 1.22 1.00 1.52 1.52 1.00 2.85 2.85 1.00 Straight up U2 U2 U2 U2
99. Bosnia and
3 776 0.0031 3.39 3.22 1.05 1.89 1.89 1.00 D1
Herzegovina
LDLK 112. Slovakia 2 296 0.0019 1.85 1.85 1.00 2.11 2.11 1.00 3.82 3.82 1.00 Straight up U2 U2
LDLK 116. Switzerland 2 048 0.0017 1.55 1.55 1.00 1.15 1.15 1.00 1.73 1.73 1.00 3.58 3.32 1.08 U-shape U1 D2 U2 U1
118. Slovenia 1 673 0.0014 2.78 2.37 1.17 3.64 2.63 1.38 4.38 2.52 1.74 Straight up U1 U3
LLDCs 119. North Macedonia 1 634 0.0013 2.64 2.64 1.00 1.72 1.72 1.00 2.13 2.13 1.00 U-shape D2 U2
127. Channel Islands 1 107 0.0009 1.30 1.30 1.00 1.81 1.81 1.00 1.65 1.62 1.02 1.35 1.35 1.00 Inverted U U2 U2 D2 D1
129. Estonia 1 090 0.0009 1.97 1.97 1.00 1.67 1.67 1.00 1.82 1.81 1.01 1.85 1.84 1.01 U-shape D2 D2 U1 U1
132. Montenegro 885 0.0007 3.16 1.91 1.66 2.87 1.59 1.81 D2
137. Latvia 717 0.0006 1.00 1.00 1.00 1.80 1.80 1.00 2.51 2.50 1.00 2.51 2.51 1.00 Inverted U U2 U2 U1 D3
149. Belgium 209 0.0002 2.50 2.50 1.00 5.15 5.15 1.00 2.42 1.22 1.99 1.00 1.00 1.00 Inverted U D2 U2 D2 D1
Africa 2 354 294 1.9206 7.92 4.95 1.60 7.65 4.93 1.55 7.88 5.50 1.43 7.00 5.61 1.25 Inverted N D3 D1 U2 D3
Northern Africa 1 632 296 1.3316 3.80 3.65 1.04 4.13 4.09 1.01 4.04 4.00 1.01 4.00 3.94 1.02 Inverted U U2 U2 D2 D2
Sub-Saharan Africa 731 848 0.5970 9.77 3.81 2.56 8.17 3.30 2.47 7.63 3.88 1.96 8.60 5.25 1.64 U-shape D3 D1 D3 U2
Eastern Africa 352 897 0.2879 3.61 1.75 2.06 2.92 1.44 2.03 4.15 1.92 2.16 5.35 2.73 1.96 U-shape U2 D1 U1 U2
Western Africa 345 674 0.2820 5.45 5.21 1.05 8.76 8.58 1.02 6.14 6.11 1.01 6.71 6.61 1.01 N-shape U2 U2 D1 U1
Southern Africa 12 920 0.0105 4.67 2.27 2.06 5.22 2.23 2.34 7.56 2.55 2.97 5.32 1.78 2.98 Inverted U U3 U3 U1 D2
Middle Africa 10 507 0.0086 2.19 2.19 1.00 1.92 1.92 1.00 2.47 2.37 1.04 3.12 2.99 1.04 U-shape U1 D2 U1 U1
8. Egypt 1 591 896 1.2987 3.27 3.27 1.00 3.95 3.95 1.00 3.90 3.88 1.01 3.77 3.73 1.01 Inverted U U1 U2 D2 D2
26. Nigeria 261 711 0.2135 4.39 4.39 1.00 6.84 6.84 1.00 5.39 5.39 1.00 5.66 5.64 1.00 N-shape U1 U2 D2 U1
LLDCs 35. Uganda 123 897 0.1011 2.75 2.75 1.00 3.43 3.43 1.00 1.96 1.96 1.00 1.89 1.89 1.00 Inverted U D2 U2 D2 D2
38. United Republic of
108 568 0.0886 1.18 1.00 1.18 1.03 1.00 1.03 1.03 1.01 1.03 1.70 1.09 1.56 U-shape U1 D3 U2 U1
Tanzania
45. Ghana 64 010 0.0522 1.70 1.70 1.00 1.90 1.90 1.00 1.39 1.39 1.00 1.36 1.36 1.00 Inverted U D2 U2 D2 D2
LLDCs 48. Zambia 45 670 0.0373 2.93 2.93 1.00 2.88 2.88 1.00 3.54 3.54 1.00 3.21 3.21 1.00 Inverted N U2 D2 U2 D2
58. Tunisia 23 486 0.0192 6.92 4.09 1.69 7.34 5.72 1.28 5.45 4.75 1.15 2.95 2.82 1.04 Inverted U D1 U2 D1 D1
61. Kenya 20 831 0.0170 4.61 2.86 1.61 3.54 3.54 1.00 2.14 2.09 1.02 2.33 1.96 1.19 U-shape D1 D3 D2 U3
LLDCs 68. Zimbabwe 15 425 0.0126 2.20 1.84 1.20 1.22 1.22 1.00 1.15 1.15 1.00 1.03 1.03 1.00 Straight down D1 D1 D2 D2
ISLD 74. Madagascar 13 550 0.0111 1.75 1.50 1.17 2.42 1.00 2.42 3.10 1.04 2.97 2.11 1.00 2.10 Inverted U U3 U3 U1 D1
78. Sudan 9 850 0.0080 1.66 1.66 1.00
79. South Africa 9 753 0.0080 4.39 2.13 2.06 4.47 1.96 2.28 5.86 2.01 2.91 4.66 1.70 2.74 Inverted U U3 U3 U1 D1
LLDCs 80. Malawi 9 393 0.0077 2.52 2.19 1.15 1.32 1.32 1.00 3.10 3.10 1.00 3.56 3.56 1.00 U-shape U2 D1 U2 U2
LLDCs 85. Mali 7 686 0.0063 1.72 1.72 1.00 1.93 1.93 1.00 2.13 2.13 1.00 1.78 1.78 1.00 Inverted U U2 U2 U2 D2
LLDCs 87. Rwanda 7 059 0.0058 1.31 1.31 1.00 1.32 1.32 1.00 1.13 1.13 1.00 1.40 1.39 1.00 N-shape U1 U2 D2 U1
92. Algeria 5 436 0.0044 3.34 2.71 1.23 2.76 2.20 1.25 5.51 5.42 1.02 4.48 3.99 1.12 Inverted N U2 D2 U2 D2
94. Côte d'Ivoire 4 620 0.0038 3.43 3.43 1.00 1.86 1.86 1.00 2.95 2.95 1.00 2.48 2.48 1.00 Inverted N D2 D2 U2 D2
101. Democratic Republic
3 590 0.0029 1.00 1.00 1.00 1.01 1.01 1.00 1.13 1.13 1.00 1.15 1.15 1.00 Straight up U2 U2 U2 U2
of the Congo
102. Cameroon 3 556 0.0029 1.99 1.99 1.00 1.65 1.65 1.00 3.96 3.96 1.00 2.12 2.12 1.00 Inverted N U2 D2 U2 D2
SIDS 104. Mauritius 3 298 0.0027 3.64 1.44 2.53 3.07 1.40 2.19 1.53 1.43 1.07 1.15 1.11 1.04 Straight down D1 D1 D3 D1
105. Mozambique 3 162 0.0026 1.00 1.00 1.00 1.84 1.08 1.71 1.00 1.00 1.00 D1
107. Benin 2 987 0.0024 1.96 1.96 1.00 2.00 2.00 1.00 U2
LLDCs 109. Lesotho 2 600 0.0021 2.08 2.08 1.00 1.00 1.00 1.00 1.01 1.01 1.00 1.00 1.00 1.00 Inverted N D2 D2 U2 D2

45
 Country or area: U1: ENS increase (both components up)
 Regional aquaculture in italic Diversity measures U2: ENS increase (only WGENS up)
 National aquaculture indexed by Aquaculture production ENS = Effective number of species (a measure of overall species diversity); WGENS = Within-group ENS (a measure of within-group diversity);
U3: ENS increase (only ENSG up)
production ranking in 2020 ENSG = Effective number of species groups (a measure of between-group diversity); ENS = WGENS × ENSG
D1: ENS decrease (both components down)
 LDLK = Landlocked D2: ENS decrease (only WGENS down)
 LLDCs = Landlocked Developing Year 2020 Year 1990 Year 2000 Year 2010 Year 2020 D3: ENS decrease (only ENSG down)
Countries
 ISLD = Island Thousand % of world Overall: 1990– 2000– 2010–
ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG ENS WGENS ENSG
 SIDS = Small Island Developing States tonnes production 1990–2020 2000 2010 2020
114. Angola 2 062 0.0017 1.00 1.00 1.00 1.15 1.15 1.00 1.08 1.08 1.00 Inverted U U2 D2
120. Morocco 1 618 0.0013 4.01 2.03 1.98 6.85 4.88 1.40 5.68 2.90 1.96 6.51 2.66 2.45 N-shape U1 U2 D2 U3
LLDCs 123. Burundi 1 450 0.0012 1.00 1.00 1.00 1.18 1.18 1.00 1.43 1.43 1.00 1.30 1.30 1.00 Inverted U U2 U2 U2 D2
126. Guinea 1 135 0.0009 1.00 1.00 1.00 1.96 1.96 1.00 3.71 3.71 1.00 U2 U2
128. Senegal 1 100 0.0009 1.82 1.00 1.82 1.41 1.05 1.34 3.30 1.52 2.18 3.67 1.84 2.00 U-shape U1 D3 U1 U2
131. Congo 900 0.0007 1.00 1.00 1.00 1.00 1.00 1.00 2.49 2.49 1.00 1.60 1.60 1.00 Inverted U U2 U2 D2
136. Togo 730 0.0006 1.00 1.00 1.00 1.00 1.00 1.00 2.13 2.13 1.00 1.31 1.31 1.00 Inverted U U2 U2 D2
LLDCs 138. Burkina Faso 651 0.0005 2.00 2.00 1.00 1.00 1.00 1.00 2.56 1.46 1.75 2.33 2.22 1.05 Inverted N U1 D2 U1 D3
LLDCs 139. Niger 649 0.0005 1.00 1.00 1.00 1.00 1.00 1.00 1.82 1.82 1.00 1.92 1.92 1.00 Straight up U2 U2 U2
LLDCs 142. Ethiopia 534 0.0004 1.69 1.69 1.00 3.75 3.75 1.00 2.96 2.96 1.00 2.17 2.17 1.00 Inverted U U2 U2 D2 D2
146. Namibia 321 0.0003 1.00 1.00 1.00 3.59 1.38 2.60 4.15 2.01 2.07 1.61 1.12 1.43 Inverted U U1 U1 U2 D1
147. Liberia 255 0.0002 3.35 3.35 1.00 1.69 1.69 1.00 1.48 1.48 1.00 Straight down D2 D2
148. Central African
LLDCs 219 0.0002 1.21 1.21 1.00 1.00 1.00 1.00 3.33 2.09 1.59 4.86 2.84 1.71 U-shape U1 D2 U1 U1
Republic
LLDCs 152. Botswana 146 0.0001 1.76 1.76 1.00
LLDCs 156. Chad 105 0.0001 1.57 1.57 1.00 3.42 2.10 1.63 U1
LLDCs 158. Eswatini 100 0.0001 1.00 1.00 1.00 4.24 3.27 1.30 1.00 1.00 1.00 1.00 1.00 1.00 U1 D1
160. Sierra Leone 85 0.0001 1.00 1.00 1.00 1.00 1.00 1.00 1.28 1.28 1.00 1.25 1.25 1.00 Inverted U U2 U2 D2
161. Gabon 60 0.0000 1.00 1.00 1.00 1.24 1.24 1.00 1.31 1.31 1.00 1.33 1.33 1.00 Straight up U2 U2 U2 U2
166. Gambia 33 0.0000 1.00 1.00 1.00 2.87 2.87 1.00 2.04 1.23 1.65 2.56 1.31 1.96 N-shape U1 U2 D2 U1
LLDCs 167. South Sudan 30 0.0000 1.28 1.28 1.00
SIDS 171. Cabo Verde 22 0.0000 1.00 1.00 1.00
ISLD 175. Mayotte 20 0.0000 1.00 1.00 1.00 1.64 1.64 1.00 1.00 1.00 1.00 Inverted U U2 D2
177. Equatorial Guinea 15 0.0000 1.96 1.96 1.00 2.69 2.69 1.00 U2
180. Libya 10 0.0000 1.00 1.00 1.00 2.95 2.95 1.00 2.55 2.55 1.00 1.00 1.00 1.00 Inverted U U2 D2 D2
ISLD 181. Réunion 10 0.0000 1.26 1.00 1.26 2.43 2.20 1.11 2.96 2.96 1.00 2.00 2.00 1.00 Inverted U U2 U2 U2 D2
Oceania 239 790 0.1956 5.87 2.23 2.64 5.79 2.16 2.68 7.56 2.65 2.85 6.26 2.22 2.82 Inverted N U3 D2 U1 D1
Australia and New Zealand 224 670 0.1833 4.43 2.65 1.67 4.06 2.16 1.88 5.66 2.52 2.25 5.06 2.13 2.37 Inverted N U3 D2 U1 D2
Pacific islands 15 120 0.0123 1.51 1.04 1.45 2.59 1.12 2.31 5.09 1.96 2.59 3.54 1.30 2.72 Inverted U U1 U1 U1 D2
ISLD 37. New Zealand 118 582 0.0967 1.74 1.29 1.35 1.53 1.18 1.29 1.60 1.12 1.43 1.57 1.07 1.47 Inverted N D2 D1 U3 D2
39. Australia 106 088 0.0865 6.24 2.77 2.25 7.61 2.83 2.69 7.19 2.91 2.47 4.43 2.14 2.07 Inverted U D1 U1 D3 D1
SIDS 89. Papua New Guinea 6 102 0.0050 1.82 1.82 1.00 1.21 1.21 1.00 2.86 2.10 1.36 2.30 1.25 1.84 Inverted N U3 D2 U1 D2
SIDS 91. Solomon Islands 5 504 0.0045 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.01 1.00 1.01 Straight up U3 U3 U3
SIDS 122. New Caledonia 1 478 0.0012 1.51 1.00 1.51 1.11 1.02 1.09 1.25 1.02 1.23 1.03 1.00 1.03 Inverted N D3 D3 U1 D1
SIDS 124. French Polynesia 1 371 0.0011 2.43 1.59 1.53 1.48 1.13 1.31 1.14 1.04 1.09 1.54 1.04 1.48 U-shape D1 D1 D1 U3
SIDS 145. Fiji 322 0.0003 1.06 1.00 1.06 2.09 1.01 2.07 1.94 1.00 1.94 3.28 1.17 2.80 N-shape U1 U1 D1 U1
SIDS 154. Guam 118 0.0001 2.50 2.50 1.00 3.01 2.14 1.41 2.85 2.09 1.36 2.79 1.94 1.44 Inverted U U3 U3 D1 D2
SIDS 155. Tonga 105 0.0001 3.13 1.28 2.44 1.01 1.00 1.01 1.01 1.01 1.00 U-shape D1 U2
164. Northern Mariana
SIDS 41 0.0000 1.63 1.00 1.63 2.21 1.10 2.00 U1
Islands
SIDS 172. Palau 22 0.0000 1.00 1.00 1.00 1.00 1.00 1.00 2.71 1.47 1.85 2.53 2.53 1.00 Inverted U U2 U1 D3
SIDS 173. American Samoa 20 0.0000 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00 1.00
SIDS 179. Samoa 10 0.0000 1.00 1.00 1.00 1.01 1.00 1.01 U1
SIDS 182. Marshall Islands 10 0.0000 1.58 1.00 1.58
SIDS 183. Cook Islands 9 0.0000 1.00 1.00 1.00 2.92 2.03 1.44 U1
SIDS 188. Vanuatu 3 0.0000 1.73 1.00 1.73 1.99 1.23 1.62 U2
SIDS 189. Tuvalu 3 0.0000 1.00 1.00 1.00 1.00 1.00 1.00
SIDS 191. Kiribati 2 0.0000 1.04 1.00 1.04 1.01 1.00 1.01 1.02 1.00 1.02 1.00 1.00 1.00 Inverted N D1 D1 U3 D3
SIDS 195. Nauru 1 0.0000 1.00 1.00 1.00

46