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1 The Living Planet Index for Global Estuarine Systems -Technical Report- © Simon Ledingham – Channel of River Wampool, Solway Estuary, Cumbria, U.K. May 2010 Report Authors: Stefanie Deinet 1 , Louise McRae 1 , Adriana De Palma 1 , Robyn Manley 1 , Jonathan Loh 1,2 and Ben Collen 1 1 Indicators and Assessments Unit, Institute of Zoology, Zoological Society of London, U.K. 2 WWF International, Gland, Switzerland Contact: Dr Ben Collen Indicators and Assessments Unit, Institute of Zoology, Zoological Society of London Regent's Park, London NW1 4RY e: [email protected]
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Page 1: The Living Planet Index for Global Estuarine Systems ...€¦ · 3 EXECUTIVE SUMMARY Estuaries are some of the most productive ecosystems in the world, playing an important role for

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The Living Planet Index for Global Estuarine Systems

-Technical Report-

© Simon Ledingham – Channel of River Wampool, Solway Estuary, Cumbria, U.K.

May 2010 Report Authors: Stefanie Deinet1, Louise McRae1, Adriana De Palma1, Robyn Manley1, Jonathan Loh1,2 and Ben Collen1 1

Indicators and Assessments Unit, Institute of Zoology, Zoological Society of London, U.K. 2 WWF International, Gland, Switzerland

Contact: Dr Ben Collen Indicators and Assessments Unit, Institute of Zoology, Zoological Society of London Regent's Park, London NW1 4RY e: [email protected]

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TABLE OF CONTENTS

TITLE PAGE ............................................................................................................................... 1

TABLE OF CONTENTS................................................................................................................. 2

EXECUTIVE SUMMARY.............................................................................................................. 3

1. INTRODUCTION.................................................................................................................... 5

1.1 ESTUARIES................................................................................................................................. 5

1.2 MEASURING CHANGE IN VERTEBRATE ABUNDANCE: THE LIVING PLANET INDEX.................... 6

1.3 INTERPRETING THE LPI.............................................................................................................. 8

2. RESULTS AND DISCUSSION ................................................................................................. 10

2.1 THE GLOBAL ESTUARINE INDEX .............................................................................................. 10

2.2 TEMPERATE AND TROPICAL ESTUARINE SPECIES ................................................................... 10

2.3 INTERPRETATION OF THE GLOBAL, TEMPERATE AND TROPICAL ESTUARINE INDICES ........... 13

2.4 CAVEATS.................................................................................................................................. 15

2.4.1 Baseline ........................................................................................................................... 15

2.4.2 Regional and taxonomic bias .......................................................................................... 15

2.4.3 Taxonomic representation .............................................................................................. 17

2.5 TAXONOMIC CLASS INDICES ................................................................................................... 19

2.5.1 Summary ......................................................................................................................... 19

2.5.2 Birds................................................................................................................................. 19

2.5.3 Fish .................................................................................................................................. 21

2.5.4 Tropical and South African fish ....................................................................................... 22

2.5.5 Temperate fish ................................................................................................................ 24

2.6 OTHER INDICES ....................................................................................................................... 24

2.6.1 Estuary type..................................................................................................................... 25

2.6.2 Drivers of change............................................................................................................. 26

2.6.3 Estuarine-dependent fish ................................................................................................ 28

2.6.4 Human population density .............................................................................................. 29

2.6.5 Income group and Human Development Index............................................................... 30

CONCLUSIONS ........................................................................................................................ 32

ACKNOWLEDGEMENTS ........................................................................................................... 32

REFERENCES ........................................................................................................................... 33

APPENDIX............................................................................................................................... 40

METHODS ...................................................................................................................................... 40

Data collection .......................................................................................................................... 40

Estuary type and drivers of change........................................................................................... 40

GROMS classification ................................................................................................................ 41

Human population density, income group and HDI .................................................................. 41

Analysis ..................................................................................................................................... 41

Baseline and end year ............................................................................................................... 42

TABLES........................................................................................................................................... 43

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EXECUTIVE SUMMARY

Estuaries are some of the most productive ecosystems in the world, playing an important

role for animals and humans alike. However, they are under threat from a range of man-

made interventions, which interfere with their natural functioning.

To assess the extent to which biodiversity is changing in estuarine ecosystems, and

whether any factors can be identified as possible causes for any such loss, changes in

vertebrate population abundance over time were assessed using the same method as is

used in the calculation of the Global Living Planet Index.

The findings of this analysis were:

- Global estuarine abundance increase by 16% over the 25 year span of this study.

- Temperate estuarine species show a near two-and-a-half-fold increase in population

abundance by 2005, with temperate fish populations doubling over the same period.

- Bird populations, especially from temperate regions, appear to be in a good state

relative to 1980 levels, with abundance more than doubling by 2005.

- By contrast, dramatic declines in abundance are apparent in tropical systems (43%

decline by 2005) and tropical fish species (74% decline by 2002), though overall

global estuarine fish populations remain steady around the baseline.

- Populations of specialist fish species, which are highly dependent on estuarine

systems, decrease by 20% between 1980 and 2002.

- Dams and water extraction are associated with a decrease in vertebrate abundance

of 75% and 43% respectively, while populations from estuarine locations listed as

not threatened quadruple in abundance by 2002.

- Estuarine wildlife populations from areas of high human population density decrease

more than those from areas of low human population density, but this is mediated by

a regional effect, with tropical abundance trends decreasing and temperate trends

increasing. An interaction was also found for region and income group, with high

income temperate populations showing the largest increase and low income tropical

populations showing the largest decline.

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This analysis represents an initial review of the potential for trends in vertebrate population

abundance to inform the status and trends of biodiversity in estuarine systems. Due to

restricted sample sizes and taxonomic and regional inequalities in the dataset, this sets a

baseline against which more data can be collected in order to build a comprehensive

picture of how different factors might be affecting population abundance, and therefore

biodiversity, in estuarine systems around the world.

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1. INTRODUCTION

1.1 Estuaries

One of the most productive ecosystems in the world (Allen 1982), estuaries are home to a

wide variety of organisms, provide important ecosystem services to humans such as

nutrient cycling and food production (Costanza et al. 1997), and a range of habitats for a

range of species. This, in combination with their role in successful protection against the

seas and oceans, makes them of utmost value to humans. Despite their importance,

estuarine systems are amongst the most modified and threatened ecosystems in the world

(Blaber et al. 2000), suffering from anthropogenic impacts that interfere with both riverine

and marine influences necessary for natural ecological functioning.

For example, only 12% of the world’s 177 longest rivers run freely from the source to the

sea (Wong et al. 2007). Dams and dikes have been constructed in most river systems

globally, and these have both direct (landward) and indirect (seaward) effects (Hood

2004). Trapping silt in reservoirs deprives downstream deltas and estuaries of

maintenance materials and nutrients that help make them productive ecosystems. In

addition, they represent barriers that reduce or altogether inhibit movement of migratory

species between freshwater and marine systems, causing extirpation or extinction of

genetically distinct stocks or species (McAllister et al. 2001). Dredging, along with

widening and straightening of channels for navigation, and water extraction for agricultural

use and the production of hydropower result in disturbance in sedimentation and water

circulation processes (Hossain et al. 2004). Resulting ecosystem losses and declines in

biodiversity caused by erosion, increased coastal inundation and landward retreat are

exacerbated by increased predation and competition from invasive species, and from

pollution and overexploitation. Although so far human interventions often outweigh climate

change in their detrimental effect on coastal systems (Chust et al. 2009), it is known that

disturbance in natural processes limits the estuarine system health and viability (Goldberg

1995), making deltas and estuaries all the more vulnerable to the consequences of climate

change. For example, rising sea levels through loss of sea ice and thawing of permafrost,

and extreme weather events such as cyclones and flooding are likely to be future threats

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to estuarine systems (Nicholls et al. 2007). Mangroves are expected to show mixed

responses to climate change based on site-specific factors, including enhanced growth

through higher CO2 levels and temperature, as well as negative responses such as

increased saline intrusion and erosion causing landward migration into adjacent wetland

communities, as has already been recorded in the Florida Everglades in the last 50 years

(Ross et al. 2000).

In 2001, over half the world's human population lived within 200 km of a coastline (Reker

et al. 2006), and was thus directly reliant on the productivity of estuaries. However, an

even greater proportion may depend on estuarine ecosystems globally, not just for

subsistence and livelihoods, but also for infrastructure. There is a well documented

economic value of coastal systems in general (Raheem et al. 2009), and of estuaries for

fish species dependent on this habitat in particular (Able 2005), making man perhaps the

most estuarine-dependent organism in the biosphere (Allanson 1980). The degradation of

estuarine ecosystems would undoubtedly have far-reaching socioeconomic impacts on

hundreds of millions of people (Nicholls et al. 2007), and it is thus of utmost importance to

restore natural processes in altered systems and preserve those systems that remain

largely intact.

1.2 Measuring change in abundance: The Living Planet Index

The Living Planet Index (LPI) is a tool originally developed by WWF for the Living Planet

Report in 1998 to quantify the loss of global biodiversity. The underlying dataset has since

been expanded for subsequent Living Planet Reports, initially by UNEP-WCMC, and

subsequently, along with further methodological developments, by the Zoological Society

of London’s Institute of Zoology (Loh et al. 2005, Collen et al. 2009). This biodiversity

measure has been adopted by the Convention on Biological Diversity (CBD) as an

indicator of change in wildlife population size, under the 2010 target focal area Status and

trends of the components of biological diversity and the headline indicator Trends in

abundance and distribution of selected species (UNEP 2006).

The LPI is used to measure global vertebrate abundance trends over time by calculating

the average change in abundance for each year compared with the preceding year, which

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is then chained to the previous average annual population change to make an index,

starting with an initial value of 1 in 1970. It can thus be thought of as a biological analogue

of a stock market index that tracks the value of a set of stocks and shares traded on an

exchange or a retail price index tracks the cost of a basket of consumer goods. More

specifically, it measures trends in the abundance of populations of species, i.e. changes in

the number of individuals within populations. As these population-based trends are

aggregated to a species level from which the overall index is calculated, the LPI can also

be said to track species abundance, i.e. the change in the number of individuals of a

particular species. The link between this population-focused abundance trend monitoring

and trends in global biodiversity is not explicit but only inferred. Population abundance is

an important measure of the state of both species and ecosystems, as populations are

sensitive to short-term changes caused by anthropogenic pressures (Balmford et al.

2003), and the loss of populations is a prelude to species-level extinctions (Collen et al.

2009). If species are lost, this necessarily implies a decrease in taxonomic, genetic and

functional diversity, as biomass in the system is reduced, and genes and ecosystem

services are lost. Abundance trends can thus be used to infer community change

(Buckland et al. 2005) and habitat change (Balmford et al. 2003), as well as the impact of

threats. There are also advantages in terms of practical applications, because of the

relatively short delay between human impact and subsequent vertebrate population

decline gives the opportunity for the implementation of proactive and targeted conservation

action (Collen et al. 2009).

The global LPI is currently calculated using time-series data on almost 7,950 populations

of over 2,500 species of mammal, bird, reptile, amphibian and fish (Loh et al. 2010). Data

are gathered from a variety of published sources; principally scientific journals, but also

government reports, wildlife and other natural resource management authority records, as

well as databases from academic organisations. The data collated comprise estimates of

total population size, density measures and, particularly for fish species, catch per unit

effort (CPUE). Depending on the species in question, a proxy of population size might be

collected, for example the number of nests of marine turtle species on specific nesting

beaches. All population time series have a minimum of two data points collected using the

same or comparable methods, with the mean length of all series being around 20 years

(Collen et al. 2009).

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1.3 Interpreting the LPI

Although the LPI was designed to be a straightforward and easily communicable measure

of biodiversity, it is nevertheless helpful to consider the use of unfamiliar methods in the

context of real-world examples. Figure 1 shows the Global LPI from the Living Planet

Report 2008 (Loh et al. 2008), which is an aggregate of equally weighted temperate and

tropical indices also shown. Due to limited data availability before 1970, the index is set to

a baseline of 1 at this point in time. According to Figure 1, vertebrate populations show an

initial rise in abundance, remaining steady until the early 1980s, after which there is a

continuous steep decline until 2005. Here, the trend has a value of around 0.70, meaning

that abundance is only 70% that of 1970 which, in turns, suggests that within the 35-year

period, there was a 30% loss in abundance of vertebrate populations.

Figure 1. The Global Living Planet Index (thick black line – bars are 95% confidence limits), its component indices for temperate (green line) and tropical (orange line) regions of the world, and the baseline (thin black line) (from Loh et al. 2008)

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When considering indices calculated using the LPI method, it is important to consider a

particular characteristic of the trends produced. Values above 1 indicate a net increase in

vertebrate abundance compared to the baseline, trend values below the baseline a net

decrease. When the trend crosses or follows the baseline, biodiversity is stable and there

is no overall change in abundance.

Knowledge of the precision with which the population trend has been estimated is

important for the interpretation of the results (Fewster et al. 2000), and the LPI offers the

possibility to assess this by examining the confidence interval, for which the desired

number of bootstraps used for resampling can be customised (Loh et al. 2005, Collen et

al. 2009). In addition, the LPI method generates inflection or change points, i.e. landmark

points in the trajectory, highlighting years in which the curvature of the index curve is

statistically significant, as indicated by the second derivative differing significantly from

zero (Fewster et al. 2000). A significantly positive derivative represents a good change for

the underlying set of populations, implying either an increase in the rate of growth or a

decrease in the rate of decline (Fewster et al. 2000). Similarly, a set of populations will

have undergone a decrease in growth rate or increase in the rate of decline if the second

derivative is significantly negative.

Although primarily a measure of global biodiversity, the LPI can be calculated for species

populations from selected regions, biomes or taxonomic groups, depending on data

availability (see METHODS in the Appendix). This report focuses on the calculation of an

index for estuarine species, i.e. populations that can tolerate a range of salinities from

freshwater to brackish water and saltwater. The aim of this project was to assess to what

extent biodiversity has changed in estuaries from 1980, and whether any identifiable trend

can be related to a variety of extrinsic, anthropogenic threats (e.g. climate change and

man-made interventions), or other factors such as estuary type or drivers of change in

brackish coastal waters.

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2. RESULTS AND DISCUSSION

2.1 The global estuarine index

Throughout this study, a concerted effort was made to increase the coverage of estuarine

species populations, especially fish, from the existing LPI dataset. A 66% increase in

coverage resulted in a total sample size of 1,224 estuarine populations. Species numbers

were boosted by the addition of 242 new species, giving a total of 500 species for the

global estuarine analysis. A detailed assessment of the improvements achieved in the

representation of the data is presented in Appendix Table 3. The number of species and

populations used for the calculations of the indices can be found in Appendix Table 5.

The final estuarine dataset comprised data from 110 estuarine locations around the globe

(Figure 2). The global index for species sampled from these locations, which is calculated

by equally weighting the sets of species from temperate and tropical regions, is shown in

Figure 3. While there is a pronounced decline of around 20% through to the mid 1990s,

the overall rate of loss subsequently slows down until recovery starts from 2001. By 2005,

population abundance has increased by 16% (95% CIs: 0.79-1.73). The mean number of

populations per year was 544, while the species split was fairly even with a total of 500

species comprising 45% fish and 52% birds.

2.2 Temperate and tropical estuarine species

The temperate estuarine index shows a consistently increasing trend from 1980 until 2005,

which is slower initially and accelerates from 1985 to slow again after 2001 (Figure 4). The

2005 value of 2.4 (95% CIs: 1.96-2.93) represents an increase in vertebrate abundance of

nearly two and a half times within this time period.

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Figure 2. The distribution of 110 estuaries in the analysis, comprising 1224 populations of 500 species.

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Figure 3. Global estuarine Living Planet Index (blue line) and 95% confidence intervals (grey vertical bars) for n = 1224 populations of n = 500 species. Full circles denote negative change points, empty circles positive change points.

Figure 4. Index and 95% confidence interval (bars) for temperate (green line, n = 335 species) and tropical (orange line, n = 202 species) estuarine populations. Full circles denote negative change points, empty circles positive change points.

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The tropical estuarine index (Figure 4) follows a very different trend. The rate of loss is

reasonably constantly declining at an average of 4.6% of vertebrate abundance per year

until 1994. After some variation between 1995 and 2001, the trend reverses to a steady

increase. But the indicator remains below 1 by 2005, and tropical estuarine populations

show an overall loss in abundance of 43% (95% CIs: 0.28-1.21) over the 25 year period.

Although it is based on a lower number of populations than the temperate index, the

tropical index had a good mean coverage of 85 populations contributing per year.

2.3 Interpretation of the global, temperate and tropical estuarine indices

The confidence interval of the global estuarine index is very wide (95% CIs: 0.79-1.73),

suggesting great variation in the trends within the underlying abundance data. This is likely

to reflect the fact that the species in this index are not homogenous: there are winners and

losers, which are responding in a variety of ways to the changes affecting estuarine

habitats.

Closer examination of the inflection points in the three indices depicted in Figures 3 and 4

reveal that the first year, in which the trajectory of the global index changes significantly,

coincides with the same trend in tropical estuarine species, with the third point matching

that of temperate species. The second change point is present in both the temperate and

tropical component indices. Interestingly, all of these change points are positive,

suggesting a significant upturn in abundance trends above 1 and a slowing rate of decline

below the baseline. The remaining two global change points – a negative change in 2000

followed by a positive change in 2001 – are only reflected in the tropical index. As the

global index is an aggregate of temperate and tropical indices, it is not unusual for it to be

influenced by these composite indices at different points in time.

The positive temperate trend and decreasing tropical trend result in a decreasing trend

globally over the first decade, albeit one which is decreasing at a smaller rate than the

tropical index. After 1989, the increase in abundance is caused by some underlying

tropical but primarily temperate increases, including a number of fish species from the

Sagavanirktok delta in Alaska (Arctic cod, Arctic grayling, Atlantic rainbow smelt, Arctic

flounder), as well as wetland birds from Donana, Ebro, S'Albufera and Llobregat in Spain,

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Merja Zerga in Morocco, and the Camargue (Red knot, Northern shoveler and Common

teal). The pronounced increase from 2001 is reflected both in temperate (Red knot and

Black bullhead) and tropical species (many fish from South Africa; see 2.5.4).

In terms of the increase in temperate species, previous analyses have also shown less of

a decline or, in some cases, an increase in abundance in temperate compared to tropical

regions (e.g. Figure 1). This could be attributable to a number of different factors. Firstly,

temperate regions may simply be less vulnerable to the effects of current threats due to

extinction filter effects (Balmford 1996), or tropical regions may be subject to greater

anthropogenic pressures (Myers 1991, Peres et al. 2006), and therefore abundance

changes. Alternatively, it may be that populations were lost pre-1980 in temperate regions

and are now increasing with concerted conservation effort, e.g. establishment of protected

areas, action plans and managed reintroductions.

Vertebrates in tropical systems are highly threatened by habitat degradation and loss

through, for example, large-scale deforestation for agricultural purposes (e.g. Geist &

Lambin 2001). Loss in population abundance in tropical estuaries is also likely to be due to

the alteration and destruction of habitat, although the drivers of change are likely to differ

from those in terrestrial systems. For example, since 1980 industrial and economical

development in tropical countries may have created the need and funds to alter waterways

to make them suitable for shipping routes. Human populations of burgeoning cities

demand large increase in fishing pressure and higher levels of aquaculture (Blaber 2002),

and agricultural intensification. This results in depletion of fish stocks, spread of diseases

through escapement from aquaculture pens, and higher nutrient load, causing harmful

algal blooms and eutrophication (FAO 1996). Reduced freshwater runoff through

damming, water extraction and more regularly occurring drought conditions may have also

contributed to a decrease in native species. While the initial declining trend in tropical

estuarine populations appears to reflect the outcome of the most recent global LPI

analysis (Figure 1), there is a much less severe decline compared to the reduction of 50%

by 2005 of the global tropical index, and its upwards sloping trend may hint at a halting of

biodiversity loss rather than the same continuing loss that is being experienced by tropical

species globally. However, there are certain caveats associated with the interpretation of

the LPI in general and the above indices in particular, which are discussed below.

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2.4 Caveats

2.4.1 Baseline

A complication in the interpretation of the LPI is that abundance change is measured

against a baseline, which, for the purposes of this analysis, was set to 1980 due to limited

data availability before this point in time. It is likely that the contrasting temperate and

tropical trends discussed above are an artefact of shifting baselines (Pauly 1995), and that

preceding events need to be taken into account. The diverging trajectories therefore do not

necessarily imply that the state of biodiversity is worse in the tropics. Declines occurred in

temperate systems before 1980 (Millennium Ecosystem Assessment 2005) due to the

earlier onset of agricultural expansion and industrialisation, and populations may now be

increasing with concerted conservation effort, e.g. the establishment of protected areas,

action plans and managed reintroductions. Indeed, the choice of start year for any LPI may

be critical, especially if it is put in a historical context and changes compared to a pristine,

historical baseline. For example, Lotze et al. (2006) examined historical data on relative

abundance of taxonomic and functional groups from several estuarine and coastal

systems and found similar trajectories of long periods of slow decline followed by rapid

acceleration in the last 150 to 300 years. Increases relative to the baseline may thus

simply be recoveries from a state of historical depletion, as appears to be the case with

many marine mammals (Lotze & Worm 2009). For this reason, although the overall

temperate trend is positive, one cannot necessarily infer that temperate populations are in

a better state than tropical populations, just that they are currently increasing in abundance

compared to 1980. In any case, it is reasonable to conclude that there is a severe and

ongoing loss of biodiversity in tropical estuarine systems.

2.4.2 Regional and taxonomic bias

To be informative, indicators must be robust, sensitive and unbiased (Collen et al. 2009).

However, because the LPI represents a summary of underlying population data, it

provides an overview which is necessarily only as representative as the data feeding into

it. As such, the estuarine indices are likely to be subject to the regional and taxonomic bias

which stems from the particulars of the monitoring projects from which they are built.

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Abundance data were not evenly distributed globally, with clusters of populations occurring

in eastern North America, and along the coasts of Europe and South Africa (Figure 2).

Although the majority of biodiversity is undoubtedly found in the tropics (Jablonski et al.

2006), 75% of all populations and over 60% of species in this analysis were from

temperate regions. This imbalance is partially addressed in the index calculation by

weighting tropical and temperate populations equally (Collen et al. 2009). But as these

data are primarily from Europe and North America, increasing temperate trends may

indeed be due to onset of recovery through habitat restoration and protection. In addition,

there was a taxonomic bias against birds, which make up the majority of both the

temperate (Figure 5) and global datasets (67% of populations and 53% of species) – likely

the result of data collected for the calculation of the Mediterranean wetland bird index

(Galewski 2008). By contrast, the tropical populations comprise a vast majority of fish

species and populations (Figure 5), possibly an artefact of focusing on closing existing

data gaps in tropical populations, especially fish, during data collection. Fish accounted for

31% of populations and 45% of species in the global dataset (see Appendix Table 5 for

breakdown of populations and species numbers by region and taxonomic class).

Despite the close resemblance in trajectories of temperate species and temperate bird

species, closer inspection of the trends by taxonomic group reveals that fish abundance is

also increasing in temperate regions (Figure 6), decreasing the likelihood of the influence

of taxonomic bias and lending support to the explanations for increasing temperate trends

presented in section 2.3 above. By contrast, the decline in tropical abundance is likely to

be attributable to a dominance of fish populations and species (accounting for 63% and

60% vs 35% and 39% birds), especially those from South African estuaries (Figure 7). For

example, the exclusion of these South African fish results in an overall increasing global

trend. However, eliminating these populations would reduce the fish and tropical datasets

considerably, and is likely to result in a highly biased sample comprising a vast majority of

temperate bird populations. Both bird and fish indices are discussed in more detail below.

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Figure 5. Distribution of populations (top) and species (bottom) by class for each biogeographic realm. Note - marine species were assigned to the equivalent terrestrial/FW realm according to location.

2.4.3 Taxonomic representation

To assess to what extent this collection of species population trend data is representative

of the taxa found globally in estuaries, a comprehensive list of all estuarine species would

have to be devised, against which the species list from this analysis (Appendix Table 4)

could be compared. While this has not been attempted here, it should be the focus of any

further study.

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Figure 6. Trends for all temperate estuarine populations (green line, n = 335), temperate birds only (light blue line, n = 216) and temperate fish only (dark blue line, n = 108).

Figure 7. Index for all tropical estuarine species (green line, n = 202), tropical fish species (blue line, n = 121) and South African fish species (orange line, n = 61). Due to a pronounced drop in the number of populations contributing to trends in tropical fish abundance between 2002 and 2003, all indices end in 2002.

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2.5 Taxonomic class indices

2.5.1 Summary

Figure 8 shows the trends for 227 estuarine fish and 260 bird species from 1980 until

2002, which was chosen due to a pronounced drop in the number of fish populations

contributing to the index after this point, with the remainder of populations completely

dominating the trend in subsequent years. Other taxonomic groups were excluded, as only

fish and birds had sufficiently large numbers of populations throughout the time series,

with a mean number of 123 and 406 contributing populations respectively. As with the

global dataset, there was a prominence of temperate rather than tropical species (n = 345

vs n = 202) and populations (n = 922 vs n = 302).

Figure 8. Trends for estuarine birds (light blue, n = 260 species) and fish (dark blue, n = 227 species) globally. Due to a pronounced drop in the number of populations contributing to trends in tropical fish abundance between 2002 and 2003, both indices end in 2002.

2.5.2 Birds

The bird index shows a consistent increase in abundance, more than doubling from its

1980 baseline over the 22 year period (95% CIs: 1.87-2.69). Its trajectory resembles that

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of the temperate index (Figure 4), the temperate bird index (Figure 6) and the index for

Mediterranean wetland birds (Galewski 2008). Taking into account the location bias in

sampled bird species – North America and Europe showing the largest clusters of

populations (Figure 2) – it is likely that this trend is being influenced by samples from these

regions. Indeed, 87% of bird populations and 73% of bird species are from temperate

locations.

What are the possible explanations for estuarine birds, especially those from temperate

regions, increasing in abundance over the last two decades? Of the reasons for temperate

systems faring better than tropical systems presented above, most may also be applicable

to bird populations from these regions: reduced vulnerability to current threats due to

extinction filter effects (Balmford 1996), less anthropogenic pressure (e.g. Myers 1991,

Peres et al. 2006), the possibility of starting from different/earlier baseline (see 2.4.1),

more protection of habitat and species (Martin-López et al. 2009) and managed

reintroductions. Overall, it may even be fair to say that, particularly in comparison to fish,

bird species have received a large amount of conservation attention (e.g. Stokes 2006),

specifically in the more affluent temperate regions, which could have possibly resulted in

the observed increase in population abundance (e.g. the conservation successes of Red

kite and Bittern in the U.K., RSPB 2010). In addition, birds may be more adaptable to

changes in environmental conditions than species from other taxonomic groups due to

their high mobility. For example, they can respond by adjusting their foraging strategies,

e.g. by using a greater area, or by leaving a habitat that has been left unsuitable for

foraging or nesting by anthropogenic changes altogether. In addition, certain bird species

such as gulls are particularly adaptable, opportunistic and gregarious, and thus highly

adapted to living in man-modified habitats (Blokpoel & Spaans 1991). Several gull species

have undergone a widespread demographic increase, particularly in Europe (Spaans et al.

1991) and in North America (Blokpoel & Scharf 1991) as a result. In the estuarine dataset

nearly a fifth of populations and over 10% of species are from the family Laridae, and

these are found to be increasing if a separate index is calculated. For bird populations

from Mediterranean wetlands, especially the Camargue, Galewski (2008) suggests a

number of contributing factors: a ban of certain pesticides, an increase in the surface area

and improvements in the management of protected wetlands, as well as the management

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of water in hunting marshes. In addition, some species are likely to have benefitted from

land use change, e.g. conversion to agricultural plots.

The increase in bird abundance, especially in temperate regions, is consistent with several

other published trends. While wetland birds are still well below historic levels, the State of

the Birds Wetland Birds Indicator has increased by 50% between 1968 and 2009 due to

management and conservation measures, particularly policy shifts to habitat protection

and restoration, which have contributed to increases in generalist species, arctic-nesting

geese and cavity-nesting ducks (NABCI 2009). Further, the Waterfowl Indicator has more

than doubled because of a curb on hunting (NABCI 2009). The Coastal Bird Indicator

(NABCI 2009) shows a 20% increase, which is driven by huge recoveries in some

generalist species such as gulls, with specialist species still in decline. In other temperate

regions, birds are either stable or decreasing, e.g. Wild Bird Indicator (Defra 2010),

European Wild Bird Indicator (EBCC 2009) and Waterbird Population Status Index

(Butchart et al. 2010). Globally, birds are reportedly in decline, as indicated by the Red List

Index for the world’s birds 1988–2008 (BirdLife International 2008). Naturally, these

indicators may comprise very different species to this estuarine analysis, so comparison

may not be appropriate. The most likely explanation for the observed increasing trends in

estuarine birds is the fact that 76% of populations and 81% of species were partially or

entirely sampled from protected areas such as Ramsar, World Heritage, and IUCN sites,

and national parks (e.g. Doñana, Camargue, Yukon-Kuskokwim delta). As this protection

status will reduce the extent of wetland habitat loss and the number of threat processes

affecting each species – two extrinsic factors that have been identified as significant

predictors of population declines in wildfowl (Long et al. 2007) – it is not surprising to

observe positive abundance trends in these samples.

2.5.3 Fish

Unlike the tropical fish index (Figure 7), the global fish index, is relatively stable around the

baseline, returning to it by 2002 (Figure 8), implying no change in abundance over the 22

year period (95% CIs: 0.66-1.53). As the split between temperate and tropical regions was

even for populations and near even for species (47% vs 53%), regional bias is unlikely to

be responsible for the trend, although it is useful to examine regional trends separately.

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2.5.4 Tropical and South African fish

The pronounced decrease of 74% (95% CIs: 0.10-0.63) in tropical fish abundance

(Figure 7) is not attributable to a decrease in native species resulting from competition of

invasive/alien species, as the latter were found to be declining. There is, however, good

evidence of a pronounced influence of South African estuarine fish populations, which

decline by 70% (95% CIs: 0.13-0.65) and account for over 25% of species and 30% of

populations. The southern US provided a further quarter of species and a fifth of

populations. Due to the similarity in trends between tropical and South African fish

(Figure 7), it is nevertheless reasonable to assume that the tropical fish trend is being

driven primarily by South African fish.

The semi-arid climate and low rainfall in South Africa, along with high evaporation rates,

result in one of the lowest conversions of rainfall to run-off in the world (Davies & Day

1998, in Grange et al. 2000). This results in naturally fluctuating environmental conditions

due to changes in the relative influence of freshwater and marine systems within the

estuary, with particular effects on salinity (Kok & Whitfield 1986) and turbidity. Many

species avoid areas of high turbidity (Cyrus 1988) because high sediment loads can be

lethal to both marine migrants and estuarine residents (Whitfield & Paterson 1995). Due to

reduced freshwater input, many estuaries are seasonally or intermittently open, and the

timing and duration of these open phases are recognised as the major determinants of fish

diversity and abundance (James et al. 2008b). Most South African estuaries are

dominated by euryhaline marine fish species that spawn at sea (Marais & Bird 1980), and

these are therefore more abundant in permanently open estuaries (Whitfield & Harrison

2003, Harrison & Whitfield 2006). There is usually an increase in abundance of these

species after mouth opening (Whitfield 1980, in Russell 1996), and declines during

extended closed phases. This is because adults within the estuary are prevented from

emigrating back to sea, which reduces the spawning population, and juveniles cannot

recruit and may search for alternative nursery habitat, e.g. in adjacent estuaries (Vivier &

Cyrus 2009). Prolonged closure of the mouth has caused a gradual decline in the

populations of estuarine-associated marine species in St. Lucia (e.g. Cyrus & Vivier 2006,

Whitfield et al. 2006) and East Kleinemonde (James et al. 2008b). While populations can

recover after breaching, artificial opening can be dangerous if a sheering effect occurs and

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upper, better oxygenated water columns flow out to sea, leaving the anoxic waters caused

by decreased freshwater input behind, resulting in mass fish kills and prompting euryhaline

species to migrate to freshwater reaches (Becker et al. 2009).

These natural fluctuations are exacerbated by anthropogenic pressures, such as water

extraction and impoundments, which greatly reduce the run-off, especially in dry years

(Whitfield & Bruton 1989). In Nhlabane estuary (Vivier & Cyrus 2002), for example, the

construction of a barrage to increase storage capacity of the associated lake, and water

abstraction from the lake and the estuary for mining purposes in addition to a natural

drought resulted in prolonged mouth closure between 1991 and 1995. This prevented the

recruitment of larvae and juveniles to their estuarine nursery habitat, and emigration of

adults to sea. Only after artificial breaching and good rainfall in late 1995 was some

recovery observed due to marine species returning to the system. In addition, over-

exploitation can greatly add to these pressures, especially if a species is slow to mature,

as is the case with the Picnic seabream (Acanthopagrus berda) in the Kosi estuarine

system (James et al. 2008a).

Although the equilibrium in estuaries is usually re-established following such events, in

recent years the decline in freshwater inflow resulting from anthropogenic influence within

the catchments has resulted in many estuaries following hypersaline trajectories (Whitfield

& Bruton 1989). The marked increase in the demand for freshwater following the rapid

growth in South Africa’s human population may cause available riverine resources to be

fully exploited within a few decades (Davies & Day 1986, in Grange et al. 2000). Climate

change may add to these pressures, as a reduction in rainfall across temperate regions

such as southern Australia and Africa have been forecast (e.g. Schulze et al. 2001,

Hughes 2003).

While these observations are specific to South African estuaries and may not be

applicable to the tropical or tropical fish indices, they do have management implications for

estuaries in this part of the world. It appears that the opening of estuaries at the right time

of year, habitat preservation and the regulation of water extraction are important factors in

counteracting more frequently occurring extreme environmental conditions such as

drought.

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2.5.5 Temperate fish

While temperate fish increase by 69% by 2005 (95% CIs: 0.75-3.76; Figure 6), 70% of

time series end in 2002, meaning that the end year should be set to this point in time. This

does, however, result in more than a doubling in abundance overall (95% CIs: 1.12-3.63).

There are a number of reasons for why temperate fish may be increasing compared to

tropical fish populations. Tropical environments are more variable (see 2.5.6, especially

South African fish), and because modifications to estuarine systems may have been made

before 1980 (see 2.4.1), the observed increases may be recoveries from a state of

depletion. Of all species and populations, 60% were from protected areas, and nearly a

third of populations were not facing any threats, both of which could explain an increase in

abundance. Overall, 80% of species and 75% populations were from the U.S., the U.K.

and the Danube delta in Romania – all regions that may have the necessary funds for

successful conservation or are the focus of international conservation attention.

2.6 Other indices

2.6.1 Estuary type

Because natural processes vary between different types of estuaries (Hansen & Rattray

1966), an estuarine system’s ecological functioning can be expected to be affected by

different drivers of change, which, in turn, should have an impact on vertebrate abundance

trends in that location. Overall, none of the indices based on estuary type suggest a

decreasing pattern in abundance, which is surprising considering, for example, the

pronounced decline of wetlands in temperate regions (60% of Europe’s wetlands are

already lost due to conversion to alternative use or lack of conservation, UNEP/DEWA

2004). However, sample sizes were often small and varied considerably, and there was an

unequal distribution of data both regionally and taxonomically, making it impossible to

draw valid conclusions from the data. An analysis of abundance trends by estuary type,

perhaps even taking into account drivers of change, would undoubtedly be of great interest

and should be undertaken in the future if the dataset can be improved through additional

data collection and data inequalities eradicated.

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2.6.2 Drivers of change

Various threats to freshwater diversity have been identified, such as overexploitation,

water pollution, flow modification including water abstraction, destruction or degradation of

habitat, and invasion by invasive alien species (Dudgeon et al. 2006, Millennium

Ecosystem Assessment 2005). These threats are likely to be exacerbated by the predicted

temperature changes and shifts in precipitation and runoff patterns associated with climate

change (Dudgeon et al. 2006).

In the present analysis, drivers of change were only considered if they yielded a sufficiently

large sample, boosted by using, for each driver category, data from all locations affected

by the driver at any rank. In addition, driver categories that were associated with increases

in abundance were excluded, as negative forces cannot feasibly have positive

consequences. See Figure 9 for indices up to 2002, Figure 10 for the percentage of

decreasing populations affected by different primary drivers of change and Appendix

Table 7 for species and population numbers for different driver categories.

Figure 9. Indices for populations from estuarine locations with the drivers Dams (light blue, n = 122 spp) and Water extraction (dark blue, n = 158 spp) at any rank, and for populations from locations that are not threatened by any driver of change (green line, n = 146 spp).

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As expected, there is an increase in species abundance from estuarine locations that are

not threatened. This may be due to either the habitat being pristine and unaltered, or

natural processes being in balance despite alterations. However, the underlying dataset is

biased towards bird species, which account for 63% and have already been shown to be

increasing globally (Figure 8) and in temperate regions (Figure 6), so trends may not be

representative of such locations.

Figure 10. Primary drivers of change for N = 523 declining estuarine populations.

Dams are said to provide green energy compared to fossil fuels (de Souza 1996, in

Fearnside 2002), and their construction is usually for the purposes of water collection in

reservoirs for crop irrigation as well as electricity generation through the exploitation of

hydropower. The index for dams shows a near 75% decline (95% CIs: 0.12-0.55), but the

dataset appears to be biased in a number of ways: towards fish species (80%) and

populations (87%), and tropical populations (57%) and species (63%). So, this decline

may simply be a reflection of declines in tropical species and fish in particular (Figure 7).

However, out of vertebrate taxa, fish will be most severely affected by barriers because

dams act as environmental filters, fragmenting habitats important in the life cycles of

migratory fishes (McAllister et al. 2001). They often present insurmountable obstacles for

migrating fish species, as supported by the finding that Agostinho et al. (2007, in

Agostinho et al. 2008) recorded four or more migratory fish species in only 5% of 75

Brazilian reservoirs, with more than 50% lacking migratory species altogether. In the

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estuarine dataset, migratory fish account for nearly 40% of populations, and because the

index for fish species only is similar to the overall index including birds, it is reasonable to

assume that it is representative. Disruptions to migrations can lead to a failure in breeding

in or recruitment to populations (McAllister et al. 2001), which will have severe knock-on

effects on local fisheries and livelihoods (e.g. Xie et al. 2007).

Populations affected by water extraction show a 43% decline overall (95% CIs: 0.41-0.78)

but there is some indication that this dataset is biased towards bird species, as well as

tropical populations and species, accounting for 61%, 74% and 83% respectively. While

birds have been shown to be on the increase (Figures 6 and 8), the decrease in

abundance may well be a reflection of tropical species trends, especially fish (Figure 7). In

addition, it is important to note that there may indeed be a close association between

dams and water extraction, which may explain similar resulting trends in species

abundance. Dams are constructed not just for the generation of hydropower but also for

the creation of reservoirs, which can provide for agricultural purposes.

There are several problems associated with the drivers approach employed in this

analysis. First, by using all ranks to increase sample sizes, it is possible that datasets will

be biased through the co-occurrence of certain drivers, leading to similar datasets. For

example, construction of dams may well be associated with a higher probability of water

extraction being a threat in the same watershed. Of course, it is also possible that the

drivers identified for this analysis are specific to the particular river or its estuary and do

not necessarily apply to the population in question, i.e. these drivers may not necessarily

pose a risk to a species at the population level. For example, dams which do not have

integrated fish ladders will have a devastating effect on fish needing to migrate upstream,

while species that migrate to brackish water from marine systems, or are permanent

residents in the estuary, may not be affected at all. So, although it is tempting to determine

a cause-and-effect relationship between drivers of change and population declines, this is

not possible, as can also be seen from the proportion of populations affected by different

primary drivers in Figure 10. Here, just fewer than 15% of populations are from locations

that are not threatened, yet these populations are decreasing in abundance. Causation

can therefore not be ascribed and it is only valid to establish a possible rather than

definitive connection between the drivers of change and changes in species abundance.

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2.6.3 Estuarine-dependent fish

Populations of fish species that can be defined as being estuarine-dependent based on

their Global Register of Migratory Species (GROMS) classification (see METHODS) were

identified in the global LPI database and indices produced for estuarine-dependent fish

(EDF) and for all estuarine-dependent fish and fish populations from estuaries combined

(EDF + EstFish) (Figure 11). Indices were cut in 2002 due to a drop in contributing

populations after this point. The EDF and EDF + EstFish indices show a decrease in

abundance of 21% (95% CIs: 0.53-1.2) and 16% (95% CIs: 0.59-1.20) over the 22 year

period. In trajectory they resemble the trend in estuarine fish populations (Figure 8),

although they are more negative and showing continuous declines in abundance.

Possible reasons for these declines are habitat changes in estuaries and associated

watersheds such as dam construction and shipping activity, which may reduce or inhibit

movement of migratory species between freshwater and marine systems (McAllister et al.

2001). However, upon examination of the threats underlying the included populations, no

clear picture emerges.

Figure 11. Indices for n = 113 estuarine-dependent fish species (EDF, green line) and n = 259 estuarine-dependent fish species and estuarine fish populations combined (EDF + EstFish, yellow line).

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Of course, there are potential problems associated with the GROMS classification

approach. For example, populations of species marked as migratory may spend their

entire life outside of brackish water and may therefore not be affected by change in these

systems. Also, populations could have been sampled from freshwater and marine

locations at great distance from the nearest brackish water body. Dependence of species

on intact natural processes being in balance in these estuaries is also difficult to assess.

As such, a merely open gateway is sufficient for some species, while others may be

flexible in their habitat choice and thus able to abandon the estuary. Yet others might find it

completely impossible to feed or breed unless their estuarine habitat is undisturbed. In

addition, any decrease in abundance cannot be readily assumed to be due to factors

causing changes in the estuary itself. For example, a diadromous fish travelling long

distances between marine feeding and freshwater spawning grounds may be threatened

by factors outside of the estuary, e.g. it might be exploited at sea. A possible way of

disentangling these different influences is to calculate indices for combinations of

population-specific threats and GROMS classification. This analysis has not been

attempted here, as this approach is likely to result in small sample sizes.

2.6.4 Human population density

Studies have shown that exposure to high human population density increases extinction

risk, particularly in the presence of certain biological traits (Cardillo et al. 2004, 2005).

Changes in estuarine species abundance in relation to Low and High human population

density (based on CIESIN/CIAT 2005) are shown in Figure 12. Due to sufficiently large

sample sizes, High density trends could be analysed for tropical and temperate species

separately.

Vertebrate abundance trends in areas less densely populated by humans increased to 2.7

times (95% CIs: 2.23-3.29) that of the baseline level and thus fare much better than those

from high density areas, which are showing relatively flat trends with an increase of 27%

(95% CIs: 0.96-1.68). However, the temperate/tropical disparity in abundance in high

density areas suggests that high human density is only a pressure on and threat to

species abundance in tropical regions, which may be explained by some of the reasons for

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the difference in tropical and temperate trends in section 2.3. The split was also apparent

for Low density populations, but as sample sizes were small, the results are not presented.

Figure 12. Indices for estuarine populations of species in areas of low human population density (grey line, n = 359 spp), and of high density overall (red line, n = 280 spp) as well as high density areas in temperate (green dashed line, n = 168 spp) and tropical regions (orange dashed line, n = 133 spp).

2.6.5 Income group and Human Development Index

The split between temperate and tropical trends was also found when data were analysed

by country income group (The World Bank 2009) (Figure 13). This is likely the result of a

correlation of low Income group ratings and tropical regions.

Populations from high income countries show highly similar abundance trends to those

from countries with a high Human Development Index (HDI) rating (UNDP 2009), which is

attributable to the fact that all bird, mammal, and reptile, and most fish populations of the

former dataset are also included in the latter. The high HDI index shows a smaller increase

over the study period, perhaps because it includes nearly all of the bird but only half of the

fish populations of the income index – estuarine birds have been shown to increase while

fish are mostly declining or increasing at a slower rate (Figures 6, 7 and 8). Overall, there

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appears to be an interaction in terms of abundance trend between the income level of the

country the population was sampled from and whether this is from a temperate or tropical

region, with positive trends in temperate high income locations, and negative trends in

tropical low income locations.

Figure 13. Indices for estuarine populations from countries of high income (dark grey dashed line, n = 324 spp) and low income (light grey dashed line, n = 268 spp), and temperate high income (dark green, n = 291 spp), temperate low income (light green, n = 129 spp), tropical high income (orange, n = 37 spp) and tropical low income (yellow, n = 172 spp).

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CONCLUSIONS

The Living Planet Index for bird, fish, reptile and mammal populations from estuarine

locations globally has increased by 16% between 1980 and 2005, with wide variation in

species responses to changing estuarine systems. Tropical populations are decreasing in

abundance but are potentially showing some signs of recovery, while a positive trend was

found for temperate estuarine locations, particularly for bird species. Temperate fish

populations are also increasing, but populations in the tropics are declining, while global

estuarine fish populations remain stable on average. Species are well adapted to the

changing conditions in estuaries, which are characterised by wide fluctuations in a variety

of factors, e.g. turbidity and salinity, and therefore retain an inherent resilience to change.

Nevertheless, as human impacts on estuarine systems grow, there are troubling negative

trends in particular groups of species, and certain areas of the world, not least tropical

ecosystems and fish populations. Of the many possible factors influencing vertebrate

population abundance in estuaries, dams and water extraction were both associated with

large declines in abundance. Worryingly, those specialist populations of fish which are

highly dependent on estuarine systems decreased by 20% between 1980 and 2002. An

increased number of targeted monitoring programmes are required to build a truly

comprehensive picture of how different factors might be affecting estuarine species

population abundance.

ACKNOWLEDGEMENTS

Thank you to Dr. Imad Cherkaoui for providing bird population data from the Sebou and

Moulouya estuaries in Morocco, and Yi Yong for data from the Yangtze River estuary.

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APPENDIX

Methods

Data collection

Data for the calculation of the estuarine Living Planet Index was collected from journals

and other publications through internet and library searches, as well as by contacting

individuals from particular locations globally. In addition, existing records in the LPI

database from estuarine locations were identified and additional information added (see

below). Locations sought were low-lying, coastal, enclosed or semi-enclosed bodies of

brackish water such as estuaries, lakes and lagoons, and the tidal areas of wetlands and

deltas. Other coastal habitats, which would be classified as either freshwater or marine,

such as freshwater estuaries, or salt marshes and saltwater mangroves, were not

considered. Search effort focused primarily on 33 coastal locations identified by WWF

Netherlands as being of particular interest (A. Berkhuysen, personal communication).

These include locations where WWF is active, deltas and estuaries that have been

identified as most vulnerable by the IPCC (Nicholls et al. 2007), and any additional

locations from WWF’s Deltas on the Move report (Reker et al. 2006). See Table 1 in the

Appendix for a comprehensive list of these locations.

Estuary type and drivers of change

For the purposes of a more detailed analysis, additional information was recorded for each

population. To establish any differences in biodiversity trends between different estuary

types, where the functioning of natural processes may be threatened by different factors,

this study distinguished between the following: Open estuary, Closed estuary, Delta,

Wetland, and Mangrove. Definitions can be found in Table 2 in the Appendix. In addition, a

driver-based approach was employed to assess the effects of anthropogenic interventions

on trends in estuarine populations of vertebrate species. The categories of drivers were

based on Wong et al. (2007) and include the following: Infrastructure – dams,

Infrastructure – navigation, Land use changes, Water extraction, Climate change,

Pollution, Invasive species, Overexploitation, Unknown drivers, and No threats. For those

estuarine locations for which drivers were ranked in Wong et al. (2007), the top three were

taken directly from the report. For other locations, main drivers of change were identified

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based on information contained in the data source or any other readily available

respectable source in order to reduce the number of populations with Unknown drivers. As

records were ranked only by driver type, no assessment was made of the impact these

may have had on the proper functioning of natural processes in that location. For the

purpose of the present study, only the primary driver was taken into account.

GROMS classification

All populations in the LPI database of fish species that could be considered estuarine-

dependent, i.e. those that are known to migrate to, from or through estuaries, were

identified based on species classification in the Global Register of Migratory Species

(GROMS, www.groms.de). Only species that were classified as diadromous, anadromous,

catadromous, amphidromous or oceano-estuarine were included in the analysis.

Human population density, income group and Human Development Index

To relate trends in estuarine biodiversity to human population density in the immediately

surrounding area, a GIS map showing human population density globally (CIESIN/CIAT

2005) was overlaid with points representing populations used for the creation of the index.

After assigning population density estimates to each population by hand, the estuarine

dataset was then divided for all species and all fish into High and Low categories using the

classification system of >99.99 people per km2 for the former, and <99.99 people per km2

for the latter. The estuarine dataset was also analysed by country income category

according to The World Bank (2009) and by Human Development Index category based

on the classification in UNDP (2009). In both cases, Middle and Low categories were

combined to yield sufficiently large datasets.

Analysis

Analysis for estuarine populations was carried out using R Version 2.5.1., package mgcv

1.3-29, following standard LPI procedure. The available data were disaggregated at a

variety of levels, to achieve as complete an analysis as possible. Annual data points were

interpolated for time series with six or more data points using generalized additive

modelling, or by assuming a constant annual rate of change for time series with less than

six data points, and the average rate of change in each year across all species was

calculated. The average annual rates of change in successive years were chained

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together to make an index. Confidence limits on all LPI graphs denote the degree of

certainty in the index: the narrower the limits, the higher the confidence. The calculations

underlying the LPI method are discussed in detail elsewhere (see Collen et al. 2008). All

indices comprised non-weighted species trends based on all populations at certain

disaggregated levels, with the exception of the global index, which is based on equally

weighted tropical and temperate indices. Here, the trend line thus represents the average

change within the entire collection of population samples, giving equal weight to each

species, whether common or rare, and to small and large populations. The tropical LPI

consists of terrestrial and freshwater species populations in the Afrotropical, Neotropical

and Indo-Pacific (Indo-Malaya, Australasia and Oceania) realms, and marine species

populations from the zone between the Tropics of Cancer and Capricorn. The temperate

LPI includes all terrestrial and freshwater species populations from the Palearctic and

Nearctic realms, and marine species populations north and south of the tropics. Unlike in

the standard LPI, in which terrestrial, freshwater and marine system trends are each given

equal weight within the tropical and temperate LPIs, this was not considered suitable for

the present analysis, as system information was deemed irrelevant for brackish water

populations. Where shown, confidence intervals were calculated using 10,000 bootstraps.

Baseline and end year

Tropical data availability was found to be relatively low in the 1970s, so indices were

calculated setting the baseline of 1 to 1980. While this reduces the number of years

covered, it results in a more robust and less biased index. Because n values were

considered unsatisfactorily small towards the present day, 2005 was chosen as the end

year for all indices, unless otherwise stated.

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Tables

Table 1. Focal areas

1. Locations where WWF is active

(A. Berkhuysen, pers. comm.)

Danube, Elbe, Ganges Brahmaputra, Mekong,

Mississippi, Rhine (Rhine–Meuse–Scheldt),

Changijang / Yangtze

2. Additional IPCC deltas

(from Nicholls et al 2007)

Amazon, Chao Phraya, Godavari, Grijalva,

Indus, Krishna, Mahakam, Mahanadi, Moulouya,

Niger, Nile, Orinoco, Red, Sao Francisco,

Sebou, Senegal, Shatt el Arab, Volta, Zhujiang

(Pearl River, Bocca Tigris estuary)

3. Additional locations from Deltas on the Move

report

(Reker et al. 2007)

Fly, Fraser, Lena, MacKenzie, Parana (Río de la

Plata), Po, Yukon (Yukon-Kuskokwim)

Table 2. Estuary types and definitions

Open estuary

A semi-enclosed body of water, which is permanently open to the sea,

where freshwater from a river mixes with seawater. Subject to both

riverine and tidal influences.

Closed estuary

An enclosed body of water, which is intermittently open to the sea,

where freshwater from a river mixes with seawater within a coastal

lagoon or lake. Also, an estuary as defined above but with a closed

river mouth. Subject to riverine influences, but lacking tidal range and

currents.

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Wetland

A low-lying coastal area that is inundated or saturated by freshwater

from rivers or other ground water, as well as occasionally flooded by

saltwater from the sea, resulting in saturated brackish soil and creating

a number of smaller brackish water bodies. Does not include swamps,

marshes, bogs, and similar areas, but should instead be used for

brackish areas where exact sampling sites are unknown or where

sampling occurred in a series of bodies of brackish water and

surrounding habitats. Introduced to distinguish these delta-like

floodplains or series of brackish lagoons from real deltas and large

brackish lagoons.

Delta

A low-lying coastal area where freshwater and saltwater meet, building

outwards from the deposition of sediments carried by the river, and

resulting in a land mass divided by a number of small streams usually

from the same river.

Brackish mangroves

Forests which grow in the intertidal areas and estuary mouths between

land and sea, comprising salt-tolerant tree and other plant species.

Found between the latitudes of 32º N and 38º S, along the tropical and

subtropical coasts of Africa, Australia, Asia, and the Americas. Only

brackish mangroves are included in the estuarine index.

Table 3. Improvements in the coverage of the estuarine LPI dataset

Old dataset Added data New dataset %

No. % of total No. % of total No. % of total increase

Fish 30 4.07 350 71.87 380 31.05 1166.67

Aves 689 93.49 129 26.49 818 66.83 18.72

Mammalia 6 0.81 3 0.62 9 0.74 50.00 Class

Reptilia 12 1.63 5 1.03 17 1.39 41.67

Total 737 100.00 487 100.00 1224 100.00 66.08

Old dataset Added data New dataset No. of %

No. % of total No. % of total No. % of total new spp increase

Fish 20 7.46 222 67.89 227 45.40 207 1035.00

Aves 237 88.43 99 30.28 260 52.00 23 9.70

Mammalia 3 1.12 3 0.92 5 1.00 2 66.67 Class

Reptilia 8 2.99 3 0.92 8 1.60 0 0.00

Total 268 100.00 327 100.00 500 100.00 232 86.57

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Table 4. List of all species included in the estuarine LPI

Fishes

Acanthogobius flavimanus Craterocephalus fluviatilis Hypseleotris compressa

Acanthopagrus australis Craterocephalus stercusmuscarum Hysterocarpus traskii

Acanthopagrus berda Crenicichla lepidota Jenynsia multidentata

Acentrogobius viridipunctatus Croilia mossambica Johnius dorsalis

Acipenser oxyrinchus Ctenogobius shufeldti Labeo umbratus

Acipenser sinensis Cynoscion arenarius Lates calcarifer

Agonus cataphractus Cynoscion nebulosus Leiognathus equulus

Alburnus alburnus Cyprinus carpio Lepidopsetta bilineata

Alosa aestivalis Dicentrarchus labrax Lepomis gibbosus

Alosa fallax Diplodus capensis Leptocottus armatus

Alosa sapidissima Diplodus cervinus Lichia amia

Ambassis productus Diplodus sargus Liopsetta glacialis

Ambassis vachelli Dorosoma petenense Liopsetta putnami

Ameiurus melas Echiichthys vipera Lithognathus lithognathus

Ammodytes hexapterus Eleginus gracilis Liza alata

Anchoa marinii Elops machnata Liza dumerili

Anchoa mitchilli Engraulis encrasicolus Liza macrolepis

Anguilla anguilla Enophrys bison Liza ramada

Anguilla rostrata Epinephelus itajara Liza richardsonii

Archosargus rhomboidalis Esox lucius Liza subviridis

Argyrosomus japonicus Eubleekeria splendens Liza tricuspidens

Ariopsis felis Eucinostomus lefroyi Luciobarbus comizo

Astyanax eigenmanniorum Gadus macrocephalus Luciobarbus microcephalus

Atherina boyeri Gadus morhua Luciobarbus sclateri

Atherina breviceps Galeichthys feliceps Lumpenus fabricii

Atherinella brasiliensis Gasterosteus aculeatus Lumpenus sagitta

Atherinosoma microstoma Geophagus brasiliensis Lutjanus argentimaculatus

Atractosteus spatula Gerres acinaces Lycengraulis grossidens

Blepsias cirrhosus Gerres filamentosus Megalops cyprinoides

Blicca bjoerkna Gerres methueni Menidia menidia

Boreogadus saida Gerres oyena Menticirrhus americanus

Brevoortia pectinata Gilchristella aestuaria Menticirrhus littoralis

Caranx sexfasciatus Gillichthys mirabilis Merlangius merlangus

Carassius carassius Glossogobius callidus Microgadus proximus

Carassius gibelio Glossogobius giuris Micropogonias furnieri

Carcharhinus leucas Gobionellus oceanicus Micropogonias undulatus

Cathorops melanopus Halobatrachus didactylus Misgurnus anguillicaudatus

Catostomus occidentalis Heteromycteris capensis Monodactylus argenteus

Chanos chanos Hexagrammos decagrammus Monodactylus falciformis

Chelonodon patoca Hexagrammos lagocephalus Morone americana

Citharichthys spilopterus Hexagrammos octogrammus Morone saxatilis

Clarias gariepinus Hexagrammos stelleri Mugil cephalus

Clupea harengus Hippocampus abdominalis Mugil curema

Clupea pallasii Hypomesus pretiosus Mugil gaimardianus

Coregonus autumnalis Hypomesus transpacificus Mugil platanus

Cottus asper Hyporhamphus capensis Myliobatis aquila

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Fishes continued Reptilia

Myoxocephalus polyacanthocephalus Rhabdosargus sarba Alligator mississippiensis

Myxocyprinus asiaticus Rhabdosargus thorpei Batagur baska

Myxus capensis Rhinobatos annulatus Caretta caretta

Nannoperca australis Rutilus rutilus Chelonia mydas

Nannoperca obscura Salmo salar Crocodylus acutus

Nuchequula gerreoides Salmo trutta Emys orbicularis

Odontesthes argentinensis Salvelinus malma Malaclemys terrapin

Oligosarcus robustus Sander lucioperca Testudo hermanni

Oncopterus darwinii Sardina pilchardus

Oncorhynchus gorbuscha Sardinella albella

Oncorhynchus keta Sarpa salpa Mammalia

Oncorhynchus nerka Scardinius erythrophthalmus Delphinapterus leucas

Oncorhynchus tshawytscha Sillago analis Lipotes vexillifer

Oreochromis mossambicus Sillago maculata Lutra lutra

Oreochromis niloticus Sillago sihama Neophocaena phocaenoides

Osmerus mordax Silurus glanis Phoca vitulina

Pachymetopon aeneum Solea bleekeri

Pallasina barbata Solea solea

Paralichthys orbignyanus Spirinchus thaleichthys

Parapimelodus nigribarbis Sprattus sprattus

Pegusa lascaris Syngnathus folletti

Perca fluviatilis Syngnathus rostellatus

Petromyzon marinus Syngnathus watermeyeri

Pholis laeta Terapon jarbua

Platanichthys platana Terapon tarbua

Platichthys flesus Thryssa vitrirostris

Platichthys stellatus Thymalllus arcticus

Platycephalus arenarius Tinca tinca

Platycephalus fuscus Torpedo fuscomaculata

Platycephalus indicus Torpedo marmorata

Pleuronectes platessa Trachinotus falcatus

Pogonichthys macrolepidotus Trachinotus marginatus

Pomadasys commersonnii Trichiurus lepturus

Pomadasys olivaceus Trichodon trichodon

Pomatomus saltatrix Triglopsis quadricornis

Pomatoschistus microps Trisopterus esmarkii

Pomatoschistus minutus Trisopterus luscus

Prionotus punctatus Trisopterus minutus

Pristis pectinata Umbrina canosai

Prosopium cylindraceum Valamugil buchanani

Psammogobius biocellatus Valamugil cunnesius

Psammogobius knysnaensis Valamugil robustus

Pseudomugil signifer Valamugil seheli

Pseudorhombus arsius

Ramnogaster arcuata

Rhabdosargus holubi

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Aves

Acrocephalus agricola Calidris acuminata Emberiza schoeniclus

Acrocephalus arundinaceus Calidris alba Erithacus rubecula

Acrocephalus schoenobaenus Calidris alpina Eurynorhynchus pygmeus

Acrocephalus scirpaceus Calidris canutus Falco columbarius

Actitis hypoleucos Calidris ferruginea Falco eleonorae

Actitis macularius Calidris minuta Falco naumanni

Aegithalos caudatus Calidris pusilla Falco pelegrinoides

Alauda arvensis Calidris ruficollis Falco peregrinus

Alca torda Calidris temminckii Falco tinnunculus

Alcedo atthis Carduelis carduelis Ficedula hypoleuca

Alectoris barbara Carduelis chloris Fregata magnificens

Amazonetta brasiliensis Casmerodius albus Fringilla coelebs

Anas acuta Certhia brachydactyla Fulica armillata

Anas clypeata Cettia cetti Fulica atra

Anas crecca Charadrius alexandrinus Fulica cristata

Anas gracilis Charadrius collaris Galerida cristata

Anas penelope Charadrius dubius Gallinago gallinago

Anas platyrhynchos Charadrius hiaticula Gallinula chloropus

Anas strepera Charadrius leschenaultii Gavia immer

Anser albifrons Charadrius melodus Gavia pacifica

Anser anser Charadrius pecuarius Gavia stellata

Anser erythropus Charadrius ruficapillus Glareola pratincola

Anthus pratensis Charadrius semipalmatus Grus canadensis

Anthus richardi Charadrius wilsonia Grus japonensis

Anthus spinoletta Chen canagica Gygis alba

Anthus trivialis Chlidonias hybrida Haematopus ostralegus

Aquila clanga Chlidonias niger Haematopus palliatus

Aramides cajanea Chloroceryle americana Haliaeetus albicilla

Ardea cinerea Ciconia ciconia Hieraaetus pennatus

Ardea cocoi Circus aeruginosus Himantopus himantopus

Ardea purpurea Circus cyaneus Himantopus mexicanus

Ardeola ralloides Circus pygargus Hippolais polyglotta

Arenaria interpres Cisticola juncidis Hirundo daurica

Asio capensis Cladorhynchus leucocephalus Hirundo rupestris

Aythya ferina Columba livia Hirundo rustica

Aythya fuligula Columba palumbus Ixobrychus minutus

Aythya nyroca Cyanopica cyanus Jacana jacana

Botaurus stellaris Cygnus atratus Jynx torquilla

Branta bernicla Cygnus columbianus Lanius excubitor

Branta canadensis Cygnus olor Lanius senator

Branta hutchinsii Delichon urbicum Larus audouinii

Bubulcus ibis Egretta caerulea Larus cachinnans

Burhinus oedicnemus Egretta garzetta Larus canus

Buteo buteo Egretta gularis Larus dominicanus

Buteo rufinus Egretta thula Larus fuscus

Butorides striata Emberiza cirlus Larus genei

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Aves continued

Larus hyperboreus Philomachus pugnax Sterna trudeaui

Larus ichthyaetus Phoenicopterus roseus Streptopelia decaocto

Larus maculipennis Phoenicurus ochruros Sturnus unicolor

Larus melanocephalus Phoenicurus phoenicurus Sylvia atricapilla

Larus minutus Phylloscopus bonelli Sylvia borin

Larus ridibundus Phylloscopus collybita Sylvia cantillans

Limosa lapponica Phylloscopus ibericus Sylvia communis

Limosa limosa Phylloscopus trochilus Sylvia hortensis

Locustella luscinioides Pica pica Sylvia melanocephala

Locustella naevia Picus viridis Sylvia undata

Luscinia megarhynchos Platalea ajaja Syrigma sibilatrix

Luscinia svecica Platalea leucorodia Tachybaptus ruficollis

Marmaronetta angustirostris Platalea minor Tadorna tadorna

Megaceryle torquata Plegadis falcinellus Tetrax tetrax

Melanitta nigra Pluvialis apricaria Threskiornis molucca

Melanocorypha calandra Pluvialis dominica Tringa erythropus

Mergus serrator Pluvialis squatarola Tringa flavipes

Mesophoyx intermedia Podiceps cristatus Tringa glareola

Miliaria calandra Podiceps nigricollis Tringa nebularia

Morus bassanus Porphyrio porphyrio Tringa ochropus

Motacilla alba Porzana porzana Tringa stagnatilis

Motacilla cinerea Porzana pusilla Tringa totanus

Motacilla flava Prunella modularis Troglodytes troglodytes

Muscicapa striata Puffinus mauretanicus Turdus merula

Myiopsitta monachus Pygoscelis papua Turdus philomelos

Netta rufina Pyrrhula pyrrhula Upupa epops

Numenius arquata Rallus aquaticus Vanellus chilensis

Numenius phaeopus Recurvirostra avosetta Vanellus spinosus

Numenius tenuirostris Regulus ignicapilla Vanellus vanellus

Nyctanassa violacea Remiz pendulinus Xema sabini

Nycticorax nycticorax Rissa tridactyla

Oxyura leucocephala Rynchops niger

Pandion haliaetus Saxicola rubetra

Panurus biarmicus Saxicola torquatus

Pardirallus nigricans Serinus serinus

Pardirallus sanguinolentus Somateria fischeri

Parus caeruleus Somateria mollissima

Parus cristatus Sterna albifrons

Parus major Sterna caspia

Passer domesticus Sterna hirundinacea

Pelecanus conspicillatus Sterna hirundo

Pelecanus crispus Sterna maxima

Pelecanus onocrotalus Sterna nilotica

Phalacrocorax brasilianus Sterna paradisaea

Phalacrocorax carbo Sterna sandvicensis

Phalacrocorax pygmeus Sterna superciliaris

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Table 5. Numbers of species and populations by region and taxonomic class

Temperate Tropical Global

Spp Pop Spp Pop Spp Pop

Fishes 108 191 121 189 227 380

Reptiles 6 11 3 6 8 17

Birds 216 711 78 107 260 818

Mammals 5 9 0 0 5 9

Total 335 922 202 302 500 1224

Table 6. Numbers of species and populations in different estuary types by taxonomic class

Open Closed Delta Wetland

Spp Pop Spp Pop Spp Pop Spp Pop

Fishes 130 191 92 145 21 27 14 17

Reptiles 1 2 0 0 4 4 3 8

Birds 112 163 82 159 151 315 131 180

Mammals 4 8 0 0 0 0 1 1

Total 247 364 174 304 176 346 149 206

Table 7. Numbers of species and populations by driver of change (all ranks) and class

Infrastructure

- dams Water

extraction No threats

Spp Pop Spp Pop Spp Pop

Fishes 98 166 60 84 53 69

Reptiles 2 2 2 2 1 2

Birds 20 20 96 145 91 92

Mammals 2 2 0 0 1 1

Total 122 108 158 231 146 164