Forest Ecology and Management 255 (2008) 3813–3818

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Forest Ecology and Management

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The effects of benthos and wetland area on shorebird abundance and species
richness in coastal mangrove wetlands of Leizhou Peninsula, China
Fasheng Zou a,*, Hanhua Zhang b, Tom Dahmer c, Qiongfang Yang a, Junxin Cai c,
Wei Zhang c, Chaoyu Liang b
a
South China Institute of Endangered Animals, 105 Xingang West Road, Guangzhou 510260, China
b
South China Sea Fisheries Research Institute, Guangzhou 510300, China
c
Sino-Dutch Integrated Mangrove Management & Coastal Protection Project, Zhanjiang, China

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

Article history: Twenty-two species shorebirds were recorded on mangrove wetlands surrounding Leizhou Peninsula,
Received 9 December 2007 Guangdong Province, China. One recorded species, Spoon-billed Sandpiper (Eurynorhynchus pygmeus), is
Received in revised form 11 March 2008 listed as endangered in the IUCN Red List (World Conservation Union) and this was the first record of the
Accepted 11 March 2008
species at Leizhou in this study. Most recorded species are listed in the Sino-Japan or Sino-Australia
Agreement on Conservation of Migratory Birds. Dunlin (Calidris alpina), Kentish Plover (Charadrius
Keywords:
alexandrinus), and Greater Sand Plover (Charadrius leschenaultii) were dominant, accounting for 34%, 32%,
Shorebirds
and 12% of total individuals, respectively. Five survey sites were designated by habitat type. There was no
Benthos
Wetland area significant difference in species, but total abundance differed significantly among survey sites. More
Mangrove wetland species were recorded in winter and spring than in summer. One hundred and sixty-five species of
Leizhou Peninsula of China benthos were recorded. Total mean biomass and density of benthos was 223.3  74.8 g/m2 (S.E.), density
210.97  67.6 ind./m2, respectively. Mollusks accounted for 83.4% of total biomass, and 75.6% of total density.
Both biomass and density of benthos varied significantly among survey sites. There was no significant
difference between spring and autumn on biomass or density of benthos. There was a significant and negative
correlation between number of shorebirds and densities of polychaetes on mudflats. Shorebird species
richness (F1,3 = 13.14, p = 0.036) and abundance (F1,3 = 12.59, p = 0.037) were significantly and positively
correlated with mudflat area. Stopping losses of mangrove habitat and tidal flats is the top priority measure
for shorebird conservation. Reducing the impacts of human disturbance on both habitats and birds is an
urgent necessity for mangrove wetland conservation management.
ß 2008 Elsevier B.V. All rights reserved.

1. Introduction Leizhou Peninsula in southwestern Guangdong Province

In temperate latitudes, the distribution of waders has been
shown to be closely related to the distribution and abundance of
the main wader food items (Puttick, 1984). Chironomid larvae
were the main prey of shorebirds at one study area in Spain where
there was a strong positive correlation between the number of
chironomid larvae available in each pond and the number of
feeding shorebirds (Sánchez et al., 2006). Sanderling (Calidris alba)
has been shown to be generalist feeders preying on crustaceans
(isopods, amphipods and mysids) and other invertebrates such as
polychaetes and bivalves that occur in the intertidal zone. The
distribution of Sanderling wintering in Japan was significantly
correlated with the distribution of Donax semigranosus, particularly
in late December 2000 and in early February 2001 (Nuka et al., 2005).
* Corresponding author. Tel.: +86 20 84182827; fax: +86 20 84183704.
E-mail address: zoufs@gdei.gd.cn (F. Zou).
supports the largest mangrove forests remaining in China and
covering around 7240 hm2. Mangrove wetlands around Leizhou
Peninsula were protected by gazettal of Zhanjiang Mangrove
National Natural Reserve (ZMNNR) in 1997. The reserve was added
to the Ramsar list of Wetlands of International Importance in 2002.
Zou et al. (2006) noted that the location of Leizhou Peninsula in the
East Asian-Australasian flyway of migratory birds contributes to its
importance as a staging site for migrants. They found that birds
were abundant on intertidal flat, which supported 69% of all birds
and 81% of all migrants. Intertidal flats proved important to
nationally and provincially protected and threatened species.
Earlier reports on birds in Chinese mangrove wetlands
provided species lists (Zhou et al., 1994; Wang et al., 1995; Zou
et al., 2001; Song and Lin, 2002). However, these researchers did
not document relationships between birds and environmental
factors. The purposes of this research were (1) to study relation-
ships between shorebirds and environmental factors, and to

0378-1127/$ – see front matter ß 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2008.03.020
3814 F. Zou et al. / Forest Ecology and Management 255 (2008) 3813–3818
determine whether bird species richness or bird abundance is
related to prey density or biomass, (2) to understand more clearly
the impacts of environmental factors on bird ecology so that more
clear guidance could be provided to policy makers with reference
to human exploitation of resources that are foods for both human
and avian users of the intertidal zone around Leizhou Peninsula.
2. Study area and methods
2.1. Study area
Leizhou Peninsula is the southernmost extent of mainland
China, located at 20–21.58N latitude. It is bounded to the west by
Beibu Bay, to the east by the South China Sea, to the south by
Qiongzhou Strait and Hainan Island (Fig. 1). The total land area of
Leizhou Peninsula is 12,470 km2 and the coastline is 1556 km
long. The area of intertidal flats is 867 km2. The Leizhou coastline
is deeply indented with more than 100 bays and estuaries. The
extensive coastline with its numerous sheltered bays and
muddy inlets creates conditions suitable for the development
of mangrove forests. Leizhou Peninsula has some of the largest
mangrove forests and the highest number of mangrove species on
China’s continental land mass. The total area of mangrove
coverage on Leizhou Peninsula was estimated in recent years as
7242 ha, and 15 species of mangrove were reported (Cai, 2004).
The government of Guangdong Province authorized establishment
of Zhanjiang Mangrove Provincial Nature Reserve on 1 January 1990
and erected the first nature reserve management station at Gaoqiao
Town of Lianjiang County. The provincial reserve was elevated to
Zhanjiang Mangrove National Nature Reserve on 7 November 1997.
Fig. 1. Sampling locations. Black shading indicates mangrove forest zone.
Table 1
Area of mangrove and intertidal flat at survey sites (hm2)
Survey sites Mangrove Intertidal flat
Gaojiao 914.2 266.6
Fucheng 285.2 933.3
Qishui 567.7 266.7
He’an 540 830
Wuli 5 35
The Zhanjiang coastal wetlands were added to the Ramsar list
of Wetlands of International Importance especially as Waterbird
Habitat in 2002. The protected area covers over 18,000 ha
(including 7242 ha of mangrove forest).
Leizhou Peninsula has a subtropical monsoon climate with an
annual average temperature of 22.3 8C and average annual rainfall
of 1500 mm. Rainfall is seasonal, and most precipitation occurs
during the summer rainy season or monsoon. The intertidal zone is
characterized by one or two tidal cycles per day. The tidal range is
approximately 2 m.
2.2. Sampling sites
Five sites were selected for sampling to test the hypothesis that
shorebird species richness and abundance depend on habitat area:
Gaoqiao and Qishui on the west coast; Fucheng and He’an on the
east coast; and Wuli on the south coast (Fig. 1). The areas of the five
survey sites are listed in Table 1. The habitats and environmental
conditions at the sampling sites were (1) Gaoqiao is largest stand of
mangrove and supports the most complete coverage of mature
mangroves in Leizhou Peninsula. Mangrove stand are segregated
by many small intertidal channels. Large areas of shrimp pond and
paddy lie adjacent to mangroves, but coastal woodlands (mainly
plantations) were far away from mangroves; (2) Fucheng supports
a small area of Sonneratia caseolaris (an introduced mangrove
species), large sandflats, and small stand of coastal forest near
mangroves; (3) Qishui is a busy commercial fishery port
characterized by large sandflats, sparse mangrove cover and
young mangrove trees; (4) He’an supports large sandflats where
the dominant mangrove was Avicennia marina. The site is
surrounded by many villages from which the human disturbance
on the intertidal zone is typically intense; (5) Wuli supports a small
stand of mangroves with dense coverage in most stands and a
small area of saltpans still used for production of sea salt.
2.3. Bird surveys
Sampling was carried out during four periods spanning four
seasons: winter (23 December 2002–13 January 2003), spring (3–
23 March 2003), summer (23 June–10 July 2003) and autumn (1–
13 November 2003) to test the hypothesis that shorebird species
richness and abundance were equal over seasons. The number of
sampling sessions was proportional to the area of mangrove and
intertidal flat at each sampling site: a combination of point counts
and transect counts for intertidal flats, and transect counts for
mangroves. Point count samplers reported all birds seen and heard
within a 150-m radius of designated center points during a 10-min
sampling periods. Transect counts were 50-m wide and 2000–
3000-m long. Average sample time for each transect was about
120 min. Bird species and numbers seen or heard were recorded
for all transects using methods described by Zou et al. (2001). At
each survey site except Wuli, where the area of mangroves and
intertidal flats was too small, each survey period included
sampling of two to three transects in mangroves and two to three
patches of intertidal flats. The time of day for sampling was
selected based on tide levels. Birds on intertidal flats were sampled
 F. Zou et al. / Forest Ecology and Management 255 (2008) 3813–3818 3815

Table 2
Checklist of shorebirds recorded at Leizhou Peninsula

Scientific name Common name Abundance Migratory status Conservation status

Rostratula benghalensis Greater Painted Snipe 1 R SJ, SA

Pluvialis squatarola Grey Plover 96 PM/WV SJ, SA
Charadrius alexandrinus Kentish Plover 1219 WV
Charadrius dubius Little Ringed Plover 7 R SA
Charadrius veredus Oriental Plover 1 PM
Charadrius leschenaultii Greater Sand Plover 435 PM/WV SJ, SA
Numenius arquata Eurasian Curlew 146 PM/WV SJ, SA
Numenius phaeopus Whimbrel 28 PM SJ, SA
Limosa limosa Black-tailed Godwit 1 PM/WV SJ, SA
Tringa ochropus Green Sandpiper 10 PM/WV SJ
Arenaria interpres Ruddy Turnstone 5 PM SJ, SA
Tringa totanus Common Redshank 213 PM/WV SJ, SA
Heteroscelus brevipes Grey-tailed Tattler 24 PM
Actitis hypoleucos Common Sandpiper 26 PM/WV SJ, SA
Tringa stagnatilis Marsh Sandpiper 76 PM/WV SJ, SA
Tringa glareola Wood Sandpiper 3 PM/WV SJ, SA
Xenus cinereus Terek Sandpiper 15 PM SJ, SA
Tringa nebularia Common Greenshank 134 PM/WV SJ, SA
Calidris abla Sanderling 32 PM SJ, SA
Eurynorhynchus pygmeus Spoon-billed Sandpiper 3 PM SJ
Gallinago megala Swinhoe’s Snipe 2 PM/WV SJ, SA
Calidris alpina Dunlin 1283 WV SJ, SA

Migratory status: WV-winter visitor, PM-passenger, R-resident. Conservation status: SJ-species in Sino-Japan Agreement on Conservation of Migratory Birds, SA-species in
Sino-Australia Agreement on Conservation of Migratory Birds.

during low tides when the flats were exposed and birds could
access them. Birds in mangrove were sampled during high tides
when birds could not access intertidal flats. Binoculars (10) and
spotting scopes (30 and 60) were used for detecting and
identifying birds.
2.4. Benthos sampling
Benthos was sampled in the high, medium and low tide zones in
March (spring) and September (autumn) of 2002. At each sample
point, three samples of mud/sand were excavated from a
50 cm  50 cm  30 cm deep quadrat. The samples were imme-
diately filtered through a series of sieves with progressively
smaller mesh sizes (first 3.35 mm, second 1 mm, and third
0.5 mm). Organisms retrieved from the mud/sand samples were
preserved in 10% formalin. All samples were identified to species
and weighed after air drying on paper. The weight of shells was
included for bivalves. Biomass (g/m2) and density (ind./m2) of each
invertebrate category were calculated using methods described by
Liang et al. (2005).
2.5. Data analysis
Dominant bird species were designated as those accounting for
over 10% of the total number of individuals recorded in all sampling
sessions (Zheng, 1995). Migratory status was based on distribution
of the avifauna of South China (Viney et al., 1994). One-way analysis
of variance (ANOVA) and post hoc multiple comparison were used to
compare shorebird abundance and density, and benthos biomass
and density between seasons or survey sites. All data were
transformed to natural logarithms for correlation analyses and
linear regression analyses. Pearson two-tailed correlation analysis
was used to analyze relationships between shorebird species
richness or abundance and habitat area, biomass and density of
benthos. Linear regression was used to explain relationships
between species richness and abundance of shorebirds and area
of habitat. We used principal component analysis (PCA) to identify
the major independent pattern of covariation in structural
attributes. All components with eigenvalues >1 were retained for
further analysis. Each component was interpreted by examining its
factor loadings. All statistical tests were conducted using SPSS
version 11.5.
3. Results
3.1. Bird species richness and abundance
Twenty-two species shorebirds were recorded in mangrove and
coastal flat habitats of Leizhou Peninsula. All recorded birds were
winter visitors or passage migrants except for Greater Painted
Snipe (Rostratula benghalensis) and Little Ringed Plover (Charadrius
dubius), which are local residents. Of the 22 recorded species, 18
and 17 are listed in the Sino-Japan Agreement on Conservation of
Migratory Birds and the Sino-Australia Agreement on Conservation
of Migratory Birds, respectively (Table 2). Dunlin (Calidris alpina),
Kentish Plover (Charadrius alexandrinus), and Greater Sand Plover
(Charadrius leschenaultii) were dominant, accounting for 34%, 32%,
and 12% of total bird numbers, respectively.
Twelve, 15, 12, 12 and 9 species of shorebirds were recorded at
Gaojiao, Fucheng, Qishui, He’an and Wuli, respectively. Species
richness did not differ significantly among survey sites (F4,15 = 1.019,
p = 0.429) but total bird numbers did (F4,15 = 3.414, p = 0.036). The
greatest number of shorebirds was recorded at Fucheng area, while
the lowest was recorded at Wuli area. More birds were recorded at
Fucheng than at Gaojiao area, and more birds were recorded at
Fucheng than at Wuli at a = 0.05 (Fig. 2).
Fifteen, 14, 7 and 13 species shorebirds were recorded in winter,
spring, summer and autumn, respectively. More species were
recorded in winter than in summer ( p = 0.016) and more species
were recorded in spring than in summer ( p = 0.031) (post hoc
multiple comparisons). Total number of individuals did not differ
significantly among seasons (F3,16 = 1.447, p = 0.266) (Fig. 3).
3.2. Species richness and abundance of benthos
One hundred and sixty-five species of benthos, including
species of Annelida, Sipuncula, Molluska, Arthropoda and Echino-
dermata were recorded. The dominant species were Cerithidea
cingulata, Batillaria zonalis, Littoraria (Palustorina) melanostoma,
Littoraria (Palustorina) articulata, Neritina (Dostia) violacea, Mono-
3816 F. Zou et al. / Forest Ecology and Management 255 (2008) 3813–3818

Table 3
Rotated component matrix of variables

Variable The first axis The second axis The third axis

Area of mangrove 0.020 0.363 0.818

Area of intermudflat 0.305 0.673 0.003
Species number of benthos 0.090 0.269 0.336
Density of benthos 0.992 0.081 0.078
Biomass of benthos 0.979 0.088 0.171
Species of polychaetes 0.773 0.297 0.518
Density of polychaetes 0.226 0.959 0.093
Biomass of polychaetes 0.678 0.633 0.339
Fig. 2. Bird species richness and abundance at five survey sites. Species of mollusks 0.148 0.018 0.532
Density of mollusks 0.995 0.073 0.032
Biomass of mollusks 0.990 0.085 0.097
Species of crustaceans 0.366 0.553 0.023
Density of crustaceans 0.263 0.095 0.957
Biomass of crustaceans 0.002 0.135 0.927
Diversity index of benthos 0.131 0.875 0.360
Evenness index of benthos 0.177 0.898 0.403

Fig. 3. Seasonality of bird species richness and abundance.
donta labio, Glauconome chinensis, Crassostrea sp., Mactra (Mectra)
venerformis, Gracilaria asiatica, Enteromorpha clathrata and Enter-
omorpha flexuosa. Total biomass of benthos was 223.3  74.8 g/m2
(S.E.). Biomass of benthos differed significantly among survey sites
(F4,5 = 12.325, p = 0.008). Greatest biomass was recorded at He’an and
lowest at Fucheng. The biomass of benthos at He’an was significantly
greater than at Gaojiao, Fucheng, Qishui, and Wuli at a = 0.05 (Fig. 4).
Biomass of benthos did not differ significantly between spring and
autumn (F1,8 = 0.000, p = 0.997). Total mean density of benthos was
211  67.6 ind./m2. Density of benthos differed significantly among
survey sites (F4,5 = 9.915, p = 0.014). Similar to biomass, greatest
density was at He’an and lowest at Fucheng. The density of benthos at
He’an was significantly higher than at Gaojiao, Fucheng, Qishui, and
Wuli at a = 0.05 (Fig. 4). Benthos densities did not differ significantly
between spring and autumn (F1,8 = 0.000, p = 0.991).
Mollusks and crustaceans were the two most important groups
of benthos in the mangrove wetlands of Leizhou Peninsula in terms
of biomass and density. Biomass of mollusks accounted for 83.4% of
total biomass, and density of mollusks accounted for 75.6% of total
density. Crustaceans accounted for 12.9% of total biomass and
18.1% of total density.
axes one, two and three had eigenvalues of 6.33, 5.01 and 3.89,
respectively. Factor loadings indicated that the first principal
component (PC1) represented biomass and density of prey
including total benthos and Molluska only. The second principal
component (PC2) explained the effect of polychaetes on shorebirds
in terms of density of polychaetes and species richness of total
benthos. The third principal component (PC3) was the density and
biomass of crustaceans, and area of mangrove. The PCA score
showed the ranking of various groups of invertebrates. Density and
biomass of benthos exerted a greater impact on shorebird numbers
than did species richness of benthos.
3.4. Relationship between shorebirds and environmental variables
There was a significant and negative correlation between number
of shorebirds and densities of polychaetes on the mudflats of Leizhou
Peninsula. The resulting regression equation was: N = 8.7 
108D 1.47 (N: number of shorebirds, D: densities of polychaetes,
F1,3 = 15.70, p = 0.03). Shorebird species richness was significantly
and positively correlated with mudflat area: S = 60.3A0.12 (S: species
richness of shorebirds, A: area of mudflat, F1,3 = 13.14, p = 0.036).
Shorebird numbers were also significantly and positively correlated
with mudflat area: N = 380.2A0.66 (F1,3 = 12.59, p = 0.037).
4. Discussion
4.1. Relationship between shorebirds and environmental variables

3.3. Principal components analysis of environment factors
In total, 16 variables were used for PCA (Table 3). The PCA of
food and area factors yielded three new, synthetic variable axes.
The first three axes of PCA accounted for 95.2% of the variance. PCA
Fig. 4. Species, biomass and density of benthos.
Benthic invertebrates are primary foods of waterbirds (Reeder,
1951; Rundle, 1982), and these invertebrates strongly influence
the distribution (Colwell and Landrum, 1993) and feeding behavior
(Murkin and Kadlec, 1986) of waterbirds. The significant and
negative correlation between number of shorebirds and densities
of polychaetes on the mudflats of Leizhou Peninsula was probably
due to sustained shorebird predation on polychaetes and resulting
declines in densities of benthic invertebrates (Mercier and McNeil,
1994). Sánchez et al. (2006) reported predation by shorebirds can
reduce benthos larval density by more than 30%, and reduce larval
biomass by 37–49%.
The density and biomass of macrobenthos at mangrove
wetlands surrounding Leizhou Peninsula were 211 ind./m2 and
223 g/m2, respectively. These values are greater than those
reported for most other coastal sites in China both south and
north of Leizhou Peninsula (Table 4). Macrobenthos is a natural
prey for shorebirds and provides the forage base for the shorebird
assemblage at Leizhou Peninsula. The density and biomass of
benthos at Leizhou Peninsula were not strongly seasonal. This
 F. Zou et al. / Forest Ecology and Management 255 (2008) 3813–3818
Table 4
Comparisons of benthos density and biomass between Leizhou Peninsula and other places in China
Sampling sites Density Biomass
DZGMW 98 105
QLGMW 352.4 198.5
QZB 144 23.13
YQB 85 41.95
YZE 150 17.57
DZGMW: Dongzhaigang mangrove wetland, Hainan Island; QLGMW: Qinglangang mangrove wetland, Hainan Island; QZB: Quanzhou Bay of Fujian Province; YQB: Yueqing
Bay of Chejiang; YZE: Yangtze Estuary; LZ: Leizhou Peninsular.
means that the shorebird prey base was available throughout the
year. The similarities in shorebird abundance across seasons at
Leizhou Peninsula supports the contention that prey biomass is
relatively constant.
Environmental factors other than prey abundance affect species
richness and abundance of shorebirds. Water depth has a direct
effect on feeding activities of shorebirds. Salinity can affect
surface-feeding shorebirds because different species of aquatic
invertebrates exhibit differences in salinity tolerance (Liang et al.,
2002). Correlation analysis revealed that shorebird species
richness at Leizhou mangrove wetlands correlated significantly
and positively with mudflat area. Shorebird numbers were also
significantly and positively correlated with mudflat area. Greater
biomass (119 g/m2) and density (145 ind./m2) of benthos were
recorded at Wuli than at Gaojiao or Fucheng. But, species richness
and abundance of shorebirds were the lowest at Wuli because of
smallest area both mangrove and intertidal flat. Surely, wetland
area was an important factor affecting shorebird distribution at
Leizhou Peninsular. Over 80% of shorebirds at Leizhou Peninsula
were recorded on intertidal flats, which proved to be the most
important habitat for shorebirds. This means that the locations of
proposed mangrove plantations must be considered with respect
to conservation of intertidal flats when planning recovery of lost
mangrove areas, as suggested by Zou et al. (2006).
Predator avoidance ability is the key parameter affecting fitness
in shorebirds (Bednekoff and Houston, 1994; Cresswell and
Whitefield, 1994; Lank et al., 2002). Shorebirds will change their
feeding behavior in order to reduce perceived risks when humans
are present (Fitzpatrick and Bouchez, 1998; Thomas et al., 2003).
Yasué (2006) reported that distance to forest cover was the only
important factor affecting bird presence. Shorebirds were more
frequently present in sections that were further from forest cover.
Greatest biomass (641 g/m2) and density (587 ind./m2) of benthos
were recorded at He’an with the second largest area, but less bird
species richness at He’an than at Fucheng, less bird abundance at
He’an that at Fucheng or Qishui. High product of benthos and large
wetland area had not resulted in richer bird species and number,
that meant factor other than prey abundance/biomass and area
appeared to affect shorebird distribution at Leizhou also. There
were many villagers around mangrove wetland at He’an for
activities including fishing, collecting of benthos, and rearing of
ducks. Human presence on the mudflats was frequent and often
intense, and appeared to be an important factor affecting shorebird
distribution. This remains to be addressed by further study.
4.2. Conservation implications
Many shorebirds protected by international migratory bird
treaties and conventions inhabit the coastal zone of Leizhou
Peninsula. Spoon-billed Sandpiper (Eurynorhynchus pygmeus), an
endangered species, only occupies four wetlands in China,
including Yancheng of Jiangsu province, Chongming Island of
Shanghai, Mai Po of Hong Kong and Lanyang of Taiwan (Birdlife
International, 2003). The first species record on Leizhou Peninsula
3817
Species Relative position to LZ References
68 South of LZ Zou et al., 1999a
45 South of LZ Zou et al., 1999b
256 North of LZ Li et al., 2006
124 North LZ Yang et al., 2007
50 North LZ Sun et al., 2007
was made during this study, demonstrating the importance of
Leizhou Peninsular is a staging area for shorebirds. Many
mangrove forests have been destroyed and fragmented to enable
economic development in Leizhou (Zou et al., 2006). Stopping the
losses of mangroves and tidal flats is the first important measure
for conservation of shorebirds. An additional important conserva-
tion measure is to reduce the impact of human disturbance to
shorebirds because survival in non-breeding habitats is thought to
limit many shorebird populations (Duffy et al., 1981; Hitchcock
and Gratto-Trevor, 1997). Local villagers engage in traditional
farming and grazing landward of the coastal zone, but also dig for
crabs and collect oysters in mangrove and mudflat/sandflat
habitats. Several species of intertidal invertebrates, including
Phascolosoma esculenta, P. scolops, Sipunculus nudus, and many
species of mollusks (Crassostrea sp., Meretrix meretrix, Solen
grandis) are considered delicious foods by local people. Many
farmers worked on the intertidal flat and collected benthos during
low tide at each survey site during this study. Qishui is also a busy
commercial fishery port. Humans probably compete with shore-
birds for food, and also impact shorebird feeding behaviors on
Leizhou Peninsula. To enhance shorebird conservation, managers
must strive to minimize human activities that disturb shorebird
feeding and thus reduce shorebird fitness.
Acknowledgements
We are grateful to officials of the Division of Wildlife Conserva-
tion and Management in Guangdong Forestry Department and
members of the Integrated Mangrove Management and Coastal
Protection Project. We thank members of the District Forestry
Bureaus, County Forestry Bureaus and Town Forestry Stations of
Leizhou Peninsular for field assistance. We also thank D. Binkley and
anonymous reviewers for comments on earlier versions of this
manuscript. Guangdong Science Fund (Nos. A3050102, 2004B3
3701015) and Sino-Dutch Integrated Mangrove Management and
Coastal Protection Project supported this project financially.
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