Coastal Zone Habitat-Use by Birds in Qatar: Insights from a Rapid Assessment Method During Spring Migration

Abstract

In the western part of the Arabian/Persian Gulf, coastal habitats such as inter-tidal wetlands and mangroves are scarce and poorly studied. We conducted a rapid assessment survey of bird species richness and abundance at the Fuwairit khor lagoon in northern Qatar, using a line transect count scheme to collect data from six different generic habitat types, repeated during five consecutive days in late April 2013. To further analyze the ecological requirements of the surveyed bird assemblage per habitat type, we assigned ecological guild categories to each species and distinguished among migrants, local, and regional breeders. Mangrove and intertidal mudflats hosted the highest bird densities and the most distinctive assemblages, while the beach habitat had high concentrations of birds but relatively few species. In contrast to the wetland habitats, near-shore marine areas and dune habitats had very limited numbers of birds and a relatively depauperate species assemblage. Employing a habitat perspective in a quantitative bird survey method shows that birds are effective biodiversity indicators for a rapid survey of coastal features in a poorly-studied region of the Arabian/Persian Gulf.

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658
Research Article
Coastal zone habitat-use by birds in Qatar:
Insights from a rapid assessment method
during spring migration
Stamatis Zogaris1* and Athanasios Kallimanis2, 3
1Hellenic Center for Marine Research, Institute of Marine Biological Resources and Inland Waters,
Anavissos, Greece; *e-mail: zogaris@gmail.com
2Department of Environmental and Natural Resources Management, University of Patras, Agrinio,
Greece
3current address: Department of Ecology, Aristotle University of Thessaloniki, Greece
Abstract
In the western part of the Arabian/Persian Gulf, coastal habitats such as intertidal wetlands and mangroves are
scarce and poorly studied. We conducted a rapid assessment survey of bird species richness and abundance at
the Fuwairit khor lagoon in northern Qatar, using a line transect count scheme to collect data from six different
generic habitat types, repeated during five consecutive days in late April 2013. To further analyze the ecological
requirements of the surveyed bird assemblage per habitat type, we assigned ecological guild categories to each
species and distinguished among migrants, local, and regional breeders. Mangrove and intertidal mudflats hosted
the highest bird densities and the most distinctive assemblages, while the beach habitat had high concentrations
of birds but relatively few species. In contrast to the wetland habitats, near-shore marine areas and dune
habitats had very limited numbers of birds and a relatively depauperate species assemblage. Employing a habitat
perspective in a quantitative bird survey method shows that birds are effective biodiversity indicators for a rapid
survey of coastal features in a poorly-studied region of the Arabian/Persian Gulf.
Titre : Utilisation des habitats côtiers par l'avifaune au Qatar: Aperçus d'une brève évaluation durant la migration
printanière
Les habitats côtiers, tels que les lagons, les zones intertidales associées et les mangroves, sont particulièrement
rares et peu étudiées dans la partie occidentale de l'Arabie/Golfe Persique. Pour la planification de la
conservation en milieu côtier, il est crucial d'effectuer un suivi de la biodiversité, et l'avifaune peut constituer un
bon bio-indicateur pour la définition des mesures de conservation prioritaires. Dans le présent cas d'étude, des
échantillonnages printaniers des populations d'oiseaux et de leur richesse spécifique par type d'habitat ont été
entrepris dans la zone côtière du lagon de Fuwairit dans le nord du Qatar. Pour cela, des comptages par transect
linéaire ont été répétés durant cinq jours consécutifs fin avril 2013. Afin d'analyser plus en détail les exigences
écologiques locales de l'avifaune, les espèces d'oiseaux ont été regroupées par guilde écologique, et une
distinction a été faite entre les espèces migrantes, celles se reproduisant localement et celles se reproduisant
dans la région. Les mangroves et les baies ensablées se sont avérées être les milieux accueillant les plus grandes
densités d'oiseaux et les assemblages les plus originaux, tandis que les plages présentaient une forte densité mais
une faible diversité d'espèces. Contrairement aux milieux aquatiques situés plus dans les terres, les habitats
marins côtiers et les dunes accueillaient un nombre très limité d'oiseaux et un assemblage d'espèces
relativement pauvre. L'utilisation d'une classification des habitats dans le cadre d'un suivi quantitatif des
populations d'oiseaux a permis de montré que l'avifaune pouvait constituer un indicateur efficace dans le but de
réaliser une évaluation rapide des caractéristiques écologiques d'une zone côtière sensible au sein d'une région
peu étudiée. Cette approche représente ainsi une procédure pertinente de suivi des populations dans le but de
mettre ne places des mesures de conservation.
Key words: Conservation assessment, birds, biodiversity, habitat, Qatar, coastal zone

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Introduction
Coastal zone wetland areas are among the most threatened landscape features in
the arid Middle Eastern countries. Wildlife-rich habitats such as intertidal lagoons
and mangroves are spatially scarce and poorly studied in the western part of the
Arabian/Persian Gulf. In several rapidly developing areas in this region, human-
induced pressures seem to especially stress coastal zone habitats [1]. Qatar is a good
example of a country where industrial and housing development is occurring at an
unprecedented rate along the coast, and biodiversity surveys are scarce and seldom
systematically updated [1,2]. The human population has boomed after industrial oil
production, and landscape conservation awareness has only recently begun to
influence protected-area policies [1,3]. Anthropogenic pressures and threats to
coastal ecosystems can only be rationally managed through strategic conservation
planning, immediate site protection, and conservation measures. Biodiversity should
be given utmost consideration, and certain indicator species groups, such as birds,
may provide useful, practical guidelines for conservation planning [4].
In areas lacking organized biodiversity inventories, rapid assessment procedures
using birds can build biodiversity knowledge baselines [4]. In our case study, we
conducted a rapid assessment of bird numbers and their habitat use in different
habitat types during a spring migration at Fuwairit, a small intertidal inlet system in
Northern Qatar. In order to best depict bird habitat use and preliminarily assess
relevant biodiversity patterns, analyses were made at three levels of organization:
habitat-based bird assemblage, food-centered ecological guild, and species residence
status.
Received: 5 January 2016; Accepted 4 April 2016; Published: 27 June 2016
Copyright: Stamatis Zogaris and Athanasios Kallimanis. This is an open access paper. We use the
Creative Commons Attribution 4.0 license http://creativecommons.org/licenses/by/3.0/us/. The
license permits any user to download, print out, extract, archive, and distribute the article, so long
as appropriate credit is given to the authors and source of the work. The license ensures that the
published article will be as widely available as possible and that your article can be included in any
scientific archive. Open Access authors retain the copyrights of their papers. Open access is a
property of individual works, not necessarily journals or publishers.
Cite this paper as: Zogaris, S. and Kallimanis, A. 2016. Coastal zone habitat-use by birds in Qatar:
Insights from a rapid assessment method during spring migration. Tropical Conservation Science
Vol. 9 (2): 658-676. Available online: www.tropicalconservationscience.org.
Disclosure: Neither Tropical Conservation Science (TCS) or the reviewers participating in the peer
review process have an editorial influence or control over the content that is produced by the
authors that publish in TCS.

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Methods
Study Area
Qatar is a small state (11,571 km2) with approximately 900 km of coastline. As a
subtropical desert peninsula, the mean annual rainfall is 81 mm, with an average
annual maximum temperature of 31oC [1]. During spring, a regionally important bird
migration takes place along the eastern coast of the Arabian Peninsula, part of the
West Asia-East Africa Flyway [5,6,7]. The study area, Fuwairit, is a small intertidal
inlet known locally as a khor lagoon system, on the north coast of Qatar (26°
1'57.02"N, 51°22'17.55"E). Fuwairit is one of the few areas along Qatar’s coast that
hosts mangrove-fringed lagoon habitats. The grey mangroves Avicennia marina,
although native to Qatar, are said to have been planted at Fuwairit during a
rehabilitation initiative in 1981 [8], but although they grow in natural stands nearby,
it is not certain whether or not mangroves existed on this site in the past. Today,
since industrial oil production dominates the economy, pressure on wood-cutting
and grazing of the mangroves has almost completely ceased, and these woodland
patches are showing signs of local expansion. For the last decade, the 2.4 km-long
marine beach of Fuwairit has been protected during the spring-summer period by
the Qatar Ministry of Environment because it hosts the second-largest known sea
turtle nesting rookery on mainland Qatar [9]. However, other biodiversity values,
such as its ornithological importance, are poorly documented, and beyond the beach
the site is not included in any protected-area designation [1].
Bird survey: A habitat approach
To record the abundance and differential use of the area’s major habitats, we
conducted a bird survey within a short time-window during the spring migration
period, from April 24th to 28th, 2013. One experienced ornithologist (the first author)
surveyed birds with binoculars (Zeiss 10x40B), using a modified line transect method
[4]; birds seen within the designated habitat areas were counted. The transect
survey followed a 3.7 km circuit route that was walked slowly with frequent stops on
each of the five consecutive days (always during near-flood tide conditions during
the early morning hours, between 5:00 and 9:00 a.m.). Bird numbers were recorded
in six constituent lists according to the generic habitat type where they were first
observed (i.e., if a bird was flushed by the observer it was only counted in the habitat
type where it was initially encountered). Total observation time amounted to 15
hours in good stable weather, most mornings being in nearly wind-still conditions.
The survey start-point was the Qatar Ministry of Environment Research Camp at the
north end of the beach, and the route always began from the inland lagoon-
mangrove side of the coastal spit and turned back along the long marine beach,
which is backed by dunes (Fig. 1). Unlike in other Eastern Middle East countries, bird
shooting was not observed during the study, and anthropogenic disturbance to birds
was minimal during the early morning hours; therefore, humans or domestic animals
did not noticeably influence bird species’ habitat use during the survey.

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Fig. 1 Left: Map of Arabian/Persian Gulf, Qatar, and the location of Fuwairit (dot). Right:
Fuwairit Khor study area. The transect route (3.7 km) for the ornithological survey taken
by one observer during five trials at Fuwairit. Six generic habitat types were surveyed
within representative area boundaries for each habitat (Satellite Image: Google Earth,
August 11th 2012).
Fuwairit’s khor lagoon system conveniently provides an opportunity to survey six
different generic habitat types nearly simultaneously during a fairly short line
transect. In the Arabian/Persian Gulf the term khor lagoon refers to a shallow tidal
inlet usually bordered by intertidal and subtidal mud and sand flats and supratidal
dune beaches and inland salt flats. These coastal zone features usually exhibit
distinct habitat formations along a marine-terrestrial gradient. In this study, a
generic habitat type unit was defined as a spatially contiguous landscape feature
that appears more or less homogenous throughout and is physiognomically
distinctive from other such units [see 10]. The generic habitat type's approximate
area boundaries visible to the observer were visually estimated. The six habitat types
surveyed were: 1) low mangrove with Avicennia marina; 2) lagoon pool (at near high
tide flood level); 3) exposed intertidal mudflat (within the khor lagoon); 4) beach
dunes; 5) beach shoreline including sandflats; and 6) near-shore marine waters (Fig.
2). Fuwairit's coastal wetland area, although not yet officially delineated, covers
approximately 150 ha with approximately 20 ha taken up by mangrove vegetation.
In order to interpret the ecological significance of the observed patterns, bird species
were used as ecosystem indicators through analyses of the collected data. Each bird
species was assigned an a priori ecological guild category on the basis of food and
foraging style following Weller’s approach [11], where ecological guild refers to a
functional trait categorization of “species (related or not) that exploit the same
resources in a similar way.” The guild assignment to species (Appendix 2) was
constructed using available knowledge of a bird’s ecology in the region [7, 12, 13]
and based on feeding-behavior ecological guild categorizations adapted from other
wetland habitat-based analyses [11, 14]. The residence status of the species
(migratory, resident breeder, breeder within Qatar) was also defined with the best

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available knowledge and corroborated by the Qatar bird checklist [12], as follows: M:
migrants, birds that only pass through the country or may also overwinter there; LB:
birds that were confirmed to breed locally in the study area or the immediate vicinity
during the study; QB: birds that are known to breed within or near the territory of
Qatar but for which no local breeding evidence was recorded during the survey.
Descriptive statistics, multidimensional scaling (NMDS) and relevant indices of
biodiversity were used to interpret ecological patterns; the data were analyzed using
MS Excel and the statistical software package R, version 3.2. [15].
Fig. 2. Low Avicennia marina mangroves within tidal lagoon mudflats and the research
camp at the north end of Fuwairit beach in the background (Photo: S. Zogaris).
Results
A total of 53 taxa (including two taxa classified to genus level) were identified at
Fuwairit; 2,163 individuals were recorded (Appendix 2). Overall there was a
significant relationship between bird species richness and abundance per habitat
type (Fig. 3), but the correlation was weak (R2=0.17, p=0.013) because of the
extremely high abundance of Lesser Crested Tern Thalasseus bengalensis roosting at
high tide on the beach habitat. Removing these samples from the analysis led to a
stronger correlation (R2=0.67, p<0.001). Despite the peak abundance of Lesser
Crested Tern, most species were sighted in relatively lower numbers. Six species
were sighted with only one recorded individual, and 13 species were recorded on
only one of the five count days.

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Habitat type was a strong discriminator of bird composition. Mangroves and
mudflats were inhabited by the most distinct species assemblages, clearly
differentiated from the other habitat types in the NMDS ordination plot (Fig. 4).
Most species concentrated on the mangrove and mudflat habitats (of the six generic
habitats surveyed). Moreover, 14 species appeared only in mangrove and nine only
in mudflats. Numbers of land birds (relative to seabirds and waterbirds) were rather
low, but the species richness within the mangroves was high (warblers, shrikes, and
several widespread land bird passerines). Numbers of seabirds and shorebirds were
relatively high (especially concentrating on the beach’s spit cape). This provides local
evidence that shallow intertidal mudflats of the khor lagoon system are an important
shorebird foraging area. As to be expected in a small wetland area along an arid
coastline in the Western Arabian/Persian Gulf, the numbers of breeding bird species
were low. Marine, dune and beach habitat exhibited very low species numbers:
there were two species recorded only in the dune habitat, one species only in the
beach habitat, and one only in the marine habitat.
In terms of residence categories, 40 species are migrants and only eight are
considered to breed locally; the remaining five are known to breed in and around the
Qatar Peninsula, but our observations did not confirm breeding on site. Local
breeders included some naturalized alien species (such as Common Myna
Acridotheres tristis) and one feral or escaped species (Rock Dove Columba livia),
perhaps due to the proximity of a small holiday home development nearby. Despite
the large difference in species richness, the total abundance of the three residence
categories are similar, but they are distributed very unevenly among the different
habitat types in both abundance and species richness, as the Kruskal Wallis results
show (see Appendix 1). Migrant species dominate in the mudflats (accounting for
76% of mudflat’s avifauna abundance and 81% of its species richness). Local
breeders are more commonly observed in the mangroves (accounting for 56% of
mangroves abundance but only 30% of its species richness). Local breeders are the
dominant species group in the dunes (where they are 88% of the avifauna
abundance and 69% of its species richness).
We designated seven different ecological guild groups as ecological functional
groups in the recorded avifaunal assemblage [11] in order to report ecological
functional diversity patterns and assess habitat conditions [16]. All 53 taxa are
assigned to the following guilds: 1) foliage-foraging insectivore (8 spp.); 2) marine
forager/fish-eater (4 spp.); 3) dry-ground forager (9 spp.); 4) flight insectivore (4
spp.); 5) raptor-like forager (6 spp.); 6) wading carnivore (3 spp.); 7) wet-surface
carnivore (19 spp). Dominant in abundance was the marine forager/fish-eating guild,
which accounted for 46% of the birds observed, but this was a species-poor group
including only terns, gulls, and one cormorant species. The most species-rich guild
was the wet surface carnivores, which accounted for 23% of the total abundance.
The distribution of these guilds among habitat types is very uneven, as is the number
of guilds observed in each habitat (Appendix 1). The most functionally diverse bird
assemblage per habitat was recorded in the mangrove habitat (both overall and in
each sample), followed by the mudflats and dunes (with the same total number of

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guilds observed but fewer groups observed in each sample). At the other end, the
lowest functional diversity was the marine waters with only two guilds recorded.
Fig. 3. Plot of species richness to
number of individuals per sample;
the beach habitat shows higher bird
number concentrations.
Fig. 4. Non-metric multidimensional
scaling plot of sites showing habitat
type categories based on recorded
bird populations per day-count
sample.

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Finally, as expected during spring migration in the region, the bird species
composition changed considerably from one day to the next. On a daily basis, we
observed approximately half the species that were recorded over the five day
period, indicating that more species would probably be recorded over a longer
period of observation. Among habitats the average number of species recorded daily
was strongly correlated with the total number of species recorded in the habitat
(R2=0.96, p<0.001). On average, the highest similarity in the avifauna species
composition among days (i.e., lowest species temporal turnover) was observed in
the mangroves, followed by the mudflats (Appendix 1); these two food-rich habitats
for birds seem to encourage some intercontinental migrants to stay longer and re-
fuel. The highest temporal turnover was observed in the species-poor marine
habitat, followed by the lagoon pool habitat. Among habitats, temporal turnover
(beta diversity) was strongly and negatively correlated with species richness at both
the sample scale (alpha diversity) and overall (gamma diversity) (spearman rank
correlation ρ>0.88 and p<0.02). Only Whittaker’s index was not significantly
correlated (Spearman p>0.2).
Discussion
Method
This study may be more valuable as a survey method exercise than as an interpretive
description of bird assemblages. Our short-term observations may overlook several
factors that influence bird habitat use, including supra-landscape factors, such as
intercontinental migration routes and migration timing [10], as well as stochastic
factors (e.g., weather events) that may also influence distributions, population
density, and habitat use [17, 18]. Further study, through a simple monitoring
scheme, is needed for a fine-scale description and interpretation of the bird
communities and their habitat use along Qatar's coastal zone.
However, our work provides one of the few systematic accounts of quantitative
habitat distribution of birds within the coastal zone in Qatar and is one of the few
published examples in the Arabian/Persian Gulf region. Since this constant-effort
survey trial was repeated over five consecutive days and there was no inter-observer
bias during the counts, the method seems adequate to provide an initial comparison
among the different habitats, especially since stable weather and similar tidal
conditions persisted during each count. In support of this notion is the fact that the
different facets of biodiversity (e.g., mean daily species richness, total species
richness, species turnover) were strongly correlated, indicating that we sampled the
study period well, and the consistency of information among the days shows that
most species were recorded. Overall, a longer period of observation during the
migration period will reveal more species [4], but we believe among-habitat diversity
patterns were satisfactorily represented during the study period. Furthermore, the
use of ecological guilds and the habitat perspective provides an important
foundation for further development of conservation-relevant monitoring and
indicator development [14, 16].

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Habitat-use insights
This study was appropriately timed since a high number of species utilized Fuwairit’s
habitats during this time. The majority of the species observed were long-distance
migrants that target what we assume are biologically productive habitat types with
important food sources (i.e., many birds were observed actively feeding). Our study
shows that two habitats had outstanding value for many bird species: mangroves
and lagoonal mudflats. These “special habitats” for conservation at this site were the
most bird species-rich habitats, with the most distinct bird assemblages, utilized by
many narrow-niche, habitat-specialist species.
Although tropical mangroves are known for their biological productivity [17], very
little ornithological research has been done on subtropical mono-species mangrove
patches such as these in Qatar [see 8, 19, 20, 21 and references therein]. Mangrove
habitat may be of significant value for biodiversity along a desert coastline such as
on the Arabian/Persian Gulf, where any woodland-like structure is extremely scarce.
In fact, the species-richness recorded at Fuwairit compares favorably to other
mangrove bird studies where longer term surveys took place [17]. In adjacent Iran,
Gashemi et al. [19] recorded the seasonal variation in the avifauna in a much larger
deltaic mangrove forest-wetland system and observed 56 waterbird species. Given
that in our case study we observed 28 species solely in the mangrove habitat in just
5 days (with an average of 15 species per day), it seems likely that with further study
the total species richness at Fuwairit may probably reach the scale of the Iranian
study. Mangroves are found in only half a dozen locations along Qatar’s coast [22],
and despite covering a small area of the coast, they mimic a woodland swamp
environment that is especially attractive to many migrant birds.
Our work also shows that khor lagoon mudflats are extremely important, since they
held the highest number of species and highest bird population densities during this
spring migration survey. Obviously many waders that use mudflats are specialized
feeders and are restricted to feeding in silty mudflats [11], which is why this habitat
is inhabited by a species-rich specialized avifauna distinct from the bird assemblages
in the other habitats. Nearly all long-distance migratory waders passing through the
area were observed only in this habitat. In our study, intertidal lagoon mudflats,
often occurring in close proximity to mangroves in the Gulf states, sustained the
highest number of "Near Threatened" bird species [23,24] (four species listed in the
IUCN Red List, see Appendix 2).
Other habitat types at Fuwairit were recorded as relatively species-poor during this
brief study, but they may also prove to have specific conservation value for birdlife.
Examples of this include the beach spit of Fuwairit, which is an important roosting
location for hundreds of terns and gulls since it provides seclusion from
predator/human disturbance and immediate access to adjacent food-rich lagoon
environments. The beach dune area is important for three locally breeding species:
Saunder’s Tern Sternula saundersi, Kentish Plover Charadrius alexandrinus, and
Greater Hoopoe-Lark Alaemon alaudipes. These ground-nesting birds are local
breeders and are probably sensitive to anthropogenic disturbance in such beach-side
environments. Fuwairit's marine habitat, although recorded as the most species-

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poor habitat type, hosted Socotra Cormorant Phalacrocorax nigrogularis, the only
species in our study area classified as globally threatened (listed as IUCN Red List
“Vulnerable” [23]).
1
2
3
4
5
6
Fig. 5. Characteristic species at Fuwairit: 1) Lesser Crested Tern, 2) Saunder’s Tern, 3) Indian Reef Heron,
4) White-eared Bulbul, 5) Kentish Plover, 6) Socotra Cormorant. (Photos: A. Vidalis).

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Implications for conservation
Qatar's baseline knowledge of its avifauna has much room for improvement; there is
no state-wide monitoring program for birds [12], and an official survey delineating
important bird areas is outdated [24, 25]. This situation of poor ornithological and
biodiversity surveying, even along quite accessible coastal areas, is by no means
unique in the states of the Western Arabian/Persian Gulf [26]. Readily documented
biodiversity indicators such as birds are important in conservation planning, but they
have been largely neglected in many parts of the Arabian Peninsula [13,27]. Our
study shows that an organized rapid survey or monitoring scheme of this type will
help identify and delineate areas with outstanding conservation values and help
associate their biodiversity with particular habitat features. Despite the lack of
organized bird survey data, the Western Arabian/Persian Gulf coast has been
considered as “one of the most important areas for wintering waders in the world”
[28], justifying more ornithological attention to Qatar’s coastal wetlands. Even today,
knowledge of migratory birds that use the East African-West Asian flyway has lagged
behind the western Europe-west Africa and the Nearctic flyways [6, 13]. Our
research exercise at Fuwairit promotes the wider development of basic survey
procedures based on rapid assessment approaches [29] in order to support policy-
relevant nature conservation in the region.
While Qatar is currently developing legislation and management agencies for nature
conservation [2], protected area creation is still in the early stages of
implementation [3]. There is abundant evidence that anthropogenic pressures and
threats to natural habitats concentrate on the country's coastal zone. Important sites
for biodiversity, such as mangroves and coastal lagoons, are under imminent threat
from poorly planned development [22]. The rapidly expanding capital city of Doha is
also likely to negatively influence Qatar's coastal natural areas with fringing holiday
home developments, road expansion, and increased recreational disturbance [30].
In our opinion, Fuwairit’s coastal zone and the surrounding landscape should be
specifically studied for inclusion within a protected area. This is clearly justified by
the site’s rich habitat composition, its scarce "special habitats" for migratory and
breeding birds, other biodiversity values such as the marine turtle rookery [9], and
its outstanding educational potential (Fig. 5).
Acknowledgements
This work was made possible by Biodiversity East, the Hellenic Centre for Marine
Research-HCMR, and through support from the Qatar Ministry of Environment. Aris
Vidalis assisted in field work and photography. Nancy Papathanassopoulou, Ali
Alhafez, ALan Rees and Nayla Beyrouthi are acknowledged for assistance in logistics
and project management.

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Appendix 1. Avifaunal diversity and abundance during the spring 2013 survey at Fuwairit.
Diversity is estimated as gamma diversity (total number of species observed throughout the
sampling period); as alpha diversity (mean number of species or mean Shannon diversity
index observed per sample, which in our case corresponds to per day of sampling); and as
beta diversity indices, which are either the ratio of gamma over alpha diversity or the mean
value of similarity indices among the samples collected in each habitat (we used Jaccard and
Sorensen similarity indices for presence/absence data and the Bray Curtis index with respect
to abundance). We classified birds according to their ecological guild (feeding-based
functional guild), residence type, and current information on their diversity and abundance.
For each mean value we also present the standard deviation. For each variable we
compared the value among the different habitats using the Kruskal Wallis test.

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Mangrove
Mudflat
Beach
Dune
Pool
Marine
Total
Kruskal
Wallis p
Total taxa richness (gama diversity)
28
29
17
12
6
9
53
Species richness per sample (mean±standard deviation alpha diversity)
15.40±3.21
13.80±3.96
6.80±2.49
5.20±1.79
3.00±0.70
2.20±1.92
7.73±5.69
<0.001
Shannon diversity index (mean±standard deviation per sample)
2.21±0.35
1.99±0.21
0.85±0.63
1.43±0.35
0.89±0.22
0.41±0.51
1.29±0.75
<0.001
Abundance (mean±standard deviation per sample)
108.2±31.1
93.6±25.7
167.8±175.7
17.8±6.4
10.6±9.2
34.6±39.4
72.1±89.4
0.005
Functional diversity (total number of different ecological guilds)
6
6
4
6
4
2
7
Functional diversity (mean±standard deviation number of different guild per sample)
5.20±0.45
4.20±0.84
3.40±0.55
3.80±0.84
2.00±0.70
1.00±0.70
3.27±1.55
<0.001
Beta diversity Whittaker index (gamma/mean alpha)
1.82
2.10
2.50
2.31
2.00
4.09
6.59
Beta diversity mean±standard deviation pairwise Jaccard similarity
0.48±0.10
0.41±0.07
0.29±0.09
0.40±0.17
0.23±0.17
0.27±0.26
0.35±0.18
0.002
Beta diversity mean±standard deviation pairwise Sorrensen similarity
0.32±0.05
0.29±0.03
0.22±0.06
0.28±0.09
0.18±0.10
0.18±0.16
0.24±0.11
0.002
Beta diversity mean±standard deviation pairwise Bray Curtis
13.96±2.33
13.99±2.15
8.42±2.03
5.35±2.13
4.33±1.31
2.77±1.31
8.18±4.88
<0.001
Distribution of species richness per guild (mean±standard deviation per sample)
foliage insectivores
4.8±1.6
0.0±0.0
0.0±0.0
0.8±0.4
0.0±0.0
0.0±0.0
1.00±1.56
<0.001
dry ground
5.0±1.0
1.2±1.3
1.4±0.5
1.8±0.8
0.0±0.0
0.0±0.0
1.57±1.85
<0.001
flight insects
1.6±1.34
0.8±0.4
0.4±0.5
0.4±0.5
0.2±0.4
0.2±0.4
0.60±0.81
0.122
fish
0.0±0.0
0.4±0.5
2.0±0.7
1.0±0.0
0.4±0.9
2.0±1.6
0.97±1.10
0.004
raptor
2.0±1.0
0.4±0.5
0.0±0.0
0.4±0.5
0.0±0.0
0.0±0.0
0.47±0.86
0.001
wading
1.6±0.55
1.2±0.4
0.0±0.0
0.0±0.0
1.4±0.5
0.0±0.0
0.70±0.79
<0.001
wet surface
0.4±0.55
9.6±2.5
2.8±1.9
0.8±0.4
1.0±1.0
0.0±0.0
2.43±3.60
<0.001
Distribution of abundance per guild (mean per sample)
foliage insectivores
16.6±10.5
0.0±0.0
0.0±0.0
0.2±0.4
0.0±0.0
0.0±0.0
2.80±7.39
<0.001
dry ground
66.2±23.9
2.2±2.1
4.8±2.7
5.6±3.8
0.0±0.0
0.0±0.0
13.13±25.88
<0.001
flight insects
9.4±11.0
2.2±1.3
2.4±4.3
0.8±1.3
0.2±0.4
0.2±0.4
2.53±5.51
0.058
fish
0.0±0.0
1.2±1.8
152.0±169.0
6.8±4.4
4.4±9.8
34.4±39.8
33.13±85.05
0.004
raptor
6.2±4.6
0.4±0.5
0.0±0.0
0.4±0.5
0.0±0.0
0.0±0.0
1.17±2.89
0.001
wading
5.8±3.4
3.6±3.2
0.0±0.0
0.0±0.0
4.8±4.2
0.0±0.0
2.37±3.43
<0.001
wet surface
1.4±2.2
83.8±25.7
8.4±10.5
3.2±2.1
1.2±1.1
0.0±0.0
16.33±32.51
0.001
Distribution of species richness per residence type (mean per sample)
migrants
9.2±3.1
11.2±2.9
2.8±2.1
1.2±0.8
2.8±0.4
0.6±0.9
4.63±4.52
<0.001
local breeders
4.4±0.5
2.6±1.5
3.2±0.8
3.6±0.9
0.2±0.4
0.8±0.4
2.47±1.71
<0.001
Qatar breeders
1.2±0.4
0.0±0.0
0.8±0.8
0.4±0.5
0.0±0.0
0.8±0.8
0.53±0.58
0.015
Distribution of abundance per migration type (mean per sample)
migrants
39.2±18.2
71.4±18.5
12.4±14.9
1.6±1.1
9.8±7.7
6.0±11.8
23.40±27.90
0.001
local breeders
65.2±22.7
22.2±10.8
10.2±3.3
15.6±6.9
0.8±1.8
2.2±1.8
19.37±24.23
<0.001
Qatar breeders
2.8±1.9
0.0±0.0
145.2±173.2
0.6±0.8
0.0±0.0
26.4±38.9
29.17±85.01
0.021

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674
Appendix 2. Species names, ecological guild category, residence status (M: Migrant, LB: Local Breeder, QB: Qatar Breeder), IUCN Red List Status (for codes see:
www.iucnredlist.org and www.birdlife.org/datazone/country/qatar) and total (summed) number of birds recorded during the five-day survey. Taxonomy and nomenclature
follows Qatar Bird Records Committee (www.qatarbirds.org/list.htm).
Scientific Name
English Name
Ecological Guild
Residence
Status
IUCN Red
List Status
Total No.
Recorded
Accipiter nisus
Eurasian Sparrowhawk
raptor-like forager
M
LC
1
Acridotheres tristis
Common Myna
dry-ground forager
QB
LC
14
Acrocephalus palustris
Marsh Warbler
foliage-foraging insectivore
M
LC
6
Acrocephalus scirpaceus
Eurasian Reed Warbler
foliage-foraging insectivore
M
LC
2
Acrocephalus sp.
Reed Warbler sp.
foliage-foraging insectivore
M
-
30
Alaemon alaudipes
Greater Hoopoe-Lark
dry-ground forager
QB
LC
3
Anthus cervinus
Red-throated Pipit
dry-ground forager
M
LC
2
Ardea cinerea
Grey Heron
wading carnivore
M
LC
11
Arenaria interpres
Ruddy Turnstone
wet-surface carnivore
M
LC
1
Butorides striata
Striated Heron
wading carnivore
QB
LC
1
Calidris alba
Sanderling
wet-surface carnivore
M
LC
6
Calidris alpina
Dunlin
wet-surface carnivore
M
LC
11
Calidris ferruginea
Curlew Sandpiper
wet-surface carnivore
M
NT
7
Charadrius alexandrinus
Kentish Plover
wet-surface carnivore
LB
LC
123
Charadrius lescheneultii
Greater Sand Plover
wet-surface carnivore
M
LC
1
Charadrius mongolus
Lesser Sand Plover
wet-surface carnivore
M
LC
155
Circus aeruginosus
Western Marsh Harrier
raptor-like forager
M
LC
2
Columba livia [var. domestica]
Rock Dove (feral/domesticated)
dry-ground forager
LB
LC
32
Egretta gularis
Indian Reef Heron
wading carnivores
M
LC
59
Falco tinnunculus
Common Kestrel
raptor-like forager
M
LC
1
Galerida cristata
Crested Lark
dry-ground forager
LB
LC
28
Haematopus ostralegus
Eurasian Oystercatcher
wet-surface carnivore
M
NT
4

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675
Hirundo rustica
Barn Swallow
flight insectivore
M
LC
60
Lanius collurio
Red-backed Shrike
raptor-like forager
M
LC
22
Lanius minor
Lesser Grey Shrike
raptor-like forager
M
LC
3
Lanius phoenicuroides
Turkestan Shrike
raptor-like forager
M
LC
6
Larus genei
Slender-billed Gull
marine forager/fish-eater
M
LC
68
Limosa laponica
Bar-tailed Godwit
wet-surface carnivore
M
NT
2
Merops apiaster
European Bee-eater
flight insectivore
M
LC
10
Motacilla flava
Yellow Wagtail
wet-surface carnivore
M
LC
18
Muscicapa striata
Spotted Flycatcher
flight insectivore
M
LC
3
Numenius arquata
Eurasian Curlew
wet-surface carnivore
M
NT
8
Numenius pheopus
Whimbrel
wet-surface carnivore
M
LC
6
Passer domesticus
House Sparrow
dry-ground forager
LB
LC
46
Phalacrocorax nigrogularis
Socotra Cormorant
marine forager/fish-eater
QB
VU
5
Phyloscopus collybita
Common Chiffchaff
foliage-foraging insectivore
M
LC
1
Phyloscopus sp.
Leaf Warbler sp.
foliage-foraging insectivore
M
-
8
Phyloscopus trochilus
Willow Warbler
foliage-foraging insectivore
M
LC
34
Pluvialis squatarola
Grey Plover
wet-surface carnivore
M
LC
37
Pycnonotus leucotis
White-eared Bulbul
dry-ground forager
LB
LC
114
Riparia riparia
Sand Martin
flight insectivore
M
LC
3
Spilopelia senegalensis
Laughing Dove
dry-ground forager
LB
LC
120
Sternula saundersi
Saunder's Tern
marine forager/fish-eater
LB
LC
69
Streptopelia decaocto
Eurasian Collared Dove
dry-ground forager
LB
LC
54
Sylvia communis
Common Whitethroat
foliage-foraging insectivore
M
LC
2
Sylvia curruca
Lesser Whitethroat
foliage-foraging insectivore
M
LC
1
Thalasseus bengalensis
Lesser Crested Tern
marine forager/fish-eater
QB
LC
852
Tringa erythropus
Spotted Redshank
wet-surface carnivore
M
LC
1
Tringa glareola
Wood Sandpiper
wet-surface carnivore
M
LC
3
Tringa nebularia
Common Greenshank
wet-surface carnivore
M
LC
2

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Tringa stagnatilis
Marsh Sandpiper
wet-surface carnivore
M
LC
4
Tringa totanus
Common Redshank
wet-surface carnivore
M
LC
11
Xenus cinereus
Terek Sandpiper
wet-surface carnivore
M
LC
90