Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 June 2021 | 13(7): 18719–18737
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
https://doi.org/10.11609/jott.6992.13.7.18719-18737
#6992 | Received 14 December 2020 | Final
received 10 April 2021 | Finally accepted 16 June 2021
Avian species richness in
traditional rice ecosystems: a case study from upper Myanmar
Steven G. Platt 1, Myo Min Win 2, Naing Lin 3 , Swann Htet Naing Aung 4, Ashish John 5 & Thomas R. Rainwater 6
1–4 Wildlife Conservation
Society–Myanmar Program, No. 12, Nanrattaw St., Kamayut Township, Yangon, Myanmar.
5 Wildlife Conservation
Society–Cambodia Program, #21 Street 21, Sangkat Tonle
Bassac, Kam Chamkamon, P.O.
Box 1620, Phnom Penh, Cambodia.
6 Tom Yawkey
Wildlife Center & Belle W. Baruch Institute of
Coastal Ecology and Forest Science, Clemson University, P.O. Box 596,
Georgetown, South Carolina 29442, USA.
1 sgplatt@gmail.com, 2 mmwin@wcs.org,
3 nilzmw@gmail.com, 4 saung@wcs.org, 5 ajohn@wcs.org,
6 trrainwater@gmail.com (corresponding author)
Editor: Anonymity
requested. Date of publication:
26 June 2021 (online & print)
Citation: Platt, S.G., M.M. Win, N. Lin,
S.H.N. Aung, A. John & T.R. Rainwater (2021). Avian species richness in
traditional rice ecosystems: a case study from upper Myanmar. Journal of Threatened Taxa 13(7): 18719–18737. https://doi.org/10.11609/jott.6992.13.7.18719-18737
Copyright: © Platt et al. 2021. Creative Commons Attribution
4.0 International License. JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: Andrew Sabin
and the Andrew Sabin Family Foundation; Critical Ecosystem
Partnership Fund; Save Our Species (IUCN).
Competing interests: The authors
declare no competing interests.
Author details: Steven G. Platt is the Associate Conservation
Herpetologist for Wildlife Conservation Society in Southeast Asia. He is
responsible for herpetological conservation projects in Myanmar, Lao, PDR, and
Cambodia. Conservation of endemic tortoises and freshwater turtles in Myanmar
is a major focus of his work. Myo Min Win is the Manager of the Burmese
Roofed Turtle Restoration Project for Wildlife Conservation Society-Myanmar
Program. He oversees all aspects of the field project on the upper Chindwin
River, including egg collection and incubation, head-starting, assurance
colonies, and community-based conservation efforts. Naing
Lin is the Wildlife Conservation Society-Myanmar Program Landscape
Coordinator for the Ayeyarwady River Landscape
Conservation Project. He is responsible for a basin-wide conservation
initiative that focuses on the regions unique biodiversity. He is currently
pursuing a graduate degree in Conservation Project Management at the University
of Kent, UK. Swann Htet Naing Aung is a Field
Research Officer with the Wildlife Conservation Society—Myanmar Program
responsible for reintroduction of the critically endangered Burmese Star
Tortoise at Shwe Settaw
Wildlife Sanctuary. Swann is also involved in various other turtle conservation
efforts, including the Burmese Roofed Turtle on the Chindwin River. Ashish
John is a Community Conservation Management Advisor with the Wildlife
Conservation Society - Cambodia Program. In this capacity he assists rural
communities with developing and implementing conservation schemes linked to
community-based natural resource management and ecotourism. He works primarily
in Cambodia and Myanmar. Thomas R. Rainwater is a Research
Scientist with the Tom Yawkey Wildlife Center and
Baruch Institute of Coastal Ecology and Forest Science of Clemson University in
Georgetown, South Carolina, USA. His
current research focuses on the ecology, ecotoxicology, and conservation of wild
reptiles, birds, and mammals.
Author contributions: Steven G. Platt—Conceived and
designed the study, conducted fieldwork, analyzed data, wrote the manuscript,
and secured funding. Myo
Min Win—Conducted fieldwork, and assisted with manuscript preparation. Naing Lin—Conducted fieldwork, assisted with
manuscript preparation, and prepared the abstract translation. Swann Htet Naing
Aung—Conducted fieldwork, prepared maps and figures, and assisted with
manuscript preparation. Ashish
John—Conducted fieldwork and assisted with manuscript preparation. Thomas R. Rainwater—Conceived and designed
the study, analyzed data, prepared figures, and assisted with manuscript
preparation. All authors reviewed and
approved the final draft.
Acknowledgements: We thank the Ministry of
Environmental Conservation and Forestry for granting us permission to conduct
research in Myanmar. Fieldwork was made possible by generous grants from Andrew
Sabin and the Andrew Sabin Family Foundation, the Critical Ecosystem
Partnership Fund, and Save Our Species (IUCN). TRR was supported by the Yawkey Foundation and Clemson University. The able field
assistance of Tun Win Zaw
and Zaw Naing Oo was
critical to the success of our project. We also thank Saw Htun
for facilitating our work in many ways, Cassandra Paul, Richard Kaminski, and
Alex Diment for providing literature, Robert Tizard, Thet Zaw Naing, and Simon Mahood for sharing their extensive knowledge of Southeast
Asian birds, and Lewis Medlock and two anonymous reviewers for insightful
comments on an early draft of this manuscript. This paper represents technical
contribution number 6925 of the Clemson University Experimental Station.
Abstract: Rice Oryza sativa
ecosystems provide foraging and nesting habitat for a variety of birds. Myanmar
is a major rice-producing nation and yet bird use of rice ecosystems remains
largely unstudied. We present the results of a case study of avian species
richness in a traditional rice ecosystem at Limpha
Village in upper Myanmar. The rice field at Limpha
occupies 17.5 ha where a single crop is produced each year without chemical
inputs (fertilizer and pesticides). Village lands are contiguous with the
buffer zone of Htamanthi Wildlife Sanctuary. We
conducted bird surveys of the rice field during dry and wet seasons (2013–20)
and documented the occurrence of 85 species (exclusive of Buttonquail these
included 58 resident species, 20 migratory species, six species with both resident
and migratory populations in upper Myanmar), including 10 species of
conservation concern. Species richness was greatest during the dry season when
an influx of Palearctic migrants was present. We ranked 52 species as Common,
23 as Uncommon, and 10 as Rare. Most birds used the rice field as foraging
rather than breeding habitat. Insectivore was the most common feeding guild (43
species), followed by Omnivore (22 species), Carnivore (12 species), Granivore (6 species), Frugivore (1 species), and
Nectarivore (1 species) guilds. We observed eight species associated with
domestic Water Buffalo Bubalus bubalis and 15 species foraging at active fires or in
burned areas in the rice field. Piles of rice straw are important foraging
sites for several species. Low intensity agricultural practices, habitat
heterogeneity, and proximity to the nearby swamp, forest, & Chindwin River
are probably responsible for the relatively high avian species richness at Limpha. Future agricultural intensification could
negatively impact avian species richness in the Limpha
rice field. Our findings suggest that traditional rice agriculture is
compatible with conservation objectives in the buffer zone of Htamanthi Wildlife Sanctuary. Our study, however, requires
replication before generalizations can be made concerning the value of
traditional rice ecosystems to avian conservation in Myanmar.
Keywords: Bird conservation, bird
diversity, buffer zone, Chindwin River, Htamanthi
Wildlife Sanctuary, Oryza sativa, rice field, Sagaing
Region, traditional agriculture, water buffalo.
Introduction
Land devoted to the production of food, fiber,
plant oils, and other resources used by human society occupies a substantial
and increasing proportion of terrestrial biomes around the world (Bennett et
al. 2006). As the extent of anthropogenically-modified landscapes expands to
meet the needs of a growing human population, the fate of global biodiversity
will increasingly depend on the quality and characteristics of farming
landscapes (Pimental et al. 1992; Pino et al. 2000;
Perfecto et al. 2009; Friskhoff et al. 2014).
Farmlands vary widely in their ability to support biodiversity with some
species being lost from agricultural landscapes, while other species persist
and can even proliferate (Friskhoff et al. 2014).
Despite the species loss that accompanies the conversion of wildlands to
farmland (Rutt et al. 2019), a growing body of
literature suggests that agricultural landscapes can make substantial
contributions to global biodiversity conservation (Pimental
et al. 1992; Jackson & Jackson 2002; Perfecto et al. 2009; Van der Weijden 2010).
Rice Oryza sativa is one of the most important food crops in the
world (Forĕs & Comín
1992; Bambaradeniya & Amarasinghe
2003). Rice is the primary source of nutrition for over half of the global
human population and constitutes one-fifth of the world’s grain supply (Elphick 2010). Rice is grown in at least 114 countries,
rice ecosystems occupy >156 million ha of land (Elphick
2010), and more land is devoted to rice than any other agricultural crop (Forĕs & Comín 1992). Because
most rice is grown under flooded conditions (Lawler 2001), rice ecosystems are
in effect, agronomically-managed freshwater marshes supporting a single species
of cultivated grass (Bambaradeniya & Amarasinghe 2003). As managed wetlands, rice ecosystems constitute
important habitat for a diverse array of wetland plants, invertebrates, and
vertebrates (Lawler 2001; Czech & Parsons 2002; Bambaradeniya
& Amarasinghe 2003; Halwart
2006; Elphick 2010). Among vertebrates, rice
ecosystems are notable for providing foraging and nesting habitat for a wide
variety of birds (Remsen et al. 1991; Dhindsa &
Saini 1994; Hohman et al. 1994; Czech & Parsons 2002; Elphick
2010), including locally rare and globally imperiled
species (Van der Weijden 2010). Furthermore, in some
areas, (particularly in Asia) waterbirds have come to
rely on rice ecosystems owing to the widespread loss of natural wetlands (Fasola & Ruiz 1996; Czech & Parsons 2002; Elphick 2010). Indeed, rice fields are often the best
remaining wetland habitat for birds in many regions of the world (Fasola & Ruiz 1996; Elphick
2010; Fujioka et al. 2010).
Despite the acknowledged importance of rice ecosystems to avian
conservation (Round 2002; Amano 2009; Van der Weijden
2010), bird use of this habitat outside of North America and Europe remains
under-studied (Czech & Parsons 2002; Elphick
2010). This is especially true in Asia where 90 % of the global rice crop is
produced (Lawler 2001; Czech & Parsons 2002), and yet information on bird
use of rice ecosystems remains surprisingly sparse (Duckworth 2007; Amano 2009;
Fujioka et al. 2010; Sundar & Subramanya 2010).
This situation is lamentable given the potentially high conservation value of
rice ecosystems (Hohman et al. 1994; Amano 2009), coupled with the need to craft
biologically-based management strategies that can maintain avian diversity
without compromising agricultural production objectives (Van der Weijden 2010; Kumar & Sahu
2020). Furthermore, an enhanced understanding of avian ecology in rice
ecosystems is critical for predicting the impacts of agricultural
intensification likely to accompany the rapid economic development now
occurring in much of southeastern Asia (e.g., Rao et
al. 2013; Clements et al. 2014; Bhagwat et al. 2017).
Myanmar is one of the largest rice-producing nations in the world (GRiSP 2013), and rice production generates direct or
indirect livelihoods for >75 % of the population (Naing et al. 2008). Rice
is grown on 8 million ha of farmland with annual production amounting to >30
million tons (GRiSP 2013). Major rice-growing areas
of Myanmar include the Ayeyarwady Delta, with
significant production also occurring in the lowlands of Mandalay, Sagaing, and Magway Regions (Hla Myo Thwe et al. 2019). Rice was
traditionally a monsoon crop until the 1970–80s when high-yielding varieties
were introduced by the Myanmar government that allow double-cropping, i.e.,
cultivation of a crop during both the wet and dry seasons, with the dry season
crop dependent on adequate irrigation (Naing et al. 2008; GRiSP
2013). Rice is typically grown on small farms (averaging 2.3 ha) by
resource-poor farmers or landless agricultural laborers (Naing et al. 2008)
Despite the large amount of land devoted to rice production and the
importance of this crop to the agricultural sector, other than passing mention
of rice fields in scattered sources (Smythies 1953; Thet Zaw Naing et al. 2017)
virtually nothing is known about bird use of rice ecosystems in Myanmar. We
here present a case of study of avian species richness in a traditional rice
ecosystem of upper Myanmar. In this study, we follow Bambaradeniya
& Amarasinghe (2003) and define a
traditional rice ecosystem as a sustainable agricultural system dedicated
primarily to the production of rice (and occasionally other crops such as fish)
that employs minimal mechanization and few if any chemical inputs. Traditional
rice ecosystems are generally assumed to support higher levels of biodiversity
than modern intensive systems of cultivation, although little empirical data
exist (Wood et al. 2010). Our objective was to determine what species of birds
are seasonally present in a traditional rice ecosystem in upper Myanmar and
their respective habitat use. To our knowledge, this is the only study (but see
also Suarez-Rubio et al. 2016) that highlights the importance of rice
ecosystems to birds in Myanmar.
Study Area
and Overview of Rice Cultivation
Our study was conducted at Limpha Village
(25.805N & 95.528E; elevation= 132m) in Sagaing
Region (formerly Division) of northwestern Myanmar.
This region experiences a tropical monsoonal climate with a wet season
extending from early June through mid-October (mean annual rainfall varies from
1,250 to 2,500 mm depending on elevation), followed by a dry season from late
October through May (Terra 1944). High diurnal temperatures (to 43 °C maximum)
are typical of the dry season with low nocturnal temperatures (to 4 °C minimum)
occurring in the winter months (January and February) (Terra 1944). Limpha is located within the Western Ornithological Region
of Myanmar as defined by King et al. (1975).
Limpha is
situated on the east bank of the Chindwin River approximately 40 km downstream
from the regional administrative center of Khamti (Image 1). Limpha is the
site of the Wildlife Conservation Society/Turtle Survival Alliance River Turtle
Conservation Project, hence our long-term (since 2008) institutional presence
in the village (Platt & Platt 2019). The village consists of 34 occupied
houses with an estimated population of 129 adults (≥18 years-old), most of whom
are ethnic Shan. Subsistence agriculture supplemented by fishing and collection
of non-timber forest products are the principal livelihoods, with many adult
males employed as laborers in distant amber, jade, and gold mines. The origin
of the rice ecosystem at Limpha is obscured by time;
the rice field has been in existence for as long as the oldest residents
(>80 years-old) of the community can remember. With the exception of the
rice field (see below), the lands surrounding Limpha
support dense tropical evergreen and semi-evergreen forest (Platt et al. 2013).
Village lands are contiguous with the buffer zone that surrounds Htamanthi Wildlife Sanctuary (2,151 km2).
The rice
field is located adjacent to the village and occupies 17.5 ha of a terraced
natural levee along the Chindwin River (Image 2a,b). The highest elevation in
the rice field is along the natural levee (elevation ca. 134 m). The rice field
slopes downwards, away from the river, and into a seasonally flooded swamp
(elevation ca. 128 m) comprising about 5 ha that is filled by backwater
flooding when river levels rise early in the wet season (July and August) and
usually has dried completely by late March. Maximum water depth (ca. 2.0 m) in
the swamp occurs in August and September. Soils under rice cultivation range
from light silt-sand at the natural levee crest to heavy clay near the swamp.
Much of the rice field is subdivided by low berms (20–30 cm high) into smaller
square and rectangular-shaped paddies (mean ±1SD= 110.2 ± 46.2 m2;
range= 9.9 to 286 m2) allotted to individual families for
cultivation (Image 2b). Unlike more extensive rice ecosystems in central and
southern Myanmar, the rice field at Limpha contains
no irrigation ditches. A hedgerow (0.9 km) along the natural levee crest
separates the rice field from the bed of the Chindwin River (Image 2c). The
hedgerow is characterized by large clumps of bamboo, small to medium-stature
trees, and thickets of the invasive perennial weed Chromolaena
odorata (L.) King & H.E. Robbins, and serves
as a source of construction materials (e.g., bamboo and timber) for the
village.
Rice
cultivation in Limpha is a subsistence activity to
produce grain for domestic consumption, and little if any of the crop is sold.
Rice is cultivated only during the wet season with a single crop being produced
each year. Planting coincides with the onset of the wet season and generally
begins in the last week of June or first two weeks of July, depending on
rainfall. Tillage is accomplished with either wooden plows
drawn by Water Buffalo Bubalus bubalis (Linnaeus, 1758) and Zebu Cattle Bos taurus indicus Linnaeus, 1758 or hand tractors; the
latter came into use only in 2014 and four are now available in the village.
Hand tractors are leased out by the hour with users responsible for the
purchase of fuel. Rice seedlings are germinated in specially prepared beds in
the village and then hand-planted into the field after the paddy substrate has
been prepared by plowing (Image 3a). Planting is a
communal activity with villagers reciprocally assisting one another as paddies
are made ready to receive seedlings (Image 3b,c). Water for irrigation is
supplied solely by rainfall and usually remains on the crop through the wet
season. As defined by Khush (1984), the rice field at
Limpha is a “rain-fed rice ecosystem”; i.e., lowland
rice ecosystem dependent on rainfall, with water depth uncontrolled but usually
shallow (1–50 cm).
Catastrophic
crop failure is rare at Limpha but has occurred in
the past when heavy rains in the headwaters caused prolonged overbank flooding
of the Chindwin River. Herbicide and pesticide use is minimal to non-existent
because villagers lack capital to purchase agrochemicals. Dung deposited by
free-ranging domestic ungulates (Water Buffalo and Cattle) that graze the
fallow rice field provides some fertilization. The rice crop is manually
harvested during late October and early November. Hand threshing takes place at
several locations scattered around the rice field. Like planting, harvesting is
a reciprocal communal activity (Image 4). Although record keeping is minimal,
villagers stated that annual rice yields can vary greatly, but average
900–1,000 kg/ha. Piles of rice straw are left at the threshing site and often
(but not always) burned during the dry season. Rice straw is occasionally used
as fodder for Water Buffalo. Rice stubble remains in the paddies to be plowed under during the next growing season.
Rice is
cultivated in about 50 % of the paddies every year, with the remainder being
left fallow for varying periods. Fallow paddies support grasses and sedges,
various herbaceous weeds, scattered perennial shrubs, and thickets of C. odorata. Berms of active and fallow paddies support
stands of high (2–3 m) grass. A herd of 20–25 Water Buffalo and two domestic
cattle are kept by villagers; domestic ungulates serve as draft animals,
provide fertilizer, and represent a capital investment that can be quickly
converted to cash if the need arises. During the fallow season (October or
early November through June) domestic ungulates graze in rice paddies, the
adjacent swamp, and surrounding forest (Image 5a). At this time, ungulates are
unrestrained and roam freely during the day, but are domiciled in the village
at night to prevent the animals from straying into the forest and becoming
feral. To protect the rice crop during the growing season, ungulates are
tethered in areas of favorable grazing and returned
to the village in the evening. Owners are financially responsible for any
inadvertent damage wrought to the rice crop by their livestock.
Grazing and
trampling by free-ranging domestic ungulates creates “lawns” (sensu Owen-Smith 1987) of closely cropped grass in
fallow paddies and around the periphery of the rice field (Image 5b). Water
Buffalo also create wallows in fallow paddies that are in effect, small
ephemeral waterholes. Wallows generally contain water throughout the wet season
but are dry by early December and remain so until the rains begin in June
(Image 5c). The rice field is burned during the dry season to kill encroaching
vegetation (particularly C. odorata) and
stimulate the growth of new grass for grazing ungulates (Image 5d). Burning
usually begins in March and continues through the dry season and seems to be a
haphazard activity with fires being opportunistically ignited when weather
conditions are favorable. The resulting
conflagrations are low intensity ground fires that often burn for >24 hours
and ultimately create a patchwork of burned and unburned vegetation. The system
of rice cultivation and domestic ungulate husbandry that we describe here
appears typical of other villages along the Chindwin River, including those
within the buffer zone of Htamanthi Wildlife
Sanctuary.
Methods
We made preliminary observations
of birds in the rice ecosystem at Limpha during our
initial, brief, and sporadic visits to the village during February–March of
2013–15. Our preliminary observations were followed by more intensive surveys
conducted during February–March 2016–20, October–November 2017, and
July–September 2020 when the bulk of fieldwork was completed. On most days we
searched for birds during the morning (0730–1100 h) and afternoon (1600–1800
h), although sampling during parts of the wet season was less frequent owing to
heavy rainfall and occasional flooding. When searching for birds, we used
footpaths that originate in the village and radiate throughout the rice field
as sampling transects. These footpaths run atop paddy berms and alongside the
hedgerow and forest edge (Image 2c). The complete study area was accessible
during the dry season, although flooding occasionally precluded access to some
areas during the wet season. We also recorded birds opportunistically
encountered in the rice field during the course of other fieldwork (e.g., Platt
et al. 2018; Platt & Duckworth 2019). We identified birds with the aid of
binoculars (Zeiss® and Nikon® 8 × 42) and occasionally by vocalizations. Our
observations were augmented by two motion-sensitive game cameras (Moultrie®
Series A programmed to take three photographs at 1-minute intervals), each
mounted on a wooden post (approximately 0.5 m above-ground) and positioned near
piles of discarded straw at two threshing areas in the rice field. Both game
cameras were continuously operational from 10 February through 31 March 2019
(98 camera-trap days).
We classified the different
habitats where birds were observed in the rice field as (1) rice paddy (paddies
under rice cultivation or where rice was cultivated within past 12 months), (2)
grass (fallow rice paddies and field margins now supporting primarily grasses),
and (3) hedgerow. We included birds that were observed aerially foraging above
the study area (e.g., swifts, swallows, and martins), but not high-flying
raptors; however, raptors perched in the hedgerow or in trees around the field
periphery, and low-flying birds obviously searching for prey were considered to
be using the rice field. We used a modification of methods outlined by Kumar
& Sahu (2020) to rank each species according to
relative abundance as Common (60–100 % of field visits), Uncommon (20–59 % of
field visits), and Rare (<20 % of field visits). We followed Sundar & Subramanya (2010) and classified birds
according to feeding guilds as Carnivore (consume mainly non-insect
invertebrates and vertebrates), Frugivore (consume primarily fruits), Granivore (consume seeds), Herbivore (consume mainly plants
and plant parts), Insectivore (consume mostly insects), Omnivore (consume
animals and plant material), and Nectarivore (consume mainly nectar). We used information provided in Smythies (1953), Robson (2008), Ali & Ripley (1989), Sundar & Subramanya (2010), and Birds of the World (www.birdsoftheworld.org),
supplemented by our personal observations to assign each species to a
particular foraging guild. We determined whether a species was resident or
migratory in the study area based on Smythies (1953),
Robson (2008), Birds of the World (www.birdsoftheworld.org), and our personal
observations. Geographic distribution records are based on comparisons with Smythies (1953), Robson (2008), and Thet
Zaw Naing (2017). Rankings of conservation threat
level are according to the IUCN Red List (2019) and Bird Conservation Society
of Thailand (BCST 2020). Our taxonomic nomenclature (common and scientific
names) follows Robson (2008) and scientific names for birds mentioned in the
text are provided in Table 1.
Results
We recorded a
total of 85 species of birds in the rice ecosystem at Limpha
in 2013–20 (Table 1). Excluding Buttonquail (see below), we recorded 58 (69.0
%) resident species, 20 migratory species (23.8 %), and six (7.1 %) species
with both resident and migrant populations in upper Myanmar (Table 1). Of the
85 species observed on our study site, 53 (62.3 %) and 14 (16.4 %) species were
recorded only during the dry and wet seasons, respectively, while 18 (21.1 %)
species were present during both seasons. Wading birds (except Cattle Egret),
kingfishers, Pheasant-tailed Jacana, and waterfowl were recorded only during
the wet season. Twelve (14.1 %) species were recorded only from the hedgerow,
while 16 (18.8 %) used the hedgerow as well as rice paddy and/or grass habitats
of our study area. Trees in the hedgerow appeared to provide important
observation sites for smaller raptors (Collared Falconet,
Amur Falcon). Six (7.0 %) species were only recorded while aerially foraging
over the study area. We confirmed nesting by four species (4.7 %) of birds
within the rice field, while four other species (4.7 %) nested in the adjacent
swamp, forest, hedgerow, and village (Table 1). We ranked 52 (61.1 %) species
as Common, 23 (27.0 %) as Uncommon, and 10 (11.7 %) as Rare (Table 2); three of
the latter were recorded only once during our study (Indian Thick-knee, Amur
Falcon, and Glossy Ibis). Indian Thick-knee and Glossy Ibis (Image 6a) have not
previously been reported from the Western Ornithological Region of Myanmar.
Buttonquail was encountered only in 2014 but observed on multiple occasions. We
were unable to confidently identify the Buttonquail to species; three species
of Buttonquail potentially occur in the area, one (Yellow-legged Buttonquail)
of which is migratory (Table 1). Spotted Dove was the most abundant species in
the study area with individual flocks often consisting of >50 birds (Image
6b,c). The Insectivore guild (43 species; 50.5 %) was the best represented
feeding guild in our study area, followed by Omnivore (22 species; 25.8 %),
Carnivore (12 species; 14.1 %), and Granivore (6
species; 7.0 %) guilds; Frugivore and Nectarivore guilds were each represented
by a single species (1.1 %) that was only recorded in the hedgerow (Table 2;
Figure 1). We recorded 8 (9.4 %) species of birds in association with domestic
ungulates (primarily Water Buffalo), including members of the Omnivore (6
species), Carnivore (1 species), and Insectivore (1 species) feeding guilds
(Table 1). We recorded 15 (17.6 %) species of birds foraging at active fires or
within recently burned areas, including members of four feeding guilds
(Insectivores= 7; Granivore= 4; Carnivore= 2;
Omnivore= 2). Our automated game cameras detected three species (Red
Junglefowl, White-breasted Waterhen, and Spotted Dove) foraging in piles of
discarded rice straw (Images 6d,e), and we directly observed three additional
species (Baya Weaver, Scaly-breasted Munia, and
White-rumped Munia) foraging in piles of rice straw.
We recorded 10 species considered to be of conservation concern by the IUCN and
BCST in the rice ecosystem at Limpha (Table 3).
White-winged Duck was the only Critically Endangered (BCST) or Endangered
(IUCN) species that we recorded in the Limpha rice
ecosystem (Image 6f). We observed White-winged Duck foraging in shallow water
only when the rice field was flooded during the late wet season; however, they
were present in the adjacent swamp throughout much of the year so long as water
was available.
Discussion
Our study
documented significant avian species richness in a traditional rice ecosystem
along the Chindwin River in upper Myanmar. In the only similar study available
for Myanmar, Suarez-Rubio et al. (2016) recorded 33 species in rice fields
along an urban-rural gradient near Mandalay. A comparison with rice ecosystems
elsewhere in Asia is challenging because most published studies are region-wide
in scope rather than focused on a single site (e.g., Fujioka et al. 2010; Sundar & Subramanya 2010; Wood et al. 2010). A limited
number of site-based studies are available, however, from rice ecosystems in
India and Sri Lanka; these found 34–65 species of birds (Nathan & Rajendran
1982; Srinivasulu et al. 1997; Borad
et al. 2000; Bambaradeniya et al. 2004) suggesting
that avian species richness at our study site is comparatively quite high, even
after removing those species (N= 12) recorded only in the hedgerow and other
species more typical of forested habitats (Red Junglefowl, White-winged Duck).
That said, among-site comparisons must be undertaken with caution given
differences in sampling methodologies, geographic location, farming intensity,
position within migratory flyways, and differing systems of cultivation (Hohman
et al. 1994; Valente et al. 2012; Cunningham et al. 2013). Most of the species
we recorded at Limpha are birds of open-country,
grassland, and early successional vegetation, which is typical of species
inhabiting not just rice ecosystems (Sundar &
Subramanya 2010), but agricultural habitats in general (Friskhoff
et al. 2014; Kumar & Sahu 2020). In common with
most studies of birds in rice ecosystems (Fasola
& Ruiz 1996; Townsend et al. 2006; Fujioka et al. 2010; Pierluissi
2010; Sundar & Subramanya 2010), the rice field
at Limpha appears to be used by birds primarily as
foraging rather than breeding habitat.
We attribute
the relatively high levels of bird species richness at Limpha
to the low intensity (i.e., non-mechanized, absence of agrochemicals) farming
practices used by villagers to produce a single crop of rice each year. Farming
intensity is known to determine the abundance and diversity of birds within
agricultural landscapes (Cunningham et al. 2013), with intensification usually
leading to declines in avian biodiversity (Maeda 2001; Ibáñez et al. 2010; Friskhoff 2014). At Limpha,
farming practices create a heterogeneous mosaic of different habitats within
the rice monoculture that includes rice paddies under cultivation, fallow rice
paddies in various successional stages, closely grazed “lawns” maintained by domestic
ungulates, tangles of weeds and high grass, and a hedgerow with vertical woody
structure. Previous studies at varying spatial scales have consistently found
that landscape heterogeneity is the single most important factor in determining
species richness of birds (Böhning-Gaese 1997; Pino
et al. 2000; Söderström et al. 2003; Tews et al. 2004). Moreover, the dearth of agrochemical
inputs at our study site probably favors the
development of speciose communities of arthropods and weeds (Fasola & Ruiz 1996; Bambaradeniya
& Amarasinghe 2003; Ibáñez et al. 2010), many of
which are important food resources for birds (Stafford et al. 2010). Finally,
the close proximity of forest, swamp, and the Chindwin River provides cover and
additional food resources for birds using the rice field at Limpha
and probably serves as a source for some species (e.g., White-winged Duck,
River Lapwing, Small Pratincole, Grey-throated Sand Martin) that would
otherwise be unlikely to occur in more expansive and homogenous rice landscapes
(e.g., Pierluissi 2010; Kumar & Sahu 2020).
We recorded
considerably more species of birds during the dry season in comparison to the
wet season, and attribute this disparity to the influx of Palearctic migrants
that occurs during the dry season in upper Myanmar; i.e., almost 25 % of the
species we recorded at Limpha were migrants. We
recorded wading birds and waterfowl at Limpha only
during the wet season, most likely because moist-soil and flooded habitat was
unavailable in the rice field during the dry season. Irrigation reservoirs and
water-filled ditches are absent from the rice ecosystem at Limpha,
and these habitats can serve as critical dry season refugia for wetland birds
when flooded fields are unavailable (Herzon & Helenius 2008; Valente et al. 2012). Although not included
as part of our study, the swamp adjacent to the rice field appears to function
in this capacity, harboring wetland birds (e.g.,
White-winged Duck, Common Moorhen, and White-breasted Waterhen) throughout most
of the dry season.
Rice seed is
perhaps the most important food resource available to birds in rice
agroecosystems (Borad et al. 2000; Stafford et al.
2010). Rice seed is a concentrated energy source made available to birds when
spilled during harvest, i.e., “waste rice” (Stafford et al. 2006), but birds
also forage on recently planted rice seeds, rice seedlings, and grains in
maturing seed heads before harvest (Stafford et al. 2010). Waste rice is most
abundant immediately after harvest and resists decomposition (Stafford et al.
2006), and in North America and Japan, the dry mass of rice seed remaining in
fields after mechanized harvest ranged from 56–627 kg/ha (Stafford et al.
2010). Because hand threshing is more efficient than mechanical threshing,
lesser but nonetheless significant amounts of rice seeds are lost to wastage in
traditional rice ecosystems (Borad et al. 2000). For
example, in India Borad et al. (2000) found the dry
mass of rice seed remaining in fields after hand threshing ranged from 60-199
kg/ha. Our observations suggest that waste rice is an abundant and important
food resource for several species of birds at Limpha,
most notably small seed-eaters, Spotted Dove, and Red Junglefowl. Additionally,
piles of rice straw left in fields after harvesting contain abundant waste rice
and arthropods (Bird et al. 2000; Lawler & Dritz
2005) and as such are important avian foraging sites in the Limpha
rice ecosystem.
Our
observations suggest that free-ranging ungulates, primarily Water Buffalo,
provide a number of benefits for birds in the Limpha
rice ecosystem. As reported for wild ungulates and birds (Heatwole
1965; Dean & MacDonald 1981; Isenhart & DeSante 1985), we observed two common interactions between
domestic ungulates and birds: 1) grazing ungulates acted as “beaters” to flush
insects towards waiting birds, and 2) cleaning symbiosis, whereby birds gleaned
nutritionally rich ectoparasites directly from ungulates. Water Buffalo also
appear to function as “ecosystem engineers” (sensu
Jones et al. 1994) in the Limpha rice ecosystem by
maintaining closely grazed “lawns” favored by some
birds (e.g., Red Junglefowl, wagtails, pipits, lapwings), and creating wallows
that harbor invertebrates, small fish, and amphibians
and serve as foraging sites for wading birds during the wet season.
Furthermore, Water Buffalo disperse seeds, especially those of small-seeded
herbs and grasses inadvertently consumed while grazing (Corlett 2017), and
possibly aid in the passive dispersal of aquatic invertebrates in the same
manner described for large wallowing mammals in Africa (Vanschoenwinkel
et al. 2011). Wild Water Buffalo once played a critical role in maintaining the
ecological integrity of wetlands in southeastern Asia
(Wharton 1968), and Grey et al. (2019) recommend using domestic Water Buffalo
as ecological surrogates for extinct (or nearly so) megafauna to replicate
historic patterns of grazing and wallowing in rewilding projects.
The effects
of anthropogenic burning on wildlife in southeastern
Asia remain largely unstudied (Rabinowitz 1990). Dry season burning at Limpha is no doubt at least partly responsible for the
heterogeneous mosaic of early successional vegetation in the rice ecosystem
(e.g., Peterson & Reich 2001). Additionally, we frequently observed birds
in association with fires and in recently burned-over areas, suggesting burning
is important in ways other than maintaining early successional habitats. Fires
can remove concealing vegetative cover and flush insects and small vertebrates,
providing foraging opportunities for insectivorous and carnivorous birds as
reported by others (Komarek 1969; Woinarski
& Recher 1997; Bonta et
al. 2017), and by incinerating ground litter, fires expose seeds that would
otherwise remain hidden and unavailable to birds (Komarek
1969; Woinarski & Recher
1997). Furthermore, arthropod abundance
is generally high in post-fire regrowth, creating foraging opportunities favorable for insectivorous birds (Woinarski
& Recher 1997). At Limpha,
fires ignited to remove piles of rice straw leftover from the harvest expose
waste rice, which is resistant to burning (Havens et al. 2009), and in turn
attracts flocks of foraging Spotted Dove and small seed-eaters. Anthropogenic
dry season burning as practiced at Limpha would seem
to pose little threat to nesting birds because most species reproduce during
the wet season when moist fuel conditions preclude ignition.
Similar to
our observations at Limpha, Sundar
& Subramanya (2010) found the guild structure of birds using rice fields in
the Indian Subcontinent was dominated by insectivorous and omnivorous species.
Although the most abundant species at Limpha (Spotted
Dove) is largely granivorous (Fujioka et al. 2010), we otherwise recorded few
granivorous birds, which is somewhat surprising given the abundance of waste
rice and weed seeds typically present in rice ecosystems (Stafford et al.
2010). Our results stand in contrast to previous mist-netting studies that
yielded primarily seed-eating birds from rice fields in Malaysia (reviewed by Bambaradeniya & Amarasinghe
2003).
The
preponderance of insectivorous species in rice ecosystems suggests this
avifauna could be at particular risk from pesticide exposure (Czech &
Parsons 2002; Ibáñez et al. 2010). Pesticides can result in direct
mortality as well as sublethal effects that include reproductive and behavioral impairment (Fry 1995; Smith et al. 2010; Parsons
et al. 2010). Pesticides can also negatively impact local avian abundance by
reducing or eliminating insect prey (Ibáñez et al. 2010; Parsons et al.
2010; Nocera et al. 2012), and widespread use of herbicides can eliminate
important food plants (Czech & Parsons 2002; Stafford et al. 2010).
Pearlstine et al. (2004) suggest that some agricultural lands could function as
population sinks by attracting birds to use habitat that is potentially
hazardous to their survival owing to the likelihood of pesticide exposure.
Pesticide and herbicide use is currently of little concern at Limpha because capital is unavailable to purchase
agrochemicals, although this situation could change as villagers become
increasingly enmeshed in the global economy.
The
importance of rice ecosystems as foraging and in some cases, breeding habitat
for threatened and endangered birds is well-documented (e.g., Pearlstine et al.
2004; Yu et al. 2006; Acosta et al. 2010; Elphick
2010; Van der Weijden 2010; Pickens & King 2011).
Although the threat status for most of the species we recorded at Limpha is listed as ‘Least Concern’ by the IUCN (IUCN 2019)
and BCST (2020), complacency is unwarranted because even common species can
undergo rapid and catastrophic declines if land-use changes or agriculture
intensifies (Newton 2004; Friskhoff et al. 2014;
Amano et al. 2010). This is certainly the case in Europe where some of the most
threatened birds were once considered common farmland species (Fuller et al.
1995; Sotherton 1998; Van der Weijden 2010).
Similarly, a trend towards “clean farming” practices (e.g., removal of
hedgerows, chemical elimination of weeds and brush, etc.) in agricultural
landscapes of the Southeastern United States is in
part responsible for declines among Northern Bobwhite Colinus
virginanus (Linnaeus, 1758) populations (Brennan
1991; Hernández et al. 2013). In rice
ecosystems, intensification usually involves a transition to mechanized,
capital-intensive production systems, the planting of rapidly maturing,
high-yielding rice varieties that require high inputs of agrochemicals, and
substantial increases in water consumption (Bambaradeniya
& Amarasinghe 2003). In Japan, several species of
once common rice field birds are now declining, largely as the result of
agricultural intensification (Amano et al. 2010; Kasahara & Koyama 2010).
Intensification of rice agriculture probably represents the single greatest
threat to avian biodiversity in traditional rice ecosystems in Myanmar and
elsewhere (Bambaradeniya & Amarasinghe
2003).
In
conclusion, our case study at Limpha demonstrates
that a relatively small traditional rice ecosystem in Myanmar can host a rich
assemblage of birds, including species of conservation concern and others that
are likely to be so in the near future. In accordance with species-area
relationships (Bennett et al. 2006), we predict that even higher levels of
avian richness will be found in larger rice ecosystems elsewhere in Myanmar.
Anecdotally, this indeed seems to be the case in an extensive (151 ha) rice
ecosystem surrounding Htamanthi Village (ca. 65 km
downstream from Limpha) where our recreational
bird-watching has documented a number of species of shorebirds, wading birds,
waterfowl, passerines, and raptors not recorded at Limpha.
Given these apparent high levels of observed avian biodiversity, traditional
rice agriculture seems compatible with conservation objectives in the
ecologically-sensitive buffer zone surrounding Htamanthi
Wildlife Sanctuary. According to Bambaradeniya & Amarasinghe (2003), traditional rice ecosystems that have
been cultivated over long periods can be considered stable, climax communities
that meet the criteria of sustainability; i.e., maintain or enhance the quality
of the environment and conserve natural resources. Finally, we close with a
cautionary caveat and emphasize that our study constitutes but a single datum
that requires replication before generalizations can be made concerning the
value of traditional rice ecosystems to avian conservation in Myanmar. To this
end, additional studies of rice field biodiversity should be undertaken,
especially in central Myanmar and the Ayeyarwady
Delta, where the bulk of the national rice crop is produced (Hla Myo Thwe
et al. 2019).
Table 1. Annotated checklist of
birds observed in a traditional rice ecosystem at Limpha
Village, Sagaing Region, Myanmar (2013–20). Season:
D= Dry; W= Wet. Asterisk denotes species observed foraging in burned areas.
Status: R= Resident; M= Migratory; R/M= Resident and Migratory populations
present in Upper Myanmar. Our taxonomic nomenclature (common and scientific
names) follows Robson (2008).
|
|
|
Habitat |
|
||
|
Species |
Season |
Rice Paddy |
Grass |
Hedgerow |
Status; notes and observations |
|
Buttonquail (Turnix sp.)
|
D |
X |
X |
– |
Observed on multiple occasions
in 2014; encountered among weeds around periphery of field and in fallow
paddies. Three species of Buttonquail known to occur in this area, including
Barred Buttonquail (T. suscitator),
Yellow-legged Buttonquail (T. tanki), and
Small Buttonquail (T. sylvaticus). |
|
Red Junglefowl (Gallus gallus) |
D,W |
X |
X |
– |
R; Occasionally feeding with
domestic ungulates; foraging in piles of discarded rice straw; nesting in
forest adjacent to rice field. |
|
White-winged Duck (Asarcornis scutulata) |
W |
– |
X |
– |
R; Observed in flooded rice
field during late wet season; occurs in adjacent swamp throughout much of the
year. |
|
Lesser Whistling Duck (Dendrocygna javanica) |
W |
X |
X |
– |
R; Nesting in flooded rice and
grass |
|
Lineated Barbet (Megalaima lineata) |
D |
– |
– |
X |
R; Fruiting trees in hedgerow
are important food resource. |
|
Common Hoopoe (Upupa epops)* |
D,W |
X |
X |
– |
R/M |
|
Indian Roller (Coracias benghalensis) |
D |
X |
– |
– |
R |
|
Plaintive Cuckoo (Cacomantis merulinus) |
D |
– |
X |
– |
R |
|
Asian Koel
(Eudynamys scolopaceus) |
D |
– |
– |
X |
R |
|
Greater Coucal
(Centropus sinensis) |
D,W |
X |
X |
– |
R; Usually encountered where
ungulate “lawns” are interspersed with high grass and scrub. |
|
White-throated Kingfisher (Halcyon
smyrnensis) |
W |
X |
– |
– |
R; Occasional in flooded rice
paddies. |
|
Common Kingfisher (Alcedo atthis) |
W |
X |
X |
– |
R/M; In flooded rice paddies
and around field margins. |
|
Chestnut-headed Bee-eater (Merops leschenaulti)* |
D |
X |
X |
X |
R; Nest burrows constructed in
fallow paddies, paddy berms, and ungulate “lawns”; large communal roost in
trees at village monastery until nesting begins. |
|
Little Green Bee-eater (Merops orientalis) |
D |
X |
X |
– |
R; Sally from small trees on
edge of field and fenceposts. |
|
Blue-tailed Bee-eater (Merops philippinus) |
W |
X |
X |
– |
R |
|
Himalayan Swiftlet (Aerodramus brevirostris) |
D |
– |
– |
– |
M; Aerial foraging |
|
Asian Palm-swift (Cypsiurus balasiensis) |
D |
– |
– |
– |
R; Aerial foraging |
|
Mountain Scops
Owl (Otus spilocephalus) |
D |
– |
– |
X |
R |
|
Spotted Dove (Streptopelia chinensis)* |
D,W |
X |
X |
X |
R; Large flocks (>50) feed
on spilled rice in threshing areas; nesting and large communal roosts in
hedgerow. |
|
Oriental Turtle-dove (Streptopelia orientalis) |
D |
X |
X |
– |
R/M |
|
Common Crane (Grus grus) |
D |
X |
X |
– |
M; Brief (< 24 hrs)
migratory stopover in 2019 and 2020. |
|
White-breasted Waterhen (Amaurornis phoenicurus) |
D,W |
X |
– |
X |
R; Feeding in straw piles and
on insects flushed by grazing ungulates; common in swamp adjacent to rice
field. |
|
Gray-headed Swamphen (Poryphyrio poliocephalus) |
W |
X |
X |
– |
R |
|
Common Moorhen (Gallinula chloropus) |
W |
X |
X |
– |
R; Common throughout year in
swamp adjacent to rice field. |
|
Pheasant-tailed Jacana (Hydrophasianus chirugus) |
W |
X |
X |
– |
R |
|
Indian Thick-knee (Burhinus indicus) |
D |
– |
X |
– |
R; Single observation (March
2013). |
|
Small Pratincole (Glareola lactea) |
D |
– |
– |
– |
R; Aerial foraging, often in
late afternoon; nesting on nearby island in Chindwin River. |
|
River Lapwing (Vanellus duvaucelii) |
D,W |
X |
X |
– |
R; Nesting on nearby island in
Chindwin River. |
|
Grey-headed Lapwing (Vanellus cinereus) |
D,W |
X |
X |
– |
M |
|
Red-wattled
Lapwing (Vanellus indicus) |
D,W |
X |
X |
– |
R; Nesting in ungulate “lawn” |
|
Pacific Golden Plover (Pluvialis fulva) |
W |
X |
X |
– |
M |
|
Little Ringed Plover (Charadrius dubius) |
D |
X |
X |
– |
M |
|
Pied Harrier (Circus melanoleucos)* |
D |
X |
X |
– |
M |
|
Collared Falconet
(Microhierax caerulescens) |
W |
X |
X |
X |
R |
|
Common Kestrel (Falco tinnunculus)* |
D |
X |
X |
– |
R/M |
|
Amur Falcon (Falco amurensis) |
W |
X |
– |
X |
M; Single record (November
2018). |
|
Black-shouldered Kite (Elanus caeruleus) |
D |
X |
X |
– |
R |
|
Eastern Cattle Egret (Bubulcus coromandus) |
D,W |
X |
X |
– |
R; Feeding on insects flushed
by grazing ungulates. |
|
Chinese Pond Heron (Ardeola bacchus) |
W |
X |
X |
– |
R |
|
Black-crowned Night Heron (Nycticorax nycticorax) |
W |
X |
X |
– |
R |
|
Glossy Ibis (Plegadis falcinellus) |
W |
X |
– |
– |
R; Single record (October
2018); foraging in water-filled buffalo wallows. |
|
Long-tailed Broadbill (Psarisomus dalhousiae) |
D |
– |
– |
X |
R; fruiting trees in hedgerow
are important food resource; common in adjacent forest. |
|
Golden-fronted Leafbird (Chloropsis aurifrons)
|
D |
– |
– |
X |
R |
|
Grey-backed Shrike (Lanius tephronotus)* |
D |
X |
X |
X |
M |
|
Long-tailed Shrike (Lanius schach) |
D |
X |
X |
– |
R |
|
Eastern Jungle Crow (Corvus levaillanti) |
D |
X |
– |
– |
R; Occasionally with domestic
ungulates; gleaning ectoparasites? |
|
Black-hooded Oriole (Oriolus xanthornus) |
D |
– |
– |
X |
R; three observations of birds
consuming large caterpillars. |
|
Hair-crested Drongo (Dicrurus hottentottus) |
D |
– |
– |
X |
R/M |
|
Black Drongo
(Dicrurus macrocercus) |
D |
X |
X |
– |
R/M |
|
Ashy Woodswallow
(Artamus fuscus) |
D |
X |
X |
X |
R; Aerial foraging; roost and
nest in village. |
|
White-throated Fantail (Rhipidura albicollis) |
D |
– |
– |
X |
R |
|
Bluethroat (Luscinia
svecica) |
D |
– |
X |
– |
M |
|
Siberian Rubythroat
(Luscinia calliope) |
D |
– |
X |
– |
M |
|
Oriental Magpie-robin (Copsychus saularis) |
D,W |
X |
X |
– |
R |
|
White-tailed Stonechat (Saxicola leucura) |
D,W |
X |
X |
– |
R |
|
Eastern Stonechat (Saxicola maurus) |
D |
X |
X |
– |
M |
|
Pied Bushchat
(Saxicola caprata)* |
D |
X |
X |
– |
R; Nesting in rice paddy berm. |
|
Daurian Redstart (Phoenicurus auroreus) |
D |
X |
X |
– |
M |
|
Black Redstart (Phoenicurus ochruros) |
D |
X |
X |
– |
M |
|
Chestnut-tailed Starling (Sturnus malabaricus)* |
D |
– |
X |
X |
R |
|
Common Myna
(Acridotheres tristis) |
D |
X |
X |
X |
R |
|
White-vented Myna (Acridotheres grandis) |
D,W |
X |
X |
– |
R; Feeding on insects flushed
by grazing ungulates; glean ectoparasites from ungulates. |
|
Collared Myna
(Acridotheres albocinctus) |
D |
X |
X |
– |
R; Feeding no insects flushed
by grazing ungulates. |
|
Asian Pied Starling (Gracucpica contra)* |
D |
X |
X |
– |
R; Feeding on insects flushed
by grazing ungulates. |
|
Grey-throated Sand Martin (Riparia chinensis) |
D |
– |
– |
– |
R; Aerial foraging; scattered
nesting colonies on banks of Chindwin River. |
|
Red-rumped
Swallow (Cecropis daurica) |
D |
– |
– |
– |
M; Aerial foraging. |
|
Red-whiskered Bulbul (Pycnonotus jocosus) |
D,W |
– |
X |
X |
R; Large communal roost in
secondary forest adjacent to rice field. |
|
Red-vented Bulbul (Pycnonotus cafer) |
D,W |
– |
X |
X |
R |
|
Striated Grassbird
(Megalurus palustris) |
D |
X |
X |
– |
R; Feeding on insects flushed
by grazing ungulates. |
|
Yellow-bellied Prinia (Prinia flaventris) |
D |
– |
X |
– |
R; In high grass of fallow rice
paddies; vocalizing males; nesting? |
|
Indian Reed-warbler (Acrocephalus brunnescens) |
D |
– |
X |
– |
M; Present in dense thickets of
Chromolaena odorata. |
|
Common Tailorbird (Orthotomus sutorius)* |
D,W |
– |
X |
X |
R |
|
Dusky Warbler (Phylloscopus fuscatus) |
D |
– |
– |
X |
M |
|
Chestnut-crowned Warbler (Seicercus castaniceps) |
D |
– |
– |
X |
R |
|
Pin-striped Tit-babbler (Macronous gularis) |
D |
– |
– |
X |
R; Often encountered in bamboo
clumps of hedgerow. |
|
Purple Sunbird (Cinnyris asiaticus) |
D |
– |
– |
X |
R |
|
Citrine Wagtail (Motacilla citreola) |
D |
X |
X |
– |
M; Frequently in mixed flocks
with White Wagtail and occasionally Red Junglefowl; present on closely
cropped lawns and in fallow rice paddies. |
|
White Wagtail (Motacilla alba) |
D,W |
X |
X |
– |
M; See comments for Citrine Wagtail. |
|
Olive-backed Pipit (Anthus hodgsoni)* |
D |
X |
X |
– |
M; Present on closely cropped
lawns and fallow rice paddies; avoid areas with thick grass. |
|
Paddyfield Pipit (Anthus rufulus)* |
D |
X |
X |
– |
R; See comments for
Olive-backed Pipit. |
|
Rosy Pipit (Anthus
roseatus) |
D |
X |
X |
– |
M; See comments for
Olive-backed Pipit. |
|
Baya Weaver (Ploceus phillippinus)* |
D,W |
X |
X |
X |
R; Feeding on waste rice in
piles of discarded straw; nesting in coconut palms in village. |
|
Scaly-breasted Munia (Lonchura punctulata)* |
D |
X |
X |
X |
R; Feeding on waste rice in
piles of discarded straw. |
|
White-rumped
Munia (Lonchura striata) |
D |
X |
X |
X |
R; Feeding on waste rice in
piles of discarded straw. |
|
Black-faced Bunting (Emberiza spodocephala)* |
D |
X |
X |
X |
M; Commonly encountered among
weeds and high grass in fallow rice paddies and in thickets on field
margin. |
Table 2. Feeding guild and
relative abundance of birds observed in a traditional rice ecosystem at Limpha Village, Sagaing Region,
Myanmar (2013–20). Feeding guild: C= Carnivore; F= Frugivore; G= Granivore; H= Herbivore; I= Insectivore; O= Omnivore; N=
Nectarivore. Relative abundance: C= Common; U= Uncommon; R= Rare.
|
|
Common name |
Scientific name |
Feeding guild |
Relative abundance |
|
1 |
Buttonquail |
Turnix sp. |
O |
U |
|
2 |
Red Junglefowl |
Gallus gallus |
O |
C |
|
3 |
White-winged Duck |
Asarcornis scutulata |
O |
R |
|
4 |
Lesser Whistling Duck |
Dendrocygna javanica |
O |
U |
|
5 |
Lineated Barbet |
Megalaima lineata |
F |
C |
|
6 |
Common Hoopoe |
Upupa epops |
I |
C |
|
7 |
Indian Roller |
Coracias benghalensis |
I |
U |
|
8 |
Plaintive Cuckoo |
Cacomantis merulinus |
I |
U |
|
9 |
Asian Koel |
Eudynamys scolopaceus |
O |
C |
|
10 |
Greater Coucal |
Centropus sinensis |
O |
C |
|
11 |
White-throated Kingfisher |
Halcyon smyrnensis |
C |
C |
|
12 |
Common Kingfisher |
Alcedo atthis |
C |
U |
|
13 |
Chestnut-headed Bee-eater |
Merops leschenaulti |
I |
C |
|
14 |
Little Green Bee-eater |
Merops orientalis |
I |
U |
|
15 |
Blue-tailed Bee-eater |
Merops philippinus |
I |
U |
|
16 |
Himalayan Swiftlet |
Aerodramus brevirostris |
I |
C |
|
17 |
Asian Palm-swift |
Cypsiurus balasiensis |
I |
C |
|
18 |
Mountain Scops
Owl |
Otus spilocephalus |
C |
C |
|
19 |
Spotted Dove |
Streptopelia chinensis |
G |
C |
|
20 |
Oriental Turtle-dove |
Streptopelia orientalis |
G |
R |
|
21 |
Common Crane |
Grus grus |
O |
R |
|
22 |
White-breasted Waterhen |
Amaurornis phoenicurus |
O |
C |
|
23 |
Gray-headed Swamphen |
Poryphyrio poliocephalus |
O |
U |
|
24 |
Common Moorhen |
Gallinula chloropus |
O |
C |
|
25 |
Pheasant-tailed Jacana |
Hydrophasianus chirugus |
C |
U |
|
26 |
Indian Thick-knee |
Burhinus indicus |
O |
R |
|
27 |
Small Pratincole |
Glareola lactea |
I |
C |
|
28 |
River Lapwing |
Vanellus duvaucelii |
I |
U |
|
29 |
Grey-headed Lapwing |
Vanellus cinereus |
I |
C |
|
30 |
Red-wattled
Lapwing |
Vanellus indicus |
I |
C |
|
31 |
Pacific Golden Plover |
Pluvialis fulva |
O |
U |
|
32 |
Little Ringed Plover |
Charadrius dubius |
O |
U |
|
33 |
Pied Harrier |
Circus melanoleucos |
C |
U |
|
34 |
Collared Falconet
|
Microhierax caerulescens |
C |
U |
|
35 |
Common Kestrel |
Falco tinnunculus |
C |
C |
|
36 |
Amur Falcon |
Falco amurensis |
C |
R |
|
37 |
Black-shouldered Kite |
Elanus caeruleus |
C |
U |
|
38 |
Eastern Cattle Egret |
Bubulcus coromandus |
C |
C |
|
39 |
Chinese Pond Heron |
Ardeola bacchusx |
C |
C |
|
40 |
Black-crowned Night Heron |
Nycticorax nycticorax |
C |
U |
|
41 |
Glossy Ibis |
Plegadis falcinellus |
I |
R |
|
42 |
Long-tailed Broadbill |
Psarisomus dalhousiae |
I |
C |
|
43 |
Golden-fronted Leafbird |
Chloropsis aurifrons |
I |
C |
|
44 |
Grey-backed Shrike |
Lanius tephronotus |
I |
C |
|
45 |
Long-tailed Shrike |
Lanius schach |
I |
C |
|
46 |
Eastern Jungle Crow |
Corvus levaillanti |
O |
C |
|
47 |
Black-hooded Oriole |
Oriolus xanthornus |
O |
C |
|
48 |
Hair-crested Drongo |
Dicrurus hottentottus |
I |
C |
|
49 |
Black Drongo |
Dicrurus macrocercus |
I |
R |
|
50 |
Ashy Woodswallow |
Artamus fuscus |
I |
C |
|
51 |
White-throated Fantail |
Rhipidura albicollis |
I |
U |
|
52 |
Bluethroat |
Luscinia svecica |
I |
U |
|
53 |
Siberian Rubythroat
|
Luscinia calliope |
I |
U |
|
54 |
Oriental Magpie-robin |
Copsychus saularis |
I |
C |
|
55 |
White-tailed Stonechat |
Saxicola leucura |
I |
C |
|
56 |
Eastern Stonechat |
Saxicola maurus |
I |
C |
|
57 |
Pied Bushchat
|
Saxicola caprata |
I |
C |
|
58 |
Daurian Redstart |
Phoenicurus auroreus |
I |
R |
|
59 |
Black Redstart |
Phoenicurus ochruros |
I |
R |
|
60 |
Chestnut-tailed Starling |
Sturnus malabaricus |
O |
C |
|
61 |
Common Myna
|
Acridotheres tristis |
O |
C |
|
62 |
White-vented Myna |
Acridotheres grandis |
O |
C |
|
63 |
Collared Myna
|
Acridotheres albocinctus |
O |
U |
|
64 |
Asian Pied Starling |
Gracucpica contra |
O |
C |
|
65 |
Grey-throated Sand Martin |
Riparia chinensis |
I |
C |
|
66 |
Red-rumped
Swallow |
Cecropis daurica |
I |
C |
|
67 |
Red-whiskered Bulbul |
Pycnonotus jocosus |
O |
C |
|
68 |
Red-vented Bulbul |
Pycnonotus cafer |
O |
C |
|
69 |
Striated Grassbird
|
Megalurus palustris |
I |
R |
|
70 |
Yellow-bellied Prinia |
Prinia flaventris |
I |
C |
|
71 |
Indian Reed-warbler |
Acrocephalus brunnescens |
I |
U |
|
72 |
Common Tailorbird |
Orthotomus sutorius |
I |
C |
|
73 |
Dusky Warbler |
Phylloscopus fuscatus |
I |
C |
|
74 |
Chestnut-crowned Warbler |
Seicercus castaniceps |
I |
U |
|
75 |
Pin-striped Tit-babbler |
Macronous gularis |
I |
C |
|
76 |
Purple Sunbird |
Cinnyris asiaticus |
N |
C |
|
77 |
Citrine Wagtail |
Motacilla citreola |
I |
C |
|
78 |
White Wagtail |
Motacilla alba |
I |
C |
|
79 |
Olive-backed Pipit |
Anthus hodgsoni |
I |
C |
|
80 |
Paddyfield Pipit |
Anthus rufulus |
I |
C |
|
81 |
Rosy Pipit |
Anthus roseatus |
I |
U |
|
82 |
Baya Weaver |
Ploceus phillippinus |
G |
C |
|
83 |
Scaly-breasted Munia |
Lonchura punctulata |
G |
C |
|
84 |
White-rumped
Munia |
Lonchura striata |
G |
C |
|
85 |
Black-faced Bunting |
Emberiza spodocephala |
G |
C |
For
figures & images - - click here
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