Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 March 2022 | 14(3): 20773–20779
ISSN 0974-7907
(Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.6985.14.3.20773-20779
#6985 | Received 13
December 2020 | Final received 15 March 2022 | Finally accepted 18 March 2022
Breeding biology of Sri Lanka
White-eye Zosterops ceylonensis
(Aves: Passeriformes: Zosteropidae)
in tropical montane cloud forests, Sri Lanka
W.D.S.C. Dharmarathne
1, P.H.S.P.Chandrasiri 2 & W.A.D. Mahaulpatha
3
1–3 Department of Zoology, University
of Sri Jayewardenepura, Sri Lanka.
1 chathurangadharma@gmail.com, 2
spchandrasiri@gmail.com, 3 mahaulpatha@sjp.ac.lk
(corresponding author)
Editor: R. Jayapal,
Sálim Ali Centre for Ornithology and Natural History,
Coimbatore, India. Date of publication: 26
March 2022 (online & print)
Citation: Dharmarathne, W.D.S.C., P.H.S.P. Chandrasiri
& W.A.D. Mahaulpatha (2022). Breeding
biology of Sri Lanka White-eye Zosterops ceylonensis (Aves: Passeriformes: Zosteropidae)
in tropical montane cloud forests, Sri Lanka. Journal of
Threatened Taxa 14(3): 20773–20779. https://doi.org/10.11609/jott.6985.14.3.20773–20779
Copyright: © Dharmarathne
et al. 2022. 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: University of Sri Jayewardenepura, Sri Lanka.
Competing interests: The authors
declare no competing interests.
Acknowledgements: We would like to thank the
University of Sri Jayewardenepura for financial support through the university
grant (ASP/01/RE/SCI/2016/20). We extend our gratitude to the Department of
Wildlife Conservation for granting permission to carry out this study (permit
no-WL/3/2/14/15).
Abstract: Breeding biology of the endemic
Sri Lanka White-eye Zosterops ceylonensis was studied from January 2017 to January
2018 in the montane cloud forests of Horton Plains National Park, in Nuwara Eliya District of Sri Lanka. A total of 46 active nests
were studied. Breeding occurred March–May. The peak egg-laying was in March and
April. Mean nest construction period 11±2.87 days (n= 17), nest height was
3.16±1.22 m from the ground. Majority of nests were built on Sarcococca brevifolia, Berberis ceylanica, and Cinnamomum ovalifolium
trees of 4–12 m tall. Most number of pen cup nests were constructed by the fine
grass stalks and mosses woven with cobwebs and internal cup lined with grass
roots. Eggs were pale blue. Mean clutch size 2.15±0.37 eggs (n= 11). The
incubation period was 10.92±0.9 days (n= 11). Nestling period was 28.33±1.55
days (n= 11). Both sexes participate in nest construction, incubation and brood
rearing. About 25.53% of nests were depredated (N= 12). The overall nest
success was 74.46%. The study reveals that undisturbed cloud forests are
critical to conservation of Sri Lanka White-eye.
Keywords: Brood rearing, eggs, endemic,
Horton Plains, incubation period, nest construction.
Information on breeding ecology
is essential for planning conservation strategies. However, the inadequacy of
scientific data on avifauna is the major problem to conserve the birds of Sri
Lanka. White-eyes (Passeriformes: Zosteropidae) are a
widely distributed family of Old World birds. They are found on the mainland of
tropical Africa, southern and eastern Asia, Australia, and on the islands of
these continents, including Indian Ocean, Indonesia, Melanesia, and parts of
Micronesia (Lack 1971). The largest of the genera is Zosterops,
with over 85 species (Clements et al. 2016). There are two species of white-eye
occurring in Sri Lanka, the Sri Lanka White-eye Zosterops
ceylonensis (Holdsworth,
1872) and Oriental White-eye Zosterops palpebrosus (Temminck, 1824).
Z. ceylonensis is a small passerine highland endemic
(Figure 1). It is a resident breeder in montane forests and usually found in
pairs during the breeding season. Outside the breeding season, it may be found
in large, scattered flocks (Ali & Ripley 2001). This species can be
distinguished by duller green mantle and more extensive yellow on the breast.
It has a darker patch between the eye and the bill (Wait 1922). Though mainly
insectivorous, also eat nectar and fruits. It is sociable, forming large
flocks. The global population size has not been quantified, climate change,
habitat alteration are current threats to the species (Birdlife international
2016).
It appears to be more sociable
than the Oriental White-eye (Henry 1971; Ali & Ripley 2001). Furthermore,
it appears to be equally at home in bushes and undergrowth as well as in the
foliage of large trees. Its food consists of nectar, berries, insects, and
caterpillars. The breeding season is bimodal. The first extends from February
to July with a peak in April. The second season is in August and September.
Investigation of breeding biology was the main objective of this study.
The nesting behavior
of many forest bird species are still completely unknown, and detailed
information concerning the breeding biology and reproductive success of most
species is lacking. Collecting information on breeding biology and ecology is
an important part of many studies of the population ecology of birds. This is
often essential in identifying effective conservation measures for threatened
and declining species (Sutherland et al. 2004). Although it is an endemic
species with a very restricted distribution, making it more susceptible to
becoming endangered, the basic natural history of Z. ceylonensis
is still poorly known and, to date, no detailed study of its breeding biology
exists.
Materials and Methods
Study site
The study was conducted in Horton
Plains National Park (HPNP) from January 2017 to January 2018. HPNP is located
at an elevation range of 2,100–2,300 m and encompasses montane grassland and
cloud forest (Gunatilleke & Gunatilleke
1986). It is rich in biodiversity and many species found here are endemic to
the region. The mean annual rainfall is greater than 2,000 mm. Frequent cloud
cover limits the amount of sunlight that is available to plants. The mean annual
temperature is 13 °C but the temperature varies considerably during a day,
reaching as high as 27 °C during the daytime, and dipping as low as 5 °C at
night. Dry season occurs from January to March. The ground frost is common in
February. Mist can persist in the day during the wet season (De Silva 2007).
The vegetation of the park is
classified into wet patana and cloud forests
(International Water Management Institute 2010). HPNP is considered as one of
the Important Bird Areas (IBAs) in Sri Lanka (BirdLife
International 2009).
Methods
Nest sites were searched on three
consecutive days in each month from March to May from 0600h to 1800h.
Nests were searched in the
interior of the forest patches. Five 25 x 25 m quadrats were marked in each
habitat using a global positioning system device (GPS). Proportioned time was
spent between habitats for nest searching (Kozma
& Mathews 1997).
Approachable nests were observed
directly. Unapproachable nests were observed through a 10 x 50 binocular. Pole
and mirror method was used to check the nest contents. Nests were monitored
until they were no longer in use. The time interval between nest checks was
optimized by fieldwork logistics. Near the dates of egg-laying and hatching,
nests were checked more frequently, when possible, to estimate the nesting
phenology precisely. Focal animal sampling method was used to study the
breeding behavior of the species (Altman 1974).
Nesting materials were identified
by observing adult birds carrying nest materials from the resources during the
nest construction period. Nesting habitat variables were recorded in each nest
site. Nest parameters such as nest length and nest width were recorded. Canopy
cover recorded using a spherical densitometer. Standard methods were used to
estimate fruit cover and flower cover (Struhsaker
1975), shrub cover (Zollner & Crane 2003) and
habitat insect availability on trunks/twigs and leaves were recorded.
Environmental variables near nesting habitats were recorded using pocket
weather meter (Krestel TM 4000, USA). Nest site characteristics, such as nest
height from the ground, height of the nesting tree and distance for the nearest
nest of same species were recorded.
Available habitats were
classified as cloud forests (CF), cloud forest die-back (CFD), and grasslands
(GL) in the HPNP. The observer stayed at a hidden position and behaviour of the
breeding couples were studied using a binocular (NikonTM
- Monarch, 10 × 42).
Surface temperatures of eggs were
measured using EXTECH Infrared thermometer initially after incubation adults
left the nest. Incubation patterns such as on-bout and off-bout duration, nest
trips rate and nest attentiveness were studied. Moreover, after the eggs
hatching on-bout and off-bout duration, feeding trips rate and nest attentiveness
of Z. ceylonensis were observed in the
nestling period separately. The diurnal period was divided as, dawn (0600–0900
h), morning (0901–1200 h), mid-day (1201–1500 h), and evening (1501–1800 h). A
nest was considered successful if at least one young fledged. Nesting
observations were made with no disturbance to the birds and nests.
Data analysis
Differences were considered at
p<0.05 significant level, mean and standard deviation (M±SD) values were
reported throughout. Microsoft Excel™ was used to store data. Principal
component analysis (PCA) was performed to analyze
nesting habitat variables of Z. ceylonensis
and graphical illustrations were performed in Minitab 17™.
Results and
Discussion
A total of 47 active nests were
recorded during the study period breeding occurred mainly from March to May
comparatively low nesting observations were recorded during the second season
from August to September. The peak egg-laying was in March and April (Figure
2).
Most of the nests were recorded
in the CF (65.95%) habitat compared to the CFD (34.05%) habitat. There were no
nests recorded in the GL habitats (Figure 3). The study revealed that CF
habitats occupied by the Z. ceylonensis for
their breeding. First two axes of the PCA analysis of habitat variables which
were significantly different from available habitat characteristics account for
85.5% of the total variance according to the Eigen analysis of the correlation
matrix.
In the first principal component
(PC1), wind speed (PC1, 0.466) contributed mostly to where the highest
contributed factors for PC1 that variable correlated positively. It correlated
negatively with temperature and (PC1, -0.528) and shrub cover (PC1, -0.504).
Hence an increase in shrub cover will lead to a decrease in wind speed.
Therefore, this habitat attributes influence on nesting habitat selection of Z.
ceylonensis.
The second component (PC2) gave
high scores to sites with high values of canopy cover (PC2, 0.562) and flower
cover (PC2, 0.572). The overall PCA result indicated that determining factors
of breeding habitat utilization in natural habitats of Z. ceylonensis in HPNP were availability of high shrub
cover, canopy cover and flower cover (Figure 4).
Mean nest construction period 11
±2.87 days (n= 17). Both male and females built the nest, during nest
construction activity. We did not observe any cases of Z. ceylonensis reusing material from an abandoned or
predated nest.
Majority of nests were built on Sarcococca brevifolia, Berberis ceylanica Cinnamomum ovalifolium trees
of 4–12 m tall. Nest height was 3.16±1.22 m from the ground. Approximate
distance for the active nests of same species was 4–6 m. There were few records
that two nests in the same tree within 4m distance.
Most of the nests (89.36%) were
built on the mosses hanging from the tree branches. Z. ceylonensis
using this Usnea barbarta
mosses as a substrate to their nests (72.4%). It will help them to conceal
their nest and avoid predation via mossy camouflage in these montane habitats.
Hammock like open cup nests constructed by mostly the fine grass stalks and
mosses woven with cobwebs and internal cup lined with grass stalks and plant
fibre. Nests were always covered by surrounding leaves. Average nest width was
6.42±0.42 cm (n= 17), and average nest length was 3.91±0.22 cm (n= 17). Eggs
were unspotted and pale blue. Mean clutch size 2.15±0.39 eggs (n= 16). One nest
was destroyed by the predators before we observe the clutch.
Nest observations revealed that
both male and female were involved in incubation, brooding and feeding of the
young at both the nestling and fledgeling stage. The
average incubation period was 10.92±0.9 days (M±SD) (n= 11) and the average
nestling period was 28.33±1.55 days (n= 11).
During the incubation period,
on-bout duration (35.41±3.28 min (M±SD)) and nest attentiveness (92.21±5.43%
(M±SD)) was higher in the evening period. Off-bout duration (22.32 ±5.21 min
(M±SD)) was higher in the morning period. Nest trips rate (4.02± 0.54 h-1
(M±SD)) was higher in the mid-day. The mean egg temperature under the parental
incubation (25.92 ±3.41 C0 (M±SD)) is significantly different from
that of the absence of parental incubation (13.17±0.54 C0 (M±SD))
(One way ANOVA, p<0.01). Both percentages of attendance and length of
sitting bouts increase in bad weather. Pair sometimes meets on a branch near
the nest to allopreening.
During the nestling period,
on-bout duration (3.14 ± 1.26 min (M±SD)) and nest attentiveness (68.96 ± 10.34%
(M±SD)) was higher in the evening period. Off-bout duration (2.49 ± 1.04 min
(M±SD)) and feeding trips rate (10.31 ± 1.54 h-1 (M±SD)) was higher
during the morning.
Z. ceylonensis single-brooded although some
pairs made re-nesting attempts after first nests failed. About 23.40% of nests
were depredated (N= 11), with evidence suggesting predation by Jungle Crow Corvus levaillantii
and Greater Coucal Centropus
sinensis. Furthermore, 2.13% of nests were broken
by rainy winds (N= 1). About 75% of nests were failed during the incubation
period while 25% nests failed during nestling stage. The overall nest success
was 74.46%.
Comparatively higher nest failure
was recorded in the CFD habitat (56.25%) compared to CF habitat (6.45%) (Figure
5). Therefore, study reveals that undisturbed cloud forests are essential to
ensuring the breeding of this endemic species.
Incubating birds face ecological
costs associated with reproductive effort during the breeding season (Conway
& Martin 2000). Studies have shown that the first step of success of
individual breeding attempts involves the location of nests being used by birds
(Krebs & Davies 2009). Generally, factors that help decide the location of
nesting site, and probably the choice of mate as well, including local food
availability, presence of suitable nest materials and shelter from the physical
environment and protection from predators (Collias
& Collias 2014).
Z. ceylonensis
occupied
different tree species to build their nest all the nests shows that nests well
protected from the heavy rains by a dense cluster of broad leave over the nest.
It is generally placed on a small branch or twig directly under a canopy of
foliage. The nest is built largely of fibres with outer surface often covered
by the mosses. the open cup nest is typical of most passerine birds, the size
of the inner cup is automatically moulded to the body size of the species
because of the typical movements used in building, pushing in the nest with
breast while rotating and pushing back with the feet (Collias
& Collias 2014).
The close fit of the nest to bird
helps make something of a seal, holding in warmth when the incubating bird is
sitting closely. Our findings indicate that Z. ceylonensis
maintained about 12 0C warmer than the surrounding
air during the incubation. One reason for the prevalence of open nests among
small birds of cool climates may be the need of the bird on the nest and its
nestlings for the warming rays of the morning sun (Collias
& Collias 2014).
Birds have evolved a variety of
anti-predator adaptations in their nest-building behavior
(Skutch 1976). When considering the Z. ceylonensis nests they are hidden in the vegetation, to
deceive predators they camouflage their nests covered with Usnea
barbarta mosses by resembling a mass of natural
vegetation. It seems that the importance of nest concealment varies with
ecological circumstances and with the type of predators at a given time and
habitat. Since there is less abundance of snakes in the Horton plains most of
the predator attacks occurred by avian predators. Therefore, nest concealment
is a very important factor for nesting success.
The Horton Plains is one of the
remaining pristine montane cloud forest habitats in Sri Lanka. Due to tourist
activities invaded the population of Jungle Crow increased at human-induced
habitat in the HPNP (Chandrasiri et al. 2017). When
considering the nest failures of Z. ceylonensis most
of the nest predation occurred due to predation by Jungle Crow. Unfortunately,
nesting colony of Jungle Crow was observed in 2018 in the CF habitat. Many crows are a major threat to endemic
animals. Increased number of crows is an indicator of pollution because they
are scavengers in the food chain. To establish the protection of Z. ceylonensis in this important Montane Cloud Forest,
admissible methods to control the number of Jungle Crow are needed.
Good nest sites are often
traditional, serving as ‘ecological magnets’ over many years and regularly
continue to attract individuals of the same species (Burnham 2007). Forest on
Sri Lanka has suffered rapid degradation and fragmentation in the past decades
through the excessive gathering of fuel-wood, clearance for permanent
agriculture, shifting cultivation, fire, urbanization and logging. It is feared
that habitat loss will continue in the hills and the status of this species
therefore requires monitoring. There is no known targeted conservation action
for this species. Therefore, conservation of breeding habitats is recommended
to protect this species.
Table 1. Nesting tress of Zosterops ceylonensis at HPNP.
Tree Species |
Number of
nests (n= 47) |
Sarcococca brevifolia |
7 |
Berberis ceylanica |
6 |
Cinnamomum ovalifolium |
6 |
Neolitsea fuscata |
4 |
Syzygium rotundifolium |
4 |
Elaeocarpus sp. |
3 |
Rhododendron arboreum |
3 |
Rubus ellipticus |
3 |
Symplocos bractealis |
2 |
Sinarundinaria debilis |
2 |
Actinoaphne speciosa |
2 |
Strobilanthes sp. |
2 |
Eugenia mabaeoides |
1 |
Rhodomyrtus tomentosa |
1 |
Vaccinium leschenaultii |
1 |
For figures &
images - - click here
References
Ali, S.L.
& S.D. Ripley (eds.) (2001). Handbook of the Birds of India and Pakistan
together with those of Bangladesh, Nepal, Bhutan, and Sri Lanka. Oxford
University Press, Delhi, India.
Altmann, J.
(1974).
Observational study of behavior: Sampling methods. Behaviour
49: 227–267.
BirdLife International (2016). Zosterops
ceylonensis. In: IUCN 2012. IUCN Red List of
Threatened Species. 2012. Date of download 2019/01/28. https://doi.org/10.2305/IUCN.UK.2016-3.RLTS.T22714023A94397877.en
Burnham, K.K.
(2007). Inter-and
intraspecific variation of breeding biology, movements, and genotype in
Peregrine Falcon Falco peregrinus and
Gyrfalcon F. rusticolus populations in
Greenland. PhD Thesis. University of Oxford.
Chandrasiri, P.H.S.P., W.D.S.C.Dharmarathne,
S. Lakmal & W.A.D. Mahaulpatha
(2017). Distribution
of the jungle crow (Corvus levaillantii lesson, 1831) and their potential threats
to biodiversity in Horton Plains National Park, Sri Lanka. Wildlanka
5(2): 073–077.
Clements,
J.F., T.S. Schulenberg, M.J. Iliff, D. Roberson, T.A.
Fredericks, B.L. Sullivan & C.L. Wood (2016). The Clements checklist of
birds of the world.
http://www.birds.cornell.edu/clementschecklist/download/Google Scholar.
Collias, N.E. & E.C. Collias (2014). Nest Building and Bird Behavior. Princeton University Press.
Conway, C.J.
& T.E. Martin (2000). Evolution of passerine incubation behavior:
influence of food, temperature, and nest predation. Evolution 54(2):
670–685.
de Silva, A.
(2007). The
Diversity of Horton Plains National Park (with special reference to its
herpetofauna). Vijitha Yapa
Publishers. Sri Lanka.
Gunatilleke, C.V.S. & I.A.U.N. Gunatilleke (1986). Horton Plains: some aspects of
its vegetation and ecology. Sri Lanka Wildlife 3(4): 9–11.
Henry, G.M.
(1971). A Guide
to the Birds of Ceylon (Second Edition). Oxford University Press, London,
UK.
Horton Plains
National Park (2010). International Water Management Institute. https://web.archive.org/web/20100805010613/http://dw.iwmi.org/wetland_profile/Horton.asp
BirdLife International (2009).
IBAs in Sri Lanka. http://www.birdlife.org/datazone/country/sri-lanka
Kozma, J.M & N.E. Mathews (1997). Breeding bird communities and
nest plant selection in Chihuahuan Desert habitats in
south-central New Mexico. The Wilson Bulletin 109(03): 424–436.
Krebs, J.R
& N.B. Davies (2009). Behavioural Ecology: An Evolutionary Approach. John Wiley &
Sons.UK.
Lack, D.L.
(1971). Ecological
Isolation in Birds. Harvard University Press, Cambridge, Massachusetts,
USA.
Skutch, A.F. (1976). Parent Birds and Their Young.
Austin and London: University of Texas Press.
Struhsaker, T.T. (1975). The Red Colobus Monkey.
University of Chicago Press, Chicago, 311pp.
Sutherland,
W.J., I. Newton & R. Green (2004). Bird Ecology and Conservation:
A Handbook of Techniques (No. 1). Oxford University Press.
Wait, W.E.
(1922). The
Passerine Birds of Ceylon. Spolia Zeylanica
12: 182.
Zollner, P.A. & K.J. Crane (2003). Influence of canopy closure and
shrub coverage on travel along coarse woody debris by eastern chipmunks (Tamias striatus). The
American Midland Naturalist 150(1): 151–157.