Diet composition of introduced barn owls (Tyto alba javanica) in urban area in comparison with agriculture settings - Oxford Academic ...
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Journal of Urban Ecology, 2020, 1–8
doi: 10.1093/jue/juz025
Research article
Diet composition of introduced barn owls (Tyto alba
javanica) in urban area in comparison with agriculture
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settings
Safwan Saufi,1 Shakinah Ravindran ,1 Noor Hisham Hamid,2
Cik Mohd Rizuan Zainal Abidin,3 Hamdan Ahmad,1,4 Abu Hassan Ahmad1
and Hasber Salim1,4,*
1Barn Owl and Rodent Research Group (BORG), School of Biological Sciences, Universiti Sains Malaysia,
Penang 11800, Malaysia, 2FGV Agri Services Sdn Bhd, Tun Razak Agricultural Research Centre, Jerantut, 27000
Pahang, Malaysia, 3FGV R&D Sdn Bhd, Tun Razak Agricultural Research Centre, Jerantut, Pahang 27000,
Malaysia and 4Vector Control and Research Unit, School of Biological Sciences, Universiti Sains Malaysia,
Penang 11800, Malaysia
*Corresponding author: E-mail: hasbersalim@usm.my
Submitted: 12 May 2019; Received (in revised form): 5 December 2019; Accepted: 16 December 2019
Abstract
This study investigated the diet of introduced barn owls (Tyto alba javanica, Gmelin) in the urban area of the Main Campus of
Universiti Sains Malaysia, Penang, Malaysia, based on collected regurgitated pellets. We also compared the diet of the intro-
duced barn owls with the diet of barn owls from two agricultural areas, i.e. oil palm plantations and rice fields. Pellet analy-
sis of introduced barn owls showed that commensal Norway rats, Rattus norvegicus, made up the highest proportion of the
diet (65.37% prey biomass) while common shrews, Suncus murinus were the second highest consumed prey (30.12% prey bio-
mass). Common plantain squirrel, Callosciurus notatus, made up 4.45% of the diet while insects were taken in a relatively
small amount (0.046% prey biomass). Introduced barn owls showed a preference for medium-sized prey, i.e. 40–120 g
(52.96% biomass and 38.71% total). In agricultural areas, rice field rats, Rattus argentiventer predominated the diet of barn
owls (98.24% prey biomass) in rice fields while Malayan wood rats, Rattus tiomanicus, were the most consumed prey in oil
palm plantations (99.5% prey biomass). Food niche breadth value was highest for barn owls introduced in an urban area
with a value of 2.90, and 1.06 in rice fields and 1.22 in oil palm plantations. Our analysis reiterates the prey preference of
barn owls in various landscapes for small mammals. Our results also indicate the suitability of utilizing barn owls as a bio-
logical control not only in agricultural areas, but also as a biological control agent for commensal rodent pests in urban
areas.
Key words: barn owl, diet, pellet analysis, urban, agriculture
C The Author(s) 2020. Published by Oxford University Press.
V
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12 | Journal of Urban Ecology, 2020, Vol. 6, No. 1
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Figure 1: Study site of introduced barn owls and location of nest boxes
agricultural areas (Hafidzi and Naim 2003b; Rizuan et al. 2017)
Introduction
and semi-urban areas (Meyer 2008) for the purpose of control-
The barn owl, Tyto alba (Tytonidae), is a common species of ling pest rodent populations. Barn owls are also translocated as
owls which occurs on almost all continents and in most open part of reintroduction programs for declining local barn owl
lands and farmlands (Bunn et al. 1982; Taylor, 1994). Like many populations (Meek et al. 2003). In this study, Southeast Asian
other cosmopolitan nocturnal raptors, barn owls display an as- barn owls, T. alba javanica, were translocated from their native
tonishing breadth of habitat association and have been able to agricultural habitats and introduced to the urban-garden area of
adapt and persist in areas that are becoming urbanized the Main Campus of Universiti Sains Malaysia to serve as a bio-
(Hindmarch et al. 2017). The diet of barn owls has been well logical control agent for the rat pest population. Here, we report
studied throughout its range because of the ease of identifying the analysis of the diet composition of introduced barn owls in
prey remnants recovered inside regurgitated pellets. Owls swal- an urban area and compare the diet of introduced barn owls to
low their prey whole and expel pellets, which are composed of the diet of barn owls in oil palm plantations and rice fields in
undigested remains such as bones, compacted in hair and Peninsular Malaysia.
feathers (Taylor 1994). Analysis of barn owl pellets has provided
information on the diet composition of owls and dynamics of
prey species communities within the owl foraging areas Methods
(Alivizatos and Goutner 1999; Kitowski 2013). Study area and introduced barn owls
Extensive studies of barn owls across their foraging range re-
port that barn owls feed primarily on small mammals, i.e. rats, In total, 24 barn owls were released intermittently from April
mice, voles and shrews, with birds, insects, amphibians, reptiles 2016 to August 2018 in the urban-garden area of the Main
and invertebrates taken in relatively small amounts (Marti 2010; Campus of Universiti Sains Malaysia Penang, Malaysia (5.3579
Paspali et al. 2013; Hindmarch and Elliot 2015). In Peninsular N, 100.2943 E). Prior to the release, 14 artificial nest boxes were
Malaysia, several studies on the food selection of barn owls in installed around the campus area. Providing nest boxes is a
major agricultural crop areas report rats as the main prey. Diet common practice to attract barn owls and increase nesting per-
analysis of the owl’s regurgitated pellets show that rats com- formance and hence sustain a barn owl population. Two types
prise >98% of the prey in oil palm plantations (Lenton, 1984) of artificial nest boxes, i.e. wooden and fibreglass, were installed
and 94.7% in rice fields (Hafidzi, Zulkifli, and Kamarudin 1999). early in January 2016 and placed around the campus in open
Barn owls are an effective small mammal predator, which areas of vegetation (Fig. 1).
has led to it being introduced into various landscapes such as The translocated barn owls were harvested from three dif-
islands (Au and Swedberg 1966; Emmerson and Ascani, 1985), ferent locations in Peninsular Malaysia; oil palm plantations atDiet composition of introduced barn owls in urban area | 3
the Tun Razak Agricultural Research Centre, Bandar Jengka to determine the size of the prey (0.01 mm accuracy). Insects
Pahang (3.777967 N, 102.517238 E), rice fields of Bumbung found in pellets were identified using Borror and White (1970),
Lima, Kepala Batas, Pulau Pinang (5.51707 N, 100.4265 E) and whereas other vertebrate prey were determined up to family
rice fields in the Kerian District, Parit Buntar, Perak (5.0081 N, level using identification keys by Beisaw (2013).
100.5394 E). Owls were harvested from areas with 100% occu- The biomass of prey items recovered from pellets were esti-
pancy of provided nest boxes, indicating a healthy population. mated using a standard log-log regression of right mandible
No more than one-third of individual fledglings and/or older length as a function of body weight (Morris 2009; Hamilton 1980;
owlets in the nest were harvested from the same nest-box. Marti 2009). The food niche breadth (FNB) of barn owls in all the
Owlets were only harvested if they were of the age of one areas was calculated to determine the dietary diversity of barn
month or more. The owls were temporarily held in the USM owls in each habitat. FNB (Levins 1968) was calculated as follows:
Aviary (5.35791944 N, 100.29416667 E) for about one month be-
fore release to allow the birds to acclimatize to their new urban 1
FNB ¼ Xn
surroundings. pi 2 ;
I¼1
All introduced barn owls were banded with customized metal
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leg bands prior to release. Transmitters were fitted to the owls us-
where p is the proportion to prey category i in the barn owl diet.
ing backpack style (Saufi, Ahmad, and Salim 2018). The transmit-
Higher values on this index represent a higher diversity of the diet.
ter and harness weighed 9 g, i.e.4 | Journal of Urban Ecology, 2020, Vol. 6, No. 1
Table 1:Diet composition of introduced barn owls and barn owls residing in agricultural areas
Prey species Urban (%) Rice field (%) Oil palm plantation (%)
Biomass Individuals Biomass Individuals Biomass Individuals
Norway rat (R. norvegicus) 65.37 45.05 0 0 0 0
Rice field rat (R. argentiventer) 0 0 98.24 96.77 0 0
Wood rat (R. tiomanicus) 0 0 0 0 94.35 90.16
House shrew (S. murinus) 30.12 35.16 1.28 2.15 0 0
Bird sp. 0 0 0 0 0.76 1.63
Reptile sp. 0 0 0 0 0.66 0.81
Amphibian sp. 0 0 0.46 1.07 0.47 1.63
Grasshopper sp. 0.11 5.49 0 0 0.02 3.27
Termites sp. 0.002 12.10 0 0 0 0
Plantain squirrel (C. notatus) 4.45 2.20 0 0 3.71 2.45
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FNB 2.90 1.06 1.22
FNB, food niche breadth.
60.00
52.96
50.00
38.17
Percentage (%)
40.00
31.18 30.89
30.00
20.00 17.20 16.00
12.90
10.00
0.15
0.00
30% of prey biomass. The extra small-
The released barn owls in the urban area recorded the highest sized prey made up 17.20% of total prey and contributed only
FNB value at 2.90, while the FNB value was second highest for 0.15% of prey biomass.
barn owls in oil palm plantations (1.22 FNB) and barn owls in As Norway rats were the most preferred prey of the barn
rice fields recorded the lowest FNB value (1.06 FNB). Thus, intro- owls, further analysis was carried out on the size of the rats.
duced barn owls had a relatively higher diet diversity compared Our analysis showed that the most consumed weight group of
with the diet of barn owls at agricultural areas. rats were medium-sized rats, i.e. individuals weighing 80–120 g
(Fig. 3). Seventeen individual medium-sized rats were con-
Prey weight of introduced barn owls sumed (44.74%). Twelve small-sized rats weighing from 40 to 80
g were the second highest weight group consumed by barn owls
The biomass of identified preys inside the collected pellets were
(31.58%) and the least consumed weight group were large-sized
estimated using a standard log-log regression of right mandible
rats weighing >120 g (9 individuals, 23.68%). Norway rats weigh-
length (mm) as a function body weight (g) as described by
ing 160 g were not found in our pellet analysis.
Hamilton (1980). Figure 2 shows the weight groups of introduced
barn owls prey by numbers and biomass. Weights of prey were
identified as extra small (Diet composition of introduced barn owls in urban area | 5
Percentage of indidivual R.norvegicus prey consumed (%) small mammal species was also reported as the second most
abundant prey species in urban and rural areas in Canada
(Hindmarch and Elliot 2015; Hindmarch et al. 2017) and the spe-
cies is more abundant in urban settings compared with agricul-
tural settings (Chang et al. 1999). During tracking of released
23.68%
(n=9) 31.58% barn owls, Norway rats and house shrews were frequently en-
(n=12) countered in residential neighbourhoods, eateries, garbage
dump areas and commercial areas within the study site (per-
sonal observation). It is however interesting to note that other
detrimental rodent pests, i.e. house mice and roof rats, were not
44.74% found in collected pellets despite being captured occasionally
(n=17) during rat trapping sessions we conducted as part of the study
on population diversity of rats in urban areas around Penang
Island (Amni et al. 2019). Timm (1994) documented that house
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Small (40-80 g) Medium (80-100 g) Large (120-140 g) mice and roof rats are typically found inside buildings and
house mice rarely travel outside, hence have low chances of
Figure 3: Percentage of size of Norway rats, R. norvegicus, consumed by intro-
falling prey to barn owls. Meanwhile, Norway rats and house
duced barn owls
shrews mainly inhabit and forage in open habitats (Timm 1994),
hence the two species inhabiting open areas and vegetation
buildings and houses, as well as abandoned structures. The
were the primary source of food for the owls (Bonvicino and
owls were also seen hunting in open grass habitats near road-
Bezerra 2003).
sides, human settlements, and backyards of shop lots. One of
Barn owls have been well documented to take advantage of
the barn owls also started occupying one of our installed nest
other temporarily abundant types of prey that are vastly differ-
boxes near the aviary, indicating the successful release of barn
ent from their usual diet, though extreme exceptions are un-
owls in an urban area. On the other hand, a substantial number
usual and usually occur in situations where small rodents are
of released barn owls in this study (21 out of 24 released owls)
absent or scarce (Taylor 1994). In Malaysia, most studies report
were untraceable a week after their release. These released
that the diet of T. alba javanica is composed >90% of rats (Smal
young barn owls dispersed further away from the release site
1990; Puan et al. 2011), with barn owls also preying on shrews,
and are probably foraging around the urban areas of Penang
squirrels, birds and lizards in smaller numbers (Smal 1990).
Island or could have travelled further to mainland Peninsular
Introduced barn owls in this study consumed small rodents
Malaysia (Saufi, Ravindran, and Salim 2019).
from the family Sciuridae. The common plantain squirrel, an un-
Similar to various studies on the diet of barn owls, our study
common barn owl prey, constituted a small fraction in the diet
reports that barn owls in agricultural areas and urban areas
of barn owls (2.20% total and 4.45% biomass). An interesting re-
prey mostly on small mammals (e.g. Marti 2010; Hindmarch and
sult from our analysis is that there were no bird remnants found
Elliot 2015; Horváth, Morvai, and Horváth 2018). Norway rats
in the pellets of introduced barn owls despite the abundant oc-
and house shrews (80.12% total and 95.49% prey biomass) were
currence of passerine birds in our study site. In contrast, several
the dominant prey group in the diet of barn owls in the urban
reports analysing the diet of barn owls in rural and urban areas
area. Our study shows that introduced barn owls were able to
document that Norway rats, R. norvegicus and birds make up a
adapt to an urban setting and consume abundant urban small
substantial proportion of the diet of owls (Salvati, Ranazzi, and
mammal species. The high number of Norway rats and house
Manganaro 2002; Teta, Hercolini, and Cueto 2012; Hindmarch
shrews consumed by barn owls indicate the owls managed to
and Elliot 2015). The pellet analysis of introduced barn owls also
hunt close to their release site and did not have to travel a great
showed that the owls preyed on insects, i.e. grasshoppers and
distance for more suitable open hunting grounds. Clark and
termites. Though infrequent, barn owls have been reported to
Bunck (1991) reported that North American barn owls do con- consume high amount of insects during insect swarms, such as
sume commensal rodents along their distribution, though only termites (e.g. Taylor 1994) and locusts (Szabo et al. 2003).
in low frequencies. Studies by Álvarez-Castan ~ eda, Cárdenas
and Méndez (2004) in Mexico and Magrini and Facure (2008) in
Brazil reported that pellets from barn owls in periurbans areas
Comparing diet of introduced barn owls in urban area
contain none to very little prey remnants from urban areas, sug- and barn owls in agricultural areas
gesting that barn owls spend more time hunting in areas away Though members of the Muridae family dominate the diet of
from human settlements. In Canada, Hindmarch and Elliot barn owls in all habitats, the main prey species differed by habi-
(2015) reported that barn owls retained their preference for tat. Rattus norvegicus were the most preyed upon by introduced
voles despite being in an urban landscape, although rats were owls while R. tiomanicus and R. argentiventer were the most
consumed in higher amounts in urban areas. preyed upon small mammal in oil palm plantations and rice
The commensal rodent pests, Norway rats, R. norvegicus, and fields, respectively. Barn owls prey-species preference in agri-
Black rats, R. rattus, are among the most widespread urban pest cultural areas from our study is similar to other reports by
species in the world and they reside in close proximity to hu- Lenton (1984) and Hafidzi and Naim (2003a) which assessed the
man habitation and are rarely found in the wild (Feng and diet of barn owls in rice fields and oil palm plantations,
Himsworth 2014). The substantial occurrence of Norway rats in respectively.
the diet of introduced barn owls show that the barn owls are FNB value of barn owls in the study was highest in the urban
taking advantage of the abundance of this pest species (Amni area when compared with agricultural areas. There was a
et al. 2019). Common house shrews were the second most con- higher component of unusual prey items in the urban area com-
sumed prey of barn owls and the second most trapped species pared with agricultural lands, with squirrels and insects ac-
in residential areas of our study site (Amni et al. 2019). This counting for 19.79% of individual prey and 4.51% of total prey6 | Journal of Urban Ecology, 2020, Vol. 6, No. 1
biomass of owls in the urban area. This observation is similar to
Conclusion
reports of Salvati, Ranazzi, and Manganaro (2002) and Teta,
Hercolini, and Cueto (2012) who report an increase in non- Our study shows that barn owls in urban and agricultural areas
rodent prey items in the pellets of barn owls as their habitat are opportunistic predators that hunt almost exclusively on
becomes more urbanized. Barn owl prey selection was showed small- to medium-sized small mammals. Our study also
by previous food-niche studies to be associated with rodent showed that barn owls can switch their prey species preference
accumulations and density (e.g. Marti 1988; Taylor 1994; Leveau in different areas according to variations in small mammal
et al. 2006; Bernard et al. 2010; Marti 2010, Milana et al. 2016). abundance. Barn owls introduced in an urban area mostly con-
Similar to other food-niche analysis of barn owls in Europe (e.g. sumed Norway rats and house shrews, which are ubiquitous
Milchev 2015; Horváth, Morvai, and Horváth 2018), North commensal small mammal pests. Squirrels and insects were
America (Marti 1988, 2010) and South America (e.g. Leveau et al. also preyed by the introduced barn owls but made up only a
2006; Teta, Hercolini, and Cueto 2012), the low values of niche small fraction of their diet. Our results strongly indicate that
breadth analysis from agricultural areas in this study reflect the barn owls introduced to urban areas have the potential to be an
high abundance of an available and profitable prey, i.e. the effective biological control agent against commensal small
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dominance of R. tiomanicus and R. argentiventer in oil palm plan- mammal pest populations following their high consumption by
tations and rice fields, respectively. It is fairly well established barn owls.
that R. argentiventer is common in rice fields (Lam 1983, 1988)
and R. tiomanicus is common in oil palm plantations (Wood and Acknowledgements
Liau, 1984).
We would like to thank Department of Agricultural,
Bumbung Lima, Kepala Batas, Penang and Department of
Prey size preference of introduced barn owls
Agricultural Kerian District, Perak for giving permission to
Barn owl diet depends on the abundance of food supply, prey carry out our sampling in their rice fields. The authors also
accessibility, which is affected by habitat characteristics, and thank FGV Agri Services Sdn. Bhd and FGV R&D Sdn Bhd for
general opportunistic feeding strategy (Taylor 1994; Bond et al. providing the necessary facilities to conduct this study in
2005; Horváth, Morvai, and Horváth 2018; Arlettaz et al. 2010). their oil palm plantations.
Morphological features, such as body size and conspicuousness,
and behaviour can also affect prey vulnerability to predation by
barn owls (Derting and Cranford 1989). Studies on differential Funding
prey selection by barn owls yield differing and often, contrast- This work was supported by the Universiti Sains Malaysia
ing results. Some studies show barn owls have a preference for Research University Grant (1001/PBIOLOGI/811270).
smaller prey weighing 50 g
(Derting and Cranford 1989; Castro and Jaksic 1995). Our analy-
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