Podoces, 2012, 7(1/2): 21–32 PODOCES 2012 Vol. 7, No. 1/2 Journal homepage: www.wesca.net Diet Composition of the Barn Owl Tyto alba (Aves: Tytonidae) and Spotted Owlet Athene brama (Aves: Strigidae) Coexisting in an Urban Environment A. Mohamed Samsoor Ali1,2*& R. Santhanakrishnan1 1) Department of Zoology, Saraswathi Narayanan College, Perungudi, Madurai - 625 022, Tamil Nadu, India 2) Present Address: Environmental Impact Assessment Division, Sálim Ali Centre for Ornithology and Natural History (SACON), Anaikatty, Coimbatore - 641 108, Tamil Nadu, India Article Info Original Research Received 17 October 2012 Accepted 7 February 2013 Keywords Arthropods Barn Owl Diet Food niche overlap Small mammal Spotted Owlet Abstract The comparative diet of the Barn Owl Tyto alba and the Spotted Owlet Athene brama were studied in an urban habitat of Madurai District, Tamil Nadu, Southern India, during January–December 2008. Regurgitated pellets of these two owl species were analysed to understand their dietary composition. The diet of the Barn Owl mainly comprised small mammals such as Suncus murinus (51.9%) and Rattus rattus (28.6%), while the diet of the Spotted Owlet comprised mostly arthropods (84.9%); Coleoptera (40.9%) and Orthoptera (32.4%). Food niche overlap between the Barn Owl and the Spotted Owlet in overall diet was 6.5%, which indicated very low degree of overlap. 1. Introduction The Barn Owl Tyto alba (Scopoli, 1769) and the Spotted Owlet Athene brama (Temminck, 1821) are known to coexist and use the same habitat in southern India. Since these two species are small mammal predators and known to live together, they are suitable species for studying resource partitioning and ecological segregation. Coexisting species usually depend on different requirements among the total resources available to them. The diversity of prey items taken by different predator species constitutes ecological studies of sympatric or coexisting species. Studies on food partitioning are relatively scarce for raptors, and especially owls in India, but considerable reports are available regarding coexisting owls in various geographical ranges (Georgiev 2005, Scheibler 2007, Riegert et al. 2009, Nadeem et al. 2012). * Corresponding: [email protected] The Barn Owl and the Spotted Owlet were studied independently by various authors in different parts of India and thus far, no attempt has been made to study the above coexisting owls in any particular area. The Barn Owl, a widely distributed, resident, and nocturnal raptor in India, is closely associated with human-made structures and agriculture (Marti et al. 1979, Kahila et al. 1994, Santhanakrishnan 1995). The Barn Owls are often found in close proximity to humans, and their occurrence is strongly associated with the presence of buildings with suitable places for roosting or nesting (de Bruijn 1994, Shawyer 1994). Its diet consists chiefly of small mammals, particularly rodents, but birds, reptiles, amphibians and insects are also eaten (Santhanakrishnan 1995, Mushtaq-ul Hassan et al. 2004, Sommer et al. 2005, Leonardi & Arte 2006, Neelanarayanan 2007a). Extensive studies have been conducted on the nesting and feeding habits of the Barn Owl in many parts of 21 Diet composition omposition of Barn Owl and Spotted Owlet – Ali & Santhanakrishnan the world (Roulin 2002, Altwegg et al. 2003, Haralambos et al.. 2005) but little is known on the ecology of this species in south outh-east Asia, particularly in India. Available literature on Indian Barn Owls focuses mainly on nest-sites nest (Nagarajan et al.. 1995), utilization of manman made nest boxes (Neelanarayanan et al. al 1995), sexual dimorphism (Kanakasabai et al al. 1996), nestling growth patterns (Nagarajan et al. 2002) and diet composition (Neelanarayanan 2007a, b). The Spotted Owlet is the most common small-sized sized (Ali & Ripley 1983) and least studied owl species in India. a. Spotted Owlets are nocturnal and roost in the day under thick, shady vegetation, cavities in tree, human-made human structures, including functional or nonnon functional irrigation wells and crevices found in rock cliffs (Santhanakrishnan et al. 2010a). It is a generalist predator that feeds on diverse prey such as rodents, bats, small birds, reptiles, amphibians and invertebrates like insects, annelids, scorpions and molluscs (Jain & Advani 1984, Kumar 1985, Jadhav & Parasharya 2003, Pande et al. 2004, 2007). The present study focused on investigating investigat the diet of the Barn Owl and the Spotted Owlet in an urban habitat simultaneously. Our aim was to compare the dietary composition and food niches of these two owl species over a period of one year (January–December December 2008) in the urban environment of Madurai District, Tamil Nadu, southern India. 2. Study Areas and Methods 2.1. Study areas This study was conducted in the Madurai District of Tamil Nadu, southern outhern India (Fig. 1). Topographically, the area is mostly flat with a few hilly tracts. Madurai, lies between 9°56' N and 78°07' E, is situated on the banks of the River Vaigai. The climate is dry and hot, with rains during October to December (northeast monsoon). During our research, research summer temperature emperature reached a maximum of 40°C and a minimum of 26.3°C, while winter temperature reached a maximum of 22.5°C and minimum of 19.7°C. The average annual rainfall is about 850 mm. Paddy is the predominantly cultivated crop in the study area; however other oth crops such as sugarcane, banana, jasmine, betle vine, groundnut and sorghum are also cultivated in different regions. 22 The regurgitated pellets of both owl species were collected in two different places situated at the middle of the city of Madurai South Taluk, aluk, surrounded by houses, shops and other human-made made structures. The roost roost/nest sites of the Barn Owl were located in Sout South Krishnan Temple (9°54'50.65˝N, 70°07'01.46˝E). A pair of the Barn Owls was spotted in the fifth opening of the temple tower. The temple tower provided a series of steps inside that provided easy access to the owls. Also a roost roost-site of the Spotted Owlet was located in Thirumalai Naikar Mahal (9°54'53.89˝N, 70°07'20.39˝E). A pair of the Spotted Owlets was located in a roof of building ng and the pellets were recovered beneath the building. Fig. 1. Map of India, Tamil Nadu and Madurai District (Map not to scale). 2.2.. Pellet collection and analysis The roost-site / nest-sites sites of both species were visited twice a month from January 2008 to December 2008 for the collection of the pellets. Fresh pellets were collected separately during each visit and pleced in plastic bags along with tags indicating the species of owl, site name and date of collection before being brought to the laboratory.. Pellets were kept at 70°C in a hot air oven for 24 hours to kill the associated invertebrates. Podoces, 2012, 7(1/2): 21–32 The length and width of owls pellets collected were first measured using a Vernier Caliper (with an accuracy of 0.05mm). The samples were then weighed using electronic balance with accuracy to 0.01 g. Pellets were analysed following Schueler (1972) and Yalden (2003). At the time of analysis, each pellet was put in warm water or 3% Sodium Hydroxide solution for softening. The pellet materials were carefully opened up using tweezers. During pellet examination, skulls, bones, feathers, beaks and insect remains were separated out first for identification. For studying seasonal variations in the diets of the two owl species, four seasons of a year namely post-monsoon (January–March), summer (April–June), premonsoon (July–September) and monsoon (October–December) were recognised. 2.3. Prey identification Mammalian prey items were identified to species level by comparing their bony remains (e.g. skulls, mandibles) with museum specimens collected from the same study area. Birds, reptiles and amphibians were identified by their feathers, beaks, skulls, synsacrum, mandibles and parasphenoid processes (Naranthiran 1989, Daniel 1992, Daniels 2005). We were not able to identify the bats, birds and amphibians down to species level. Arthropod prey items were identified up to order level on the basis of undigested anatomical pieces such as heads, mandibles, wings, legs, stings and body rings (Naranthiran 1989, Yalden 2003, Asokan et al. 2009). Most of the remains of bats, birds, lizards and frogs were not intact. 2.4. Biomass estimation The biomass of the prey of all mammalian species recovered from the pellets were estimated using a standard log-log regression of right mandible length as a function of body weight following Hamilton (1980), Santhanakrishnan (1987, 1995) and Neelanarayanan (2007b). For this study, the locally available mammalian species such as House Rat Rattus rattus, Lesser Bandicoot Rat Bandicoota bengalensis, Indian Field Mouse Mus booduga and Grey Musk Shrew Suncus murinus and their corresponding log-log regression values for the calculation of biomass were used as described by Santhanakrishnan (1995). 2.5. Data analysis Magurran (1988) was followed to assess and compare the diversity in the diet of two owl species by using species richness (S), Shannon’s index (H’), and Peilou’s evenness (E) indices. Food niche breadth (NB) was estimated according to Levins (1968): NB = (∑Pi2)–1, where, Pi is the proportion of prey i in the food. The food niche overlap values between two owl species were calculated by Pianka’s index (Pianka 1973): O = ∑PijPik/ √∑Pij2 Pik2 where, Pi is the proportion of prey species i in the diet of predator j and k respectively. A value of 0 means no overlap and 1 means complete overlap. Kruskal Wallis ANOVA was used to estimate the variation of the prey remains between seasons. Data are presented as mean ± SD. The MINITAB (version 13.1) statistical software was used for the statistical analyses. 3. Results 3.1. Pellets Fresh Barn Owl pellets were dark green in colour and covered with mucous. Pellets were compact and had many bones (Fig. 2). A total of 402 Barn Owl pellets were collected during the study period. The average pellet length (N=213) was found to be 3.7 ± 0.88cm (range: 1–8cm), width (N=213) of pellet was 2.5 ± 0.53cm (range: 0.8–4cm) and the average dry weight of pellet (N=98) was found to be 3.3 ± 1.23g (range: 1.2–6.9g). The number of prey items per pellet varied from one to seven with the mean of 1.3±0.98 prey per pellet. The regurgitated pellets of the Spotted Owlets were in different size and shapes (Fig. 3). The fresh pellets were compact and dark in colour because they contained masses of undigested material, mostly insects. The average length of the Spotted Owlet pellet (N=310) was 2.7 ± 0.23cm (range: 2.2–3.3cm) and width (N=310) was 1.4 ± 0.13cm (range: 1.1–1.8cm). The mean weight of the 111 pellets was 0.39 ± 0.77g (range: 0.11–0.89g). The mean number of prey items per pellet was 8.3 ± 2.45 (range: 3–19). 23 Diet composition of Barn Owl and Spotted Owlet – Ali & Santhanakrishnan 3.2. Diet of the Barn Owl From all collected pellets, a total of 567 prey items could be distinguished with a total mammalian biomass of 30,153 g (Table 1). The Barn Owl fed on insectivore mammals, rodents, bats, lizards, frogs and insects. Six small mammalian species comprised 89.7% of all prey recorded from the study area. Among the mammals, the Suncus murinus (51.9%) was the most dominant prey species in the diet of the Barn Owl, followed by rodents, Rattus rattus (28.6%) and Mus booduga (4.8%). The proportion of insects in the diet was 3.4% and other prey items were formed less than 3%. Among the mammalian biomass consumed by the Barn Owl, rodents constitute 58.4% of the diet and that of Suncus murinus was 41.6% (Table 1). Fig. 2. Various shapes and sizes of the Barn Owl pellets recorded in Madurai District, Tamil Nadu, Southern India. Fig. 3. Various shapes and sizes of the Spotted Owlet pellets recorded in Madurai District, Tamil Nadu, Southern India. 24 3.3. Seasonal diet of the Barn Owl A total of 131 prey items were identified from the analysis of 89 pellets in the post-monsoon. Small mammals were main prey items (90.2%) of the Barn Owl. The Suncus murinus was predominant prey species among small mammals which accounted for 63.6% of the total number of the prey individual and 67.2% of their biomass. The Suncus murinus was followed by Rattus rattus (15.0%, 30.1%), Mus booduga (9.3%, 2.7%) and frogs (3.1%). The contribution of other prey species such as bats, birds, lizards and insects was less than 3%. The Bandicoota bengalensis was absent in the post-monsoon diet of the Barn Owl (Table 2). Podoces, 2012, 7(1/2): 21–32 Table 1. Prey frequencies (%) and biomass (%) consumed by the Barn Owl and Spotted Owlet in Madurai District, Tamil Nadu, Southern India. Prey items Small mammals Insectivores Suncus murinus Rodents Bandicoota bengalensis Rattus rattus Mus booduga Bats (Chiroptera) Birds Lizards Frogs Arthropods Coleoptera Orthoptera Hemiptera Diptera Hymenoptera Lepidoptera Arachnida (Scorpions) Unidentified items Total Number and Estimated biomass (g) Barn Owl (N=402) % % Total Number Biomass Number Spotted Owlet (N=310) % % Total Number Biomass Number 51.9 41.6 294 2.8 34.7 72 2.3 28.6 4.8 2.1 1.8 0.4 1.8 3.4 53.5 1.5 ----- 13 162 27 12 10 2 10 0.5 3.1 1.2 --4.5 -- 3.6 58.9 2.8 ---- 13 79 31 --115 -- 3.4 ------3.2 100 --------30153 19 ------18 567 40.9 32.4 3.9 1.7 2.3 2.2 1.5 3.0 100 --------4125 1053 836 100 44 60 58 38 78 2577 Table 2. Seasonal changes in frequency (N%) and biomass (B%) of consumed prey items in the diet of Barn Owl in Madurai District, Tamil Nadu, Southern India. Prey items Small mammals Insectivores Suncus murinus Rodents Bandicoota bengalensis Rattus rattus Mus booduga Bats (Chiroptera) Birds Lizards Frogs Insects Coleoptera Unidentified items Number of pellets Total prey items Mean prey/pellet Estimated biomass (g) Post-monsoon %N %B Summer %N %B Pre-monsoon %N %B Monsoon %N %B 63.6 67.2 36.2 48.5 52.1 39.3 56.4 28.9 0.0 15.0 9.3 2.3 1.6 0.4 3.1 0.0 30.1 2.7 ----- 2.9 41.3 0.0 2.2 2.9 0.0 2.9 3.3 48.2 0.0 ----- 0.7 32.5 5.6 0.0 0.7 0.0 1.4 1.0 58.4 1.3 0.0 ---- 5.2 25.0 4.5 3.8 1.9 0.0 0.0 7.2 61.8 2.1 ----- 0.8 3.9 --- 7.2 4.3 --- 5.6 1.4 --- 0.0 3.2 --- 89 131 1.1 ± 0.25 5151 95 138 1.2 ± 0.19 7056 89 142 1.1 ± 0.35 7851 129 156 1.3 ± 0.21 10095 25 Diet composition of Barn Owl and Spotted Owlet – Ali & Santhanakrishnan Values During the summer, Prey species Richness Diversity Index (H') Evenness Index (E) 95 Barn Owl pellets were analysed, yielding 138 9 prey items. The 8 consumption of small 7 mammals was decreased 6 (82.6%) in the summer 5 diet. The most preferred prey both in number and 4 biomass was Rattus rattus 3 (41.3%, 48.2%) and 2 Suncus murinus (36.2%, 1 48.5%). The Bandicoota 0 bengalensis, bats, birds and frogs were consumed Post-monsoon Summer Premonsoon Monsoon almost equally. The Seasons proportion of coleopteran Fig. 4. Prey diversity in the diet of the Barn Owl in Madurai District, Tamil Nadu, insects increased (7.2%) Southern India. in summer, and Mus booduga and lizards were 3.4. Diet of the Spotted Owlet absent in the summer diet. The remains of 2,577 prey items were found in During the pre-monsoon, 89 pellets were analysis of 310 pellets of the Spotted Owlet. examined and again small mammals (90.9%) The Spotted Owlet had eaten most frequently were most dominant in the diet of the Barn on arthropods (84.9%) followed by small Owl. Numerically, Suncus murinus (52.1%) mammals (7.6%) and lizards (4.5%). The was most dominant prey species but Rattus arthropods preyed upon are exclusively insects, rattus was more important in term of the predominantly Coleoptera (40.9%) and biomass (58.4%). The consumption of Mus Orthoptera (32.4%) followed by Hemiptera booduga increased (5.6%) and the proportion of (3.9%), Hymenoptera (2.3%), Lepidoptera other prey items were low. (2.2%) and Diptera (1.7%). Small mammals Totally, 129 pellets were analysed during were represented by two orders and four monsoon, yielding 156 prey items. Like other species. Among the mammalian prey, Rattus seasons, small mammals were the principal diet rattus was the dominant (3.1% in numbers and and represented 94.9%. Among the small 58.9% in biomass) followed by Suncus murinus mammals Suncus murinus numerically (2.8% in numbers and 34.7%in biomass) (Table dominated (56.4%) in the diet but Rattus rattus 1). contributed 61.8% biomass. The consumption of Bandicoota bengalensis, Mus booduga, bats 3.5. Seasonal diet of the Spotted Owlet and birds were increased during the monsoon. A total of 511 prey items were recovered from No lizards and insects were consumed during the analysis of 60 Spotted Owl pellets in the the monsoon. post-monsoon. Arthropods were predominant The consumption of Suncus murinus (84.8%) in the diet while rest of it comprised (Kruskal Wallis ANOVA: H=56.21, P<0.001), small mammals (7.6%) and lizards (4.8%). Mus booduga (H=123.5, P<0.0001), bats Among the arthropods, the Spotted Owlet (H=94.2, P<0.001) and birds (H=110.2, P< preyed mainly on the Coleoptera (47.2%) and 0.0001) was significantly different between the Orthoptera (24.4%). The Suncus murinus seasons. Both prey diversity index (H') and (3.6%) was a frequent mammalian prey species evenness index (E) had its lowest value in in the diet of the Spotted Owlet but Rattus summer and its highest value was in monsoon rattus contributed more biomass (50.3%; Table (Fig. 4). 3). 26 Podoces, 2012, 7(1/2): 21–32 Table 3. Seasonal changes in frequency (N%) and biomass(B%) of consumed prey items in the diet of Spotted Owlet in Madurai District, Tamil Nadu, Southern India. Prey items Post-monsoon Summer Pre-monsoon Monsoon %N %B %N %B %N %B %N %B 3.6 40.2 0.9 55.6 4.9 27.7 3.6 38.0 Bandicoota bengalensis Rattus rattus Mus booduga 0.4 2.2 1.4 6.0 50.3 3.5 0.5 3.2 2.3 4.2 40.2 0.0 0.0 1.5 0.7 0.0 68.5 3.8 0.0 6.9 0.0 0.0 62.0 0.0 Lizards 4.8 -- 4.9 -- 1.2 -- 5.2 -- Arthropods Coleoptera Orthoptera 47.2 24.4 --- 39.6 36.7 --- 35.4 40.1 --- 41.7 28.1 --- Hemiptera Diptera Hymenoptera Lepidoptera 4.0 2.2 2.0 2.6 ----- 3.1 1.2 0.9 1.7 ----- 4.6 2.8 3.3 2.1 ----- 3.8 0.9 2.7 2.6 ----- Arachnida (Scorpions) Unidentified items Number of pellets Total prey items 2.4 2.8 --- 1.2 3.8 --- 0.8 2.6 --- 1.7 2.9 --- Small mammals Insectivores Suncus murinus Rodents Mean prey/pellet Estimated biomass (g) 60 511 83 572 86 724 81 770 8.3 ± 3.25 984 6.9 ± 2.51 1008 8.4 ± 4.21 962 9.5 ± 4.17 1171 Overall, 572 prey items were identified in the analysis of 83 pellets during summer. Again arthropods remained dominant by numbers (84.4%) but the biomass was mostly due to Suncus murinus (55.6%) and Rattus rattus (40.2%). Coleoptera (39.6%) and Orthoptera (36.7%) were the most frequent among arthropods. The proportion of Suncus murinus, Coleoptera, Hemiptera, Diptera, Hymenoptera, Lepidoptera and Arachnida decreased, while that of Rattus rattus, Mus booduga, lizards and Orthoptera increased in the diet. Analysis of 86 pellets of the Spotted Owlet during pre-monsoon revealed the presence of 724 prey items. The consumption of Arthropods increased (89.1%) as compared to other seasons. The proportions of Rattus rattus, Mus booduga, lizards and Coleoptera decreased and the proportions of Suncus murinus (4.9%) and Orthopteran insects (40.1%) increased comparatively. The Rattus rattus (68.5%) had the greatest portion in its diet in terms of biomass. No Bandicoota bengalensis was eaten in this season. During the monsoon, 81 pellets were examined, arthropods remained dominant by numbers (81.5%), but the biomass was mainly because of the presence of two mammalian species, Rattus rattus (62.0%) and Suncus murinus (38.0%). The proportions of Orthoptera (28.1%) decreased and the proportions of Coleoptera (41.7) and lizards (5.2%) increased greatly. No Bandicoota bengalensis and Mus booduga were consumed during the monsoon season. There was significant seasonal variations in the diet of the Spotted Owlet for the prey items such as Coleoptera (Kruskal Wallis ANOVA: H=85.06, P<0.001), Orthoptera (H=185.3, P<0.0001), Hymenoptera (H=158.3, P<0.0001), lizards (H=65.52, P<0.001), Rattus rattus (H=156.41, P<0.0001) and Mus booduga (H=85.65, P<0.001). Diversity indices (H') revealed that the greatest and lowest diet diversity was during the summer and postmonsoon, respectively. The diet evenness index (E) had the highest value in summer and lowest value in monsoon (Fig. 5). 27 Diet composition of Barn Owl and Spotted Owlet – Ali & Santhanakrishnan Food Overlap Values 3.6. Comparison of diets of two sympatric owl Prey species Richness Diversity Index (H') Evenness Index (E) species 14 The Barn Owl and the Spotted Owlet consumed a 12 wide variety of prey items, 10 but the dominant prey taxa were different between the 8 two owl species. The diet composition of the Barn 6 Owl indicated that small mammals remained the 4 dominant prey, making up to 89.7% of their diet. 2 Among these, the 0 insectivore mammal Suncus murinus was the Post-monsoon Summer Premonsoon Monsoon major prey species Seasons (constituting 51.9%), Fig. 5. Prey diversity in the diet of the Spotted Owlet in Madurai District, Tamil followed by the rodent Nadu, Southern India. species Rattus rattus (28.6%). Arthropods 0.1 clearly dominated the diet of the Spotted Owlet, 0.08 constituting 84.9%, along with a lesser amount of 0.06 small mammals (7.6%) and lizards (4.5%). Among 0.04 arthropods, the Coleopteran beetles and 0.02 Orthopteran insects were the most dominant prey 0 items throughout the year. Post-monsoon Summer Pre-monsoon Monsoon Both species were dependent significantly Seasons more on the biomass of Fig. 6. Seasonal changes in food overlap of the Barn Owl and Spotted Owlet Rattus rattus. The niche inhabiting in Madurai District, Tamil Nadu, Southern India. breadth of the Barn Owl was considerably wider the pellets were different than the reports by (2.8) than the Spotted Owlet (2.3). The food Buxton & Lockley (1950; 110×50 mm), Glue niche overlap between the Spotted Owlet and (1967; 45×26 mm) and Nadeem et al. (2012; the Barn Owl in overall diet composition was 48.2×38.2 mm). In agreement, Hardy (1977) 6.5%, which indicated very low degree among stated that the size of the pellets significantly both species. The low degree of food overlap varied, which depended on the composition of was also found in different seasons in both the diet and the size as well as nutritive value of species (Fig. 6). taken prey. The mean number of prey per pellet 4. Discussion The size of the Barn Owl pellets in this study area were similar to those reported by other workers (Glue 1967, Santhanakrishnan 1995, Alvarez-Castaneda et al. 2004, Mahmood-ul Hassan et al. 2007a). The length and width of 28 averaged 1.3 for the Barn Owl, which was similar to the average given by other workers (Neelanarayanan 2007a, Travaini et al. 1997, Bellocq 1998, Charter et al. 2007, Magrini & Facure 2008). The average measurements of pellet sizes reported for the Spotted Owlet in Podoces, 2012, 7(1/2): 21–32 other studies (Tariq et al. 2003, Pande et al. 2004, Nadeem et al. 2012) were different from our findings. Mean number of prey items per pellet was 8.3 for the Spotted Owlet, which compares favourably with mean given by Naranthiran (1989) and Pande et al. (2007). In contrast, Mahmood-ul Hassan et al. (2007b) reported the mean number of prey as being only 1.6 prey items in Pakistan. The number of prey per pellet of the Spotted Owlet was higher than that of the Barn Owl possibly because much larger numbers of insects were consumed by the Spotted Owlet. Several investigations on the diet of the Barn Owl in different geographical regions confirmed that small mammals were the main components of the diet (Marti 1974, Gubanyi et al. 1992, Pezzo & Morimando 1995, Travaini et al. 1997, Alivizatos & Gounter 1999, Love et al. 2000, Bose & Guidali 2001, Mushtaq-ulHassan et al. 2004, Alivizatos et al. 2005, Leonardi & Arte 2006, Mahmood-ul-Hassan et al. 2007c, Magrini & Facure 2008, Nadeem et al. 2012) and our results also indicated a similar pattern. Numerically, the Suncus murinus (51.9%) was the main contributor while in terms of biomass, the Rattus rattus (53.5%) was the main prey in the diet of the Barn Owl. In contrast to our findings, Santhanakrishnan (1995) and Neelanarayanan (2007a) reported that field rodents of agricultural value such as Bandicoota bengalensis, Tatera indica, Millardia meltada and Mus spp. were consumed predominantly and that Suncus murinus constituted only 20% of the diet in the Cauvery Delta region, Tamil Nadu, India. In the present study, the regurgitated pellets of both owl species were collected in an urban environment (dominated by human habitations). Therefore, Suncus murinus and Rattus rattus were the predominant prey in the Barn Owl diet. Barn Owls consume negligible amounts of birds. As other studies indicate, this may be because Barn Owls may concentrate on birds when other prey are scarce (Alivizatos et al. 2005, Leonardi & Arte 2006). Small proportions of lizards and frogs were recorded in the diet of the Barn Owl in this study. Some authors report these prey in the diet of the Barn Owls (Sommer et al. 2005, Neelanarayanan 2007a). The presence of insects in the diet of the Barn Owls have been described by many authors in different locations (Travaini et al. 1997, Alvarez-Castaneda et al. 2004, Alivizatos et al. 2005, Charter et al. 2007). Small fractions of Coleopteran insects were recorded in the diet of the Barn Owl in the present study. The consumption of small mammals in the diet of the Barn Owls among the seasons was more or less similar, but changes were noticed among the prey species within the small mammals. The main prey component of the Barn Owl diet, Suncus murinus was taken at lowest frequency during the summer season. In this period, the proportion of Rattus rattus, Bandicoota bengalensis, birds and insects increased. This seasonal trend was also documented by other authors (Barbosa & Benzal 1996, del Guasta 1999, Debrot et al. 2001, Mahmood-ul-Hassan et al. 2007c). The changes in prey composition in our study area may reflect the opportunistic hunting strategy of the Barn Owl in relation to the abundance and availability of prey items (Marti 1972, Barbosa & Benzal 1996, del Guasta 1999, Bose & Guidali 2001, Mahmood-ul-Hassan et al. 2007c). The pellet analysis demonstrates that the Spotted Owlet is not a specialist, with a diet composition of 84.9% invertebrates and 12.1% vertebrates. In general, insects (83.4%) play an important part of the Spotted Owlet diet. Among insect prey, Coleoptera (40.9%) and Orthoptera (32.4%) formed important insect prey. Our results were similar to those documented for the Spotted Owlet in India and Pakistan (Jain & Advani 1984, Kumar 1985, Naranthiran 1989, Beg et al. 1990, Jadhav & Parasharya 2003, Pande et al. 2004, 2007, Shah et al. 2004, Mahmood-ul-Hassan et al. 2007b). Mammals constituted only 7.6% of the total prey items. Rattus rattus and Suncus murinus were the most common prey among mammals and thus the Spotted Owlet plays an important role in pest control in human environments. Similar small mammal prey was reported for the diet of the Spotted Owlet by other authors (Jain & Advani 1984, Kumar 1985, Naranthiran 1989, Pande et al. 2004). Lizards and scorpions were hunted throughout the year. The Spotted Owlet foraged more on Coleopteran beetles throughout the year, with its lowest proportion during the pre-monsoon season. The decrease was compensated by an increased proportion of Orthoptera, Hemiptera, Diptera, Hymenoptera, 29 Diet composition of Barn Owl and Spotted Owlet – Ali & Santhanakrishnan Lepidoptera and Suncus murinus. During the monsoon season, the consumption of Coleoptera, Rattus rattus and lizards increased while a decrease in Orthoptera and other insect prey was observed. In conclusion, the diet composition of the Barn Owl and the Spotted Owlet varied throughout the year. The low dietary overlap between the two owl species may be due to their different body sizes. The Spotted Owlet is smaller (80–100g) than the Barn Owl (400– 500g) and seems to prefer comparatively smaller prey. The prey size selection of small mammals in the study area indicated that the Spotted Owlet had consumed 68% of the mammals in the 1–20g weight class interval, whereas the Barn Owls had selected mammals with a preference on 1–60g weight class interval up to 73% (Santhanakrishnan et al. 2010b, 2011). The observed differences between the Barn Owl and the Spotted Owlet diet in the study area may be caused by several factors. Taken collectively, their body size, prey availability, habitat type used for hunting prey, and hunting techniques allow these owls to minimise potential competition for food. Acknowledgements We thank the Ministry of Environment and Forests, Government of India, New Delhi for providing financial support. We are grateful to the Principal and the Management of Saraswathi Narayanan College, Madurai for having rendered facilities and encouragements. We are also thankful to Mr. U. Anbarasan and Mr. P. Muthukumar who assisted with the fieldworks. We thank David H. Johnson (Director – Global Owl Project) for his thoughtful edits to an earlier version of this manuscript. References Ali S. & Ripley S.D. (1983). Handbook of Birds of India and Pakistan. Oxford University Press, Bombay. Alivizatos H. & Gountner, V. (1999). Winter diet of the Barn Owl (Tyto alba) and Long-eared Owl (Asio otus) on northeastern Greece: a comparison. Journal of Raptor Research, 33, 160–163. Alivizatos H., Gounter V. & Zogaris S. (2005). Contribution to the study of four owl species (Aves, Strigiformes) from mainland and island areas of Greece. Belgian Journal of Zoology, 135(2): 109–118. Altwegg R., Roulin A., Kestenholes M. & Jenni L. (2003). Variation, covariation in survival, 30 dispersal and population size in Barn Owls Tyto alba. Journal of Animal Ecology, 72, 391–399. Alvarez-Castaneda S.T., Cardenas N. & Mendez L. (2004). Analysis of mammal remains from owl pellets (Tyto alba), in a suburban area in Baja California. Journal of Arid Environment, 59, 59– 69. Asokan S., Ali, A.M.S. & Manikannan R. (2009). Diet of three insectivorous birds in Nagapattinam District, Tamil Nadu, India – a preliminary study. Journal of Threatened Taxa, 1(6), 327–330. Barbosa A. & Benzal J. (1996). Diversity and abundance of small mammals in Iberia: peninsular effects or habitat suitability. Zeitschrift fur Saeugertierkunde, 61, 236–241. Beg M.A., Maqbool M. & Mushtaq-ul-Hassan M. (1990). Food habits of spotted owlet Athene brama. Pakistan Journal of Agricultural Science, 27, 127–131. Bellocq M.I. (2000). A review of the trophic ecology of the Barn Owl in Argentina. Journal of Raptor Research, 34(2), 108–119. Bose M. & Guidali F. (2001). Seasonal and geographic differences in the diet of the Barn Owl in an agro-ecosystem in Northern Italy. Journal of Raptor Research, 35(3), 240–246. Buxton J. & Lockley R.M. (1950). Island of Skomer. London. Charter M., Izhaki I., Shapira L. & Leshem Y. (2007). Diets of urban breeding Barn Owls (Tyto alba) in Tel Aviv, Israel. Wilson Journal of Ornithology, 119(3), 484–485. Daniel J.C. (1992). The Book of Indian Reptiles. Bombay Natural History Society and Oxford University Press, Mumbai. Daniels R.J.R. (2005). Amphibians of Peninsular India. Universities Press (India) Private Limited, Hyderabad. de Bruijn O. (1994). Population ecology and conservation of the barn owl Tyto alba in farmland habitats in Liemers and Achterhoek (The Netherlands). Ardea, 82, 1–109. Debrot A., Defreitas O. J.A., Brower A. & Kooy M.M. (2001). The curaco barn owl: status and diet 1987–1989. Caribbean Journal of Science, 37, 187–193. del Guasta M. (1999). The diet of barn owl (Tyto alba) of Mugello (Flourence) in relation to environmental factors. Museo Regionale di Scienze Natural Bullettino, 16, 39–58. Georgiev D.G. (2005). Food niche of Athene noctua (Scopoli, 1769) and Tyto alba (Scopoli, 1769) (Aves: Strigiformes) co-existing in one region of the upper Tracian Valley (South Bulgaria). Animalia, 41, 115–122. Glue D.E. (1967). Prey taken by the barn owl in England and Wales. Bird Study, 14, 169–183. Podoces, 2012, 7(1/2): 21–32 Gubanyi J.A., Case R.M. & Wingfield G. (1992). Diet and nesting success of Barn Owls breeding in western Nebraska. American Midland Naturalist, 127, 224–232. Hamilton K.L. (1980). A technique for estimating Barn Owl prey biomass. Journal of Raptor Research, 14(2), 52–55. Haralambos A., Gounter V. & Zogaris S. (2005) Contribution to the study of the diet of four owl species (Aves, Strigiformes) from mainland and island areas of Greece. Belgian Journal of Zoology, 135, 109–118. Hardy A.R. (1977). Hunting ranges and feeding ecology of owls in farmland. Ph.D. Thesis, University of Aberdeen, Aberdeen. Jadhav A. & Parasharya B. M. (2003). Some observations on the nesting behaviour and food of the Spotted Owlet Athene brama. Zoos’ Print Journal, 18(8), 1163–1165. Jain P. & Advani R. (1984). Winter food of Spotted Owlet Athene brama indica. Journal of the Bombay Natural History Society, 80(2), 415–416. Kahila G., Aviel S. & Tchernov E. (1994). Reproductive cycle of the Barn Owls (Tyto alba) in nesting boxes. Israel Journal of Zoology, 40, 100. Kanakasabai R., Neelanarayanan P. & Nagarajan R. (1996). Sexual dimorphism in Barn Owl (Tyto alba). Newsletter for Birdwatchers, 36, 55. Kumar T.S. (1985). The Life History of the Spotted Owlet (Athene brama brama, Temminck) in Andhra Pradesh. Raptor Research Centre, Heyderabad, India. Leonardi G. & Arte G.L.D. (2006). Food habits of the Barn Owl (Tyto alba) in a steppe area of Tunisia. Journal of Arid Environment, 65, 677– 681. Levins R. (1968). Evolution in Changing Environments. Monograph in Population BiologyVol . 2. Princeton University Press, Princeton, NJ. Love R.A., Webon C., Glue D.E., Harris S. & Harris S. (2000). Changes in the food of the British barn owls. Mammal Review, 30, 107–129. Magrini L. & Facure K.G. (2008). Barn Owl (Tyto alba) predation on small mammals and its role in the control of hantavirus natural reservoirs in a periurban area in southeastern Brazil. Brazilian Journal of Biology, 68(4), 733–740. Magurran A.E. (1988). Ecological Diversity and Its Measurements. Croom Helm Ltd., London. Mahmood-ul-Hassan M., Beg M.A. & Mushtaq-ul Hassan M. (2007a). Locality related changes in the diet of Barn Owl (Tyto alba stertens) in agroecosystem in Central Punjab, Pakistan. Wilson Journal of Ornithology, 119(3), 479–483. Mahmood-ul-Hassan M., Beg M.A., Mushtaq-ulHassan M. & Rana S.A. (2007b). Food habits of the spotted owlet Athene brama in central Punjab, Pakistan. Acta Zoologica Sinica, 53(1), 64–68. Mahmood-ul-Hassan M., Beg, M.A. & Ali H. (2007c). Seasonal variation in the diet of the barn owl Tyto alba stertens in central Punjab, Pakistan. Acta Zoologica Sinica, 53(3), 431–436. Marti C.D. (1972). The years of barn owl prey data from a Colorado nest site. Wilson Bulletin, 85, 85– 86. Marti C.D. (1974). Feeding ecology of four sympatric owls. Condor, 76, 45–61. Marti C.D., Wager P.W. & Denne K.W. (1979). Nest boxes for the management of barn owls. Wildlife Society Bulletin, 7, 145–148. Mushtaq-ul-Hassan M., Raza M.N., Shahzadi B. & Ali A. (2004). The diet of Barn Owl from canal bank, canal rest house and graveyard of Gogra. Journal of Research (Science), 15(3), 291–296. Nadeem M.S., Imran S.M.K., Mahmood T., Kayani A.R. & Shah S.I. (2012). A Comparative study of the diets of Barn Owl (Tyto alba) and Spotted Owlet (Athene brama) inhabiting Ahmadpur East, Southern Punjab, Pakistan. Animal Biology, 62(1), 13–28. Nagarajan R., Thiyagesan K., Natarajan R. & Kanakasabai R. (2002). Patterns of growth in nestling Barn-owls. Condor, 104, 885–890. Nagarajan R., Neelanarayanan P. & Kanakasabai R. (1995). Descriptions of nesting sites of Barn Owl (Tyto alba) in man-made structures. Zoos’ Print Journal, 10(5), 15. Naranthiran G. (1989). Ecology of Southern Spotted Owlet Athene brama brama (Temminck) in Thanjavur District, Tamil Nadu with special reference to its food habit and ecotoxicity. M.Phil. Thesis, Bharathidasan University, Trichy, India. Neelanarayanan P. (2007a). Diet of Barn Owl Tyto alba stertens Hartert, (1929) in a portion of Cauvery delta, Tamil Nadu, India. Zoos’ Print Journal, 22(8), 2777–2781. Neelanarayanan P. (2007b). Technique for estimation of barn owl (Tyto alba stertens Hartert, 1929) prey biomass with special reference to mandible length body weight ratio of small mammals. Zoos’ Print Journal, 22(1), 2519–2521. Neelanarayanan P., Nagarajan R. & Kanakasabai R. (1995). Scope for the utilization of man-made nest boxes to Barn Owl, Tyto alba. Zoos’ Print Journal, 8(11), 3. Pande S., Pawashe A., Murlidhar M.N., Joglekar C. & Mahabal, A. (2007). Effect of food and habitat on breeding success in Spotted Owlets (Athene brama) nesting in villages and rural landscapes in India. Journal of Raptor Research, 41(1), 26–34. Pande S., Pawashe A., Bastawade D.B. & Kulkarni P.P. (2004). Scorpions and molluscs: Some new dietary records for Spotted Owlet Athene brama in India. Newsletter for Ornithologist, 1(5), 68–70. 31 Diet composition of Barn Owl and Spotted Owlet – Ali & Santhanakrishnan Pezzo F. & Morimando F. (1995). Food habits of the barn owl, Tyto alba, in a Mediterranean rural area: Comparison with the diet of two sympatric carnivores. Italian Journal of Zoology, 62, 369– 373. Pianka E.R. (1973). The structure of lizard communities. Annual Review of Ecology and Systematics, 4, 53–74. Riegert J., Lovy M. & Fainova D. (2009). Diet composition of Common Kestrels Falco tinnunculcus and Long-eared Owls Asio otus coexisting in an urban environment. Ornis Fennica, 86, 1–8. Roulin A. (2002). Offspring desertion by doublebrooded female Barn Owls (Tyto alba). Auk, 199, 515–519. Santhanakrishnan R. (1987). Studies on population, food habits and nesting of Barn Owl, Tyto alba (Scopoli) in a portion of Cauvery river basin. M.Phil. Thesis, Bharathidasan University, Trichy, India. Santhanakrishnan R. (1995). Ecology of Barn Owl, Tyto alba (Scopoli) with special reference to its population, feeding and breeding in Mayiladuthurai, Tamil Nadu, South India. Ph.D. Thesis, Bharathidasan University, Trichy, India. Santhanakrishnan R., Ali A.M.S. & Anbarasan U. (2010a). Roost-site preference of Spotted Owlet Athene brama (Aves: Strigiformes) in Madurai District, Tamil Nadu, India. Global Journal of Environmental Research, 4(3), 161–167. Santhanakrishnan R., Ali A.M.S. & Anbarasan U. (2010b). Diet Variations of the Barn Owl Tyto alba (Scopoli, 1769) in Madurai District, Tamil Nadu, Southern India. Podoces, 5(2), 95–103. Santhanakrishnan R., Ali A.M.S. & Anbarasan U. (2011). Food habits and prey spectrum of Spotted Owlet (Athene brama) in Madurai District, Tamil 32 Nadu, southern India. Chinese Birds, 2(4), 193– 199. Scheibler D.R. (2007). Food partitioning between breeding White-tailed Kites (Elanus leucurus, Aves, Accipitridae) and Barn Owls (Tyto alba, Aves, Tytonidae) in southern Brazil. Brazilian Journal of Biology, 67(1), 65–71. Schueler F.W. (1972). A new method of preparing owls pellets boiling in NaOH. Bird Banding, 43, 142. Shah Z.A., Beg, M.A. & Khan A.A. (2004). Prey preferences of the spotted little owl (Athene brama) in the croplands near Faisalabad-Pakistan. International Journal of Agriculture and Biology, 6, 278–280. Shawyer C. (1994). Barn owl Tyto alba. In: Tucker, G.M. and Heath, M.F. (eds.), Birds in Europe. Their conservation status. BirdLife Conservation Series No 3, BirdLife International, Cambridge, U.K. Sommer R., Zoller H., Kock D., Bohme W. & Griesau A. (2005). Feeding of the barn owl, Tyto alba with first record of the European free-tailed bat, Tadarida teniotis on the island of Ibiza (Spain, Balearics). Folia Zoologica, 54(4), 364– 370. Tariq S.A., Ghazi R.R. & Nisa N. (2003). Study on the food of Spotted Owlet (Athene brama) at an Agricultural Farm Gharo, Lower Sindh, Pakistan. Journal of Natural History and Wildlife, 2(1), 31– 33. Travaini A., Donazar J.A., Ceballos O., Rodriguez A., Hiraldo F. & Delibes M. (1997). Food habits of common Barn-owls along an elevational gradient in Andean Argentine Patagonia. Journal of Raptor Research, 31(1), 59–64. Yalden D.W. (2003). The Analysis of Owl Pellets. The Mammal Society, London.
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