A Comparative Study on the Nest Site Selection and Cavity Availability of the

Forest owlet Heteroglaux blewetti and other Sympatric Owls in Khandwa

District, Madhya Pradesh
A Dissertation Submitted to Pondicherry University in Partial Fulfilment of the
Requirement for the Award of the Degree of Masters in Ecology and
Environmental Sciences
By Akshay V Anand
Registration Number: 17371005
Under the Guidance of
Dr. S. Jayakumar
Professor
Department of Ecology and Environmental Science
Pondicherry University, Pondicherry
And
Dr. Prachi Mehta
Senior Scientist and Executive Director, Research
Wildlife Research and Conservation Society, Pune

Department of Ecology and Environmental Sciences

School of Life Sciences
Pondicherry University, Puducherry.
April, 2019

1
A Comparative Study on the Nest Site Selection and Cavity Availability of the
Forest owlet Heteroglaux blewetti and other Sympatric Owls in Khandwa
District, Madhya Pradesh
A Dissertation Submitted to Pondicherry University in Partial Fulfilment of the
Requirement for the Award of the Degree of Masters in Ecology and
Environmental Sciences
By Akshay V Anand
Registration Number: 17371005
Under the Guidance of
Dr. S. Jayakumar
Professor
Department of Ecology and Environmental Science
Pondicherry University, Pondicherry
And
Dr. Prachi Mehta
Senior Scientist and Executive Director, Research
Wildlife Research and Conservation Society, Pune

Department of Ecology and Environmental Sciences

School of Life Sciences
Pondicherry University, Puducherry.
April, 2019

2
Acknowledgements

This work was a team effort, several people have provided their encouragement and
suggestions without which the thesis would not have been completed successfully.

I thank

Dr. Prachi Mehta for her help in study designing as well as support in field work and timely
and useful suggestions and comments which helped me throughout the project.

Dr. S. Jayakumar, my faculty advisor for guiding me and supporting my autonomy during the
course of the study and encouraging me at all times.

Mr. Jayant Kulkarni for helping me design the study and for his constant inputs during the
study period.

Raptor Research and Conservation Foundation (RRCF) Mumbai for generously funding my
project.

Madhya Pradesh Forest Department for their permissions and logistic support during the
field work

My field assistant, Mr. Jabbar Chauhan for his loyalty and integrity during the field service as
well as his sleight of hand in the kitchen.

Mr. Narendra Yadav, deputy range officer, for his support in carrying out the field work.

Mr. Satyanarayanan Yadav, beat officer, for his constant interests and for all the healthy
discussions.

The villagers of Awalia for their hospitality as well as complete acceptance and trust in me
and my work.

Last but not the least my family and friends for their constant love, support and
encouragement.

3
 A Comparative Study on the Nest Site
Selection and Cavity Availability of
the Forest owlet Heteroglaux blewetti
Title
and other Sympatric Owls in Khandwa
District, Madhya Pradesh.

Name of the Student Akshay Vinod Anand

Faculty Supervisor Dr. S Jayakumar

Principal Investigator Dr. Prachi Mehta

Institute Pondicherry University.

Wildlife Research and Conservation

Supported By
Society, Pune, India.

Raptor Research & Conservation

Funded By
Foundation, Mumbai, India.

Field Support Madhya Pradesh Forest Department

4
Table of Contents Page
1. Introduction…………………………………………………………………………………………………… 8-13
1.1 Background of the Study…………………………………………………………………………. 8-9
1.2 Study Species……………………………………………………………………………………………. 9-12
1.3 Research Questions and Study Objectives………………………………………………… 13
2. Review of Literature………………………………………………………………………………………. 14-16
3. Methodology…………………………………………………………………………………………………. 17-23
3.1 Study Area………………………………………………………………………………………………. 17-18
3.2 Vegetation Quadrats………………………………………………………………………………… 18-19
3.3 Cavity Availability……………………………………………………………………………………… 19-21
3.4 Nest Box Acceptance………………………………………………………………………………… 22
3.5 Data Analysis……………………………………………………………………………………………. 23
4. Results…………………………………………………………………………………………………………… 24-32
4.1 Nest Site Selection between the three Owls……………………………………………. 24-30
4.2 Cavity Availability Between the Three Owls………………………………………………. 31-32
4.3 Nest Box Acceptance………………………………………………………………………………… 32
5. Discussion………………………………………………………………………………………………………. 33-36
5.1 Nest Site Selection of the Three Owls………………………………………………………. 34
5.2 Cavity Availability……………………………………………………………………………………… 35
6. Conclusion……………………………………………………………………………………………………… 36
7. References……………………………………………………………………………………………………... 37-40

5
List of Abbreviations
Km2 – Square Kilometer

°C – Celsius

m - Meter

amsl – Above Mean Sea Level

GBH – Girth at Breast Height

cm – Centimeter

SI – Suitability Index

HSI – Habitat Suitability Index

SPO – Spotted Owlet

FO – Forest Owlet

JO – Jungle Owlet

List of Tables Page

1. Mean, Standard Deviation, Standard Error, F and P values of measured Habitat

Variables……………………………………………………………………………………………………………. 30

List of Figures
1. Forest Owlet, Heteroglaux blewitti………………………………………………………………………. 10
2. Spotted Owlet, Athene brama……………………………………………………………………………… 11
3. Jungle Owlet, Glaucidium radiatum……………………………………………………………………… 12
4. Map of Study Area………………………………………………………………………………………………. 18
5. Map of Forest Owlet Nest Locations……………………………………………………………………. 20
6. Map of Spotted Owlet Nest Locations…………………………………………………………………. 21
7. Map of Jungle Owlet Nest Locations……………………………………………………………………. 21
8. Artificial Nest Box………………………………………………………………………………………………… 22
9. Spotted Owlet: Frequency Distribution of Tree Species………………………………………. 24

6
10. Forest Owlet: Frequency Distribution of Tree Species…………………………………………. 25
11. Jungle Owlet: Frequency Distribution of Tree Species…………………………………………. 25
12. Average Tree GBH………………………………………………………………………………………………… 26
13. Average Tree Height……………………………………………………………………………………………. 27
14. Average Shrub Cover……………………………………………………………………………………………. 27
15. Average Forest Floor Vegetation…………………………………………………………………………. 28
16. Average Canopy Cover…………………………………………………………………………………………. 28
17. Total Basal Area……………………………………………………………………………………………………. 29
18. Total Number of Available Cavities………………………………………………………………………. 31
19. Number of Suitable Cavities…………………………………………………………………………………. 32

7
1. Introduction

1.1 Background of the Study

Habitat is the sum of resources available for an organism (Thomas, 1979). These include
food, shelter, water and any other unique factor that is required by the species for its
survival (Leopold, 1933). When an animal shows a preference to a particular habitat type
then the animal is selecting the habitat rather than using habitats at random. Habitat use is
the proportion of their time that animals spend in a particular habitat. Selection of habitat is
the process by which an animal actually chooses habitat (Johnson, 1980). Use is considered
selective if habitat is used disproportionately compared with its availability, the latter being
the amount of that habitat accessible to the animal. In controlled environments, preference
is the likelihood of an animal selecting a given item when offered alternative choices on an
equal basis (Johnson, 1980). When the availability of habitat is varied, habitat preference is
the use of habitat relative to its availability in the environment.

Nest site selection in birds is a widely studied aspect in open nesting and cavity nesting
species. Quality of nest sites can be affected by microclimate, food availability, and nest
predation. Cavity orientation can determine internal microclimate effects of the nesting
cavity (Relier 1972, Conner 1975, Stauffer & Best 1982). Cavity-nesting birds commonly
choose nests in areas where foraging perches are dense (Mannan et al. 1980, Raphael and
White 1984, Swallow et al. 1986). However, density of live trees could also affect risk of nest
predation. Nest predation is usually the primary source of nest mortality for both open- and
cavity-nesting birds (Lack 1954; Nice 1957; Ricklefs 1969; Nilsson 1984). As a result, choice
of nest sites with reduced risk of nest predation and more foraging substrates should be
favoured. Probability of predation may decrease with increasing abundance of potential
nest sites (cavities) because predators must search more empty sites to find an occupied
site (Martin and Roper 1988). Therefore, higher the number of cavities the lower will be the
predation risk.

In India there have been a limited number of studies on the nest site selection of owls based
on habitat variables. The nest site selection of the Forest owlet (Heteroglaux blewitti) has
been studied in Toranmal Reserve Forest (Jathar & Rahamani, 2004, Ishtiaq & Rahmani
2005), in East Kalibhit forests, Madhya Pradesh (Mehta et al. 2017) and In Melghat Tiger

8
Reserve (Mehta et al ongoing). The nests sites of Spotted Owlet (Athene brama) has been
studied in Madurai district of Tamil Nadu (Santhanakrishnan et al, 2011), Andhra Pradesh
(Pande et al, 2006), East Kalibhit Forests in Madhya Pradesh (Mehta et al. 2017) and
Pakistan (Mahmood-ul-Hassan et al, 2007). Jungle Owlet nest site selection has been
studied in the East Kalibhit Range of Madhya Pradesh (Mehta et al 2017) In this study I
aimed at understanding the nest site selection of the Forest Owlet, Spotted Owlet and the
Jungle Owlet based on habitat variables in a 150m radius of the nest sites as well as cavity
availability around nest sites.

1.2 Study Species

The Forest Owlet is listed as endangered in the IUCN Red List of Threatened Species (Birdlife
International, 2019). Thought to be extinct for 113 years this elusive owl was historically
rediscovered in 1997 (Rasmussen & Collar, 1998). Post its rediscovery, the Forest Owlet has
been the centre of attention for many raptor researchers. A study on the vocalisation and
breeding biology of the Forest Owlet was conducted in the Taloda reserve forest
(Rasmussen & Ishtiaq, 1999). This study gave us the first insight into the behaviour and
ecology of this secretive owl. Following this was a study on the distribution and breeding
biology of the Forest Owlet in the Toranmal reserve forest. This study brought to light the
basic ecology of the Forest Owlet (Jathar & Rahmani, 2004). A further study on the
vocalisation of the Forest Owlet was conducted by Ishtiaq in the Toranmal reserve forest
yielding interesting insights into the vocal range and calling behaviour of the bird (Ishtiaq &
Rahmani, 2005). A five-state survey in 2005-2007, yielded new locations of the Forest Owlet
in Madhya Pradesh, these were in Khandwa district as well as in Burhanpur district (Mehta
et al, 2009). A few more new records of Forest Owlets were from Betul district in Madhya
Pradesh (Mehta et al 2015), Tansa Wildlife Sanctuary (Laad & Dagale, 2015) and Gujarat
(Patel et al. 2015). This discovery is an anomaly when it comes to the known habitat
preference of the Forest Owlet, as till this study the bird had been found mainly in teak
dominant, dry deciduous forests. A recent systemic grid wise survey of northern
Maharashtra yielded a new record of the Forest Owlet in the Navapur range of the
Nandurbar District (Mehta et al. 2017a). A comprehensive ecological study of Forest Owlet
was carried out in East Kalibhit Forests in Khandwa district of Madhya Pradesh that yielded
insights in to demography, diet, breeding ecology and threats on the endangered species

9
(Mehta et al. 2017b). The diet of the Forest Owlet, Spotted Owlet, and Jungle Owlet have
been compared in the East Kalibhit Range of Madhya Pradesh (Mehta et al. 2018) and their
nest site selection has been studied (Mehta et al in review).

Figure 1: Forest Owlet, Heteroglaux blewitti.

10
The Spotted Owlet (Athene brama) is widely distributed all over the subcontinent and south
east Asia. It is categorised as least concerned in the IUCN Red List of Threatened Species
(Birdlife International, 2019). The extent of its occurrence is estimated to be 10,800,000
km2. The breeding biology and nest site selection of the spotted have been studied in
Madurai district of Tami Nadu (Santhanakrishnan et al, 2011). The food habits and prey
spectrum of the Spotted Owlet have also been studied (Santhanakrishnan et al, 2011). In the
study area the Spotted Owlet is seen around human settlements and agricultural fields. It
breeds during February to April (Mehta et al, 2017b), in tree cavities that are formed
naturally by branch fall or natural excavators (Mehta et al, in review). They are crepuscular
in habit and are seen foraging mainly during dawn and dusk. They have a sharp call which
comprises of repetitions of a sharp “Chirr-Chirr”.

Figure 2: Spotted Owlet, Athene brama.

11
The Jungle Owlet (Glaucidium radiatum) is distributed over the Indian subcontinent, Bhutan,
Myanmar, Nepal and northern Sri lanka (Birdlife International, 2019). It is classified as least
concerned in the IUCN Red List of Threatened Species (Birdlife International, 2019). The
Jungle Owlet although widely distributed has been hardly studied. Only basic information on
its distribution and breeding season are available. Apart from the diet and nest sites of
Jungle Owlet in East Kalibhit forests (Mehta et al. 2017b, 2018), there have been no other
studies on the breeding biology, nest site selection, diet and habit of the Jungle Owlet. In
the study area the Jungle Owlet is seen in forest patches or in forest borders. They breed
between April and June in tree cavities formed by branch fall or by natural excavators. The
calls of the Jungle Owlet have been described by Neelakantan in the Journal of the Bombay
Natural History Society in 1971 (Neelakantan, 1971).

Figure 3: Jungle Owlet, Glaucidium radiatum.

12
1.3 Research Questions and Study Objectives
This study was motivated by a need to understand the nest site selection of these three
sympatric owls in the rapidly changing landscape of Khandwa District, Madhya Pradesh. The
inspiration of the study raised from the varied distribution of the study species and the
endemism of the Forest Owlet. The study area is one of the few locations where the three
owl species nest in close proximity with each other. This made it an idea location to study
the nest site selection and habitat preference of these three species and compare them
among one another.

Cavity availability was another aspect of interest in nest site selection. A previous study on
the preferred cavity dimensions of these three species was carried out in the study area
(Mehta et al. 2017b, Mehta et al. in review) which furthered the need to understand the
availability of suitable cavities for nesting within the study area. Nest box acceptance was
also tested as the study area is managed for timber logging by the Madhya Pradesh Forest
Department. Timber logging could affect the number of suitable cavities for nesting for the
three study species and hence as a conservation strategy nest box acceptance was tested.

The objectives of the study were three-fold

1. To compare nest site selection between the Spotted Owlet, Forest Owlet and Jungle
Owlet based on habitat variables surrounding nest sites.
2. To understand cavity availability and suitability of available cavities as a factor in nest
site selection of the three study species.
3. To test for nest box acceptance of the three study species.

13
2. Review of Literature
There have been only a handful of studies on nest site selection of owls around the world.
The Northern Spotted Owl (Strix occidentalis caurina) is probably one of the most studied
species of owl around the world. A study in North Western California on the roost and nest
site selection of the Northern Spotted Owl (Blakesley et al, 1991) studied habitat use in
relation to habitat availability during the breeding season. The study found that the Spotted
Owl used large diameter conifer trees in higher proportion than they were available. Stands
dominated by smaller hardwood trees or conifers were used less than, or in proportion to
their availability. The owls avoided the upper third of slope for both nesting and roosting.
These results were in compliance with results of earlier studies on the Northern Spotted
Owl who used radio tagged owls to assess habitat selection (Forsman et al. 1984, Carey et
al. 1990, Sisco 1990, Solis & Gutikrrez 1990).

A study on the nest site selection of the Great Horned Owl (Bubo virginianus) in New York,
New Jersey and Connecticut in the United States of America compared nest sites and
habitat selection between rural and urban nest sites (Smith et al, 1999). The habitat use was
compared with available habitat in both categories of the nest sites. The study found that
urban nest trees were significantly larger in diameter and taller than rural nest trees, and
accordingly, nests were higher in urban nest trees as well. Both urban and rural Great
Horned Owls demonstrated habitat selection. Although urban owls showed a stronger
degree of selection, probably because of the more complex nature of habitats available in
the urban areas.

Another study in Kentucky on Eastern Screech-Owl (Otus asio) nests compared used nest
sites to randomly chosen unused nest sites (Belthoff & Ritchson, 1990). This was done to
determine which features of the nest tree or cavity and surrounding vegetation influenced
nest-site selection. Eastern Screech-Owls selected nest sites based on the depth of the
cavity, cavity height and entrance size. Neither tree species nor entrance orientation of the
cavity hole were important in nest selection. Similarly, nest-site vegetation parameters were
not significant in nest-site selection. They theorised that if, suitable cavities are limited in
supply, unsuitable cavities reduce protection from predators and decrease reproductive
success, then the availability of suitable cavities can limit Eastern Screech-Owl populations.

14
Nest site selection has been studied for the Forest Owlet in Toranmal Reserve Forest,
Maharashtra (Jathar & Rahmani, 2004, Ishtiaq & Rahmani, 2005) and in East Kalibhit Range
of Khandwa District of Madhya Pradesh (Mehta et al, 2017b, Metha et al in review). The
above studies found that the Forest Owlet was preferring nest sites in Teak (Tectona
grandis) dominated forest stands.

The breeding biology and nest site selection of the Spotted Owlet have been studied in
Andhra Pradesh (Pande et al, 2006), south India (Santhanakrishnan et al. 2011) in east
Kalibhit forests (Mehta et al. 20017b) and Pakistan (Mahmood-ul-Hassan et al, 2007) all
these studies focused on nest tree and nest cavity parameters and did not quantify the
vegetation surrounding nest sites.

The nest site selection of the jungle owlet in the East Kalibhit forests of Madhya Pradesh
showed that it was preferring nest sites in old stands of forests with tall trees. The Jungle
owlet in East kalibhit forests also preferred areas of highest canopy cover and intermediate
forest floor vegetation as compared to the Forest Owlet and Jungle Owlet (Mehta et al. In
Review).

Nest box acceptance of the Spotted Owlet was studied in Punjab (Kumar, 2012). Here a total
of 62 nest boxes were set up in four habitat types including agricultural fields, agricultural
fields with tree line, small vegetable fields and agroforestry areas. The dimensions of the
nest boxes were fixed. Out of the total number of nest boxes deployed only 10 were
occupied by spotted owlet pairs.

The sampling protocol used was the tenth acre method. Tenth acre (0.04 ha) (Li & Martin,
1991) methods of vegetation sampling are used regularly in several studies on nest success
of birds. Tenth acre sampling method has been suggested to be most efficient in its ability to
predict the actual density of trees at the end of a specified period of field effort amongst
several other measures like the quarter method, wandering quarter method and hundredth
acre rectangle method (James & Shugart, 1970).

There are large research gaps in the nest site selection based on habitat variable in studies
on owls in India. Further cavity availability and nest box acceptance have rarely been
studied in the country. This study aims to work towards filling these research gaps and set
the course for further research on these unexplored aspects.

15
Wildlife Research and Conservation Society (WRCS) has been working on the ecology of Forest Owlet
and other owls in East Kalibhit Range in Khadnwa District of Madhya Pradesh since 2013 (Mehta et
al. 2017b). I chose to carry out my M. Sc dissertation in the same area considering my previous work
in WRCS on the project as it provided me basic understanding of the species ecology in the area.

16
3. Methodology.

3.1 Study Area

The study was carried out from December 2018 to April 2019. The study area was East
Kalibhit Reserved Forests (21.8887, 76.9673) located in Khandwa district of Madhya Pradesh
(Figure 4). It extends for 40 km2 from the village of Awalia in the west to Jhinjhri in the east.
The area was characterised by Teak (Tectona grandis) dominant dry deciduous forest. The
weather conditions are widely varied with hot dry summers, temperatures sometimes
exceeding 48°C, and cold dry winters, minimum temperatures of 7°C. Monsoons are from
the middle of June to the middle of September, with average rainfall of 770 mm. The
elevation ranges from 300-700 m amsl (in field measurements). The region is characterised
by gently sloping terrain with moderate to high altitude hills. The soil type is classified as
medium black soil with alluvial deposits of clay and silt along river basins which snake their
way through the region. Inside the study area there are three tribal villages of moderate size
with populations of 800-1000. Due to their presence, a considerable portion of this area was
under cultivation. Most of the cultivated area is centred around the habited area and hence
there are still large tracts of undisturbed forest area. Overall the area appears as a mosaic of
forest area interspersed with agricultural lands. The forest cover ranges from agricultural
open forest to dense deciduous forest. Predominantly the area is Teak (Tectona grandis)
dominant other tree species include Gakhad (Bridelia retusa), Palas (Beautea monosperma),
Sajad (Terminalia tomentosa), Arjun (T. Arjuna) Salai (Boswellia serrata), Mouha (Madhuca
latifolia), Tendu (Diospyros melanoxylon) and Amla (Phyllanthus emblica).

17
Figure 4: Map of the Study Area.

3.2 Vegetation Quadrats

To understand nest site selection based on the surrounding habitat I collected vegetation
data for 11 nest sites of Forest Owlet, Spotted Owlet and Jungle Owlet that were detected
during the previous study (Mehta et al, 2017b). This was done by plotting one 0.04 ha
square quadrat (James & Shugart, 1970) centred around the nest tree (Belthoff and
Ritchson, 1990). An additional 4 satellite quadrats were plotted at a distance of 150m from
the nest tree in all four cardinal directions. Inside each quadrat we recorded tree species,
tree GBH and height of all trees greater than 10 cm in GBH (Girth at Breast Height). To
calculate the density of shrubs within the quadrat we used the line intercept method (James
& Shugart, 1970). In this we plotted two perpendicular transects, from the north to south
and east to west corners of the plot and measured the intercepts of all woody stems
touching the transect that were less than 10 cm in GBH. Total shrub percentage was
calculated for each quadrat, from which shrub density was obtained. Canopy cover was
measured using a 96-point densitometer. Readings were taken in all four corners of each

18
quadrat. Average canopy percentage was calculated for each nest site and pooled for each
species to get total canopy percentage for the nest sites of each of the study species. We
also quantified ground cover. This was done by walking 100 steps in each of the cardinal
directions classifying each step into vegetation and bare soil. From this we calculated
average percentage of ground cover surrounding each active nest.

All quadrat data was pooled for each of the study species and the pooled data was analysed
and compared between the three species. Descriptive statistics and inferential statistical
tests were performed to test for significant differences in the collected data.

3.3 Cavity Availability

To understand the relationship between nest site selection and the availability of cavities I
carried out a count of cavities around active nest locations of all three species. The study
was carried out from the beginning of February 2019 to the end of March 2019. A total of 11
active nests of each species were identified in the study area between the period of 2013-
2017 (Mehta et al, 2017b, Mehta et al, in review) (Figure 5,6 & 7). A cavity was considered
as active if egg laying and incubation were carried out within the cavity. These 33 nests were
considered for my study. To assess the availability of cavities around the nest site a 200 m
square buffer was plotted around each of the nest trees of each species. This was done
using QGIS Ver. 2.14.9 Essen. Belt transects of 400x20 m were carried out within the
buffered area. Twenty such belts were sampled around each nest accounting for 100 %
sampling to obtain a total count of available cavities in the vicinity of the nest sites. Within
each belt visual scanning of each tree was carried out to locate cavities. Visual scanning was
carried out by two observers on either side of the belt transect at a distance of 10m from
one another. Once cavities were located the tree was climbed and the following cavity
parameters were measured for each located cavity (Belthoff and Ritchson, 1990, Mehta et
al. 2017b)

1. Nest tree species

2. Nest tree Girth at Breast Height (GBH) (cm)
3. Nest tree height (m)
4. Height of the cavity from the ground (m)
5. GBH at the cavity (cm)

19
 6. Height of the cavity entrance/opening (cm)
7. Width of the cavity entrance/opening (cm)
8. Internal diameter of the cavity (cavity opening to back wall) (cm)
9. Internal depth of the cavity (top to bottom) (cm)
10. Orientation of the cavity in degrees
11. Altitude of the nest tree (amsl)
12. Presence of a branch adjacent to the cavity opening

Figure 5: Forest Owlet Nests Locations

20
Figure 6: Spotted Owlet Nest Locations

Figure 7: Jungle Owlet Nest Locations

21
3.4 Nest Box Acceptance

Artificial nest boxes were constructed using the preferred nesting parameters of each
species (Mehta et al, in review). These were constructed to be slightly varied from one
another in their dimensions based on the range of nesting characteristics of the bird. The
nest boxes were set up within the home range of 3 breeding pairs of each species. Each pair
was observed to see if they are preferring the artificial nest boxes to naturally available
cavities. This was done in one-week intervals for each pair. Camera traps were also set up at
each nest box location to check for use of the nest boxes. These were checked with a three-
day interval. The nest boxes were set up at 3 Forest Owlet locations by the beginning of
January 2019. Six nest boxes were set up in three Spotted Owlet and three Jungle Owlet
locations respectively by the end of January 2019. All nest boxes were left till the end of
March 2019. Figure 8 shows the artificial nest box deployed in the field.

Figure 8: Artificial nest box

22
3.5 Data Analysis

3.5.1 Vegetation Data

Descriptive statistics were carried out on the raw data. Firstly, the data was pooled per
species and then was analysed. Mean, Standard Deviation and Standard Error was
calculated for each of the measured parameters using Microsoft Excel 2019. The inferential
statistical test used to test for significant difference in means between the habitat variables
of the three study species was a one-way Analysis of Variance (ANOVA). This was done by
grouping the variables per species and testing them for each variable separately. ANOVA
was carried out on R software version 3.5.3.

3.5.2 Cavity Suitability

To test for suitability of cavities the Habitat Suitability Index (Oldham et al, 2000) was used.
In this an index of suitability for each cavity was created and ranked on a scale of 0.1-1, 1
being the optimum value. Each measured parameter (except tree species, orientation,
altitude and presence of branch) was ranked from 0.1-1 this is known as a Suitability Index
(SI). A total of 8 SI’s were used to test for suitability. This was performed for each cavity
located. To calculate overall suitability of the cavity for nesting the following formula was
used:

HSI= (SI1XSI2XSI3X. . . . SIN)1/N

This yielded the number of suitable cavities available for nesting, from the total cavities
available, surrounding nest sites of each species. This was then compared between each
species to test if total number of cavities and total number of suitable cavities were differing
between the three study species.

3.5.3 Map Creation

All Maps were created on QGIS version 3.14.9.

23
4. Results

4.1 Nest Site Selection between the three Owls

A total of 119 vegetation quadrats were plotted during the study (Spotted Owlet=43, Forest
Owlet=43, Jungle Owlet=33). Teak (Tectona grandis) was the dominant species within all
quadrats with and average value of 59.17% of the total number of trees measured, this was
followed by Wrightia tinctoria with 8.81% coverage. The average woody stem density was
87.93 stems/acre, with the Jungle Owlet having the highest woody stem density of 102.42
stems/acre.

Tree Species

There was a total of 32 tree species recorded from all the 119 plots. The maximum number
of tree species was found in the Spotted Owlet nest sites (23 species). Teak was dominant in
all nest sites with the Jungle Owlet having the highest teak percentage of 59.17%.

Figure 9: shows the number of stems of each tree species recorded around the nest sites of
the Spotted Owlet. Tectona grandis was dominant with 57.83% followed by Wrightia
tinctorial with 16.41%.

Spotted Owlet
229
Number of Stems

65

17 15 14 13 7
5 5 3 3 3 3 2 2 2 1 1 1 1 1 1 1 1

24
Figure 10: shows the number of stems of each species recorded around the Forest Owlet
nest sites. Here again Tectona grandis was the dominant species with 54.03% this was
followed by Wrightia tinctorial with 8.38%.

Forest Owlet
161
Number of Stems

25 20
14 13 8 7 7 7 6 5 5 5 3 3 2 2 2 1 1 1

Figure 11: shows the number of stems of each tree species in the vicinity of Jungle Owlet
nest sites. Tectona grandis was dominant here as well with 59.17%, this was followed by
Lagerstromeia parviflora with 4.14%.

Jungle Owlet

200
Number of Stems

22 14 14 14
11 9 8 8 6 5 5 4 4 3 2 2 2 1 1 1 1 1

25
Tree GBH:

The mean GBH of the nest sites of the Forest Owlet were the highest (71.95 cm ± 41.68),
followed by the Jungle Owlet (66.86cm ± 34.62) and the Spotted Owlet (58.77cm ± 37.47)
(Figure 12).

Figure 12: Average tree GBH.

80

70

60

50
Spotted Owlet
40
Forest Owlet
Jungle Owlet
30

20

10

0
TREE GBH

Tree Height:

The Forest Owlet selected nest sites with tallest trees (8.62m ± 3.23). The Jungle Owlet
preferred stands of forest with intermediate height of trees (8.22m ± 2.79) when compared
to the other two study species. The Spotted owlet had selected stands of forest with the
shortest trees (7.29m ± 3.57) (Figure 13).

26
 Figure 13: Average Tree Height

10

6
Spotted Owlet
5
Forest Owlet
4 Jungle Owlet

0
TREE HEIGHT

Shrub Cover:
With regards to shrub cover surrounding the nest site the Jungle Owlet selected areas with
highest shrub cover of 7.28%, this was followed by the Spotted Owlet with an average shrub
cover percentage of 6.7%. The Forest Owlet showed preference for stands of forest with
least shrub cover with 4.93% (Figure 14).

Figure 14: Average Shrub Cover

5 Spotted Owlet
Forest Owlet
4
Jungle Owlet
3

0
SHRUB COVER

27
Forest Floor Vegetation: There did not exist a large variation in the forest floor vegetation
among the nest sites of the three study species (Figure 15). The Jungle Owlet showed the
highest forest floor vegetation percentage with 65.62%, followed by the Forest Owlet
(60.35%) and the Spotted Owlet (56.85%).

Figure 15: Average Forest Floor Vegetation

80

70

60

50
Spotted Owlet
40
Forest Owlet

30 Jungle Owlet

20

10

0
FOREST FLOOR VEGETATION

Canopy Cover: The canopy cover for the Jungle Owlet was the highest with 76.67%. This
was followed by the Forest Owlet with 68.65% and the Spotted Owlet with 63.41% canopy
cover (Figure 16).

Figure 16: Average Canopy Cover

90

80

70

60

50 Spotted Owlet
Forest Owlet
40
Jungle Owlet
30

20

10

0
CANOPY %

28
Basal Area:
The basal area was calculated from the GBH of trees for each of the study species.
Therefore, the trend was similar to that of the tree GBH mentioned above, with the Forest
Owlet showing highest basal area of 18.11 m2/ha. This was followed by the Spotted Owlet
with a total basal area of 15.34m2/ha. The total basal area of the Spotted Owlet was not
much varied from that of the Jungle Owlet, with a value of 15.24m2/ha.

Figure 17: Total Basal Area.

18.50

18.00

17.50

17.00

16.50
Spotted Owlet
16.00
Forest Owlet
15.50 Jungle Owlet

15.00

14.50

14.00

13.50
TOTAL BASAL AREA

29
 One-way Analysis of Variance (ANOVA):
The Results of the one-way ANOVA showed that three of the six measured habitat
parameters were significantly different from one another. The Forest Owlet preferred nest
sites with trees of highest GBH (P < 0.05) and with the tallest trees (P < 0.05). The canopy
cover percentage was the highest for the Jungle Owlet (P < 0.05). The other three measured
parameters i.e. shrub cover, forest floor vegetation and basal area, were not significantly
different (P > 0.05). The Results of the ANOVA are given in Table 1.

Table 1: Table 1: Mean (X), Standard Deviation (SD), Standard Error (SE), F and P values of
measured Habitat Variables.
SPO FO JO
Parameters
X SD SE X SD SE X SD SE F P
Tree GBH
58.77 37.47 1.88 71.95 41.68 2.88 66.86 34.62 1.88 9.384 0.00009144
(cm)
Tree Height
7.29 3.57 0.17 8.62 3.23 0.19 8.22 2.79 0.15 15.71 0.00000019
(m)
Shrub Cover
6.7 4.32 0.66 4.93 3.31 0.5 7.28 4.66 0.81 2.964 0.0555
%
Forest Floor
Vegetation 56.85 22.81 3.48 60.35 23.09 3.52 65.62 20.34 3.54 1.45 0.2388
%
Canopy
63.41 37.35 2.85 68.65 31.34 2.39 76.67 27.42 2.39 6.163 0.0023
Cover %
Basal Area
0.039 0.043 0.002 0.046 0.111 0.006 0.038 0.044 0.002 1.271 0.281
(m2)

30
4.2 Cavity Availability Between the Three Owls

The second objective of my thesis was to understand the relationship between the
availability of cavities on the nest site selection of the three study species. The total count of
cavities was the highest for the Forest Owlet (34 cavities). The Spotted Owlet and Jungle
Owlet had a total of 14 and 12 cavities around their nest sites, respectively. Figure 18 shows
the total number of cavities surrounding the nest sites of all three study species.

Figure 18: Total number of available cavities surrounding nest sites of each of the study
species.

40
34

35

30

25
SPO
20
FO
JO
14

15
12

10

0
AVAILABLE CAVITIES

The suitability of each cavity for species specific nesting was carried out to understand the
portion of available cavities that were suitable for nesting for each of the study species. This
yielded interesting results, in which out of the total 34 cavities in the vicinity of Forest Owlet
nest sites, 12 cavities were suitable for Forest Owlet nesting. This was considerably higher
than the number of suitable cavities of the other two study species. The Jungle Owlet had
only three suitable cavities and the Spotted Owlet had no suitable cavities. Figure 19 shows
the number of suitable cavities of the three study species.

31
 Figure 19: Number of suitable cavities of the three study species.

14

12
12

10

8 SPO
FO
6
JO

3
2
0

0
SUITABLE CAVITIES

4.3 Nest Box Acceptance

The final objective of my thesis was to test for nest box acceptance between the three
species. A total of 9 nest boxes were deployed in the field, three for each study species.
Unfortunately, all 9 nest boxes were rejected by the study species. Camera trap footage
confirmed the rejection of all deployed nest boxes.

32
5. Discussion

The nest site selection of the Forest Owlet, Spotted Owlet and the Jungle Owlet in the East
Kalibhit Reserved Forests in Khandwa district of Madhya Pradesh has been studied earlier
(Mehta et al 2017b). A comparative study has been conducted between these three
sympatric owls based on cavity dimensions and vegetation within a 20m radius of the nest
tree (Mehta et al, in review). In this study I examined the vegetation within an 150m radius
from the nest tree and compared them between the three species.

The study in the Toranmal Reserve Forest in Maharashtra showed that the nesting habitat of
Forest Owlet had Tectona grandis as the dominant species followed by Lagerstroemia
parvifolia, Anogeissus latifolia, Butea monosperma, Acacia catechu, Diospyros melanoxelon,
Boswellia serrata, Bombax ceiba, Soymida febrifuga, Adina cordifolia, Dalbergia sissoo and
Lania grandis Ishtiaq & Rahmani, 2005). An earlier study on the habitat use of the Forest
Owlet in the Toranmal Reserve Forest yielded similar results, with Tectona grandis being the
most dominant tree species in occupied nesting sites, this was followed by Anogeissus
latifolia, Lagerstroemia parvifolia, Ougenia oogenensis, Butea monosperma, Boswelia
serrata and Wrightia tinctorial (Jathar & Rahmani, 2004). In my study as well Tectona
grandis accounted for 54.03% of the total number of stems in the Forest Owlet nest sites.
Other abundant species were, Wrightia tinctorial, Lagerstroemia parvifolia, Terminalia
tementosa, and Diospyros melanoxelon.

A study on the nest site selection of the Spotted Owlet showed that the bird was preferring
sites with Ficus benghalensis, Enterolobium saman, Cocos nucifera and Tamerindus indicus,
in the Madurai District of Tamil Nadu (Santhanakrishnan et al, 2011). Another study in
Andhra Pradesh showed that the spotted Owlet was using nest locations with Ficus
religiosa, Tamarindus indicus, Mangifera indica, Ficus benghalensis, Moringa oleifera and
Melia azaderach (Pande et al, 2006). In Pakistan showed that the Spotted Owlet was
preferring nest sites with Dalbergia sissoo, Acacia nilotica, Butea monosperma,
Ficus benghalensis, Mangifera indica and Eucalyptus citriodora (Mahmood-ul-Hassan et al
2007). The results of the present study showed that the spotted Owlet was preferring nest
locations in which Tectona grandis was the dominant species, accounting for 57.83% of total
stems, this was followed by Wrightia tinctorial, Lagerstroemia parvifolia, Butea

33
monosperma and Diospyros melanoxelon. The variation in species composition between the
three species could be due to the regional species variation. That is to suggest that since the
Spotted Owlet is widely distributed over India and South east Asia the bird has adapted to
different habitat types with different species composition.

The present study showed that the Jungle Owlet was preferring nest sites in Tectona grandis
dominant forest stands, accounting for 59.17% of total woody stems. This was the highest
among the three study species.

The current study also quantified shrub cover percentage, forest floor vegetation and
canopy cover percentage within a 150m radius of the nest sites of each study species. These
factors were not considered for any of the previous studies.

5.1 Nest Site Selection of the Three Owls

The two studies in Toranmal Reserve Forest along with my study showed that the Forest
Owlet was selecting Teak dominant forest stands for nesting. The Forest Owlet being an
endemic species to central India is not as widely distributed as the Spotted Owlet and the
Jungle Owlet. As the Forest Owlet is preferring nest sites which are dominant in teak the
distribution of the species could be limited by this factor.

The earlier studies on the Spotted Owlet show a varied tree species composition among the
three study sites. The Spotted Owlet is also more widely distributed over the Indian sub-
continent as well as south east Asia occupying a wide range of habitat types. This speaks
towards the adaptability of the Spotted Owlet to varied habitats. This can be substantiated
by comparing the results of my study to the results of the earlier studies, which all show a
different tree species preference.

The present study showed that the Jungle Owlet was preferring areas with higher shrub
cover and higher forest floor vegetation. The jungle owlet feeds mainly on insects and other
invertebrates from the forest floor and the canopy (Mehta et al. 2018). Their diet includes a
relatively lower number of small mammals as compared to the other two study species
(Mehta et al, 2018). This could be a possible reason as to why the Jungle Owlet is selecting
nest sites with higher shrub cover and forest floor vegetation as the density of invertebrates
might be higher in these locations. The Forest Owlet and Spotted Owlet on the other hand

34
have a larger portion of small mammals in their diet as compared to the Jungle Owlet
(Mehta et al, 2018). This could be a reason why they are selecting nest sites with lower
shrub cover and forest floor vegetation in which small mammals would be more visible on
the forest floor.

The one-way Analysis of Variance showed that three of the measured parameters were
significantly different between the three owls. Therefore, the nest site assessment showed
that the Forest Owlet is preferring older stands of forest with taller trees and intermediate
canopy cover. The Jungle Owlet selected nest sites in middle aged forest stands with
medium sized trees and highest canopy cover, whereas the Spotted Owlet selected young
forest stands with shortest trees and lowest canopy cover.

5.2 Cavity Availability

The presence of cavities in the vicinity of the nest sites are of paramount importance to the
nest site selection of these three sympatric owls. Apart from nesting the three study species
use cavities for caching of prey and protection from avian predators (Akshay V Anand,
personal observation). The present study showed that there does exist a dependence on
presence of cavities and nest site selection of all three owls. The Forest Owlet showed the
highest number of cavities around the nest site as compared to the other species. Of these
41% proved to be suitable for Forest Owlet nesting, which was much higher than the other
two species. The Forest Owlet has very specific nesting requirements (Mehta et al in
review). This could be a factor as to why the Forest Owlet are selecting location where there
are more suitable cavities for nesting as small changes in the cavity dimensions, natural or
anthropogenic, could alter the suitability of the cavity. Therefore, the availability of suitable
cavities around Forest Owlet nest sites are a factor in its nest site selection.

The study showed that there does exist a relationship between available cavities and nest
site selection. Therefore, I assumed that increasing the number of available cavities around
the nest sites would aid the owls in selecting suitable nests. This was done by setting up
artificial nest boxes in the home ranges of breeding pairs. The nest boxes were deployed for
a period of three months. They were designed to be suitable for each of the owl species
(Mehta et al. In Review). Unfortunately, none were occupied by any of the three owl
species. This could be because the nest boxes were set up only for a small period of time

35
and it takes the owls a longer time to get used to the artificial nest box. This can only be
examined through long term projects which study nest box acceptance of the three owl
species.

6. Conclusion

The present study has not only shown the nest site selection based on habitat preference of
the three study species but also the importance of cavities and cavity bearing trees in the
nest sites of the three species. The study area is managed for timber logging by the Madhya
Pradesh Forest Department. In a managed forest the Forest Department must take
adequate steps to exempt cavity bearing trees from felling. Further in the study area illegal
tree felling by local tribes is rampant. This is a disturbance to nesting birds not only because
cavity bearing trees are being cut but also due to the constant human presence in the
nesting sites. Timber logging by the Forest Department is also a disturbance as mud roads
are made deep inside the forest area and large trucks filled with logged wood are carried
through the forest. These are major disturbances to cavity nest birds. The Forest Owlet is
very specific in its nest site selection (Mehta at al. in review). Therefore, the protection of
their nest sites is required throughout their home range. Major disturbances like timber
logging and illegal felling of trees have to be monitored in and around their nest sites.

Long term monitoring of the nest sites and effective monitoring of breeding pairs of these
three species of owl will yield a better understanding on their nest site selection and
breeding biology. The clumped distribution of the Forest Owlet needs to be further explored
through large scale systematic grid surveys to understand the species total range
distribution. Comparative studies need to be carried out in different locations where these
three species are nesting in close proximity to understand differences in breeding success
and nest site selection in different habitat types. In depth research and analysis is required
to be carried out on the habitat selection, nest site preference, breeding biology and range
distribution of the Forest Owlet in order to formulate effective conservation action plans.
This can be achieved by increasing awareness on the status of the species and generating
interest on varied research questions that for be studied on the species.

36
7. References
Belthoff, J. R., And G. Ritchison. 1989. Natal dispersal of Eastern Screech-Owls. Condor
91:254- 265.
Belthoff, J. R., And G. Ritchison. 1990. Roosting behaviour of post fledging Eastern Screech-
Owls. Auk 107:567-579.
Belthoff, J. R., Richson, G. Nest-site selection by Eastern Screech-Owls in Central Kentucky.
1990. The Condor 92:982-99.
BirdLife International (2019) IUCN Red List for birds. Downloaded from
http://www.birdlife.org on 28/04/2019.
Blakesley, J. A. 1991. Spotted Owl Roost and Nest site Selection in north-western California.
Journal of Wildlife Management. 56(2):388-392.
Bolan, J., Sinclair, R., 2014. Developing age-size relationships for long lived tree species.
Journal of Biological Systems 22(2):309–326. 10.1142/S0218339014400087.
Brush, T. 1983. Cavity use by secondary cavity- nesting birds and response to manipulations.
Condor 85:461-466.
Carey, A. B., J. A. Reid, & S, P. Horton. 1990. Spotted owl home range and habitat use in
southern Oregon coast ranges. Journal of Wildlife Management. 54: 1 l-17.
Conner, R. N. (1975). Orientation of entrances to woodpecker nest-cavities. Auk 92: 371-
374.
Devereux, J. G., And J. A. Mosher. 1984. Breeding ecology of Barred Owls in the central
Appalachians. Raptor Research 18:49-58.
Forsman, E. D. & H. M. WIGHT. 1984. Distribution and biology of the spotted owl in Oregon.
Wildlife Monographs. 87: l-64.
Forsman, E. D., E. C. Meslow, And H. M. Wight. 1984. Distribution and biology of the Spotted
Owl in Oregon. Wildlife Monographs. 87: 3-64.
Goad, M. S., R. W. Mannan. 1987. Nest site selection by Elf Owls in Saguaro National
Monument, Arizona. Condor 89:659-662.
Goodenough, A.E., Maitland, D. P., Hart, A. G., & Elliot, S. L. 2008. Nestbox orientation: a
species-specific influence on occupation and breeding success in woodland passerines. Bird
Study 55: 222-232.DOI: 10.1080/00063650809461526.
H. Van Balen, J & J. H. Booy, C & Van Franeker, Jan & R. Osieck, E. (2002). Studies on Hole-
Nesting Birds in Natural Nest Sites. Ardea. 38-90. 1-24. 10.5253.
Ishtiaq, F. & Rahmani, A. R. 2005. The Forest Owlet Heteroglaux blewitti: vocalization,
breeding biology and conservation. Ibis 147: 197–205.

37
James, F. C., & Shugart Jr., H. H. (1970). A quantitative method of habitat description.
Audubon Field Notes, 24 (6), 727-736.
Jathar, G. A., & Rahmani, A., 2004. Ecological studies of the Forest Spotted Owlet Athene
(Heteroglaux) blewitti. Final report. Bombay Natural History Society, Mumbai, India. Pp. 1–
77.
Johnson, D. H. (1980). The Comparison of Usage and Availability Measurements for
Evaluating Resource Preference. USGS Northern Prairie Wildlife Research Centre. 198.
Jones, J. 2001. Habitat Selection Studies in Avian Ecology: A Critical Review. Auk 118:557-
562.
Kler, T. K. & Kumar, M. 2012. Breeding of Spotted Owlet Athene brama in Nest Boxes and
Conservation Aspects. Journal of the Bombay Natural History Society, 109(1 & 2).
Laad, Sunil & Dagale, Rohidas. (2014). First Report of Forest Owlet Heteroglaux blewitti from
Tansa Wildlife Sanctuary (Western Ghats), Maharashtra, India. Bombay Natural History
Society (BNHS). 111. 134. 10.17087/jbnhs/2014/v111i2/72238.
Lack, D. 1954. The natural regulation of animal numbers. Oxford, Clarendon Press.
Leopold, A. (1933). Game Management. Charles Scribner’s Sons, New York
Li, P., & Martin, T. E. (1991). Nest-Site Selection and Nesting Success of Cavity-Nesting Birds
in High Elevation Forest Drainages. The Auk, 405-418.
Mahmood-ul-Hassan, M., Beg, M.A., Mushtaq-ul-Hassan, M., Rana, S.A. (2007). Nesting and
breeding habits of the Spotted Owlet (Athene brama) in Punjab, Pakistan. Journal of Raptor
Research, 41(1), pp 50-52.
Mannan, R. W., E. C. Meslow, & H. M. WIGHT. 1980. Use of snags by birds in Douglas-fir
forests, western Oregon. Journal of Wildlife Management. 44: 787-797.
Martin, T.E., & J. J. Roper. 1988. Nest predation and nest site selection of a western
population of the Hermit Thrush. Condor 90: 51-57.
Mehta P., Kulkarni, J., & Patil, D., 2008. A survey of critically endangered Forest Owlet
Heteroglaux blewitti in Central India. Birding ASIA 10: 77–87.
Mehta P., Kulkarni, J., Patil, D., Kolte, P., & Khatavkar, P., 2007. A survey of critically
endangered Forest Owlet Heteroglaux blewitti in 5 states of India. Envirosearch. Final
Report submitted to the Ministry of Environment and Forests, New Delhi.
Mehta, P., & Kulkarni, J., Mukherjee, S., Chavan, S., Anand, A. V. 2017a. A distribution survey
of the Forest Owlet Heteroglaux blewitti in north-western Maharashtra. Indian BIRDS 13
(4):103-108.
Mehta, P., Kulkarni, J., Talmale, S. & Chandarana, R. 2018. Diets of sympatric Forest Owlets,
Spotted Owlets, and Jungle Owlets in East Kalibhit Forests, Madhya Pradesh, India. Journal
of Raptor Research. 52: 338-348.

38
Mehta, P., Prasanna N. S., Nagar, A. K., & Kulkarni, J., 2015. Occurrence of Forest Owlet
Heteroglaux blewitti in Betul District, and the importance of its conservation in the Satpura
landscape. Indian BIRDS 10 (6): 157–159.
Neelakantan, K. K (1971). Calls of the Malabar Jungle Owlet (Glaucidium radiatum
malabaricum). Journal of the Bombay Natural History Society 68 (3): 830–832.
Nice, M. M. 1957. Nesting success in altricial birds. Auk 74: 305-321.
Nilsson, S. G. 1984. The evolution of nest-site selection among hole-nesting birds: the
importance of nest predation and competition. Ornis Scandinavica 15: 165-175.
Nilsson, S. G. 1986. Evolution of hole-nesting in birds: On balancing selection pressures. Auk
103: 432-435.
Oldham R.S., Keeble J., Swan M.J.S. & Jeffcote M. (2000). Evaluating the suitability of habitat
for the Great Crested Newt (Triturus cristatus). Herpetological Journal 10(4), 143-155.
Pande, S., Pawashe, A., Mahajan, M.N., Mahabal, A. (2006). Changing nest site preference
for holes in earth cuttings in Spotted Owlet Athene brama. Indian Birds, 2(1), pp 78.
Patel, J. R., Patel, S. B., Rathor, S. C., Patel, J. A., Patel, P. B., & Vasava, A. G., 2015. New
distribution record of the Forest Owlet Heteroglaux blewitti Hume, 1873, (Aves:
Strigiformes: Strigidae) in Purna Wildlife Sanctuary, Gujarat, India. Journal of Threatened
Taxa 7 (12): 7940–7944.
Purcell, K. L., Verner, J, And Oring, L. W. 1997. A comparison of the breeding ecology of birds
nesting in boxes and tree cavities. Auk 114(4):646-65.
Raphael, M. G., & M. Wight. 1984. Use of snags by cavity-nesting birds in the Sierra Nevada.
Wildlife Monographs. No. 86.
Rasmussen, P C., & Collar, N.J. 1998. Identification, distribution and the status of the Forest
Owlet Athene (Heteroglaux) blewitti. Forktail 14: 43–51.
Rasmussen, P. C. & Ishtiaq, F. (2019). Vocalizations and behaviour of the Forest Owlet
Athene (Heteroglaux) blewitti.
Rasmussen, P. C., & Collar, N. J., 1998. Identification, distribution and the status of the
Forest Owlet Athene (Heteroglaux) blewitti. Forktail 14: 43–51.
Rellier, A. W. (1972). Aspects of behavioural ecology of Red-headed and Red-bellied
woodpeckers. Am. Midl. Nat. 88: 270-290.
Ricklefs, R.E. 1969. An analysis of nesting mortality in birds. Smithsonian Contribution to
Zoology. 9: 1-48.
Ripley S. D., 1976. Reconsideration of Athene blewitti (Hume). Journal of Bombay Natural
History Society 73 (1): 1–4.
Robinson, B., & Davis, C. 2014. A Communal Roosting of the Great Horned Owl (Bubo
virginianus). Journal of Raptor Research .48: 88-89.

39
Santhanakrishnan, R., Ali, M. S., & Anbarasan, U. 2011. Breeding biology of the Spotted
Owlet (Athene Brama) in Madurai district, Tamil Nadu, India. International Journal of
Environmental Science. 1: 1682-1691.
Sisco, C. L. 1990. Seasonal home range and habitat ecology of spotted owls in north-western
California. M.S. Thesis, Humboldt State Univ., Arcata, Calif. 81pp.
Smith, D. G., Bosakowski, T., Devine, A. 1999. Nest site selection by urban and rural Great
Horned Owls in the Northeast. Journal of Field Ornithology. 70(4):535-542.
Solis, D. M., Jr., And R. J. Gutierrez. 1990. Summer habitat ecology of northern spotted owls
in north-western California. Condor 92:739-748.
Stauffer, D. F., & L. B. BEST. 1982. Nest site selection by cavity-nesting birds of riparian
habitats in Iowa. Wilson Bull. 94: 329-337.
Swallow, S. K., R. J. Gutierrez, & R. A. Howard. 1986. Primary cavity-site selection by birds.
Journal of Wildlife Management 50: 576-583.
Thomas, J. W. (1979). Wildlife habitats in managed forests-the Blue Mountain of Oregon
and Washington.
Titus, K., & Mosher, J. A. 1981. Nest-site habitat selected by woodland hawks in the Central
Appalachians. Auk 98: 270 -281
Vanitha, V., Kumar, S., & Thiyagesan, K. 2014. Roost and diet selection by southern Spotted
Owlet Athene brama brama (Temminck, 1821) in the Cauvery Delta of Nagapattinam
District, Southern India. Journal of Threatened Taxa. 6 :5845–5850.
Zar, J. H. 1996. Biostatistical Analysis, Third Ed. Prentice Hall, Upper Saddle River, NJ, USA.

40