




































Ecology, Economy, and Society–the INSEE Journal 7(2): 29-60, July 2024 

RESEARCH PAPER 
 

Coexistence of Wild Fauna in the City: A Case Study 
of the Golden Jackal (Canis aureus Linn.) in Central 
Ridge, Delhi 
 

Ajay Immanuel Gonji, Sonali Chauhan, Suresh Babu 
 
Abstract: Rapid urbanization across the world has drawn attention to the unique 
state of urban woodlands. New Delhi is one of the world’s most populous cities; 
yet, it harbours several woodlands that support a variety of wildlife. The persistence 
of mesopredators like the golden jackal (Canis aureus Linn.) in the city is intriguing 
and provides an opportunity to explore coexistence in an urban context. Using a 
combination of camera trapping and occupancy modelling, our study aims to 
understand the habitat use, distribution, and urban adaptations of the jackal in 
Delhi’s Central Ridge Reserve Forest. Preliminary analysis shows extensive habitat 
use and sophisticated adaptations including adjustments in activity patterns in 
response to human activity and competition from feral dogs. The study also 
demonstrated behavioural adaptations, particularly in terms of foraging, denning, 
and coexistence with other species, which enable these urban populations of golden 
jackals to persist in the city. The study indicates the need for newer frameworks for 
conservation of synanthropic wild fauna that persist in calorie-rich urban 
environments.   
 
Keywords: Urbanisation, Delhi Ridge, Golden Jackals, Mesopredator, Behavioural 
Adaptations, Synanthropic Fauna, Urban Adapters 

 

1. INTRODUCTION 

Urban ecosystems are unique combinations of the natural sphere, with 
abiotic and biotic components, and the anthroposphere, which exists as a 
result of the socio-politico-economic actions of humans (Endlicher et al. 

                                                      
 Ph.D. Scholar, School of Human Ecology (SHE), Dr. B. R. Ambedkar University Delhi, 
Lothian Road, Delhi, India 110006 (AUD); aj.immanuel3@gmail.com 

 PhD Scholar, SHE, AUD;. sonalichauhan7@gmail.com 

 Professor, SHE, AUD; suresh@aud.ac.in  

Copyright © Gonji, Chauhan and Babu 2024. Released under Creative Commons 
Attribution © NonCommercial 4.0 International licence (CC BY-NC 4.0) by the author.  

Published by Indian Society for Ecological Economics (INSEE), c/o Institute of Economic 
Growth, University Enclave, North Campus, Delhi 110007.  

ISSN: 2581–6152 (print); 2581–6101 (web).  

DOI: https://doi.org/10.37773/ees.v7i2.1192  

mailto:aj.immanuel3@gmail.com
mailto:sonalichauhan7@gmail.com
mailto:suresh@aud.ac.in
https://doi.org/10.37773/ees.v7i2.1192


Ecology, Economy and Society–the INSEE Journal [30] 

2007; Gehrt et al. 2010). Human activities are central to the creation of new 
patterns and processes, leading to novel forms of nature in cities (Shochat et 
al. 2006). Urban landscapes have always hosted, and continue to cater to, a 
unique set of species and novel biological communities that are shaped by 
the loss of native species and the introduction of alien species (Grimm et al. 
2015; Kowarik 2011). Urban ecology thus becomes a study of habitat 
fragmentation (Schilthuizen 2018). Most urban landscapes today are 
undergoing rapid compartmentalization into mosaics of areas constructed 
for residential, commercial, industrial, and transportation purposes, 
interspersed with green spaces (Venn and Niemelä 2004; Bateman and 
Fleming 2012; Forman and Alexander 1998). These landscapes may be 
composed of disconnected subpopulations or metapopulations of plants 
and animals that have limited interaction between them (Rodewald and 
Gehrt 2014; Andrén 1994; Bennett and Saunders 2010; Fahrig 2003). 

Urban landscapes are fascinatingly heterogeneous, including niches that 
have resulted from the development of built infrastructure and modified 
natural areas (Thomas 2017; Schilthuizen 2018). These new habitats, some 
of which are abundant in resources, provide opportunities for species to 
optimize behaviourally and physiologically to these “super-rich patches” 
(Bateman and Fleming 2012; Prange et al. 2004). Several species 
continuously adapt to transforming city environments and undergo a 
process of “synurbization”, taking advantage of novel opportunities in cities 
(Luniak 2004). Urban wildlife populations may therefore be significantly 
different from their counterparts in the wild, having distinctive life histories, 
behaviours, and physiological traits that enable them to navigate urban 
environments; but these differences are poorly understood (Rodewald and 
Gehrt 2014). Further, Blair (2001) suggests that species that are successful 
in urban settings—“urban adapters”—respond differently in terms of 
behavioural adaptations despite being challenged by a similar set of 
environmental factors.  

Urban adapters demonstrate a wide array of responses to urban 
environmental conditions. Besides making evolutionary adaptations to 
altered environmental conditions, organisms are capable of non-
evolutionary adaptations by adjusting their physiology, morphology, and 
behaviours (McDonnell and Hahs 2015). In the case of plants and 
microbes, physiological and genetic adaptations, which are well known and 
widely studied, dominate. In the case of animals, however, behavioural 
adaptations seem to play a major role. This could be the case because 
animals, being mobile, can avoid unfavourable environmental conditions by 
relocating. Behaviours that manifest as a result of such factors indicate 
which animals are more likely to survive in a given environment—“survival 



[31] Gonji, Chauhan and Babu 

value” or adaptation (Tinbergen 1963; Dawkins 2008; McDonnell and Hahs 
2015). These behavioural adaptations, whether innate or learned, are a 
product of genetic and environmental factors (Jensen 2007; Sol et al. 2013).  

Among urban adapters, mesopredators or medium-sized predators are 
extremely flexible in terms of their behaviour, diet, and ability to adapt to 
highly fragmented and human-dominated landscapes (Ćirović et al. 2016; 
Crooks 2002). Mesopredators play an important role in urban ecosystems 
(Morell 2013) because of their position in the food chain (Hepcan 2012). 
The establishment of coyotes in cities in the United States (Gehrt 2007) and 
the expansion of golden jackals through Europe (Trouwborst et al. 2015) 
are compelling examples of mesopredator adaptations in urban 
environments. Although humans have made intensive efforts to control 
populations of canids such as red foxes, coyotes, golden jackals, and feral 
dogs, these species have managed to not only survive but thrive 
(MacDonald et al. 2019). 

The focal species of this study—the golden jackal (Canis aureus Linn.)—is a 
mesopredator of the family Canidae and one of the most widespread canids 
across the globe (Markov 2012). Indian golden jackals form the oldest 
haplogroup in the world (Ibiş et al. 2015), with the northern and west Indian 
populations being the oldest genetic stock from which jackals radiated all 
over the planet after the Last Glacial Maximum (Yumnam et al. 2015). 
Presently, the golden jackal is listed as a species of Least Concern by the 
International Union for Conservation of Nature (IUCN) (Hoffmann et al. 
2018). Through a recent amendment to the Wild Life (Protection) Act in 
2022, the golden jackal has been given the highest level of protection in 
India as a Schedule I species (Government of India 2022).  

In Delhi, the golden jackal is found in parks and woodlands of the Delhi 
Ridge and has recently been sighted in other conservation areas such as the 
Yamuna Biodiversity Park (Gandhiok 2023). Golden jackal populations that 
thrived in most parts of the city a century ago are now restricted to the 
forest fragments of the Delhi Ridge and a few tracts of the floodplains. The 
rest of the land has been taken over by the built infrastructure of the 
National Capital Territory (NCT) of Delhi. 

The objectives of this investigation are twofold: (i) to study the distribution 
and habitat use of the golden jackal in the Delhi Ridge, and (ii) to study the 
behavioural adaptations of the golden jackal and other common faunal 
species in the Ridge—specifically in terms of temporal and spatial responses 
to life in the city. 

 



Ecology, Economy and Society–the INSEE Journal [32] 

2. STUDY SITE 

Spread over an area of 1,483 km2 and with over 30 million residents, Delhi 
is the most populous city in India. The city is projected to become the most 
populous in the world, with a population of 39 million, by 2030 (United 
Nations 2019; Census of India 2011). Delhi is unique in its rich 
biogeographical heritage, and in comparison to other regions in the country, 
it has a remarkable number of trees and a large area of dense forests 
(Crowley 2015). Delhi’s urbanization has historically been characterised by 
major landscape-level changes, modernization, sprawl, and the consequent 
transformation of its two major ecological features: the Aravallis and the 
Yamuna floodplains. Aravalli is the oldest mountain range in the country, 
traversing Gujarat, Rajasthan, Haryana, and Delhi (Varsha 2014). Although 
“ridge” is a colonial term used to describe the hilly topography of Delhi, 
when people speak of the Ridge, they are usually referring not to its rocky 
outcrops but its forests (Crowley 2015; Baviskar 2020). 

Figure 1: Physical Layout of the Study Site 

Source: Authors’ analysis 

Note: The study was conducted on 464 hectares (4.64 km2) of the Central Ridge 
Reserve Forest, which is under the jurisdiction of the Department of Forests and 
Wildlife, Government of the NCT of Delhi. 

The Central Ridge (formerly called the Southern Ridge) is a reserved forest 
spread over an area of 864 hectares (8.64 km2), situated in the 
administrative centre of Delhi (Sud 2017; Varsha 2014). It was accorded 
protection in 1913 by British rulers under Section 4 of Act VII of the 



[33] Gonji, Chauhan and Babu 

Indian Forest Act, 1878, through the appropriation and afforestation of the 
land of about 25 villages (then listed as 788 hectares) (Varsha 2014; Sud 
2017; Crowley 2015). The present Central Ridge falls under the jurisdiction 
of a surprisingly large number of state agencies including the Department of 
Forests and Wildlife, the Indian Army, the Delhi Development Authority 
(DDA), the Central Public Works Department (CPWD), the New Delhi 
Municipal Corporation (NDMC), the Ministry of Home Affairs (MHA), the 
Municipal Corporation of Delhi (MCD), the Indian Railways, the Land and 
Development Office (L&DO), and the Sports Authority of India (Solanki 
2014; Crowley 2015). As can be seen in Figure 1, the Central Ridge is 
subject to multiple land uses. The various state and private agencies 
occupying the Central Ridge often have competing agendas and policies, 
making the space highly contested and politicized (Sud 2017). 

The principal vegetation type of the Ridge is northern tropical thorn forests 
(Champion and Seth 1968). The dominant tree groups found are Acacia 
and Mimosa, with naturally occurring populations of native tree species 
such as Acacia leucophloea, Acacia nilotica, Butea monosperma, Anogeissus pendula, 
Diospyros montana, Cordia dichotoma, Cordia rothii, and Ehretia laevis, and native 
shrubs such as Grewia tenax and Capparis sepiaria (Krishen 2006; Maheshwari 
1963). As far as faunal species are concerned, golden jackals and rhesus 
macaques (Macaca mulatta) are commonly seen in large numbers, especially 
at feeding sites—locations in the Ridge where people come to feed animals. 
Sizeable populations of feral cattle (Bos indicus) and feral pigs (Sus domesticus) 
are also frequently spotted at these feeding sites. Other wild species that are 
observed include nilgai (Boselaphus tragocamelus), Indian hare (Lepus nigricollis), 
common palm civet (Paradoxurus hermaphroditus), small Indian civet 
(Viverricula indica), Indian grey mongoose (Urva edwardsii), and several other 
species of rodents, reptiles, amphibians, and birds, including the commonly 
occurring Indian peafowl (Pavo cristatus) (ZSI 1997). 

 

3. METHODOLOGY 

In pursuance of the objectives of our study on golden jackals and other 
associated faunal elements, we developed a study design that used a mixed 
methods approach, including a grid-based design, systematic sampling, the 
use of camera traps, occupancy/habitat use analysis, quantitative estimation 
of habitat variables or covariates, direct and indirect observation of animal 
behaviours, and analysis of secondary data. 

3.1 Camera Trapping  

Camera traps can be used to document species’ presence in an area and to 
study activity patterns and certain aspects of behaviour. Camera trap 



Ecology, Economy and Society–the INSEE Journal [34] 

surveys are extensively used in wildlife surveys and have been standardized 
to fit into statistical models for population studies (McCallum 2013; 
Rowcliffe and Carbone 2008). Besides documenting species’ presence and 
obtaining information on activity patterns and habitat preferences, camera 
traps can allow for estimating density and occupancy when used in 
scientifically robust, inferential sampling studies (Rovero et al. 2010).  

Figure 2: Grid Network and Placement of Camera Trap Stations in the Central 
Ridge Reserve Forest 

Source: Authors’ analysis 

Our camera trapping study design incorporated standard methods and 
recommendations as per Nichols et al. (2011). The study site was overlaid 
with 400 × 400 m grids, and each grid was considered a sampling unit. 
Grids that had 40% or more of built-up area were excluded from the 
sampling, resulting in 24 grids. A tracks and signs survey using the line 
transect method was conducted to find suitable spots to set up camera traps 
within each grid. Prior knowledge of golden jackal ecology and behaviours 
(i.e., preferred habitat, den characteristics, and visual appearance of spoor 
and scat) also helped with locating potential camera trap stations (Ancrenaz 
et al. 2012). 



[35] Gonji, Chauhan and Babu 

To maximize the probability of detecting a jackal in a grid, we deployed 
three camera traps per grid. As such, the entire study site had 72 camera 
trap stations (Figure 2). The study used 10 units of the Cuddeback 123 IR 
black flash camera trap—a passive sensor infrared camera system. All 
camera traps were operational for approximately 24 hours over three 
surveys. After three months of camera trapping, the survey effort amounted 
to a total of 5,184 camera trap hours (24 hours × 72 camera trap stations × 
3 repeated surveys). 

3.2 Patch Occupancy/ Habitat Use 

Estimating population parameters such as abundance or density has long 
been the focus of animal population studies (O’Connell and Bailey 2011). 
However, estimating parameters such as abundance requires that individuals 
are identifiable (by coat colour and/or pattern) to keep accurate records of 
the number of encounters for each animal (Mackenzie and Nichols 2004). 
The basic sampling protocol commonly used for occupancy estimation 
involves visiting sites and spending time in each one, looking either for 
individuals of the species of interest or for evidence that the species is 
present (Mackenzie et al. 2006). Mackenzie et al. (2006) developed models to 
estimate site occupancy and detection probability—the probability of 
detecting at least one individual of a species on one sampling occasion, 
given that the species is present in the sampling area—based on repeated 
presence/absence surveys of multiple sites. 

Our study was based on the single-season occupancy modelling approach 
designed by Mackenzie et al. (2006). We used camera traps to establish the 
presence/absence of the golden jackal in the sampling grids. Occupancy 
and detection probability were modelled by coding photographic count data 
in a binary (1/0) format, describing detection (1) or non-detection (0) of the 
animal during the three repeated surveys. Due to the paucity of baseline 
data and home range of jackals in the urban setting, the results of our patch 
occupancy study are being interpreted as “habitat used” instead of “patch 
occupied” (Mackenzie and Royle 2005)*. 

The probability of occupancy or habitat use (ψ or psi) was modelled as a 
function of site-specific covariates or spatial variables, such as tree density, 
shrub density, dominant vegetation, and normalized difference vegetation 
index (NDVI), to make an inference about factors determining the golden 
jackals’ habitat use. Meanwhile, detection probability (p) was modelled as a 

                                                      
* The 400 m x 400 m size we fixed for the sampling units was based on observations and not 
based on recorded home range data for urban conditions. Therefore, this assumption had to 
be relaxed by interpreting results not as patch ‘occupancy’ but as patch ‘use’ (Mackenzie and 
Royle, 2005), or habitat use. 



Ecology, Economy and Society–the INSEE Journal [36] 

function of site-specific covariates, such as tree density and shrub density, 
and survey-specific covariates or temporal variables, such as maximum and 
minimum temperatures, to determine how likely a species is to be detected 
when present (Sollmann 2018). 

The single-season occupancy modelling approach and detection history 
were analysed using the package unmarked in R (Fiske and Chandler 2011) 
to generate maximum likelihood estimates for occupancy/habitat use and 
detection probabilities. Further, all possible combinations of site-specific 
and survey-specific covariates as influences on occupancy/habitat use and 
detection probabilities were modelled using the logit link function. For the 
interpretation of results, only model(s) with the least AIC (Akaike 
information criteria) were considered (Burnham and Anderson 2002). 

3.3 Behavioural Adaptations of Golden Jackals  

The section below outlines methods used to survey specific behavioural 
adaptations examined in the study, including temporal activity patterns, 
foraging, and denning. 

3.3.1 Temporal Activity Patterns 

Temporal activity data for golden jackals in the Central Ridge Reserve 
Forest were derived from camera trap images that were obtained from the 
patch occupancy or habitat use study of the species. Additionally, we 
obtained data on the temporal activity of feral dogs and humans, since these 
species seem to influence golden jackal activity, especially in the urban 
context.  

Images were analysed for the presence of golden jackals, feral dogs, and 
humans, and when detected, the timestamps were recorded. Once all this 
data was collected, following the method used by Gil-Fernández et al. 
(2020), we employed the package overlap in R (Meredith and Ridout 2014) 
to determine the overlap between two activity patterns based on kernel 
density estimates. We compared the temporal activity patterns of jackal–
dog, jackal–human, and dog–human across the study site. To interpret the 
overlap in activity patterns, we used the overlap estimator Dhat4, which is 
ideal for samples with >50 observations per species. 

Temporal activity data for all other species found in the Central Ridge 
Reserve Forest was also derived from the same camera trapping exercise. 
Besides golden jackals, feral dogs, and humans, we detected the common 
palm civet, small Indian civet, rhesus macaque, feral cattle, feral pig, Indian 
peafowl, Indian hare, Indian grey mongoose, and nilgai. Detections of each 
species were analysed and activity pattern graphs were generated using the 
package overlap in R. 



[37] Gonji, Chauhan and Babu 

3.3.2 Foraging Behaviour 

Golden jackal foraging behaviour, especially for anthropogenic food, was 
directly and indirectly observed at a major feeding site in the Central Ridge 
Reserve Forest. Feeding sites are locations around the forest where people 
come to feed the animals, although not jackals specifically. People usually 
bring chapati, bread, grains, and fruits that are tossed away or kept on the 
ground to feed the animals. A variety of animals—macaques, cows, pigs, 
dogs, and jackals—congregate at these locations during feeding sessions.  

We used opportunistic photographs and video clips, as well as direct visual 
sightings that were obtained throughout the study period and across all 
seasons, to understand the foraging behaviour of golden jackals at the 
feeding site. Using a Nikon P900 ultrazoom handheld camera, we captured 
the feeding behaviour of the animal. We used binoculars for direct visual 
sightings, while also making note of behavioural information. Our camera 
trap data facilitated an analysis of foraging for naturally occurring foods. 
Whenever possible, we conducted interviews with people who visited the 
Ridge and feeding site to offer food to the animals, to learn about their 
interactions with jackals and to gather other anecdotal information.  

3.3.3 Reproduction and Denning Behaviour 

Data on the denning behaviour of golden jackals was collected using the 
focal-group sampling method (Altmann 1974; Dawkins 2008). Focal-group 
sampling involved focusing on one family unit of the target species, 
recording everything they were doing, and generating statistics that were 
recorded as a single data point among a series of observations (see Dawkins 
2008). Dens were opportunistically encountered during the fieldwork, and 
camera traps were deployed at active den sites. During our fieldwork in the 
Central Ridge Reserve Forest, we located two dens (Figure 6), one of which 
was occupied for consecutive seasons. The first den (henceforth D1) was 
located on 30 April 2022, and the second den (henceforth D2) was first 
located on 25 May 2022 and was abandoned and occupied again on 20 May 
2023. Extensive data collection on denning behaviour was possible only for 
D2 from 25 May 2022 to 28 June 2022. The survey effort for D2 amounted 
to 22 calendar days or, more precisely, 415 camera trap hours. 

3.4 History and Management of the Ridge 

In addition to the primary survey, we consulted secondary sources to 
understand the history and management of the Delhi Ridge. These sources 
included archival records; popular articles in print and electronic media; 
reports of non-governmental organizations (NGOs); gazettes; planning 
documents such as the Delhi Master Plan; annual reports of government 



Ecology, Economy and Society–the INSEE Journal [38] 

departments, the National Green Tribunal (NGT), and the Ridge 
Management Board; and other published work on regional history. 

 

4. RESULTS 

The results of our observations on the golden jackal’s habitat use, temporal 
activity patterns, foraging behaviours, reproduction and denning, and the 
implications of the history of management of the Delhi Ridge, are 
presented in the following section. 

Table 1: Comparison of Models to Explain Golden Jackal Habitat Use in the 
Central Ridge Reserve Forest 

Model K AICc ΔAICc 
AICc 

Weight 
Model 

Likelihood 

psi (.), p (.) 2 80.85 0.00 0.27 1.00 

psi (.), p (tree den.) 3 82.35 1.50 0.13 0.47 

psi (.), p (max. temp.) 3 83.28 2.43 0.08 0.29 

psi (.), p (min. temp.) 3 83.41 2.56 0.08 0.27 

psi (.), p (shrub den.) 3 83.44 2.59 0.07 0.27 

psi (dist. road), p (.) 3 83.48 2.63 0.07 0.26 

psi (tree den.), p (.) 3 83.48 2.63 0.07 0.26 

psi (shrub den.), p (.) 3 83.48 2.63 0.07 0.26 

psi (dom. veg.), p (.) 3 83.48 2.63 0.07 0.26 

psi (NDVI), p (.) 3 83.48 2.63 0.07 0.26 

Source: Authors’ analysis 

Notes: psi = occupancy; p = detection probability; (.) = null. Covariates: tree den. 
= tree density; shrub den. = shrub density; max. temp. = maximum temperature; 
min. temp. = minimum temperature; dom. veg. = dominant vegetation; NDVI = 
Normalized Difference Vegetation Index; dist. road = distance to the nearest road. 

4.1 Patch Occupancy/ Habitat Use 

Analyses of our patch occupancy/habitat use data revealed that golden 
jackals in the Central Ridge Reserve Forest do not favour any particular 
habitat type, as they were detected in all our study grids or patches (Figure 
3). While the naïve occupancy estimate was a perfect 1, indicating 100% 
usage of the study site by the golden jackal, the detection probability 
estimate was 0.78, i.e., there was a 78% chance of detecting the species 
given that it was present. However, the naïve occupancy does not consider 
imperfect detection. Yet, as evident in Table 1, the null model [psi (.), p (.)] 
is the top-ranking model, with the least AICc and maximum likelihood. In 
other words, none of the site-specific covariates (dominant vegetation, tree 
density, shrub density, NDVI, and distance to the nearest road) seem to 
influence the probability of a patch being used by the golden jackal. 



[39] Gonji, Chauhan and Babu 

Similarly, the probability of detecting a golden jackal in our patches does 
not appear to be affected by survey-specific covariates such as maximum 
and minimum temperatures, and site-specific covariates such as tree density 
and shrub density. Although tree density seems to slightly influence the 
probability of detecting a golden jackal in a patch given its presence 
(ΔAICc: 1.50; model likelihood: 0.47), it is nowhere close to the best-
performing model, i.e., the null model. 

Figure 3: Golden Jackals Were Found in All of the 24 Study Grids; Some Grids 
Recorded More Detections than Others 

 
Source: Authors’ analysis 

Among the 72 camera trap stations spread across the 24 study grids, golden 
jackals were detected at 60 stations (Figure 2). As can be seen in Figure 3, 
golden jackals were detected in all 24 grids, although varyingly. After 3 
repeated surveys, we made 387 independent detections of golden jackals in 
the Central Ridge Reserve Forest, with 159 detections in the first survey, 
126 in the second, and 102 in the third. Further, in the 387 independent 
detections, jackals were seen in trios in 11 instances, in pairs in 61 instances, 
and as solitary individuals in 232 instances. As expected, the highest number 
of captures were in grids near the feeding trail. We made 66 detections in 
one of these grids and 34 in the other. Of the remaining 22 grids that were 
surveyed for habitat use, 45.45% had 1–10 detections; 27.27% had 10–20; 
and 27.27% had 20–30.  

According to one of the few published studies on golden jackals in the 
urban Indian context, the population density of jackals in a 100-hectare (1 
km2) golf club was 43 individuals (Sanyal et al. 2011; Gupta et al. 2016). A 
much lower population density of approximately 15 jackals per km2 was 
reported by Singh et al., (2016) in the Keoladeo National Park, India. In the 
metropolis of Delhi, with its forest patches rich in anthropogenic resources 



Ecology, Economy and Society–the INSEE Journal [40] 

Figure 4: Temporal Activity Pattern Graphs for Different Species in the Central 
Ridge Reserve Forest 

 
Source: Authors’ analysis 

Note: Graphs (a), (b), and (c) show the overlap in activity patterns between jackal 
and dog (53%); jackal and human (53%); and dog and human (66%), respectively. 
Graphs (d) to (l) depict the activity patterns of other species. 



[41] Gonji, Chauhan and Babu 

and jackals showing extensive habitat use, our estimate of golden jackals in 
the 4.64 km2 Central Ridge is 120–150 individuals, i.e., a population density 
of 26–32 individuals per km2. 

4.2 Temporal Activity Patterns 

Results of the temporal activity pattern analysis of golden jackals, feral dogs, 
and humans in the Central Ridge Reserve Forest reveal a somewhat clear 
separation in temporal use between the three species (Figures 4a–c). While 
golden jackals use the same areas as feral dogs and humans, unlike the 
latter, golden jackals are most active at night, with activity peaking between 
19:00 and 02:00. The mean overlap between golden jackal and feral dog 
activity was 0.53 (95% CI = 0.44–0.62); between golden jackal and human 
activity 0.53 (95% CI = 0.42–0.65); and between feral dog and human 
activity 0.66 (95% CI = 0.55–0.77). It is important to note, however, that 
the total number of sightings, depicted by the colour-coded bars at the base 
of the graphs, was highly variable. The highest number of sightings were of 
golden jackals; there were fewer sightings of feral dogs and humans. 

When comparing the activity patterns of other faunal species (Figures 4d–l) 
found in the Central Ridge, we observed differences in their movement and 
activity. Temporal partitioning seems to exist between species—some are 
nocturnal, others diurnal, and still others exhibit cathemeral behaviours. 
The common palm civet and small Indian civet are primarily arboreal and 
rarely seen during the day; they come down to the forest floor at night in 
search of food. Therefore, the activity pattern graphs in Figures 4d–e are 
indicative of their nightly movements on the forest floor. The Indian hare is 
also predominantly active at night, although it can occasionally be sighted 
during the day (Figure 4j). In the case of the nilgai, the largest wild mammal 
found in the Ridge, sightings were restricted to 2 of the 24 grids, with most 
occurring after sundown. While occasional sightings of the nilgai are 
possible in the Central Ridge during the day, our data (Figure 4l) provides 
clear evidence of increased movement and activity at night. Our data 
suggests that species such as the common palm civet, small Indian civet, 
Indian hare, and nilgai are mostly nocturnal, at least in the Central Ridge. 

Among the strictly diurnal species found in the Central Ridge are the rhesus 
macaque, Indian peafowl, and Indian grey mongoose. As can be seen in 
Figures 4f, 4i, and 4k, none of these three species show any significant 
activity from dusk to dawn (approximately 18:00–06:00). Similar to civets, 
macaques and peafowls exhibit a mix of arboreal and terrestrial behaviours; 
therefore, their activity pattern graphs only depict their movements on the 
forest floor. Feral cattle and feral pigs exhibit cathemeral behaviours, with 
both species being detected across the 24-hour cycle (Figures 4g–h). Both 



Ecology, Economy and Society–the INSEE Journal [42] 

these species spend a significant amount of time close to feeding sites 
during the day, especially in the early mornings when feeders visit the 
forest. However, during the twilight hours and sometimes the night, they 
travel from one part of the forest to another. This is evidenced by the 
movement of large herds of cattle recorded by our camera traps. 

4.3 Foraging Behaviour 

While jackals are generally nocturnal (Patil and Jhala 2008), especially in the 
wilderness, in the Central Ridge Reserve Forest, they have also taken to 
opportunistic feeding on human-provisioned food during the day, especially 
in the early mornings. The food items golden jackals feed on in the Central 
Ridge include bananas, melons, bread, chapati, cake rusks (Figure 5), 
groundnuts, chickpeas, maize, and the occasional carcasses and offal 
material dumped by butchers. Jackals also drink water from the many 
troughs that line the feeding trail. Every morning, a few feeders come in 
cars with jerrycans of fresh water to refill the water troughs. 

Figure 5: A Golden Jackal Grabbing Cake Rusks from the Pavement, Close to the 
Feeding Site 

 
Source: Authors’ compilation 

At feeding sites, jackals often expectantly wait for food to be handed to 
them by people who come in cars and on bikes; honking from these 
automobiles often incites curiosity, excitement, and hope among the 
animals. In one instance, we observed an adult jackal resting a little distance 
away from the main feeding trail, running to the trail at every sound of a car 
honking, and returning to its resting place once the car had passed. This 
happened at least three times after which the jackal moved away from its 
resting place. On several other occasions, jackals were seen trotting behind 
cars in the hope of receiving food. Such behaviours are a testament to the 



[43] Gonji, Chauhan and Babu 

fact that jackals are opportunists and that they express certain novel 
behaviours in urban environments. 

Our camera trapping data reveals the golden jackals’ dependence on 
naturally occurring foods such as birds and insects; this is confirmed by 
other studies (e.g., Mukherjee et al. 2004; Baskaran et al. 2020). In India’s 
Sariska Tiger Reserve, a scat analysis showed that 45% of the golden jackal’s 
diet comprises mammals (of which 36% are rodents), 20% vegetable 
matter, 19% birds, 8% reptiles, and 8% invertebrates (Mukherjee 1998).  

4.4 Reproduction and Denning Behaviour 

Our observations on the denning behaviours and reproduction of the 
golden jackal in the Central Ridge along with details on feeding behaviours 
during denning are presented in this section.  

4.4.1 Denning Season 

In India, golden jackals are typically monogamous, forming lifetime pair 
bonds and parenting offspring once a year, as the females are monestrous, 
i.e., they have one estrous cycle per year (Jhala and Moehlman 2004; Negi 
2014). However, the social unit of jackals may also consist of “helpers”—
offspring from the previous litter (11–20 months old) who help raise their 
younger siblings (Fuller et al. 1989). Carnivores such as wolves have highly 
regularized breeding and denning periods (Fuller et al. 2010), and as our 
research suggests, this may also be the case for jackals. Golden jackals begin 
exhibiting reproductive behaviour, i.e., mating, in February–March. This is 
followed by den selection/excavation in April–May. Finally, whelping 
occurs after a gestation period of 63 days, which often coincides with an 
abundance of food supply, i.e., at the beginning of the monsoon (Jhala and 
Moehlman 2004).  

In 2022, D1 contained a single pup, which we estimated to be less than a 
week old when first detected. We triangulated this based on the fact that 
golden jackal pups are born blind and open their eyes only after about nine 
days (Jhala and Moehlman 2004). In the same season, D2 contained four 
pups, which we estimated to be between four to five weeks old when first 
detected. In 2023, D2 contained four pups which we estimated to be 
between three to four weeks old when first detected. Thus, there is a strong 
indication that, in all three cases, mating likely occurred in the last week of 
February and whelping in the last week of April. 

4.4.2 Den Location and Characteristics 

Canids are a group of mammals which do not build their own burrows as 
primary excavators do but are secondary modifiers that inhabit and 



Ecology, Economy and Society–the INSEE Journal [44] 

maintain burrows built by primary excavators (Mukherjee et al. 2018). Dens 
are intrinsic to the lives of most wild canids and may be decisive factors in 
their abundance and distribution (Nurvianto et al. 2015). A denning site 
determines access to critical resources, provides protection from predators, 
acts as a safe haven from environmental vagaries, and offers conducive 
ambient conditions for the optimal development of offspring (Majumder et 
al. 2016). 

Figure 6: Golden Jackal Dens in the Central Ridge Reserve Forest 

 
Source: Authors’ compilation 

Note: (a) D1 contained a single pup in 2022; (b) D2 contained four pups each in 
2022 and 2023. 

As can be seen in Figure 6, D1 and D2 are very different types of natal 
shelters. D1 was located in a rocky habitat, with even elevation and 
characteristics of open-scrub forests. It was situated in a single-opening 
rock shelter in a crevice and had a shallow depression below the surface of 
the ground. D1 was surrounded on all sides by vegetation such as Carissa 
spinarum, Abutilon indicum, and grasses. The inside of the den had a bed of 
loose sand that probably provided cushioning for the pups. D2, on the 
other hand, was located in a rocky habitat, with uneven elevation and 
characteristics of a mixed forest. It was situated at the base of an Azadirachta 
indica tree and surrounded by other species such as Holoptelea integrifolia and 
Ehretia laevis, although the dominant vegetation in this patch was Acacia 
catechu. Some of the shrub species found around D2 were Adhatoda vasica, 
Capparis sepiaria, and Carissa spinarum. D2 also had a single opening, but 
unlike D1, it was under a living tree and had a mound-like appearance. At 
the entrance of the den were the roots of the tree, and the den was on 
slightly higher ground in relation to the natural objects in its immediate 
vicinity. D2 was located a few metres from the boundary wall along a main 
road and a busy intersection, but the breeding pair in both pup-rearing 
seasons (2022 and 2023) still chose this natal shelter. 

 

 



[45] Gonji, Chauhan and Babu 

4.4.3 Litter Size 

Depending on their geographical location, golden jackals have varying litter 
sizes. In Tanzania, litter sizes vary from 1 to 8, the average being 5.7 
(Kebede 2017), and in the Bhal region of India, litter sizes range from 2 to 
5, with an average of 3.6 (Jhala and Moehlman 2004). In 2022, D1 
contained a single pup, while D2 contained four pups. Similarly, in 2023, 
D2 contained four pups from the same breeding pair that occupied D2 in 
2022. We also captured two other females and their respective pups on 
camera traps during the patch occupancy surveys. Both groups were led by 
the breeding female on foraging bouts. While one of the females had a 
single pup, the other had four. Therefore, with some level of confidence, 
we estimate the litter size of golden jackals in the Central Ridge Reserve 
Forest to range from one to four pups. 

4.4.4 Feeding Behaviour 

Canid pups consume only their mother’s milk for the first few weeks after 
birth, after which, along with milk, they start consuming regurgitated food 
brought by the adults. According to MacDonald et al. (2019), regurgitation 
in canids is an evolutionary adaptation in monogamous social systems that 
facilitates parental care, alloparental care, and/or cooperative breeding. 
Regurgitating food provides an economical and safe way for parents and 
alloparents to bring large quantities of food to their pups (Moehlman 1987). 
During our research, we found that besides hunted food items such as small 
mammals (piglets, mongoose, etc.), golden jackal pups were also fed 
anthropogenic food such as chapati, bananas, and other high-calorie foods 
(Figure 7).  

4.2 History and Management of the Ridge 

In the last century, and more so in the post-Independence period, the 
urbanization processes of Delhi has led to a progressive fragmentation of 
the city’s natural habitats; this fragmentation was particularly accelerated in 
the case of the Delhi Ridge (Baviskar 2018; Mohan 2002; Planning 
Commission 2009). As a remnant of the Aravalli mountains, which are rich 
in quartzite, the Ridge provided the rocks and other materials for the 
construction of Lutyens’ Delhi. It also created an “evergreen 
background”—with the introduction of Prosopis juliflora—for the president’s 
estate and several other government establishments built in and around the 
Central Ridge Reserve Forest (Mann and Sehrawat 2009; Bowe 2009; 
Baviskar 2018).  

In the post-Independence period, despite its status as a reserved forest, the 
Central Ridge was fragmented both for legal construction and illegal 



Ecology, Economy and Society–the INSEE Journal [46] 

settlements, and its forests were cleared whenever there was a need to 
expand the city. Despite citizen movements in the 1980s, the fragmentation 
of the Ridge initially proceeded to create refugee settlements after the 
partition of India in 1947, and subsequently, to construct government 
establishments, military infrastructure, schools, and gardens, and to expand 
transport networks and various illegal encroachments (Baviskar 2018; Arora 
et al. 1991). More recently, developmental activities and the building of 
roads, metro corridors, mega malls, residential complexes, corporate offices, 
luxury hotels, and farmhouses have all encroached upon forested land 
(Crowley 2015). 

Figure 7: Golden Jackal Pups Were Provided a Combination of Natural and 
Anthropogenic Food Items by Both the Adults (Breeding Pair) and Sub-adults 
(Helpers); (a) Piglet; (b) Mongoose; (c) Chapati; (d) Regurgitated Banana 

 

Source: Authors’ compilation 

To address the issue of fragmentation of the Ridge, the Supreme Court of 
India had to intervene.  Following the directions of the top court, in 1995, 
the Ridge Management Board was constituted with a mandate to protect 
and conserve the Ridge (Sinha 2014). However, when it comes to the 
management of the Ridge, state agencies in charge of the Ridge have always 



[47] Gonji, Chauhan and Babu 

favoured the conversion of wild forests into semi-wild parklands (by cutting 
the undergrowth) or totally new parklands (by planting ornamental trees 
and shrubs after removing most of the original vegetation) (Arora et al. 
1991). In other words, Ridge management has often meant turning wild 
forests into tamed parks (Crowley 2015). Presently, within the Ridge is a 
wildlife sanctuary, biodiversity parks, a citizens’ forest, restoration projects, 
and a sizeable number of manicured parks (Sud 2017). Taming the forests 
became a mandate after the Ridge was perceived as a place of danger and 
criminal activity (Baviskar 2020). Further, although the Delhi Master Plan 
2021 lays special emphasis on the conservation of the 77.77 km2 Ridge, 
there are discrepancies between the area notified and the physical 
boundaries of the total area owned by various state agencies, the Forest 
Department, and the Ministry of Defence (DDA 2010). 

The Ridge was once known for its abundant and diverse flora and fauna 
(Punjab Government 1883). However, pressure from hunting and massive 
fragmentation in the post-Independence era led to a sharp decline in 
wildlife populations, resulting in the local extinction of several species 
(Kumar 1997). For instance, archival records reveal that the golden jackal in 
Delhi was previously the target of post-colonial extermination campaigns 
and fight-to-the-finish drives because the animal was negatively portrayed as 
a rapidly multiplying species, a disturbance to the peace and tranquillity of 
the city, a danger to citizens, and a source of rabies for street dogs (The 
Times of India 1950). However, despite such campaigns, the rapid scale of 
urban expansion and the resulting fragmentation of natural habitats, the 
Ridge continues to harbour diverse fauna. Indeed, the species found in the 
Ridge have not formed habitat associations over a short span of time but 
gradually settled over a few centuries (Mann and Sehrawat 2009), in 
response to a changing, urbanising environment. However, being subject to 
certain combinations of ecological stresses, disturbances, structures, and 
functions (sensu Pickett et al. 1997), some native species that could not adapt 
to the urban landscape were exterminated or perished. Meanwhile, other 
native and non-native species that were able to adapt and coevolve, 
particularly in terms of behaviour, were able to survive.  

 

5. DISCUSSION 

One of the highlights of our research is the discovery of the ubiquitous 
presence and habitat use of the golden jackal in the Central Ridge Reserve 
Forest. Our findings reemphasize the fact that jackals are urban adapters, 
fitting into the spatial and temporal matrix of the city while performing 
their life functions. Although there were differences in habitat variables 



Ecology, Economy and Society–the INSEE Journal [48] 

such as tree density, shrub density, dominant vegetation type, and NDVI 
across the study site, as well as the presence of multiple land uses—from 
parks and gardens to cultural, religious, and government establishments—
the golden jackal was found in all 24 study grids. Typically, when examining 
wildlife habitat associations, a proper characterization of environmental and 
other potential explanatory variables or covariates at spatial scales pertinent 
to the species being researched is necessary (Niedballa et al. 2015). Yet from 
the outcomes of our study, it would seem that small habitat variations do 
not affect the distribution and habitat use of golden jackals in the urban 
context. While there were jackal detections in all our study grids, there were 
some grids, such as the one near the feeding trail, which made as many as 
66 detections. However, the activity pattern of a particular species in a 
single season may not be a true reflection of its overall activity pattern 
across seasons. Indeed, several studies indicate that activity patterns are 
strongly influenced by factors such as seasonality, circannual rhythms (for 
instance, breeding and non-breeding seasons), diurnal variations, and 
vegetation structure (Iannarilli et al. 2021; Kays et al. 2020). However, within 
the constraints of the city, we find that habitat use and activity patterns of 
species do not demonstrate these typical relationships. We recognize, 
therefore, the need for a long-term study to monitor seasonal changes in 
activity patterns as well as the distribution and habitat use of the golden 
jackal in the Central Ridge. To make more accurate estimations of golden 
jackal numbers and deepen our understanding of foraging behaviour in the 
Central Ridge, further research involving radio-collaring, scat analysis, and 
genetic sampling is necessary. 

There is a clear separation in time use between jackals, dogs, and humans in 
our Central Ridge Reserve Forest site. The co-occurrence of species, 
especially those belonging to the same guild, may lead to high niche overlap 
and the possibility of conflictual interactions such as competition or even 
exclusion (Gómez et al. 2018; Pascual-Rico et al. 2020). Studies have shown 
that, when dogs and native predators compete over human-derived 
materials, dogs may be more effective competitors with native carnivores 
(Vanak and Gompper 2009). But species may manage to avoid competition 
by evolving specialized ways of living, such as adopting a novel diet, 
occupying other habitats, or foraging at different times (Southern 1955). 
Therefore, temporal partitioning—using the same space at different 
times—facilitates the coexistence of species and access to available habitats 
and key resources (Gaynor 2018; Haswell 2020). In our study, jackals and 
feral dogs used the same habitat patches but separated their habitat use 
temporally, with jackals being predominantly nocturnal and dogs being 
predominantly diurnal.  



[49] Gonji, Chauhan and Babu 

Similarly, the majority of human detections occurred outside the peak 
activity hours of the golden jackal. Several studies have demonstrated that, 
where humans and wild animals coexist, animals adopt nocturnality to 
reduce contact with humans, who are mostly active during the day (Benítez-
López 2018; Grinder and Krausman 2001). Our observations are consistent 
with studies (Gehrt 2007; Patil and Jhala 2008) that suggest that 
mesopredators such as coyotes and jackals become increasingly nocturnal as 
they come closer to human habitation and cities. Therefore, the shift to 
nocturnality among golden jackals in the Ridge is a coevolutionary outcome 
of a multispecies interaction.  

Past studies on mesopredators have shown that the availability of a 
sustained supply of anthropogenic food in cities leads to a change in the 
home range behaviour of the mesopredators because they can spend less 
time foraging and simultaneously have access to highly nutritious food 
(Ryan and Partan 2014). Similarly, personal observations of early-morning 
jackal roadkills and other direct observations of jackals indicate that they 
move in relation to the spatial arrangement of resources such as food, 
water, and denning sites, and some individuals get killed while attempting to 
access these resources (Coffin 2007). Regardless, the novel behaviours of 
urban adapters point to their propensity to negotiate, learn, and adapt to the 
complex and dynamic nature of the city (Sinha and Barua 2020). This is 
consistent with our temporal activity pattern analysis which suggests that by 
following varied behavioural strategies, species have learnt to coexist in the 
small and fragmented habitats of the Ridge, synchronizing their life history 
strategies with what the city offers. 

An important enquiry for biologists has been how some species are able to 
behaviourally adjust and adapt to cities while others are not, especially when 
resources are not a limiting factor (Lowry et al. 2013). The changes urban-
dwelling wild canids have made to their reproductive behaviour and the 
successful raising of their offspring, both central to their life history 
strategies, are evidence that they are thriving in the city. Shrinking habitats 
results in the decline of denning sites (Majumder et al. 2016), leading 
mesopredators like jackals to take to human habitations and even 
abandoned or less used built infrastructure to bring up their offspring 
(Marks and Bloomfield 1999). One of the denning sites in this study was 
located close to a major traffic intersection, despite the noise from 
continuous vehicular movement and the blaring of horns. Such responses 
of urban adapters may also be an indication of their propensity for risk-
taking behaviours, whereby they display a high disturbance tolerance or 
bold temperament (Lowry et al. 2013). In this study, we also recorded that 
jackal pups were being fed anthropogenic food regurgitated by the breeding 



Ecology, Economy and Society–the INSEE Journal [50] 

pair and helpers, suggesting that even the pups are being reared through 
human provisioning. Traits like these may be game changers in the urban 
context for species to gain access to critical resources that can enhance not 
just their survivability but also that of their offspring. 

On the notion of “wild” in the urban context, we observe that, while 
domesticated species are integral to daily city life and their presence is not 
just tolerated but also encouraged by human beings, the presence of wild 
animals in the city is often unacknowledged, especially because their cryptic 
biology and behavioural adaptations render them invisible. A 1999 
publication by the Delhi government features maps that label large swathes 
of the Ridge forest as a wasteland, completely overlooking their ecological 
functions (Crowley 2015). Such a view has been instrumental in the 
“parkification” of the Ridge forest—systematic intervention by the 
management has led to the conversion of native vegetation patches into 
manicured and aesthetically pleasing recreational spaces (Baviskar 2018). 
Nature in the city is being reimagined to conform with ideas of order, 
beauty, and safety. In other words, “natural” has to be managed within an 
acceptable range, a point made clear in the Delhi Master Plan’s 
categorization of land use as “green areas” but with an emphasis on not 
allowing these areas to take the form of unmanaged wilderness (Baviskar 
2020). By deeming natural spaces in the city as “wild”, “wasteland”, 
“degraded”, “invaded”, or in need of “development”, urban planners 
overlook the many nonhuman creatures that occupy these habitats, thus 
increasing their precarity (Palmer 2003). 

Still, conservation biologists continue to view urban wildlife as ‘threatened’ 
by shrinking habitats and local extinction (Magle et al. 2010). The Ridge is 
now a Reserved Forest, and this title offers a layer of protection to the area. 
There is also ongoing work by the forest department to eradicate the 
invasive Prosopis juliflora vegetation and restore native forests. As these forest 
patches have been increasingly brought under strict protection, the fate of 
the fauna becomes uncertain. As noted in this study, the wild urban fauna 
are adapters that have attuned to human provisioning for sustenance. With 
tall walls and fences around the forest for protection, their mobility and 
ability to access resources from neighbourhoods are becoming increasingly 
difficult. There are obvious challenges in the development of effective 
conservation strategies in these novel ecosystems, primarily because 
conventional conservation biology is unfamiliar with the urban landscape 
(Parker 2015). The idea that “disturbed” urban ecosystems can nevertheless 
support wildlife populations is noteworthy because it opens up possibilities 
for research on how these populations survive in such ecosystems. Our 
study indicates a need for alternate frameworks to describe habitats that are 



[51] Gonji, Chauhan and Babu 

wild and urban, and composed of novel plant associations (Chauhan et al. 
2022). There is a growing trend in historical ecology to break the discourse 
around the pristine idea of nature and view urban habitats not as isolated 
natural habitats but as socio-ecological landscapes embedded in burgeoning 
metropolises (McDonnell 2011; Cavin and Kull 2017). 

Is it necessary to view urban wilderness and wildlife as dwindling, degraded, 
and in urgent need of conservation focus? Are there alternative frameworks 
that visualize these landscapes as coevolutionary and adaptive frontiers, 
where species are not perceived as victims but as successful urban adapters? 
Jamie Lorimer (2015) expresses dissatisfaction with a cross-section of 
studies on wildlife across urban landscapes using present-day urban 
conservation paradigms and emphasizes the need for “anticipatory 
semantics”. He espouses “cosmopolitics for wildlife” and highlights the 
need for “new forms of interspecies responsibility” (pp. 179). 

While this study provides insight into the pressing concerns surrounding 
the persistence of urban wildlife, the debate is far from settled. Frameworks 
based on pure conservation, especially with the establishment of protected 
area systems, may not be appropriate for such landscapes. Instead, these 
urban spaces need to be seen as adaptive margins of the conservation 
realm, where species have coevolved and adapted to the unique conditions 
and resources that these urban habitats provide. As our study indicates, the 
species that presently inhabit the Ridge have lived through several changes 
over the years—they have survived, responded, and adapted to these 
changes. This study accentuates the need to seriously rethink urban nature 
in general and urban planning and to incorporate critical insights and 
explore alternate frameworks that liberate us from the tyranny of green 
constructs. As Schilthuizen (2018) and Lorimer (2015) propose, urban 
wildlife habitats can be reimagined with porosity and some inviolate 
pockets, allowing for further adaptation, with as large a reservoir of genetic 
diversity as possible. With present rates of urbanization—both in terms of 
the number of cities and geographical extent—such a rethinking is crucial. 

 

ACKNOWLEDGEMENTS 

We thank the Department of Forests and Wildlife, Government of the 
NCT of Delhi, for granting us the necessary permits to carry out research in 
the Central Ridge Reserve Forest. We are also grateful to the Centre for 
Urban Ecology and Sustainability (CUES) for providing all the survey 
equipment we used for the study. We thank Shiwani for helping us with 
GIS work, particularly the planning of field surveys and depiction of 
collected data. We are grateful to past and present researchers at CUES for 



Ecology, Economy and Society–the INSEE Journal [52] 

their help and support at various stages of the study. Finally, we thank our 
field assistant, Saurabh Kumar, for his valuable support, hard work, and 
dedication for the entire duration of the fieldwork. 

Ethics Statement: I hereby confirm that this study complies with 
requirements of ethical approvals from the institutional ethics committee 
for the conduct of this research.  

Data Availability statement: The data used to support this research is 
available in a repository and the hyperlinks and persistent identifiers (e.g. 
DOI or accession number) are stated in the paper. 

Conflict of Interest Statement: No potential conflict of interest was 
reported by the author. 

REFERENCES 

Altmann, Jeanne. 1974. “Observational Study of Behavior: Sampling Methods.” 
Behaviour 49 (3): 227–67. https://doi.org/10.1163/156853974X00534  

Ancrenaz, Marc, Andrew J Hearn, Joanna Ross, Rahel Sollmann, and Andreas 
Wilting. 2012. Handbook for Wildlife Monitoring Using Camera-traps. Kota Kinabalu: 
BBEC Publication. 

Andrén, Henrik. 1994. “Effects of Habitat Fragmentation on Birds and Mammals 
in Landscapes with Different Proportions of Suitable Habitat: A Review.” Oikos 71 
(3): 355 66. https://doi.org/10.2307/3545823. 

Arora, Aditya, Pallava Bagla, Nandita Hazarika, Ashish Kothari, Ranjit Lai, 
Subhadra Menon, and Ghazala Shahabuddin. 1991. “The Delhi Ridge Forest: 
Decline and Conservation.” New Delhi: Kalpavriksh. 

Baskaran, Nagarajan, Ganesan Karthikeyan, and Kamaraj Ramkumaran. 2020. 
“Golden Jackal Canis Aureus Linnaeus, 1758 (Mammalia: Carnivora: Canidae) 
Distribution Pattern and Feeding at Point Calimere Wildlife Sanctuary, India.” 
Journal of Threatened Taxa 12 (11): 16460–68. 
https://doi.org/10.11609/jott.4489.12.11.16460-16468.  

Bateman, Philip W, and Patricia Anne Fleming. 2012. “Big City Life: Carnivores in 
Urban Environments.” Journal of Zoology 287 (1): 1–23. 
https://doi.org/10.1111/j.1469-7998.2011.00887.x.  

Baviskar, Amita. 2018. “Urban Jungles: Wilderness, Parks and Their Publics in 
Delhi.” Economic & Political Weekly 53 (2): 46–54. 

———. 2020. Uncivil City: Ecology, Equity and the Commons in Delhi. New Delhi: Sage 
Publications. 

Benítez-López, Ana. 2018. “Animals Feel Safer from Humans in the Dark.” Science 
360 (6394): 1185–6. https://doi.org/10.1126/science.aau1311.  

Bennett, Andrew F, and Denis A Saunders. 2010. “Habitat Fragmentation and 
Landscape Change.” In Conservation Biology for All, edited by Navjot S Sodhi and 

https://doi.org/10.1163/156853974X00534
https://doi.org/10.2307/3545823
https://doi.org/10.11609/jott.4489.12.11.16460-16468
https://doi.org/10.1111/j.1469-7998.2011.00887.x
https://doi.org/10.1126/science.aau1311


[53] Gonji, Chauhan and Babu 

Paul R Ehrlich, 88–106. Oxford: Oxford University Press. 
https://doi.org/10.1093/acprof:oso/9780199554232.003.0006.  

Blair, Robert B. 2001. “Birds and Butterflies Along Urban Gradients in Two 
Ecoregions of the United States: Is Urbanization Creating a Homogenous Fauna?” 
In Biotic Homogenization, edited by Julie L Lockwood and Michael L McKinney, 33–
56. New York: Kluwer Press. https://doi.org/10.1007/978-1-4615-1261-5_3  

Bowe, Patrick. 2009. “The Genius of an Artist: William R Mustoe and the Planting 
of the City of New Delhi and Its Gardens.” Garden History 37 (1): 68–79. 

Burnham, Kenneth P, and David R Anderson, eds. 2002. “Model Selection and 
Multimodel Inference: A Practical Information-theoretic Approach.” New York: 
Springer. 

Cavin, Joëlle Salomon, and Christian A Kull. 2017. “Invasion Ecology Goes to 
Town: From Disdain to Sympathy.” Biological Invasions 19 (12): 3471–87. 
https://doi.org/10.1007/s10530-017-1588-9.  

Champion, Harry G, and S K Seth. 1968. A Revised Survey of the Forest Types of India. 
Dehradun: Natraj Publishers. 

Chauhan, Sonali, Gitanjali Yadav, and Suresh Babu. 2022. “Ecological Networks in 
Urban Forest Fragments Reveal Species Associations between Native and Invasive 
Plant Communities.” Plants 11 (4): 1–17. https://doi.org/10.3390/plants11040541.  

Ćirović, Duško, Aleksandra Penezić, and Miha Krofel. 2016. “Jackals as Cleaners: 
Ecosystem Services Provided by a Mesocarnivore in Human-dominated 
Landscapes.” Biological Conservation 199 (1): 51–5. 
https://doi.org/10.1016/j.biocon.2016.04.027.  

Coffin, Alisa W. 2007. “From Roadkill to Road Ecology: A Review of the 
Ecological Effects of Roads.” Journal of Transport Geography 15 (5): 396–406. 
https://doi.org/10.1016/j.jtrangeo.2006.11.006.  

Crooks, Kevin R. 2002. “Relative Sensitivities of Mammalian Carnivores to Habitat 
Fragmentation.” Conservation Biology 16 (2): 488–502. 
https://doi.org/10.1046/j.1523-1739.2002.00386.x.  

Crowley, Thomas. 2015. Fractured Forest: The Political Ecology of the Delhi Ridge. New 
Delhi: Intercultural Resources. 

DDA. 2010. “Master Plan for Delhi – 2021.” New Delhi: Delhi Development 
Authority. 

Endlicher, Wilfried, Marcel Langner, Markus Hesse, Harald A. Mieg, Ingo 
Kowarik, Patrick Hostert, Elmar Kulke et al. “Urban ecology-definitions and 
concepts.” In Shrinking Cities: Effects on Urban Ecology and Challenges for Urban 
Development, pp. 1-15. Bern, Switzerland: Internationaler Verlag der Wissenschaften, 
2007.  

Fahrig, Lenore. 2003. “Effects of Habitat Fragmentation on Biodiversity.” Annual 
Review of Ecology, Evolution, and Systematics 34 (1): 487–515. 
https://doi.org/10.1146/annurev.ecolsys.34.011802.132419. 

Fiske, I. J., & Chandler, R. B. (2011). Unmarked: An R package for fitting 

https://doi.org/10.1093/acprof:oso/9780199554232.003.0006
https://doi.org/10.1007/978-1-4615-1261-5_3
https://doi.org/10.1007/s10530-017-1588-9
https://doi.org/10.3390/plants11040541
https://doi.org/10.1016/j.biocon.2016.04.027
https://doi.org/10.1016/j.jtrangeo.2006.11.006
https://doi.org/10.1046/j.1523-1739.2002.00386.x
https://doi.org/10.1146/annurev.ecolsys.34.011802.132419


Ecology, Economy and Society–the INSEE Journal [54] 

hierarchical models of wildlife occurrence and abundance. Journal of Statistical 
Software, 43(10), 1–23. https://doi.org/10.18637/jss.v043.i10.  

Forman, Richard T T, and Lauren E Alexander. 1998. “Roads and Their Major 
Ecological Effects.” Annual Review of Ecology and Systematics 29 (1): 207–31. 
https://doi.org/10.1146/annurev.ecolsys.29.1.207.   

Fuller, Todd K, Audrone R Biknevicius, Pieter W Kat, Blaire van Valkenburgh, and 
Robert K Wayne. 1989. “The Ecology of Three Sympatric Jackal Species in the Rift 
Valley of Kenya.” African Journal of Ecology 27 (4): 313–23. 
https://doi.org/10.1111/j.1365-2028.1989.tb01025.x.  

Fuller, Todd K, Stephen DeStefano, and Paige S Warren. 2010. “Carnivore 
Behavior and Ecology, and Relationship to Urbanization.” In Urban Carnivores: 
Ecology, Conflict, and Conservation, edited by Stanley D Gehrt, Seth P D Riley, and 
Brian L Cypher, 13–19. Baltimore: The Johns Hopkins University Press. 

Gandhiok, Jasjeev. 2023. “Delhi: Golden Jackals Spotted at Yamuna Biodiversity 
Park.” Hindustan Times, April 11, 2023. 
https://www.hindustantimes.com/cities/delhi-news/golden-jackals-sighted-for-
the-first-time-in-delhi-s-yamuna-biodiversity-park-indicating-successful-rewilding-
efforts-101681151965374.html.  

Gaynor, Kaitlyn M, Cheryl E Hojnowski, Neil H Carter, and Justin S Brashares. 
2018. “The Influence of Human Disturbance on Wildlife Nocturnality.” Science 360 
(6394): 1232–35. https://doi.org/10.1126/science.aar7121    

Gehrt, Stanley D, Seth P D Riley, and Brian L Cypher, eds. 2010. Urban Carnivores: 
Ecology, Conflict and Conservation. Maryland: The Johns Hopkins University Press. 

Gehrt, Stanley D. 2007. “Ecology of Coyotes in Urban Landscapes.” 12th Wildlife 
Damage Management Conference Proceedings 63: 303–11.  

Gil-Fernández, Margarita, Robert Harcourt, Thomas Newsome, Alison Towerton, 
and Alexandra Carthey. 2020. “Adaptations of the Red Fox (Vulpes Vulpes) to 
Urban Environments in Sydney, Australia.” Journal of Urban Ecology 6 (1): 1–9. 
https://doi.org/10.1093/jue/juaa009.  

Giménez Gómez, Victoria C, José R Verdú, Andrés Gómez-Cifuentes, Fernando Z 
Vaz-de-Mello, and Gustavo A Zurita. 2018. “Influence of Land Use on the Trophic 
Niche Overlap of Dung Beetles in the Semideciduous Atlantic Forest of 
Argentina.” Insect Conservation and Diversity 11 (6): 554–64. 
https://doi.org/10.1111/icad.12299.  

Government of India. 2011. Census of India 2011. New Delhi: Government of India. 

———. 2022. “The Wild Life (Protection) Amendment Act.” New Delhi: Ministry 
of Law and Justice. 

Grimm, Nancy B, Stanley H Faeth, Nancy E Golubiewski, Charles L Redman, 
Jianguo Wu, Xuemei Bai, and John M Briggs. 2015. “Global Change and the 
Ecology of Cities.” Science 319 (5864): 756–60. 
https://doi.org/10.1126/science.1150195.  

https://doi.org/10.18637/jss.v043.i10
https://doi.org/10.1146/annurev.ecolsys.29.1.207
https://doi.org/10.1111/j.1365-2028.1989.tb01025.x
https://www.hindustantimes.com/cities/delhi-news/golden-jackals-sighted-for-the-first-time-in-delhi-s-yamuna-biodiversity-park-indicating-successful-rewilding-efforts-101681151965374.html
https://www.hindustantimes.com/cities/delhi-news/golden-jackals-sighted-for-the-first-time-in-delhi-s-yamuna-biodiversity-park-indicating-successful-rewilding-efforts-101681151965374.html
https://www.hindustantimes.com/cities/delhi-news/golden-jackals-sighted-for-the-first-time-in-delhi-s-yamuna-biodiversity-park-indicating-successful-rewilding-efforts-101681151965374.html
https://doi.org/10.1126/science.aar7121
https://doi.org/10.1093/jue/juaa009
https://doi.org/10.1111/icad.12299
https://doi.org/10.1126/science.1150195


[55] Gonji, Chauhan and Babu 

Grinder, Martha I, and Paul R Krausman. 2001. “Home Range, Habitat Activity of 
Use, and Nocturnal in an Urban Environment.” The Journal of Wildlife Management 65 
(4): 887–98. https://doi.org/10.2307/3803038  

Gupta, Sreetamaa, Aritri Sanyal, Goutam K Saha, and A K Ghosh. 2016. “Diurnal 
Activity Pattern of Golden Jackal (Canis Aureus Linn.) in an Urban Landscape of 
Kolkata, India.” Proceedings of the Zoological Society 69 (1): 75–80. 
https://doi.org/10.1007/s12595-014-0119-2.  

Hepcan, Çiǧdem Coşkun. 2012. “Identifying and Assessing of the Red Fox (Vulpes 
Vulpes) Habitats in Çeşme-Urla Peninsula.” Journal of Food, Agriculture and 
Environment 10 (1): 726–31. 

Hoffmann, Mike, Janosch Arnold, J W Duckworth, Yadvendradev V Jhala, Jan F 
Kamler, and Miha Krofel. 2018. “Canis Aureus – Global Assessment.” IUCN Red 
List of Threatened Species 8235. 

Iannarilli, Fabiola, John Erb, Todd W Arnold, and John R Fieberg. 2021. 
“Evaluating Species-specific Responses to Camera-trap Survey Designs.” Wildlife 
Biology 2021 (1): 1–12. https://doi.org/10.2981/wlb.00726.  

İbiş, Osman, Eren Aksöyek, Servet Özcan, and Coşkun Tez. 2015. “A Preliminary 
Phylogenetic Analysis of Golden Jackals (Canis Aureus) (Canidae: Carnivora: 
Mammalia) from Turkey Based on Mitochondrial D-loop Sequences.” Vertebrate 
Zoology 65 (3): 391–7. https://doi.org/10.3897/vz.65.e31532.  

Jensen, Per. 2007. The Behavioural Biology of Dogs. Trowbridge: Cromwell Press. 
https://doi.org/10.1079/9781845931872.0000  

Jhala, Yadvendradev V, and P D Moehlman. 2004. “Golden Jackal Canis Aureus.” 
In Canids: Foxes, Wolves, Jackals and Dogs Status Survey and Conservation Action Plan, 
edited by Claudio Sillero-Zubiri, Michael Hoffmann, and David W MacDonald, 
156–60. Gland: IUCN/SSC Canid Specialist Group. 

Kays, Roland, Brian S Arbogast, Megan Baker-Whatton, Chris Beirne, Hailey M 
Boone, Mark Bowler, Santiago F Burneo, et al. 2020. “An Empirical Evaluation of 
Camera Trap Study Design: How Many, How Long and When?” Methods in Ecology 
and Evolution 11 (6): 700–13. https://doi.org/10.1111/2041-210X.13370.  

Kebede, Yigrem. 2017. “A Review On: Distribution, Ecology and Status of Golden 
Jackal (Canis Aureus) in Africa.” Journal of Natural Sciences Research 7 (1): 32–43. 

Kowarik, Ingo. 2011. “Novel Urban Ecosystems, Biodiversity, and Conservation.” 
Environmental Pollution 159 (8–9): 1974–83. 
https://doi.org/10.1016/j.envpol.2011.02.022.  

Krishen, Pradip. 2006. Trees of Delhi: A Field Guide. New Delhi: Dorling Kindersley 
(India) Pvt Ltd. 

Kumar, Arun. 1997. “Fauna of Delhi: An Overview.” In Fauna of Delhi, edited by 
The Director, Zoological Survey of India, 1–16. Calcutta: Zoological Survey of 
India. 

Lorimer, Jamie. 2015. Wildlife in the Anthropocene: Conservation after Nature. 
Minneapolis: University of Minnesota Press. 
https://doi.org/10.5749/minnesota/9780816681075.001.0001  

https://doi.org/10.2307/3803038
https://doi.org/10.1007/s12595-014-0119-2
https://doi.org/10.2981/wlb.00726
https://doi.org/10.3897/vz.65.e31532
https://doi.org/10.1079/9781845931872.0000
https://doi.org/10.1111/2041-210X.13370
https://doi.org/10.1016/j.envpol.2011.02.022
https://doi.org/10.5749/minnesota/9780816681075.001.0001


Ecology, Economy and Society–the INSEE Journal [56] 

Lowry, Hélène, Alan Lill, and Bob B M Wong. 2013. “Behavioural Responses of 
Wildlife to Urban Environments.” Biological Reviews 88 (3): 537–49. 
https://doi.org/10.1111/brv.12012.  

Luniak, Maciej. 2004. “Synurbization: Adaptation of Animal Wildlife to Urban 
Development.” Proceedings 4th International Urban Wildlife Symposium.  

MacDonald, David W, Liz A D Campbell, Jan F Kamler, Jorgelina Marino, 
Geraldine Werhahn, and Claudio Sillero-Zubiri. 2019. “Monogamy: Cause, 
Consequence, or Corollary of Success in Wild Canids?” Frontiers in Ecology and 
Evolution 7 (September): 1–28. https://doi.org/10.3389/fevo.2019.00341. 

Mackenzie, D.I., and J.A. Royle. 2005. ‘Designing Occupancy Studies: General 
Advice and Allocating Survey Effort’. Journal of Applied Ecology 42: 1105–14. 
https://doi.org/10.1111/j.1365-2664.2005.01098.x.   

Mackenzie, D I, and James D Nichols. 2004. “Occupancy as a Surrogate for 
Abundance Estimation.” Animal Biodiversity and Conservation 27 (1): 461–7. 

Mackenzie, D I, James D Nicolas, Jeffrey Andrew Royle, K H Pollock, Larissa L 
Bailey, and James E Hines. 2006. Occupancy Estimation and Modeling: Inferring Patterns 
and Dynamics of Species Occurrence. Burlington: Elsevier. 

Magle, Seth B, Perla Reyes, Jun Zhu, and Kevin R Crooks. 2010. “Extirpation, 
Colonization, and Habitat Dynamics of a Keystone Species along an Urban 
Gradient.” Biological Conservation 143 (9): 2146–55. 
https://doi.org/10.1016/j.biocon.2010.05.027.  

Maheshwari, J K. 1963. Flora of Delhi. New Delhi: Council of Scientific and 
Industrial Research. 

Majumder, Sreejani Sen, Manabi Paul, Shubhra Sau, and Anindita Bhadra. 2016. 
“Denning Habits of Free-Ranging Dogs Reveal Preference for Human Proximity.” 
Scientific Reports 6 (August): 1–8. https://doi.org/10.1038/srep32014.  

Mann, Michael, and Samiksha Sehrawat. 2009. “A City With a View: The 
Afforestation of the Delhi Ridge, 1883–1913.” Modern Asian Studies 43 (2): 543–70. 
https://doi.org/10.1017/S0026749X07002867.  

Markov, Georgi. 2012. “Golden Jackal (Canis Aureus L.) in Bulgaria: What Is 
Going On?” Acta Zoologica Bulgarica 64 (1): 67–71. 

Marks, Clive A, and Tim E Bloomfield. 1999. “Distribution and Density Estimates 
for Urban Foxes (Vulpes Vulpes) in Melbourne: Implications for Rabies Control.” 
Wildlife Research 26 (6): 763–75. https://doi.org/10.1071/WR98059  

McCallum, Jamie. 2013. “Changing Use of Camera Traps in Mammalian Field 
Research: Habitats, Taxa and Study Types.” Mammal Review 43 (3): 192–206. 
https://doi.org/10.1111/j.1365-2907.2012.00216.x.  

McDonnell, Mark J, and Amy K Hahs. 2015. “Adaptation and Adaptedness of 
Organisms to Urban Environments.” Annual Review of Ecology, Evolution, and 
Systematics 46 (1): 261–80. https://doi.org/10.1146/annurev-ecolsys-112414-
054258.  

https://doi.org/10.1111/brv.12012
https://doi.org/10.3389/fevo.2019.00341
https://doi.org/10.1111/j.1365-2664.2005.01098.x
https://doi.org/10.1016/j.biocon.2010.05.027
https://doi.org/10.1038/srep32014
https://doi.org/10.1017/S0026749X07002867
https://doi.org/10.1071/WR98059
https://doi.org/10.1111/j.1365-2907.2012.00216.x
https://doi.org/10.1146/annurev-ecolsys-112414-054258
https://doi.org/10.1146/annurev-ecolsys-112414-054258


[57] Gonji, Chauhan and Babu 

McDonnell, Mark J. 2011. “The History of Urban Ecology An Ecologist’s 
Perspective.” In Urban Ecology: Patterns, Processes, and Applications, edited by Jari 
Niemelä, 5–13. Oxford: Oxford University Press. 
https://doi.org/10.1093/acprof:oso/9780199563562.003.0002  

Meredith, M. and Ridout, M.S. 2014. “The Overlap Package”. Central R Archive 
Network Online. https://kar.kent.ac.uk/41474/  

Moehlman, Patricia D. 1987. “Social Organization in Jackals.” American Scientist 75 
(4): 366–75. 

Mohan, Madan. 2001. ‘Gis-Based Spatial Information Integration, Modeling And 
Digital Mapping: A New Blend of Tool for Geospatial Environmental Health 
Analysis for Delhi’. International Archives of Photogrammetry and Remote Sensing 34: 69–
70. 

Morell, Virginia. 2013. “Predators in the “Hood.’” Science 341 (6152): 1332–5. 
https://doi.org/10.1126/science.341.6152.1332.  

Mukherjee, Aditi, Honnavalli Nagaraj Kumara, and Subramanian Bhupathy. 2018. 
“Golden Jackal’s Underground Shelters: Natal Site Selection, Seasonal Burrowing 
Activity and Pup Rearing by a Cathemeral Canid.” Mammal Research 63 (3): 325–39. 
https://doi.org/10.1007/s13364-018-0356-2.  

Mukherjee, Shomita, S P Goyal, A J T Johnsingh, and M R P Leite Pitman. 2004. 
“The Importance of Rodents in the Diet of Jungle Cat (Felis Chaus), Caracal 
(Caracal Caracal) and Golden Jackal (Canis Aureus) in Sariska Tiger Reserve, 
Rajasthan, India.” Journal of Zoology 262 (4): 405–11. 
https://doi.org/10.1017/S0952836903004783.  

Mukherjee, Shomita. 1998. “Habitat Use by Sympatric Small Carnivores in Sariska 
Tiger Reserve, Rajasthan, India.” PhD diss., Saurashtra University. 

Negi, Tripti. 2014. “Review on Current Worldwide Status, Distribution, Ecology 
and Dietary Habits of Golden Jackal, Canis Aureus.” Octa Journal of Environmental 2 
(4): 338–59. 

Nichols, James D, Allan F O’Connell, and K Ullas Karanth. 2011. “Camera Traps 
in Animal Ecology and Conservation: What’s Next?” In Camera Traps in Animal 
Ecology: Methods and Analyses, edited by Allan F O’Connell, James D Nichols, and K 
Ullas Karanth, 253–63. Tokyo: Springer. https://doi.org/10.1007/978-4-431-
99495-4_14  

Niedballa, Jürgen, Rahel Sollmann, Azlan bin Mohamed, Johannes Bender, and 
Andreas Wilting. 2015. “Defining Habitat Covariates in Camera-trap Based 
Occupancy Studies.” Scientific Reports 5 (1): 1–10. 
https://doi.org/10.1038/srep17041.  

Nurvianto, Sandy, Muhammad Ali Imron, and Sven Herzog. 2015. “Activity 
Patterns and Behaviour of Denning Dholes (Cuonalpinus) in a Dry Deciduous 
Forest of East Java, Indonesia.” Bulletin of Environment, Pharmacology and Life Sciences 4 
(12): 45–54. 

 

https://doi.org/10.1093/acprof:oso/9780199563562.003.0002
https://kar.kent.ac.uk/41474/
https://doi.org/10.1126/science.341.6152.1332
https://doi.org/10.1007/s13364-018-0356-2
https://doi.org/10.1017/S0952836903004783
https://doi.org/10.1007/978-4-431-99495-4_14
https://doi.org/10.1007/978-4-431-99495-4_14
https://doi.org/10.1038/srep17041


Ecology, Economy and Society–the INSEE Journal [58] 

O’Connell, Allan F, and Larissa L Bailey. 2011. “Inference for Occupancy and 
Occupancy Dynamics.” In Camera Traps in Animal Ecology: Methods and Analyses, 
edited by Allan F O’Connell, James D Nichols, and K Ullas Karanth, 191–205. 
Tokyo: Springer. 

Palmer, Clare. 2003. “Colonization, Urbanization, and Animals.” Philosophy and 
Geography 6 (1): 47–58. https://doi.org/10.1080/1090377032000063315.  

Parker, Sophie S. 2015. “Incorporating Critical Elements of City Distinctiveness 
into Urban Biodiversity Conservation.” Biodiversity and Conservation 24 (3): 683–700. 
https://doi.org/10.1007/s10531-014-0832-1.  

Pascual-Rico, Roberto, José Antonio Sánchez-Zapata, Joan Navarro, Sergio Eguía, 
José Daniel Anadón, and Francisco Botella. 2020. “Ecological Niche Overlap 
between Co-occurring Native and Exotic Ungulates: Insights for a Conservation 
Conflict.” Biological Invasions 22 (8): 2497–508. https://doi.org/10.1007/s10530-
020-02265-x. 

Patil, Vinayak Krishna, and Yadvendradev V Jhala. 2008. “Movement Patterns and 
Habitat Use of Golden Jackal Canis Aureus in Bhal Region of Gujarat.” Journal of 
the Bombay Natural History Society 105 (2): 209–43. 

Pickett, Steward T A, William R Burch Jr, Shawn E Dalton, Timothy W Foresman, 
J Morgan Grove, and Rowan Rowntree. 1997. “A Conceptual Framework for the 
Study of Human Ecosystems in Urban Areas.” Urban Ecosystems 1 (1): 185–99. 
https://doi.org/10.1023/A:1018531712889  

Planning Commission. 2009. Delhi Development Report. New Delhi: Government of 
India. 

Prange, Suzanne, Stanley D Gehrt, and Ernie P Wiggers. 2004. “Influences of 
Anthropogenic Resources on Raccoon (Procyon, Lotor) Movements and Spatial 
Distribution.” Journal of Mammalogy 85 (3): 483–90. https://doi.org/10.1644/BOS-
121.  

Punjab Government. 1883. “Gazetteer of the Delhi District.” Punjab: Punjab 
Government. 

Rodewald, Amanda D, and Stanley D Gehrt. 2014. “Wildlife Population Dynamics 
in Urban Landscapes.” In Urban Wildlife Conservation, edited by Robert A McCleery, 
Christopher E Moorman, and M Nils Peterson, 117–47. New York: Springer. 
https://doi.org/10.1007/978-1-4899-7500-3_8  

Rovero, Francesco, Mathias Tobler, and James Sanderson. 2010. “Camera-trapping 
for Inventorying Terrestrial Vertebrates.” In Manual on Field Recording Techniques and 
Protocols for All Taxa Biodiversity Inventories and Monitoring, edited by Jutta Eymann, 
Jérôme Degreef, Christoph Häuser, Juan Carlos Monje, Yves Samyn, and Didier 
VandenSpiegel, 100–128. Brussels: ABC Taxa.  

Rowcliffe, J Marcus, and C Carbone. 2008. “Surveys Using Camera Traps: Are We 
Looking to a Brighter Future?” Animal Conservation 11 (3): 185–6. 
https://doi.org/10.1111/j.1469-1795.2008.00180.x  

 

https://doi.org/10.1080/1090377032000063315
https://doi.org/10.1007/s10531-014-0832-1
https://doi.org/10.1023/A:1018531712889
https://doi.org/10.1644/BOS-121
https://doi.org/10.1644/BOS-121
https://doi.org/10.1007/978-1-4899-7500-3_8
https://doi.org/10.1111/j.1469-1795.2008.00180.x


[59] Gonji, Chauhan and Babu 

Ryan, Amy M, and Sarah R Partan. 2014. “Urban Wildlife Behaviour.” In Urban 
Wildlife Conservation, edited by Robert A McCleery, Christopher E Moorman, and M 
Nils Peterson, 149–73. New York: Springer. https://doi.org/10.1007/978-1-4899-
7500-3_9  

Sanyal, Asok K, J K Dey, and P L Kankane. 2011. “The Jackals of Tollygunge 
Club, Kolkata.” Records of the Zoological Survey of India 111 (1): 37. 
https://doi.org/10.26515/rzsi/v111/i1/2011/158890.  

Schilthuizen, Menno. 2018. Darwin Comes to Town: How the Urban Jungle Drives 
Evolution. London: Quercus Editions Ltd. 

Shochat, Eyal, Paige S Warren, Stanley H Faeth, Nancy E McIntyre, and Diane 
Hope. 2006. “From Patterns to Emerging Processes in Mechanistic Urban 
Ecology.” Trends in Ecology and Evolution 21 (4): 186–91. 
https://doi.org/10.1016/j.tree.2005.11.019.  

Singh, Aakriti, Aditi Mukherjee, Sumit Dookia, and Honnavalli Nagaraj Kumara. 
2016. “High Resource Availability and Lack of Competition Have Increased 
Population of a Meso-carnivore—A Case Study of Golden Jackal in Keoladeo 
National Park, India.” Mammal Research 61 (3): 209–19. 
https://doi.org/10.1007/s13364-016-0267-z.  

Sinha, Anindya, and Maan Barua. 2020. “Nonhuman Lifeworlds in Urban India.” 
The Philosopher 108 (1): 22–27. 

Sinha, G N, ed. 2014. An Introduction to the Delhi Ridge. New Delhi: Department of 
Forests and Wildlife, Government of NCT of Delhi. 

Sol, Daniel, Oriol Lapiedra, and Cesar González-Lagos. 2013. “Behavioural 
Adjustments for a Life in the City.” Animal Behaviour 85 (5): 1101–12. 
https://doi.org/10.1016/j.anbehav.2013.01.023.  

Solanki, Srishti. 2014. “Evolution of Statutory and Management Framework.” In 
An Introduction to the Delhi Ridge, edited by G N Sinha, 29–46. New Delhi: 
Department of Forests and Wildlife, Government of NCT of Delhi. 

Sollmann, Rahel. 2018. “A Gentle Introduction to Camera-trap Data Analysis.” 
African Journal of Ecology 56 (4): 740–9. https://doi.org/10.1111/aje.12557.   

Southern, H N. 1955. “Nocturnal Animals.” Scientific American 193 (4): 88–99. 
https://doi.org/10.1038/scientificamerican1055-88  

Stamp Dawkins, Marian. 2008. Observing Animal Behaviour: Design and Analysis 
of Quantitative Data. Oxford: Oxford University Press. 
https://doi.org/10.1093/acprof:oso/9780198569350.001.0001.  

Sud, Megha. 2017. Political Ecology of the Ridge: The Establishment and Contestation of 
Urban Forest Conservation in Delhi. Stuttgart: Franz Steiner Verlag. 
https://doi.org/10.25162/9783515117159  

The Times of India. 1950. “Delhi Diary.” ProQuest Historical Newspapers. 

Thomas, Chris D. 2017. Inheritors of the Earth. London: Penguin. 

Tinbergen, Niko. 1963. “On Aims and Methods of Ethology.” Zeitschrift fu  r 
Tierpsychologie 20 (1): 410–33. https://doi.org/10.1111/j.1439-0310.1963.tb01161.x  

https://doi.org/10.1007/978-1-4899-7500-3_9
https://doi.org/10.1007/978-1-4899-7500-3_9
https://doi.org/10.26515/rzsi/v111/i1/2011/158890
https://doi.org/10.1016/j.tree.2005.11.019
https://doi.org/10.1007/s13364-016-0267-z
https://doi.org/10.1016/j.anbehav.2013.01.023
https://doi.org/10.1111/aje.12557
https://doi.org/10.1038/scientificamerican1055-88
https://doi.org/10.1093/acprof:oso/9780198569350.001.0001
https://doi.org/10.25162/9783515117159
https://doi.org/10.1111/j.1439-0310.1963.tb01161.x


Ecology, Economy and Society–the INSEE Journal [60] 

Trouwborst, Arie, Miha Krofel, and John D C Linnell. 2015. “Legal Implications of 
Range Expansions in a Terrestrial Carnivore: The Case of the Golden Jackal (Canis 
Aureus) in Europe.” Biodiversity and Conservation 24 (10): 2593–610. 
https://doi.org/10.1007/s10531-015-0948-y.  

United Nations. 2019. “World Urbanization Prospects: The 2018 Revision.” New 
York: Department of Economic and Social Affairs, Population Division.  

Vanak, Abi Tamim, and Matthew E Gompper. 2009. “Dogs Canis Familiaris as 
Carnivores: Their Role and Function in Intraguild Competition.” Mammal Review 
39: 265–83. https://doi.org/10.1111/j.1365-2907.2009.00148.x  

Varsha. 2014. “The Delhi Ridge.” In An Introduction to the Delhi Ridge, edited by G N 
Sinha, 1–28. New Delhi: Department of Forests and Wildlife, Government of NCT 
of Delhi. 

Venn, Stephen J, and Jari K Niemelä. 2004. “Ecology in a Multidisciplinary Study 
of Urban Green Space: The URGE Project.” Boreal Environment Research 9 (1): 479–
89. 

Yumnam, Bibek, Tripti Negi, Jesús E Maldonado, Robert C Fleischer, and 
Yadvendradev V Jhala. 2015. “Phylogeography of the Golden Jackal (Canis Aureus) 
in India.” PLoS ONE 10 (9): 1–18. https://doi.org/10.1371/journal.pone.0138497.  

ZSI. 1997. Fauna of Delhi. Calcutta: Zoological Survey of India. 

 

https://doi.org/10.1007/s10531-015-0948-y
https://doi.org/10.1111/j.1365-2907.2009.00148.x
https://doi.org/10.1371/journal.pone.0138497

