Temples and bats in a homogeneous agriculture landscape: Importance of microhabitat availability, disturbance and land use for bat conservation.

Cave-dwelling bats widely use anthropogenic structures such as temples in south Asia as roosting and nursery sites. Such roosts are constantly under threat, even more so after the COVID-19 pandemic. Despite the importance of such roosts, there is no detailed understanding of what makes temples favorable for bats and the critical factors for their persistence. Here we relate temple microhabitat characteristics and land use around ancient temples (>400 years) to bat species richness and abundance in the Tamiraparani river basin of south India. Temples were selected for sampling along the river basin based on logistics and permission to access them. We counted bats at the roost in the mornings and late afternoons from inside the temples. Temple characteristics such as dark rooms, walkways, crevices, towers, and disturbances to the roosts were recorded. Based on European Space Agency land use classifications, we recorded land use such as crops, trees, scrub, grassland, urban areas, and water availability within a 5 km radius of the temple. Generalized Linear Mixed Models were used to relate the counts in temples with microhabitats and land use. We sampled 59 temples repeatedly across 5 years which yielded a sample of 246 survey events. The total number of bats counted was 20,211, of which Hipposideros speoris was the most common (9,715), followed by Rousettus leschenaultii (5,306), Taphozous melanopogon (3,196), Megaderma lyra (1,497), Tadarida aegyptiaca (303), Pipistrellus sp. (144) and Rhinopoma hardwickii (50). About 39% of the total bats occurred in dark rooms and 51% along walkways. Species richness and total abundance were related to the availability of dark rooms and the number of buildings in the temple. Land use elements only had a weak effect, but scrub and grassland, even though they were few, are critical for bats. We conclude that retaining undisturbed dark rooms with small exits in temples and other dimly lit areas and having natural areas around temples are vital for bat conservation.

Introduction

Many studies suggest that human-modified landscapes with a heterogeneous mosaic of different land uses can preserve species-rich bat assemblages [1-7]. This is possible with the concurrent availability of microhabitats that bats use as roosting sites ranging from caves, crevices, tree cavities, foliage, and several types of anthropogenic structures, including road bridges and buildings [8, 9]. Although these microhabitats are important features of a bat’s environment, the selection of specific roost sites can strongly impact the survival and fitness of bats [10-12]. However, emerging challenges such as changing agricultural practices and climate change pose significant threats to bat assemblages and populations in human-dominated landscapes [13, 14]. They could negatively impact human populations by altering the ecosystem services provided by bats [15, 16]. Moreover, bats that roost in anthropogenic structures are vulnerable to disturbance since, just like cave roosting species, they often form large, concentrated aggregations attracting attention [17]. Most studies on bats using anthropogenic structures have come from urban areas and under temperate conditions [9, 15, 18–20]. While bats are still poorly studied in the Paleotropics [21], the selection of microhabitats and landscape-level influences on their population remains unknown [22-24]. Moreover, most studies on bat–habitat relationships are conducted at fixed spatial scales [25], while recent studies have shown the importance of scale for bats [26, 27].

In South Asia, several species of both insectivorous and frugivorous bats use temples as roosting sites [28, 29]. In south India, numerous very old (>400y) temples dot the landscape and form important bat habitats [30]. These temples are built of granite stones with several enclosures and have towers (Gopuram) made of bricks. The temples are not air-conditioned, and the temperature inside is stable and warm (320–330 C) and relatively constant across the day (personal obs T.Ganesh). Bats use these temples as roost and nursery and enter and leave through numerous open vents that circulate air inside the temple.

Bat use of anthropogenic structures, such as temples, brings them into conflict with humans and makes bat conservation in the human-dominated landscape a challenge. These temples dominate the human-agricultural landscape where water-intensive paddy and banana cultivation have replaced traditional crops. Such agricultural practices are considered a major threat to bats [14]. With irrigated agriculture spreading to dry regions and with a predicted increase in rains [31], threats to the bat population can rise in the future. Bats in many temples have dwindled [32, 33], and many are driven away during temple renovation [34]. The changing attitude of people towards bats has led many temples to be made "bat-proof" by placing nets and power washing roost areas with chlorinated chemicals to prevent bats from colonizing. Further, the recent COVID-19 pandemic has made them more vulnerable to disturbance and direct persecution. In India, only two out of 120 species of bats come under the strict Wildlife Protection Act 1972, while others under the International Union for Conservation and Nature (IUCN) category remain unprotected.

Management decisions for bats and buildings, including the timing of maintenance activities, restriction of human use, and bat deterrence or exclusion, require information on the roosting habits of temple-roosting bats. In addition, studies in temperate regions have shown how features of bridges and buildings can help identify sites for conservation and thereby reduce bat-human conflict [9, 20]. Such efforts to conserve bats in tropical regions are needed. It’s therefore essential to understand what attributes of temples are critical for bats and how such information can help bat conservation efforts in temples.

More than 50% of the 35 bat species found in Tamil Nadu state dwell in caves and old buildings [28]. All bats are insectivores, except Rousettus leschenaultii, a frugivore. In the paddy agriculture-dominated landscape of south India, ancient temples are often the only habitat for several bat species. They provide critical ecosystem services for agriculture and, in south Asia, can save up to US$1.2 million per year from insect depredation [35] and consume large quantities of mosquitoes, thus providing human health benefits [36]. However, despite the importance of bats in providing key ecosystem services in human-dominated landscapes [34, 37], no systematic study on the ecology of bats in temples exists, while behavioral studies have been carried out on them in South India [38, 39]. Moreover, with populations of bats dwindling, no attempt to monitor bats in the temples or how important temples are in an agricultural matrix to bats has been investigated. Much of the bat diversity in human-dominated landscapes can be attributed to habitat diversity and roost site availability in an area. However, in tropical agricultural regions with rice paddies and bananas dominating the landscape providing low habitat heterogeneity, it would be interesting to compare the role of land use elements and availability of microhabitats on bat species richness and abundances.

We initiated a bat monitoring program in 2012 in temples of south India to understand why bats use temples and how the population is responding to disturbance and land use. In this study, we identify temples used by bats for roosting in a semi-arid region characterized by the winter monsoon (Oct to Dec) and agriculture dominated by irrigated rice paddy and banana for 6–8 months a year. The rest of the time, the land is fallow in most places. We recorded the species composition, rarity, and abundance of bats in temples and identified factors that affect bat colonization, such as microhabitat availability, disturbance in temples and landscape features surrounding the temples. This paper tests three major hypotheses: 1. Disturbance at the roost negatively affects species richness and abundance of bats, 2. Roost characteristics are more important than landscape features for bats to colonize temples, and 3. In a homogeneous monoculture cropping, species occurrence and abundance are independent of the spatial availability of habitats around roosts.

Materials and methods

Study area

The study was conducted between lat 8.739, long 77.443 and lat 8.306, long 77.913 in Tirunelveli & Thoothukudi districts of Tamil Nadu state in South India. The semi-arid region experiences the northeast winter monsoons between Oct-Dec while the rest of the months are dry. Several ancient temples built during the Chola and Pandya dynasties 500 to 1000 years back exist in the districts along the perennial Tamiraparani river and its tributaries. The river and numerous interconnected village ponds are the lifelines for paddy and banana-dominated agriculture, supporting a dense human population in the region. The ancient temples are built of large granite stones and provide a habitat for several bat species. There are over 4041 temples [40] in the study area, with 2–3 villages having one ancient temple. A typical temple structure includes towers at the four cardinal entrances and one over the deity. The deity chamber is lit by an oil lamp and an electric bulb, a walkway surrounding the deity chamber for circumambulation by devotees, and several rooms, including storerooms and a kitchen, which are dark and rarely frequented by people.

Temple selection

We selected 59 temples from the Tamiraparani basin and surveyed them with 2–4 personnel between 06:00–12:00 h and again between 16:00 to 17:00 h to detect the presence of bats. The survey was done from May to Dec in 2012, 2013, 2014, 2018 and 2019, totaling 246 survey efforts, with most temples revisited each year. All species of Yinpterochiroptera and Yangochiroptera bats found roosting inside the temples were recorded. First, the presence of bats in the temple was confirmed by several signs such as droppings and smell, interviewing local people, temple priests, and caretakers. Later, using a red LED light, we scanned the dark corners, temple roof, abandoned & isolated rooms inside the temple and the temple tower for chiropteran presence. Once bat colonies were detected, we counted the number of individuals, and in some cases, especially in closed buildings, we took photographs to estimate them.

Temple features

a. Microhabitat. There were multiple microhabitats inside the temples where bats could roost. Dark rooms: closed rooms with few small openings, low natural light and not frequently used by people; Crevices: stone pillars that supported the roof created gaps between the joints for some species to roost; Temple towers: conical structures built over the entrance or the deity and Walkways: wide walking space around the deity room covered by a stone roof. We also counted the number of independent buildings in the temple complex, which bats use to move between them during the day due to disturbance. In 2012 we did a one-time measurement of temperature and humidity while sampling for bats and found no difference between temples and therefore did not consider them in the analysis. We could not set dataloggers for continuous monitoring because of objections by the temple authorities to placing any devices inside temples. We estimated temple size based on visual observation; those with deity rooms and walkways were considered small, those with deity rooms, walkways and towers considered medium-sized, and those having deity rooms, multiple towers, and two walkways (inside and outside) were considered large. We also recorded the number of trees in the temple complex and open wells used by bats.

b. Disturbance. We classified disturbances such as renovation, construction, power washing, and other maintenance activities as 0 or 1. A temple received a value of 0 when there was no disturbance and a value of 1 when there was a disturbance. We also collected information on the number of temple visitors and classified temples on a scale of 1 to 3, with 1 indicating few visitors and 3 indicating having large crowds.

Land-use features

We used the composite analysis of land cover available using high-resolution (10m) European Space Agency (ESRI) data for 2020 [41] to identify major land uses around temples in the region. Unfortunately, cloud cover prevented us from deciphering land use change across each sampling period. Water, scrub, grassland, bare ground, flooded area, crop, urban and tree-covered areas were the primary land use identified [41]. We combined bare ground and grassland into grassland habitat and water and flooded area into water habitat after verifying the land use on the ground and using Google Earth. Crop availability each season was obtained from Google Earth images when available and further substantiated by informal interviews with farmers around each temple. If crops were present, we used the ESRI calculated value of crop area in the analysis, and if not, we considered it zero. The extent of other land uses has not changed much during the sampling period based on the data available with the district administration (Statistical handbook of Tirunelveli district 2009–2019).

Since the spatial scale is a driver for bat foraging [27] we categorized land use at 500m, 1km, 3km and 5km from the temple based on the foraging distance of bats from their roost [38]. We also recorded trees and water availability (open wells) used by bats in the temple complex for land-use analysis at 0km level.

Species identification and abundance

We identified bats to the species level using the bat identification guide by Bates and Harrison (1997). Bats were counted at the roost when they were least active (late morning and early evenings) using a flashlight with red filters. We took photographs as soon as bat aggregations were noticed in a particular microhabitat and later counted the bats in the picture to estimate abundance [42]. Several images taken in quick succession gave a complete count. We could not do a roost exit count in the evening as it was not logistically possible to be in each temple at the right time for the count. Binoculars helped to see the species’ features for identification.

Data analysis

We used total roost count (abundance) and total species richness at each temple for the analysis. We constructed a linear model to explain the species richness and abundance of bats in the temples based on the microhabitat and landscape features. We based the variables used in our linear models on our observations made in 2012. We eliminated strongly correlated variables (>0.70) with other variables to reduce variable redundancy and retained ones that were biologically meaningful. We used mixed models to relate the total species richness and abundance of bats with temple variables and landscape elements. Since temples were sampled more than once and repeated measurements on the same temple are often correlated, we accounted for this by using random effects in the mixed models [43]. We used lmer package in R for normally distributed species richness and glmmadmb package with a negative binomial function for overdispersed bat abundances [44]. We considered temple I.D. and year as a random factors in the model and temple characteristics, disturbance and land use elements as fixed factors. We used the function dredge in MuMIn package (version 1.43.17) in R to obtain all possible combinations of models using the select variables. We fitted two models, one for temple characteristics with disturbance and the other for land use elements at 5 spatial scales; 0m, 500m, 1km, 3km and 5km. The models were ranked using Akaike’s Information Criterion (AICc) corrected for sample size and model weights. Model parameters were averaged for closely related models [45] using model selection criteria in MuMIn package (version 1.43.17) in R. We calculated model-averaged parameter estimates and unconditional standard errors to assess the relative importance of each variable and account for model uncertainty [46]. We only considered parameters with 95% confidence intervals not overlapping with 0 to be informative [47]. We tested for differences in microhabitats and land use between species using Kruskal Wallis and Mann Whitney tests. Data analysis and statistical tests were done using Microsoft Excel®, PAST® and R version 4.

Results

Species richness and abundance

The total roosts sampled across 5 years was 351 in 246 temples. Of the 351 temples, 286 had bats and 16% of these had multiple roosts. Seven species of bats were recorded, of which six were Yinpterochiroptera and one Rousettus leschenaultii was a Yangochiroptera (Table 1). Among the Yinpterochiroptera, Hipposideros speoris was the most frequently seen species found in 135 (38%) roosts sampled, followed by Taphozous melanopogon (58), Megaderma lyra (29), Tadarida aegyptiaca (26) Rousettus leschenaulti (22), Pipistrellus sp. (9) and Rhinopoma hardwickii (7). Forty-five percent of the temples had only one species, 34% had two species, 15% had three species, and 6% had four species of bats. Hipposideros speoris was the most common (9,715, range 2–1150) across temples followed by R. leschenaultia (5306, 1–2946), T. melanopogon (3196, 1–1165), M. lyra (1497, 20–605), T. aegyptiaca (303, 3–84), Pipistrellus sp. (144, 2–70) and R. hardwickii (50,1–15). None of the bat species we detected are "threatened" as per the International Union for Conservation of Nature (IUCN).

Table 1

Species characteristics and IUCN status of bats recorded in temples.

The breeding season in parenthesis is based on the present study.

Species name	IUCN	Diet	Breeding season	Roosting habitat
Rousettus leschenaultii	L.C.	Flowers and fruits	Nov-March	Temples, old buildings, caves
Rhinopoma hardwickii	L.C.	Moths, neuropteran insects and beetles	Feb-Mar	Crevices, roof, houses, temples, boulders
Hipposideros speoris	L.C.	Beetles, termites, low flying insects and flies	Dec-Mar (Dec)	crevices in hills, caves, caverns, disused buildings, tunnels and temples
Megaderma lyra	L.C.	Insects, reptiles, fishes and birds	Nov-Apr	caves, temples, forts, dilapidated old buildings, underground tunnels, old cow sheds, grain godowns, shallow soapstone mines and attics of houses
Taphozous melanopogon	L.C.	Insects	Jan-June (Dec-June)	ruins temples and caves, dark dungeons in the old fort
Tadarida aegyptiaca	L.C.	I Caterpillar, spider, water beetle, ground-dwelling insects	June-Sep	Crevices in cliff faces, crevices in large piles of rocks and boulders, narrow spaces formed by slabs of stone leaning against walls, the expansion joints at the top of supporting pillars in modern grain store, narrow cracks in a pillar of old mosques and crevices in buildings and old forts
Pipistrellus tenuis	L.C.	Small insects, beetle, cockroach, termite, grasshopper	Feb-march, July-Aug (Dec)	Roofs of bungalows, holes and crevices in walls, hollow branches of trees, dead leaves of trees

Bats and roost characteristics

Seven species of bats in the temple used four microhabitats inside the temple (Fig 1). About 39% of the total abundance was in dark rooms, while 51% used walkways around the deity room, the main microhabitat for 5 out of 7 species. Hipposideros speoris occupied three out of four microhabitats, but most commonly dark rooms and walkways when dark rooms were not available or disturbed. Likewise, M. lyra, T. melanopogon and R. hardwickii were found in two microhabitats (walkways and crevices) while T. aegyptiaca was found only between pillars (crevices), and Pipistrellus sp. only in roof crevices. The fruit bat R. leschenaultii preferred walkways.

Fig 1

Percentage of microhabitats used by the different species of bats for roosting.

Temple and disturbance

The number of buildings and dark rooms significantly affected species richness and bat abundance in temples (Fig 2, Table 2). The top lmer model explained 69% of the variation in species richness, of which the fixed factors explained 58%.

Fig 2

Effect plots showing the relationship between species richness, abundance and the number of buildings and dark rooms in temples.

Table 2

Effects of temple characteristics on species richness (lmer model) and bat abundances (glmmadmb model).

Temple id and year are included as random factors. The top models’ average estimates and confidence intervals are given, and the individual model details are included as (S1 Table).

Species richness	Estimate	Std. Error	CI 2.500	CI 97.500
(Intercept)	-0.052	0.122	-0.291	0.187
No. buildings	0.674	0.044	0.587	0.761
Dark rooms	0.246	0.103	0.044	0.448
Renovation	-0.364	0.081	-0.524	-0.204
Bat Abundances
(Intercept)	1.817	1.177	-0.498	4.132
No. buildings	0.802	0.206	0.397	1.206
Dark rooms	1.597	0.536	0.541	2.654
Walkway	1.678	1.057	-0.404	3.761
Renovation	-0.319	0.354	-1.016	0.378
Visitors	-0.869	0.307	-1.473	-0.265
Tower	0.349	0.447	-0.532	1.230
Hipposideros speoris
(Intercept)	-0.923	2.296	-5.439	3.594
No. buildings	0.769	0.345	0.090	1.448
Dark rooms	2.731	1.200	0.367	5.095
Renovation	-0.323	0.602	-1.509	0.863
Visitors	-0.559	0.669	-1.877	0.759
Walkway	3.631	2.324	-0.948	8.209
Tower	-0.748	0.922	-2.566	1.069
Megaderma lyra
(Intercept)	-10.230	3.243	-16.610	-3.849
No. buildings	0.982	0.581	-0.162	2.126
Renovation	-3.462	1.326	-6.074	-0.850
Visitors	-1.769	2.205	-6.113	2.575
Tadarida aegyptiaca
(Intercept)	-6.086	3.413	-12.775	0.603
Renovation	-3.723	1.782	-7.216	-0.230
Visitors	-2.231	1.942	-6.036	1.575
No. buildings	1.083	0.479	0.145	2.021
Taphozous melanopogon
(Intercept)	-14.690	5.538	-25.594	-3.786
No. buildings	0.758	0.318	0.132	1.384
Renovation	-1.181	0.750	-2.659	0.297
Visitors	2.312	1.983	-1.594	6.217
Dark rooms	2.576	3.516	-4.351	9.503
Rousettus leschenaultii
(Intercept)	-9.636	3.318	-16.172	-3.100
No. buildings	1.163	1.248	-1.296	3.622
Visitors	-0.913	1.561	-3.988	2.161
Dark rooms	0.687	2.822	-4.871	6.245
Renovation	0.667	1.681	-2.645	3.979

Renovation of temple structures had a strong negative influence on bat species richness, while the scale of visitors to the temple negatively affected abundance (Fig 3). Species response to microhabitat and disturbance varied (Table 2). Hipposideros speoris abundance is influenced positively by dark rooms and the number of buildings but not by any disturbance factors. The abundance of T. melanopogon and T. aegyptiaca were also affected positively by the number of buildings but negatively by renovation. Megaderma lyra responded negatively to renovation, while R. leschenaultii was not affected by any temple characteristics or disturbance.

Fig 3

Effects plots showing the relationship of disturbance (Renovation and number of visitors) on species richness and abundance.

Landscape

The percentage of land use around the temples changed with distance. At 5km from the temple, crops (63%) dominate, followed by urban areas (13%), scrub (12%), trees (7%), water (5%), grass and bare areas (0.23%). At 500m crops (54%) continued to dominate followed by urban areas (36%), scrub (0.7%), trees (3%), water (6%) grass and bare areas (0.09%). Since temples are close to settlements, urban areas dominate near the temple, and crops, trees, scrub and grass areas are fewer (Fig 4). Grasslands positively affected species richness at more than 3km from temples than other land uses, while trees in the temple complex positively affected species richness and abundance (Table 3). As one moved away from the temple, total abundance was positively affected by scrub availability and negatively by trees, water, and urban areas. At the species level, scrub availability positively affected the abundance of H. speoris across the distance intervals, while trees had a negative effect (Table 3). Megaderma lyra and aegyptiaca were negatively affected by crops up to 5km, while T. melanopogon and R. leschenaultii did not show any effect of land use on their roost abundance.

Fig 4

High resolution (10m) land-use around the temples sampled.

Land use classification is based on Karra et al. 2021. See data analysis for more details.

Table 3

Species response to land-use elements.

Model averaged parameter estimates of top models (<2 Δ AICc) with non-overlapping confidence intervals are shown in the last column* The top models’ details are included as (S2 Table).

Species/Abundance	Distance (m)	Parameters	Estimate	CI 2.50%	CI 97.50%	SE	Strong association
Species richness	0	Intercept	0.829	0.503	1.155	0.166
		Water	0.029	-0.144	0.202	0.088
		Trees	0.006	0.002	0.010	0.002	*
	500	Intercept	0.937	0.694	1.180	0.123
		Grassland	-0.071	-0.398	0.255	0.166
	1000	Intercept	0.940	0.695	1.184	0.124
		Grassland	-0.197	-0.982	0.588	0.399
	3000	Intercept	0.793	0.541	1.071	0.128
		Grassland	0.653	0.190	1.120	0.235	*
	5000	Intercept	0.861	0.548	1.173	0.159
		Grassland	0.618	-0.205	1.441	0.418
Abundance	0	Intercept	2.917	1.963	3.870	0.484
		Trees	0.017	0.000	0.033	0.008	*
		Water	-0.174	-0.804	0.456	0.320
	500	Intercept	3.307	2.587	4.027	0.366
		Trees	-0.032	-0.097	0.032	0.033
		Grassland	0.184	-1.330	1.698	0.769
		Water	-0.001	-0.061	0.059	0.030
		Scrub	0.142	-0.088	0.371	0.116
	1000	Intercept	3.229	1.964	4.495	0.643
		Scrub	0.176	0.057	0.294	0.060
		Crop	0.002	-0.007	0.011	0.005
		Trees	-0.144	-0.237	-0.051	0.047	*
		Urban	0.016	-0.018	0.051	0.018
		Water	-0.019	-0.089	0.051	0.035
		Grassland	-0.094	-3.954	3.767	1.960
	3000	Intercept	3.282	2.007	4.557	0.648
		Scrub	0.066	-0.004	0.136	0.036
		Trees	-0.064	-0.133	0.004	0.035
		Grassland	0.291	-1.387	1.968	0.851
		Water	-0.084	-0.289	0.120	0.104
		Urban	0.035	-0.059	0.129	0.048
	5000	Intercept	7.065	3.306	10.824	1.913
		Water	-0.498	-0.864	-0.132	0.186	*
		Urban	-0.139	-0.267	-0.010	0.065	*
		Trees	-0.077	-0.137	-0.017	0.030	*
		Crop	0.001	-0.008	0.011	0.005
		Grassland	-0.023	-3.133	3.087	1.579
		Scrub	0.069	0.002	0.137	0.034	*
Hiposederous speoris	0	Intercept	1.591	0.171	3.011	0.724
		Trees	-0.004	-0.032	0.025	0.015
	500	Intercept	1.681	-0.070	3.431	0.890
		Grassland	1.091	-1.123	3.304	1.124
		Urban	-0.036	-0.080	0.008	0.022
		Scrub	0.246	-0.128	0.621	0.190
		Water	0.017	-0.085	0.119	0.052
		Trees	-0.005	-0.112	0.102	0.054
	1000	Intercept	1.583	-0.295	3.460	0.954
		Scrub	0.275	0.059	0.492	0.110	*
		Crop	0.001	-0.014	0.016	0.007
		Trees	-0.173	-0.333	-0.013	0.081	*
		Urban	-0.026	-0.089	0.037	0.032
		Water	0.015	-0.106	0.135	0.061
		Grassland	0.444	-5.734	6.622	3.136
	3000	Intercept	0.672	-0.668	2.012	0.684
		Scrub	0.124	0.009	0.240	0.059	*
	5000	Intercept	2.098	-2.593	6.789	2.389
		Grassland	-0.595	-6.040	4.850	2.765
		Scrub	0.112	-0.007	0.231	0.060
		Water	-0.275	-0.972	0.423	0.354
		Urban	-0.197	-0.446	0.051	0.126
		Trees	-0.002	-0.101	0.097	0.050
Megaderma lyra	0	Intercept	-10.086	-14.399	-6.069	2.219
		Trees	0.006	-0.057	0.068	0.032
		Water	-0.064	-1.373	1.648	1.252
	500	Intercept	-9.165	-12.658	-5.673	1.773
		Grassland	-1.701	-22.185	18.783	10.399
		Crop	-0.038	-0.074	-0.002	0.018
		Water	-0.053	-0.387	0.282	0.170
	1000	Intercept	-8.910	-12.676	-5.143	1.912
		Grassland	-2.381	-29.684	24.921	13.860
		Crop	-0.031	-0.060	-0.001	0.015
		Water	-0.130	-0.769	0.509	0.324
	3000	Intercept	-9.571	-13.736	-5.407	2.115
		Grassland	-0.325	-9.084	8.435	4.447
		Crop	-0.028	-0.059	0.003	0.016
		Water	-0.168	-1.223	0.887	0.536
	5000	Intercept	-9.432	-14.681	-4.182	2.666
		Grassland	-1.791	-16.318	12.736	7.375
		Crop	-0.027	-0.061	0.006	0.017
		Water	-0.103	-1.593	1.386	0.756
		Trees	0.008	-0.220	0.236	0.116
Rousettus leschenaultii	0	Intercept	-10.086	-14.457	-5.715	2.219
		Trees	0.006	-0.057	0.068	0.032
		Water	-0.064	-2.531	2.403	1.252
	500	Intercept	-9.441	-13.064	-5.819	1.839
		Trees	-0.354	-1.823	1.115	0.746
		Grassland	-2.717	-28.261	22.827	12.968
		Water	0.029	-0.167	0.224	0.099
	1000	Intercept	-9.219	-13.146	-5.291	1.994
		Trees	-0.381	-1.805	1.043	0.723
		Water	0.013	-0.251	0.277	0.134
	3000	Intercept	-9.805	-14.066	-5.543	2.164
		Trees	-0.159	-0.928	0.610	0.391
		Urban	0.078	-0.220	0.375	0.151
		Grassland	1.390	-4.757	7.536	3.120
	5000	Intercept	-10.148	-15.311	-4.984	2.622
		Water	0.241	-0.913	1.395	0.586
		Grassland	2.023	-8.862	12.908	5.526
		Trees	-0.049	-0.391	0.294	0.174
		Crop	-0.010	-0.054	0.035	0.023
		Scrub	0.054	-0.207	0.315	0.132
Tadarida aegyptiaca	0	Intercept	-9.608	-14.070	-5.146	2.266
		Water	0.555	-1.653	2.763	1.121
		Trees	0.001	-0.057	0.059	0.030
	500	Intercept	-8.367	-12.152	-4.581	1.922
		Grassland	-1.819	-19.858	16.221	9.158
		Crop	-0.023	-0.046	0.000	0.012	*
		Urban	-0.010	-0.109	0.088	0.050
		Trees	-0.317	-1.849	1.215	0.778
		Water	-0.085	-0.484	0.314	0.203
	1000	Intercept	-7.652	-11.239	-4.065	1.821
		Grassland	-24.772	- 149.996	100.453	63.572
		Crop	-0.022	-0.043	-0.001	0.010
		Water	-0.081	-0.506	0.344	0.216
	3000	Intercept	-8.235	-13.475	-2.995	2.663
		Grassland	0.812	-5.570	7.193	3.240
		Crop	-0.022	-0.040	-0.004	0.009
		Water	-0.487	-1.728	0.755	0.630
		Urban	0.061	-0.228	0.350	0.147
		Trees	-0.031	-0.344	0.283	0.159
	5000	Intercept	-8.291	-14.453	-2.129	3.130
		Grassland	-0.488	-12.427	11.452	6.061
		Crop	-0.024	-0.044	-0.004	0.010	*
		Water	-0.368	-1.794	1.058	0.724
		Urban	0.117	-0.338	0.572	0.231
		Trees	-0.022	-0.264	0.220	0.123
Taphozous melanopogon		Intercept	-8.476	-12.411	-4.542	1.998
		Water	-0.997	-3.967	1.973	1.508
		Trees	0.010	-0.051	0.071	0.031
	500	Intercept	-9.273	-12.846	-5.699	1.815
		Trees	-0.096	-0.837	0.645	0.376
		Water	0.029	-0.176	0.233	0.104
		Grassland	0.322	-3.117	3.760	1.746
		Urban	0.047	-0.052	0.147	0.051
		Crop	-0.007	-0.036	0.021	0.015
	1000	Intercept	-9.137	-12.374	-5.901	1.644
		Grassland	1.637	-5.136	8.410	3.438
		Water	0.056	-0.195	0.308	0.128
		Trees	-0.032	-0.425	0.362	0.200
		Crop	-0.012	-0.030	0.006	0.009
		Urban	0.029	-0.100	0.157	0.065
	3000	Intercept	-8.635	-13.813	-3.456	2.633
		Grassland	7.892	-1.886	17.670	4.964
		Urban	-0.208	-0.693	0.276	0.246
		Water	0.302	-0.337	0.940	0.324
		Scrub	0.050	-0.162	0.261	0.107
		Trees	-0.059	-0.502	0.384	0.225
	5000	Intercept	-9.437	-15.329	-3.546	2.994
		Grassland	5.918	-7.637	19.473	6.883
		Trees	-0.091	-0.514	0.332	0.215
		Water	0.198	-0.856	1.252	0.535
		Crop	-0.009	-0.025	0.006	0.008
		Urban	-0.262	-0.930	0.405	0.339
		Scrub	0.081	-0.171	0.333	0.128

The availability of roost microhabitats across temples with and without bats was not significantly different (Kruskal Wallis test; D = 0.4, p = 0.689), Fig 5). Of the total temples sampled, 80% of them had bats across all the years, and in only 4 of 57 temples, we found bats abandoning the temple. Renovation levels were higher in temples with no bats 44% (15/34) compared with temples having bats 35% (16/46). Visitor levels were also marginally higher in no bat (x±se, 2.11 ± 0.79) compared to temples with bats (1.89 ± 0.71). Land use between bat-occupied and not-occupied temples showed no differences except for trees in the vicinity of temples being higher in bat-occupied sites (Mann-Whitney U = 3917 p<0.01) (Fig 6), which corroborates with the land use analysis.

Fig 5

Percentage availability of habitats in temples with and without bats.

Fig 6

Mean land-use elements around temples with and without bats at various distances.

* indicates significant differences at p< 0.05.

Discussion

Why do bats prefer some temples? Bats occupied 80% of the temples sampled and over 20,211 individuals were counted. Though only seven species were recorded, the most common species Hipposideros speoris (9,715), accorded for nearly half of the population and found in more than 50% of bat-inhabited temples. Hipposideros speoris is a widespread species across India and Sri Lanka and occurs in large congregations with high fidelity to roost sites [48]. The other 6 species are also not rare or threatened but use many anthropogenic structures for roosting [28]. Except for T. aegyptiaca, Pipistrellus sp. and Rhinopoma hardwickii, species counts of all others ranged between 200 to 450 in each temple and appear to follow the general tendency of bats that roosts in caves and human-made structures to be highly gregarious [49]. Crevice roosting T. aegyptiaca and Pipistrellus sp. species may have been underestimated because they are hard to count inside the narrow crevices.

Several species that roost and breed in temples across India are cave-dwelling species that have found temples as suitable alternative habitats [28, 39]. Our results show that temples offer dark, undisturbed roosts and crevices similar to caves for seven species of insectivorous bats that account for 33% of the bat species found in the plains and hills of Tamil Nadu state [29]. Such dark areas in old temples have several openings through which bats can quickly fly out, a likely reason for their selection by most bat species. Further, granite stones used to build the ancient temples are stacked together and offer crevices for some species, such as the free-tailed bats (T. aegyptiaca), to roost, thereby providing a safe roost. Walkways are long corridors with a high roof with vents that remain dark during the day and provide a safe roost for bats like H. speoris. The high roof help bats remain unnoticed and therefore not disturbed during the morning hours when there is a movement of people. A similar response of cave-dwelling bats from the genus Hipposideros was found in Malaysia, where they roost in very dark corners and high places [50]. We did not take species roost height measurements, so it’s not evident if species segregate roost according to the height, which is also unlikely as the height inside the temple is the same. However, unlike in a cave, many species that roost in dark corners in temples seem to segregate spatially, which varies between temples but would require more study.

This study demonstrates that disturbance to roosts is a major factor affecting roosting bats, as other similar studies have shown in old buildings under temperate conditions [20, 51]. Disturbance in the temple was less impactful on the abundant H. speoris, but other species were negatively affected by renovation and visitor levels. Large temples usually have more visitors but also more buildings with more potential roosts that may buffer disturbance effects. Therefore, even when disturbance levels increase, bats might persist by temporarily moving to other temple buildings. In many cases, these disturbance activities are routine in temples and bats might have become accustomed to these threats. However, several other factors that may affect bats in temples and not measured in the study, such as festivals that increase lighting, sound levels, human activity and prayer-related smoke, can impact bat abundance. The recent increase in the renovation of old temples where dark areas are lit, walls are pressure cleaned to remove the odor of bats, painted and openings ’bat proofed’ to keep away bats and pigeons can effectively prevent bats from colonizing temples [48, 52]. Bats have come under even greater threat and surveillance because of local media misreporting the current coronavirus and bat association.

While temple characteristics and disturbances are significant drivers of bat abundances and occupancy, land use plays a less critical role across species. Several studies have shown the importance of landscape elements for the presence of bats [27, 53–55], but in our case, there seems to be no single habitat effect. Fine-scale (10m) mapping of land use around temples revealed differential use of habitat elements by species, but these associations were either weak or had high variations in the effect. The landscape, as mentioned earlier, is highly homogeneous; crops and urban areas together constitute about 70–80%, with scrub, grasslands, water bodies and trees comprising the rest in small proportions. Even though crops were everywhere, they did not have much effect while trees and water availability seem to have a consistent negative influence across species. The availability of scrub and grasslands is critical for species richness and the overall abundance of bats. These are the only two natural habitats available, with the rest being tree plantations or occupied by invasive Prosopis juliflora and artificial wetlands. Even though scrub and grasslands occupy less than 5% of the area, it has a disproportionately positive effect on all the species except M. lyra. The radio-tracking study in south India of the M. lyra showed it has a very flexible diet and can take prey both from air and ground, enabling it to use multiple habitats but prefers to forage primarily within 500m from temples [38]. The importance of natural habitat is emphasized for many species, including bats [26] and this is evident in this study. Water availability is vital for many species of bats and may drive bat movement and activity in some landscapes [25, 56]. The effect of water was variable, with species like T. aegyptiaca preferring water points in the temple’s vicinity while others did not. Studies in Thailand on similar species have shown that insect diversity is less influential than water availability [57]. Moreover, water is not a constraint in the region because of the perennial river and extensive irrigation network. Also, the water requirements of many of the species may be low as they are adapted to arid conditions.

Scale is vital for many species, including bats [27]. In this study, at 5km from the roost, most land-use effects became negligible, or the direction of the effects changed from positive to negative or vice versa. The extent of natural habitats such as grassland and scrub that increases with distance may influence bats to forage at larger spatial scales. For example, temples are located mainly close to human habitations and rivers while natural open dry habitats are further away and bats may be making regular forays to them, as observed in England [26].

This study provides a broad understanding of land use effect on bats. A more nuanced study and analysis will need radio telemetry and acoustic sampling to see how land-use elements influence bats. Foraging bats are most strongly associated with variables measured at small spatial scales and distance measures [27]. For instance, the only study on radio telemetry of temple bats shows 41% of the time, the bats foraged 500m from the temples but ranged up to 4km over the night during the summer months. In rainy months this may be different when the general productivity of the habitat is higher, and more extensive use of the landscape can happen as scrub and grassland habitats also become more productive. Further, with more advanced remote sensing tools, more nuanced land-use change analysis could be related to seasonal changes in bat occurrence. For example, a recent study in the same area using radar images has shown greater grassland area than conventional methods [58]; such an approach would also enable mapping across seasons irrespective of cloud cover.

The current study relies on the 2020 high-resolution land-use analysis [41], but this study predates that in 2012–2019. This mismatch, as mentioned earlier, may not be very high given the land use has remained essentially the same over this period. However, for natural habitats with less than 5% of the area and critical for bats, even small changes can significantly affect a foraging animal. Further, the dynamics of wetlands are not incorporated in this study since it was mostly done in the wet season. However, a more nuanced classification of these dynamics around each temple might be necessary as irrigation systems and water availability are affected by regional water distribution [59].

The presence of bats is linked to microhabitat or stand-level structural conditions. However, Lewis [60] found that roost-site fidelity for bats varied depending on the availability and permanency of the roosts used. For these reasons, he suggested that fidelity was high in buildings. Supporting this, a study on banded H. speoris bats from the study area showed 82% roost fidelity throughout the year [48]. Therefore, the temples that have remained unchanged for centuries provide conditions for high roost fidelity despite changes in the landscape and therefore are more important for bats to persist in the area.

Conservation of bats in temples

The study has shown that microhabitats within temples, such as dark corners that are relatively less disturbed and rooms rarely used, are essential for the persistence of bats in the temples. The effect of disturbance on bats is strongly negative and can override the availability of microhabitats in temples and is also species-specific. Most of the landscape factors were surprisingly not important though the availability of natural areas around temples is crucial. While bats are known to provide critical pest control services in paddy agriculture, the role of paddy in sustaining the bat populations is not clear as crops did not have any significant effect on bat species richness and abundance. This calls for a more nuanced study using radio telemetry and acoustic sampling to understand the use of the landscape by bats since natural areas are likely to be converted to irrigated agriculture in the future. Since temples were built from breaking granite hillocks that would have harbored caves and bats in the region, temples and other old buildings ironically are the only alternative habitats. Bat conservation in temples and historical buildings is possible if small dark corners are allowed to exist along with some abandoned areas. If this is not possible, we need to think of building alternate habitats like bat houses in the landscape and develop a comprehensive plan to let bats persist in the landscape and provide ecosystem services.

Top models of species abundance response to microhabitat and disturbance.

Model abbreviations-BLD: No.buildings, DR: Dark rooms, WAY: walkway, TWR: tower, REN: Renovation, VIS: visitors.

(DOCX)

Click here for additional data file.

Top models of species response to land-use elements.

(DOCX)

Click here for additional data file.

16 Jun 2021

PONE-D-21-13889

Temples the last abode for bats in a homogeneous agriculture matrix: Importance of microhabitat availability and land use factors for bat conservation.

PLOS ONE

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Reviewer #1: There is some clarification needed to determine if statistics were statistically sound. Including which data were used in which analysis. The authors sampled 59 temples repeatedly, and it is unclear if they treated each survey as an independent sample, or if data from only one survey year were used, or if it was averaged across the 5 survey periods for each temple.

There are multiple grammatical and spelling errors throughout that do not detract from the quality of research, but would need to be corrected before publication.

Overall this study would be a valuable contribution to the literature on bats using anthropogenic structures, and my specific comments for improving the manuscript are attached.

Reviewer #2: Overall, this paper has significant importance for publishing by emphasizing the importance of temples as habitat worthy of conservation especially at a time when bats are at a high risk of exclusion from places because of the pandemic. Some modifications are necessary to clarify the paper and its findings, however they do not detract from the importance of the study.

Reviewer #3: Specific suggestions in attached document.

Methodology needs explanation - 59 temples surveyed over 5 years would be 295 surveys. Not sure how the seasonality is included. Were temples surveyed over the years in each season? How did year influence species richness and abundance? Were morning and afternoon counts added together?

Needs more grammatical review - some misspellings, incomplete and run-on sentences

Passive voice is used throughout manuscript

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Figures need major edits - proper labeling of variables, Figure 6 has no y-axis, land use x-axis should be 0, 0.25, 0.50, 0.75, 1

Why on Figure 5 and 6 are species abundance and richness only 10-30? Did you only use means? Using raw data would be much more informative.

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Submitted filename: Review_bats_temples.docx

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Submitted filename: Review.docx

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Submitted filename: PONE-D-21-13889_reviewer (2).docx

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7 Dec 2021

All responses to reviewers are marked as authors response(AR) in red.

Editor

The reviewers agree and the AE concurs that there is a degree of opacity in the analytical approaches that require elucidation before the manuscript can be fully vetted (at least with respect to the analyses, their results, and the inferences drawn from them). The reviewers identify a number of areas in the analytical methods that require additional detail and clarification.

AR: We have re written the Material and methods section giving clarity in the methods used and the data used in the analysis

the manuscript requires substantial editing to correct considerable errors in grammar and syntax.

AR: This has been addressed to the extent possible

Reviewer 1.

Review: Temples the last abode for bats in a homogeneous agriculture matrix: Importance of

microhabitat availability and land use factors for bat conservation.

Ganesh et al studies historic temples as roosting habitat for bats in India. This work highlights the importance of anthropogenic structures as essential roosts, particularly as temples are places where humans are actively working to remove bats. While focusing on one region in India, the authors conducted roost counts and identified species of bats over several years while also assessing each temple for several microhabitat and landscape factors. They conducted linear models to compare species richness and abundance to the landscape and microhabitat variables and calculated niche breadth and overlap. The authors found seven species of bat across forty seven out of fifty-four temples that they surveyed. Dark rooms and corridors were found to be the most important microhabitats for roosting, while landscape effects had little overall effect albeit with some species-specific instances. Bat abundance varied by species in season, but overall was lowest during the monsoon season. There was little species overlap in temples and little niche overlap. The authors attributed a lot of the microhabitat preferences to species-specific life history and roosting preferences. The seasonality differences correlated with the parturition and lactation of many of the species. While the landscape effect had a weak response, the importance of scrub and open landscape is still noted. Overall, this paper lays the foundation for further studies and emphasizes the importance of an anthropogenic roost for bats. This paper certainly merits publication with additional clarification and editing.

My first concern overall for the paper is some grammatical and formatting issues. I have listed specific examples of reoccurring problems throughout at the end of this review, however there are additional grammar edits that need to be fixed.

My second concern is with the landscape ecology analysis and discussion throughout the paper. The analysis of the landscape features was measuring presence/absence of landscape features over a significantly sized area. This broad measuring does not get at some of the heterogeneity of the landscape and may explain some of the lack of effect of the landscape features in the model. A more rigorous measurement of the landscape types could be better such as using the total amount or percentage of land cover type in the 5km buffer. As the remote sensing data may be unable to provide this level of detail (as mentioned in the methods), a mention of this drawback in the discussion would be warranted. Additionally, the discussion section touched on the landscape feature briefly but the middle of the paragraph still discussed temple characteristics such as festivals and renovations and the ending sentences focused on site fidelity. The importance of open spaces was stated but never explained or elaborated on. A deeper discussion with a paragraph focused solely on landscape characteristics while moving the still-relevant topics to more appropriate places would improve the analysis especially since the third hypothesis on landscape features was disproven.

Overall, this paper has significant importance for publishing by emphasizing the importance of temples as habitat worthy of conservation especially at a time when bats are at a high risk of exclusion from places because of the pandemic. Some modifications are necessary to clarify the paper and its findings, however they do not detract from the importance of the study.

Authors response(AR):

The landscape analysis has been completely revised. We have used recently released ESA landcover classes at 10m resolution for the analysis (https://tiledimageservices.arcgis.com/P3ePLMYs2RVChkJx/arcgis/rest/services/Esri_2020_Land_Cover_V2/ImageServer). However, this approach allows us to estimate the landcover class for a particular period and not across seasons because of cloud cover as mentioned in the MS earlier. We have therefore assumed, based on government literature that landcover has not drastically changed during the study period which is also substaintiated by authors observations. We analysis the % landcover under each class to species richness and abundance of bats at 3 spatial scales;500m, 1km and 3km. The variable in the landcover class is crop which we determine based on google earth images as earlier and interviews with farmers around temples. Some periods of the sampling May and Sep do not have any crops and there fore we have considered crop area in such times as zero and considered it as fallow agriculture. More details in the MS.

Reviewers: A deeper discussion with a paragraph focused solely on landscape characteristics while moving the still-relevant topics to more appropriate places would improve the analysis especially since the third hypothesis on landscape features was disproven.

AR: Our discussion has changed and improved given the changes in the analysis. We have as suggested given more space for discussing landscape characterstics.

Grammatical and Formatting Comments:

Check the spacing throughout the article. There should be a space in between the last word and opening parentheses. On the last sentence of page 11, there should be a space after the comma and between each year. The same issues occur in the caption for Table 4 with the list of symbols. Lastly, there should be a space in between genus and species in the species names in the results - I.e. M. lyra, not M.lyra.

In the first two paragraphs of the introduction, there are a few words and several citations where the year is in blue. They should all be in black since they are not hyperlinks.

A w is bolded in the word “with” in the beginning of the full paragraph on page 10.

Throughout the entire paper, halls is followed by (corridors) or vice versa each time. I think saying “halls and corridors” is appropriate the first time and then use one consistently for the rest of the paper.

AR: We agree on the English and have tried our best to improve. Some of the issues have also arised due to formatting.

We have modified halls to a more appropriate word ‘walkway’

Figure 1 appears blurry, so I would recommend the resolution to be larger. Partially because of the resolution, the temples and town dots are hard to tell apart. It would be easier to read if one of the dots was hollow.

AR: We have redone the figure

Figure 3 does not have axis labels which helps the reader.

Corrected

In Figure 4, the dark room and hall patterns are very hard to tell apart between the bars. A different pattern would make the graph easier to read.

We have redone the figure

Figure 5 has extra ticks around the outside of the graph.

We have redone the figure

In Figure 8, the no-village bar is distracting and confusing as it overlaps with the error bars.

Changed the fig as we are using landcover maps

Reviewer 2

Reviewer Summary of Paper: This is a study of factors important for bats using ancient temples and overlap of roost use in bats using ancient temples. The authors investigate roost characteristics of temples (microhabitat) and surrounding land use (landscape-scale habitat at 5 km scale). They determine that roost characteristics (mainly presence of dark rooms and halls) influence bat richness/abundance in temples. Landscape variables at the 5 km scale were not important for bat abundance and bat presence.

Overall impression: The subject of managing historic/ancient buildings and bat conservation will appeal to many readers. The study would add to the literature of managing ancient/historic buildings for bats. However, the manuscript would need major revision before publication in PLoS One. The discussion is the strongest section of this paper, and other sections could be improved by incorporating more literature into the introduction to set up the hypotheses, clarifying the hypotheses, organizing the data analysis section by hypothesis, adding more information about which data were used in which analysis, and clarifying some of the figures.

My specific comments to the authors to improve the manuscript are outlined below:

Title:

1. The term “matrix” implies a heterogenous landscape of multiple land use/cover types not a homogenous one (only one cover type). I suggest rewording or clarifying the title.

Author response AR: replaced “matrix” with “landscape”

Abstract:

2. Please specify which pandemic.

3. AR: Mentioned as COVID19 pandemic

4. “We sampled 59 temples repeatedly across 5 years which yielded a sample of 256 temple.” I believe that 59 temples sampled repeatedly is a sample size of 59 temples. I suggest saying “which yielded 256 survey events” or rephrasing another way. Was every temple sampled 5 times?

AR:Yes sample size is 59 temples and these were repeatedly sampled. Suitably modified in the abstract. The number of times each temple was sampled varied between 1 to 5. Have mentioned the methods.

Introduction:

The introduction should be supported with more literature to justify the research question. I suggest the authors incorporate the literature on bats using anthropogenic structures and introduce and explain the issues here for bats, humans managing structures, and bat-human conflicts. A good paper to read is Fagan et al. 2018. “Roost selection by bats in buildings, Great Smoky Mountains National Park” in the Journal of Wildlife Management 82: 424-434, as they evaluated bat roost selection of historic buildings in a non-urbanized area and also measured landscape variables. If the literature is limited for bats in buildings, the authors could draw on the literature of bats using bridges or other anthropogenic transportation structures that humans have to manage. One example is Meierhofer et al. 2019. “Structural and environmental predictors of presence and abundance of tri-colored bats in Texas culverts” in the Journal of Mammalogy 100: 1274-1281. One of the more interesting aspects of this subject is that bats using historical buildings or other human infrastructure makes it difficult to balance bat conservation with human-bat conflicts. And this could be applied beyond ancient temples.

More information on the types of bat species that use these temples, why we should care about them, and current conservation status would greatly improve this manuscript. For example, the authors could mention the importance of bats to agricultural pest suppression (if these bats are insectivores) or seed dispersal (if these bats are frugivores) or providing other examples of ecosystem services. The authors should explain what they mean by the pandemic negatively affecting bats. Future readers may not know which pandemic the authors are referring to, and why it affected bats. If any species are “data deficient” or of conservation concern on the IUCN Red List of Threatened Species (https://www.iucnredlist.org/), this should also be mentioned in this introduction. Additionally, if there have been previous studies involving these species, i.e., what they usually use to roost if not temples, what they eat, where they forage, etc. this would be helpful to include some basic background information to set up the hypotheses. Are they mostly fruit bats or are they insectivores? Some species background information is provided in the discussion but is really needed in the introduction to support the justification of these research questions.

Similarly, more information on temple structure, e.g., What materials are they made out of? How do bats get into the temples (through doors or cracks in the walls or unknown?) Are they air conditioned or are they about the same temperature/hotter/cooler than outside?) Additionally, introduce readers to the monsoon seasons briefly in the introduction and more in the study area section.

AR: More information on temple structure given in the MS itself

The authors should be careful about using the term “niche” when only referring to roosting habitat. A niche encompasses all needs of a species (roosting habitat, foraging habitat, all biotic and abiotic factors (climate), etc.). In some parts of this manuscript, I believe “niche” should be replaced with “roost habitat”.

The objectives of this paper need to be clarified if they are investigating what microhabitat and landscape factors determine roost use and bat abundance in order to inform bat conservation in ancient temples, and/or if they are investigating which bats have roost overlap needs (what I think the authors are referring to when they say “niche overlap and breadth”). What is important about quantifying niche overlap/breadth?

AR: we have removed niche analysis from the results.

5. “These are situated in a human-dominated agricultural matrix that is changing due to several economic reasons and climate change effects (Fischer et al. 2009, 2010; Jones et al. 2009).” Please describe the climate change effects affecting bats in this area. Do you mean the monsoons? I also suggest including literature on anthropogenic disturbance, agriculture, and habitat/fragmentation threats to support your justification for this research. I suggest developing those ideas more with Frick et al. 2019 “A review of the major threats and challenges to global bat conservation” paper (available at: https://www.researchgate.net/publication/332146982_A_review_of_the_major_threats_and_challenges_to_global_bat_conservation ) or other studies

AR: The role of climate and habitat transformation in the region has been stated in the context of Frick et al 2019 paper. We have also added some available literature for the region.

6. “The recent pandemic has further made them more vulnerable to disturbance and direct persecution.” Please describe which pandemic and why this is an issue for bats.

AR: COVID 19, Done

7. “ 1. Given that the temple bats are mostly cave-dwelling species we expect larger

Yinpterochiroptera occupying similar micro-habitat, show greater niche overlap and similar niche breadth.” Please introduce the species before stating the hypotheses. This is the first time the bats and some traits are mentioned. Please describe niche breadth and overlap in this context. Does this mean the authors are trying to determine overlap in roost needs/preferences of species using the temples and that they expect Yinpterochiroptera to have overlapping needs with each other? Why does this matter? Please clarify this hypothesis. Are Yinopterochiroptera the only group that use the temples? Are they all cave-dwelling?

AR: we have modified this hypothesis. We have removed niche overlap from the paper. Mention of species is done earlier and most of the species are insectivorous expect for one

8. “2. Microclimate conditions inside the temples are very stable and any influence on

bat abundances could be a reflection of food availability or disturbance at roost and therefore we expect bats to show seasonal changes in niche overlap and breadth due to variability in crop and insect availability.”

This hypothesis is confusing to me. I’m not sure what you are hypothesizing. Would it not be better to test abundance changes seasonally rather than niche overlap and breadth? The climate/monsoon seasons could also be described before introducing this hypothesis. Additionally, it is unclear how disturbance at roost and food availability were measured.

AR: we have modified this hypothesis and have made a separate result for roost disturbance which is mentioned in the introduction and methods.

9. “3. Since the structure of ancient temples are all similar, we do not expect any

change in microhabitat availability in temples with and without bats, instead we expect bat abundances driven by the availability of landscape features especially seasonal crops around the temples.” Please describe how the temples differ or are similar to each other and the seasonality of crops before introducing this hypothesis. Also, is it not true that the temples are structurally different? Some are smaller/larger and more complex than others?

AR:We have described the temples in detail and the type of crop and their availability in the section. Temple did vary in size but structurally they were more or less similar.

Materials and methods:

“The ancient temples(>400 years) are built of large granite stones and provide a niche for several bat species.” I think the authors mean that the temple structures provide roost habitat, not a full niche.

AR: Agreed and we have clarified in the text

“The surveyed temples are situated in an agricultural matrix surrounded by rice paddies or bananas” Are the temples completely surrounding in the whole 5 km radius landscape or just mostly surrounded in the immediate area around the temple? What is the extent of the surrounding agriculture? An agricultural matrix implies mixed cover types so this sentence is somewhat confusing.

AR: The entire area even beyond 5km are dominated by paddy. We have given more information about the landscape in the introduction and study area section

“stone pillars that supported the stone roof was considered a separate habitat as they provided a niche for few species that roost in the gaps created between the stone joints this was categorized as crevices” This sentence is a little confusing as it reads. Were the stone joints in the pillars categorized as crevices? The stone pillars provided a roost, not a niche.

AR: Yes, they provide crevices and therefore a roost habitat

“In 2012 we measured temperature and humidity while sampling for bats and found no difference between temples and therefore not considered in the analysis.” What are the temperature and humidity conditions in these temples and is it similar to cave conditions used by these bats?

AR: We are not aware of temperature and humidity conditions inside caves in the region as caves are non-existent because the rocky hills have been quarried. Conditions inside the temples are mentioned though, as stated in the methods section, we could not monitor these continuously

“Since we did not have information on how far the different bat species forage in the landscape we took 5 km based on the study on the large bat Megaderma lyra (Audet et al 1991).” Did the authors consider testing multiple spatial scales? Many bat studies have determined that landscape scale factors are important at different scales. See the discussion in Bellamy et al. 2013 “Multiscale, presence-only habitat suitability models: fine resolution maps for eight bat species” in the Journal of Applied Ecology 50: 892-901 (available at: https://besjournals.onlinelibrary.wiley.com/doi/full/10.1111/1365-2664.12117) about importance of factors at multiple landscape scales. In the case of not knowing foraging distance for all species, it would improve this study to quantify landscape availability at more than one scale to determine if it is important (For example, maybe test 500 m, 1 km, and 5 km or landscape scales that are relevant to bats in other parts of the world). What kind of bat is the Megaderma lyra? Is it an insectivorous species?

AR: Thank you for the paper link. We have used one-time remote sensing classification of the land use available at ESA website and reanalyzed the data at 4 different spatial scales as suggested. Since such information was available only at one particular year for the entire region we considered the land use not to have changed during the sampling period and when the image was taken. This we justify based on government records of area under agriculture. We also tried to do a fine level classification by gridding the area around temples and finding the land use within the grid cells using google earth. This was a herculean task as the grid cells for each temple at 3km was over 1000 and was not possible to do that for each temple across years and is the reason we resorted to remote sensing analysis for fine scale analysis

.

AR: Megaderma lyra is an insectivorous species and we have mentioned the details of all the species in a separate Table 1..

“If a land use element is present we gave a value of 0.25 in each quadrant and calculated the frequency of each land use for the entire 5km radius area.” Is 25% (i.e, 0.25) sensitive enough to detect variation? Why 25% and not 10%? If a 5 km radius landscape only has a little bit of banana, but it is present, then it would still get a value comprising 25% of the landscape? This may be a limitation of the study to mention in your discussion. Would the results be different if the 5 km landscape was divided by 10% increments?

AR: This is a very valid point and we agree on the bias and as mentioned above we have done our best under the current conditions. Keeping all the comments related to landuse we have changed the analysis as mentioned before.

Did land use around each temple differ or was it correlated among the 5 sampling years?

AR: Yes, the land use did not change much across years

“The best model was selected using.” Please finish this sentence and indicate what AICc weights and delta AIC value criteria were used to determine plausible models.

AR: done

“We constructed a linear model to explain the species richness and abundance of bats in the temples based on the microhabitat and landscape features. We based the variables used in our linear models on our observations made in 2012.” Does this mean that only the landscape features from 2012 and the bat counts from 2012 were used? I think it would be statistically incorrect if all survey years at the same temples were treated as independent temples in the models.

AR:We have use mixed model approach to repeated sampling of temples across years following Bolker et al. 2009 wherein they say “Repeated measurements on the same individual, the same location, or observations taken at the same point in time are often correlated; this correlation can be accounted for by using random effects in generalized linear mixed models”. Since there were variations both within the sampled temples and across years, making temple and years as random factors accounts for this variation and gives a more robust relationship between the response variable (species richness and abundance of bats) with explanatory variables like temple features and land use elements.

Please organize the data analysis section to be more clear which analysis you are doing to answer which question/hypothesis and clarify which data were used to examine microhabitat and landscape factor influence on abundance and richness. In other words, did you use only data from 2012 or did you treat each repeated survey as a separate “temple”? If only used 2012 data, what were the monsoon conditions at that time?

AR: Temples were sampled for its features in 2012 and if any new temples were added in later years their features was recorded then. Temple features are subjected to modifications and therefore we have included renovation of temples and visitors to the temple as variables that could affect bats. Renovation is a dynamic variable that can differ between years and therefore we have measured it in all sampled years in all temples compared to more static ones like number of buildings, tower etc which do not change across years.

We have now tried to bring in more clarity in the data analysis section as suggested

Results:

“The total number of roosts sampled was 351 across 246 temples in 5 years.” This sentence is a little confusing. There were only 59 temples. Were any of the roosts that were repeatedly sampled the same roosts as previous years?

AR: We have modified the sentence and yes, same roosts were sampled across years and some temples also had multiple roosts

“None of these bat species is threatened as per IUCN category.” Are any of them considered “data deficient”? Please move this species information to the introduction to introduce the species that use the temples.

AR: Done. No bat was data deficient

“The lmer model explained 67% of the variation in species richness of which 55% were explained by the fixed factors.” Do you mean the model averaged mixed model comprising all models with AICc <5 or the top model or something else?

AR: The % values pertain to the top model. The models have changed after reanalysis. The delta AICc was kept at <2 unless mentioned specifically. We have corrected for any discrepancies

Discussion:

The authors are effective at relating the results to the previous literature and this is the strongest section of the paper.

Mention the limitations of the study (e.g., temples were chosen based on logistics rather than randomly, only one landscape scale was considered, land use in the surrounding 5 km radius was considered at a course resolution (0.25))

AR: The landscape section has been reanalyzed completely as mentioned earlier

From your results, are their certain species that may be more impacted by temple renovation/disturbance?

AR: We have now made renovation as an objective as mentioned earlier and identify species that are affected, most species are.

Figures:

Fig 1 map is a bit blurry. I suggest increasing dpi/resolution. Did there appear to be any spatial pattern of temples that did not have bats vs. those that did? If so, mention in discussion.

AR: The fig has been redone. We did not test for spatial correlations

Fig 4 bar graph gray colors are difficult to differentiate between dark room and hall.

AR: Corrected

Fig 5 is this a bar graph of all temples, including the ones that were resampled? If so, perhaps plot the average species richness across the 5 years against the no. of buildings in each temple. It is difficult to interpret as is.

AR: These are not bar graphs, they are effect plots from the mixed model showing the relationship between number of buildings and species richness keeping other factors constant in the model. Such effect plots are routinely used to plot the results of mixed models and are considered valid to predict the relationships between variables.

We have corrected the graph and included datapoints along with the prediction line.

Same comment as last figure (which is also labeled Fig 5. Is that intentional?) What is DR? Define in figure or in parentheses in caption.

AR: expanded and clarified

Fig 6 I suggest the axis title say “Proportion of rice paddy in 5 km radius” or similar

AR: After the new analysis using remote sensing categorization we could not do the analysis at individual crop level so this is excluded

Fig 8 This figure is difficult to read with No. villages plotted on top of the land use classes.

AR: Changed completely

Table 1: Nicely presented. Please indicate in the caption at what values are there considered an overlap? Higher values mean more overlap?

AR: We have removed this analysis

Table 4: Missing a variable after the “+” under Hiposederous speoris? Did you mention that you also had species-specific models in the main text?

AR: Modified

Reviewer 3

Made comments in the MS itself which is addressed therein

Submitted filename: Response to Reviewers.docx

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10 Mar 2022

Submitted filename: PONE-D-21-13889_R1_Feb2022.docx

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Submitted filename: PONE-D-21-13889_R1_ReviewerEdits.pdf

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13 Jun 2022

Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1: It is obvious that the authors have spent a lot of time and effort in revising this manuscript. The details on the analyses have been provided and the methods are sound. The information presented is very interesting. There is not a data availability statement in the manuscript, but the authors do indicate in the submission form that they are available upon request. There are still many grammatical errors, but I have provided specific suggestions for the authors to improve the grammar and readability. My suggestions are included in the attached PDF and the authors can use Adobe Acrobat Reader DC to view the edits.

Authors response (AR)

Thank you for the reviews and edits on the manuscript. We have incorporated your edits and have gone through the MS thoroughly to correct for grammatical errors.

Reviewer #2: Review: Temples the last abode for bats in a homogeneous agriculture landscape: Importance of microhabitat availability, disturbance and land use factors for bat conservation.

The edits to this article made the methods much more robust and clarified many of my previous concerns, especially the updated methods for landcover classification. There remain some spacing and grammatical issues some of which I lined out below. With minor grammatical and editorial revisions, this paper will meet publication criteria.

Revisions

Ensure there is a space between every genus and species- M. lyra, not M.lyra

Ensure there is a space between each paratheses and last word in each instance- i.e. temples (Audet et al 1991) or trees (3%)

AR:We have corrected for space

Figure 6’s font is quite hard to read. I think the resolution of the image needs to be increased.

AR:We have changed the resolution. The actual image is clearer than the embedded one.

Double check the style of your citations. The format differs between citations with respect to spacing and comma use between the last name and year.

In the last paragraph, two species mentioned need to have the Genus capitalized and the species italicized.

AR:We have used the Plos One reference format

The last sentence in the Bats and Roost Characteristics paragraph is incomplete or contains an extra word.

The list of years- 2012,2013,2014,2018,2019. – should be 2012, 2013, 2014, 2018, 2019 with spaces in between and no period at the end. Similar spacing should be maintained in the list of distances surveyed in the Data Analysis paragraph.

AR: Rectified

I would recommend rephrasing your title and your introductory sentence of the Conservation of bats in temples paragraph. You’ve shown that many bats consistently utilize temples as roosts and that they have preferred microhabitats, but I don’t think you can infer that temples are essential for the persistence of bats. To do this, you would need telemetry data/population-level data for species to show that temples are the majority of roosting locations and reducing temples reduces bat populations because of limited roosting which may not be true for common species. There may be enough other buildings nearby to sustain bat populations without temples.

AR: We agree. Accordingly have modified the title “Temples and bats in a homogeneous agriculture landscape: Importance of microhabitat availability, disturbance and land use for bat conservation”.

Reviewer #3: There are still many grammatical corrections to make for publication (attached Word document with track changes). Also the in-text citations are not consistent with regard to semicolons and commas between citations and periods after et al. There are still species abbreviations missing the space between genus and species as mentioned by reviewer 1 previously.

AR: We have now addressed these issues. We have removed species abbreviations.

Submitted filename: author-responses.docx

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22 Jun 2022

Temples and bats in a homogeneous agriculture landscape: Importance of microhabitat availability, disturbance and land use for bat conservation

PONE-D-21-13889R2

Dear Dr. T,

We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.

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Additional Editor Comments (optional):

Thank you for your diligence in revising this manuscript. While a few minor errors in grammar, syntax, and formatting remain, I cam confident that they can be dealt with in the proofs. I look forward to seeing this article published.

Reviewers' comments:

30 Jun 2022

PONE-D-21-13889R2

Temples and bats in a homogeneous agriculture landscape: Importance of microhabitat availability, disturbance and land use for bat conservation

Dear Dr. T:

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