bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

1 How Fishing viverrinus : Describing a felid’s strategy to hunt aquatic 2 prey 3 *Divyajyoti Ganguly AND Tiasa Adhya 4 Department of Zoology, Maulana Azad College, Calcutta University, 8, Rafi Ahmed Kidwai Rd, 5 Taltala, , West 700013(DG). 6 Centre for Conservation of Natural Resources, The University of Trans-disciplinary Health 7 Sciences and Technology, 74/2, Post Attur via Yelahanka, Jarakabande Kaval, Bengaluru, 8 Karnataka 560064 (TA). 9 The Fishing Project, Small Wild Cat Conservation Foundation, 90 Rincon Rd, Corrales, 10 NM,87048-7619USA(TA). 11 *Correspondent: [email protected] 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29

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37 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

38 Hunting strategies are key to carnivore survival (Krebs and Davies 2009;Kamil et al.

39 2012;Michalko and Pekar 2016). Fishing Cat’s (Prionailurus viverrinus) persistence in the

40 ‘semi-aquatic niche’ (Kitchener et al. 2010) despite felids being terrestrial carnivores in general

41 (>95%) (Hunter 2019) suggests the evolution of a successful hunting strategy. Its morphological

42 adaptations further suggest selection for hunting in . What energy conserving strategies

43 did the Fishing Cat borrow from its family and how were these adapted to optimize energy

44 gained from hunting fish, its primary prey? We attempt to answer this by analyzing 197 video

45 footages collected opportunistically from a participatory science initiative conducted over 2.5

46 years. We found that the felid switches between stationary and active modes of searching for

47 prey depending on the depth of water and the corresponding loss of body heat/energy. For

48 example, diving in deeper waters requires the submergence of the upper portions of the body and

49 loss of more body heat/energy. Our analysis shows that the cat spent much of its time (~52%)

50 sitting and waiting for prey (fish) to come nearer and then took limited attempts to dive into

51 deeper water (2.78%). We suggest that this is a strategy to optimize the net energy gain. In

52 shallow waters where the cat could forage without submerging the upper body it adopted a

53 predominantly active mode of hunting (~96%) to flush out prey. Thus, prominent hunting

54 strategies in the small cat lineage like ‘sit-and-wait’ and ‘active foraging’ is adapted to hunt in

55 the water. We recorded a 60% hunting success in deeper waters but did not detect a successful

56 hunt in shallow waters due to the low sample size of data from shallow water hunting. The major

57 caveat in our study is the post-hoc analysis of opportunistically collected data as opposed to data

58 derived from a structured design with predefined objectives. With more sampling from various

59 seasons and landscapes, finer details can be explored which would have conservation

60 implications. For example, we would expect variations in ‘attempts to hunt’ during cold and bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

61 warm seasons because heat loss might be less of a challenge in the latter. Quantifying ‘attempts

62 to hunt’ and ‘successful hunts’ across seasons could help focus management interventions to

63 minimize negative interactions between fish cultivators and Fishing Cat. The strictly nocturnal

64 activity of fishing cat as demonstrated in our study could be a strategy taken by the cat to avoid

65 humans. Our approach of using participatory-science is relevant for conducting research on

66 behavior in human dominated landscapes.

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68 Keywords: camera-trap, ethogram, felids, Fishing Cat, hunting, participatory-science

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85 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

86 Successful hunting strategies allow the persistence of efficient predators through natural

87 selection (Krebs and Davies 2009;Kamil et al. 2012;Michalko and Pekar 2016). The global

88 carnivore guild exhibits a diversity of such hunting strategies that optimize prey capture while

89 overcoming foraging constraints (Hayward et al. 2006;Carbone et al. 2007;Williams et al.

90 2014;Rizzuto et al. 2018). For example, some members of hunt in groups to subdue

91 larger prey by coordinating a prolonged chase in open areas (Kruuk and Turner 1967;Murray et

92 al. 1995;Padilla and Hilton 2015). In contrast, mostly has solitary hunters that ‘stalk’ to

93 reduce distance to prey before a successful ambush (Carbone et al. 2007;Sunquist and Sunquist

94 2013). They do this with the help of cover in their immediate surroundings (Kruuk and Turner

95 1967;Elliot et al. 1977;Corbett 1979;Thiel 2011). With the exceptions of Cheetah (

96 jubatus) (Sunquist and Sunquist 2013) and Snow Leopard ( uncia) (Sunquist and

97 Sunquist 2017), felids do not engage in long chases.

98 A brief chase is present in some large felids while others hunt by pouncing or with a short rush

99 (Kruuk and Turner 1967;Sunquist and Sunquist 2017). Large felids in general hunt prey similar

100 to their body size (Carbone et al. 2007;Sunquist and Sunquist 2017). Chase is still less significant

101 in small felid (<25 kg) hunting strategies (Table 1) including in the larger and heavier ones (15-

102 25 kg) that can opportunistically take prey similar to their body size (Carbone et al.

103 2007;Sunquist and Sunquist 2017;Hunter 2019). Their diet is primarily composed of much

104 smaller prey such as , lagomorphs, , and invertebrates (Sunquist and

105 Sunquist 2017;Hunter 2019) that require lesser handling time, i.e., less energy (Carbone et al.

106 2007). Hence, capturing prey through short bursts of chase or by concentrating efforts in a final

107 attempt to hunt such as through leap/dash/pounce is common (Table 1). This counters the greater

108 amount of energy spent in searching and procuring smaller prey by small carnivores (Carbone et bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

109 al. 2007;Rizzuto et al. 2018). In addition, they lose more body heat (energy) due to a greater

110 surface area/body mass ratio (Tracy 1977).

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112 Small felids have evolved different hunting strategies depending on the prey taken. For

113 instance, small like rodents and hares are ambushed by sitting and waiting near

114 burrows or along trails frequented by prey or even by digging at their burrows (Sunquist and

115 Sunquist 2017). Birds on the ground are either flushed out from vegetation or stalked upon to

116 reduce distance (Olbricht and Sliwa 1997;Sunquist and Sunquist 2017). Arboreal primates are

117 hunted by climbing on trees and scaring them to lower branches and then to the ground as seen in

118 Ocelot ( pardalis) (Bianchi and Mendes 2007) whereas Margay (Leopardus wiedii)

119 uses vocal mimicry to lure prey (de Oliveira Calleia and Gordo 2009). Kitchener et al. (2010)

120 sequentially divide a typical felid hunt into ambush, detecting prey, stalking and running,

121 catching, killing, processing and eating. While ambush, detecting prey and stalking are typically

122 low-cost activities, running and catching are associated with higher costs (Williams et al. 2014).

123 The cost of killing, processing and eating differs with the type and size of prey. Small rodents or

124 birds can be easily subdued and consumed whole while large prey like large birds are required to

125 be processed by removal of feathers before eating (Sunquist and Sunquist 2017).

126

127 Fishing Cat (Prionailurus viverrinus) and Flat-headed Cat (Prionailurus planiceps) are

128 exceptional because of their choice of aquatic prey (Kitchener et al. 2010) as against terrestrial

129 prey taken by most felids. Further, morphological adaptations such as semi-retractile claws to

130 grip slippery aquatic prey and a double-coated fur to prevent the body from getting wet suggest

131 strong selection for hunting in wetlands (Hunter 2019). However, dentition in Fishing Cat bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

132 suggests a more eclectic diet when compared to Flat-headed Cat’s (Kitchener et al. 2010;Hunter

133 2019). The muscular build of Fishing Cat is suitable for hunting down similar sized terrestrial

134 prey (Sunquistand Sunquist 2017;Hunter 2019). Despite a broad dietary spectrum, fish is the

135 major dietary component (Haque and Vijayan 1993;Cutter 2015;Hunter 2019). Examining the

136 hunting strategy of the cat could provide insight on why such a preference could have evolved.

137 The elusive nature of the cat however is an impediment for studying their behavior in the wild

138 (Sunquist and Sunquist 2017). Only a few observations on their hunting exist (Prater

139 1965;Sunquist and Sunquist 2017;Hunter 2019). Technological advances such as motion

140 detecting camera traps have however facilitated non-invasive studies on their ecology (Nair

141 2012;Das et al. 2017;Malla 2016;Malla et al. 2019). But it is difficult to conduct such studies in

142 human dominated landscapes due to the issues of theft and requirement of greater manpower to

143 prevent the same.

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145 Here, we used data collected through a participatory-science initiative. Such initiatives

146 are targeted at achieving answers to real world questions under limited resources, in a cost-

147 effective way through a partnership between volunteers and scientists where volunteers

148 contribute data and/or assist with its analysis (Miller-Rushing et al. 2012;Lawson et al. 2015).

149 Such initiatives find use in a wide variety of wildlife disciplines. For example, ornithology has

150 benefited the most from citizen science initiatives with data on relative abundance and

151 distribution of species (Lawson et al. 2015). Participatory-science has also found use in

152 studies on wildlife management (van Vliet et al. 2018), land-use change (Johansson and

153 Isgren2017), wildlife monitoring (de Mattos Vieira et al. 2015), and wildlife disease surveillance

154 (Lawson et al. 2015). bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

155 We engaged residents sharing space with Fishing Cat in documenting and monitoring the

156 cats in their backyard through a camera trap exercise (http://thelastwilderness.org/know-thy-

157 neighbours-the-spectacular-fishing-cat/). We used opportunistically collected camera-trap data of

158 over 2.5 years from this program along with field observations. All the behavior types from the

159 Fishing Cat footages were first identified and used in ethogram construction. An ethogram is a

160 formal description in the form of a table of all behavior types observed in a species or of

161 particular functional classes of behavior(McDonnell and Poulin 2002).

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163 The major objectives of our study were to:

164 165 (a) describe major behavioral activities of Fishing Cat around waterbodies and channels through 166 ethogram 167 168 (b) examine activity budget in order to describe the hunting strategy of Fishing Cat 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

192 MATERIALS AND METHODS 193 Study area.––– Our study area consists of 3 sites, the district in the lower

194 Gangetic floodplains of and two coastal wetlands in the Mahanadi floodplains

195 along the Eastern coast - Paradip and Chilika (Fig. 1).

196 The (22°27'40.66"N, 88° 0'32.71"E) lies in the lower Gangetic

197 floodplains and is drained by the Hooghly and the Rupnarayan rivers. Our study was conducted

198 in fragmented marshy patches of Howrah district interspersed with croplands, small scale

199 , human settlements and industrial complexes. Human density of Howrah district is

200 also highest among all sites and due to the sub-urban nature of the populace, human activity is

201 high till 10 pm.

202 In Paradip (20°18'50.63"N, 86°36'45.49"E) data was collected from a marshy area of 9

203 sq. km inside the Paradeep Phosphates Limited campus. Small-scale aquaculture ponds

204 intersperse the marshes. Fishing activity by humans take place during day-time. After dark,

205 human activity is minimized since the area is accessible to campus residents only.

206 The study site in Chilika (19°51'39.66"N, 85°25'32.95"E) located along the lake’s north

207 east sector constitutes coastal wetlands formed by the Mahanadi floodplains in the Eastern

208 coast and has a contiguous marshland of 115 sq. km., crisscrossed by canals. Here local

209 fishermen catch wild fish during the day or anchor their boat in water channels at night after

210 laying nets and/or fish-baskets. The area is not inhabited by humans and all human activity stops

211 save for travelling fishing boats early morning. Human activity and presence is minimal in

212 Chilika when compared to Howrah and Paradip.

213 214 Data collection.––– We initiated a participatory-science initiative in the study sites

215 among interested and enthusiastic members of the local community who were given camera traps bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

216 to monitor Fishing Cat in their neighborhood. The initiative was named ‘Know Thy Neighbour’.

217 20 camera traps (14 Cover Illuminator 9 and 6 Browning IR) were deployed opportunistically

218 over 2.5 years starting from November 2016 across the different study sites. Camera traps were

219 deployed to maximize the probability of capturing Fishing Cat. We consulted local people to

220 decide where to deploy the camera traps. As suggested by them we searched for probable sites to

221 place camera traps along shallow ponds or water channels. All camera trap deployment sites

222 were chosen after locating scats and/or tracks of felids. They were set on video mode and were

223 active from 5:30 p.m. - 6:00 a.m. Each trained volunteer picked up traps early morning and

224 repositioned them at the same location at night to reduce chances of misplacement/theft. We

225 assumed that we would not fail to miss fishing cat activity when camera traps were non-active

226 since Fishing Cats are primarily nocturnal (Hunter 2019) and villagers mostly reported seeing

227 them after dark.

228 Data analysis.––– We collected over 200 camera trap video footages from the participatory

229 science initiative to analyze hunting behavior of Fishing Cat. All videos were analyzed by the

230 first author to avoid introducing observer bias. Consecutive videos from the same camera trap on

231 the same night which were taken less than a minute apart were considered as single data, as we

232 assumed lack of independence between closely occurring activities. ‘Behavior States’ were first

233 identified which represents behavioral patterns of relatively long duration (Martin and Bateson

234 1993). These were further divided into ‘behavior sub-states’. Behavioral patterns of relatively

235 short duration were classified as Events (Martin and Bateson 1993). We calculated the

236 proportion of time spent performing each State and the frequency of each Event using an

237 ethogram. bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

238 To examine activity pattern, videos (n=98) were selected such that they differed by at

239 least 30 minutes to ensure independence between data points. Such videos were then used to

240 generate line graphs to understand peak activity pattern of Fishing Cat. If two separate

241 individuals were detected in the videos, these were considered independent behavioural events

242 and analysed as two different samples.

243 In addition to these, any visible social behavior involving interactions between two or

244 more individuals was also noted.

245 In total, we analyzed 139.92 minutes from 197 camera trap video footages constituting

246 198 datapoints from all three study sites combined.

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260 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

261

262

263 RESULTS

264 This resulted in the identification of 12 distinct ‘Behavior States’ with their components,

265 i.e., behavior sub-types. The 12 ‘Behavior States’ are ‘Stationary Searching’ (with behavior sub-

266 types - ‘Standing’, ‘Sitting’, ‘Crouching’), ‘Mobile Searching’ (behavior sub-types: ‘Slow

267 Walking’, ‘Fast Walking’), ‘Approaching’, ‘Stalking’ (behavior sub-types: ‘Stationary Stalking’,

268 ‘Mobile Stalking’), ‘Attempt to hunt/Attacking’, ‘Eating Prey’, ‘Pheromone Spray’,

269 ‘Defecating’, ‘Sniffing’, ‘Self-Grooming’, ‘Swimming’ and ‘Yawning’. We found 7 States and 5

270 Events (Table 3 and 4). The detailed ethogram is presented in Table 2.

271 The felid spent much of its time (89.86%) searching for prey around resource patches

272 (Fig. 2).

273 274 275 Strategy in Deeper Water (n=180) 276 Searching.––– Here it walked either fast or at a slower pace (‘Mobile Searching’ =

277 32.79%) along the banks of water bodies, stopping intermittently to observe the water surface

278 and on a few occasions (n=9) to spray urine on bank vegetation. It also kept observing the edge

279 of the water body, which was usually fringed with floating/submerged/emergent vegetation, from

280 time to time while walking. On finding a suitable spot, it slowed down, sniffed the ground or air

281 (4.63%) and approached (5.15%) closer towards the water body. ‘Approaching’ was sometimes

282 followed by waiting (‘Stationary Searching’ = 51.65%). While waiting, which includes standing,

283 sitting and crouching, the cat remained in ambush, keeping mostly still and carefully watching

284 the water near the edge for signs of prey. Fishing Cat was also found to perch on beds of dried bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

285 emergent vegetation formed on banks presumably to catch fish entrapped in ‘dhauris’ or bamboo

286 fish-baskets set by artisanal fishermen.

287 288 Stalking.––– ‘Approaching’ is occasionally followed by ‘stalking’ (4.05%). When prey is

289 detected, it became alert and stalked with sight and ears focused in prey’s direction. While

290 stalking, the cat either a) kept still and waited for prey to swim within striking distance

291 (‘Stationary Stalking’), or, b) advanced closer on ground to reach within striking distance

292 (‘Mobile Stalking’). In doing so floating vegetation might have provided some concealment to

293 the cat.

294 Attacking.––– At striking distance the fishing cat plunged/dived (depending on depth)

295 into water (n=5) with forelimbs stretched followed by its head to catch prey. If the attack was

296 unsuccessful, the cat immediately came out of water, shook off excess water from the body and

297 in might take position immediately at a different spot.

298

299 300 Strategy in Shallow Water (n=18) 301 Searching.––– In knee-deep/paw-deep water the cat resorted to a more active form of

302 search (‘Active Searching’, 96.32%) whereby it walked in the shallow water, pawing at pebbles

303 or grass perhaps to flush out prey from hiding.

304 Stalking.––– The cat stalked (3.68%) by keeping still for prey to come within striking

305 distance, i.e., it only employed ‘Stationary Stalking’.

306 Attacking.––– The cat pounced (n=6) after prey either post stalking or immediately on

307 detecting movements while in ‘Active Searching’ mode. We did not record a successful attempt.

308 309 Eating and Killing bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

310 In a successful attempt(n=3), the cat grabs hold of the prey and emerges out of water onto

311 land. Small prey got consumed quickly and wholly along the bank while keeping vigilant.

312 However, large prey was dragged away from the water body as was observed in the field when a

313 -head presumably around 2 kilograms in weight was caught by a female Fishing Cat and

314 dragged into the nearby reeds. According to observations in the field, the cat consumed the flesh

315 of larger fish while the head, midrib, intestines and egg-sac are left behind.

316 Hunting Success 317 Overall, we recorded a naive hunting success rate of 27.27% (i.e., 3 successful attempts

318 out of 11 events). Frequency of attempt to hunt were recorded directly from camera trap records

319 and inferred from videos where the cat had fish-in-mouth. We assumed that an event of fish-in-

320 mouth must have been preceded by a successful hunting attempt. ‘Attempt to hunt’ in deeper

321 water was 2.78% and the same in shallow water was found to be 33.33%. Hunting success in

322 deeper water was 60% (i.e., 3 successful hunts out of 5 attempts). No successful hunt was

323 recorded from shallow water.

324

325 We found very little proportion of swimming behavior (0.35%). 326 327 328 Multiple individuals were found to use the same resource patch and were also found to

329 patrol the same camera trap site in the same night. Individuals were also found to spray

330 pheromones along vegetation, or, prominent structures like a stack of hay near the waterbody.

331

332 Cats engaged in ‘Self-Grooming’ sometimes (i.e., 1.25% of the time). Self-grooming is a

333 maintenance behavior after feeding (Rochlitz 2014).

334 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

335 We also recorded instances of social interactions between fishing cat individuals. In one

336 such instance, a male and a female were observed walking a trail, one behind another, along a

337 pond (Supplementary Data SD 16). In another instance, two individuals were observed

338 seemingly to communicate vocally by the side of a pond (Supplementary Data SD 14). A mother

339 with two sub-adult kittens was also observed to interact in a series of video clips while foraging

340 in shallow wetlands (Supplementary Data SD 15).

341

342 Nocturnal activity pattern of the fishing cat showed a gradual shift towards midnight with

343 increasing levels of human presence in the area. In Howrah district, the area with most human

344 presence and activity, Fishing Cat hunting times peak between 12 p.m. - 4 a.m. (higher peak;

345 n=37; Fig. 4a). Paradip area with intermediate human presence, showed Fishing Cat activity

346 peaking between 10-12 p.m. (n=4) and 2-4 a.m. (n=5) (Fig. 4b) and in Chilika area with the least

347 amount of human presence, activity peaked between 6-8 p.m. (n=8) and 10-12 p.m. (n=7) (Fig.

348 4c). All three areas combined, Fishing Cat activity peaked between 12 p.m.- 6 a.m. (n=59; Fig.

349 4d).

350

351 352 353 354 355 356 357 358 359 360 361 362 363 364 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

365 366 367 368 369 370 371 DISCUSSIONS 372 373 Most of Fishing Cat’s foraging time was spent in searching for prey (89.86%) in and

374 around resource patches. While searching for prey in deeper waters which required submergence

375 of the upper body, the cat alternated between mobile (37.94%) and stationary searching modes

376 (51.65%). When no prey was detected the cat shifted to mobile searching again. In the event of

377 an unsuccessful hunt, Fishing Cat came out of the waterbody, shook off the excess water and

378 took position immediately at a different spot. Sometimes it also returned to the same spot. These

379 findings are similar to an anecdote from Keoladeo Ghana Bird Sanctuary where it was observed

380 to wait along the water’s edge concentrating for signs of prey and was found to change positions

381 roughly every 15 minutes (Sunquist and Sunquist, 2017). In contrast, our study shows that it

382 employs a different strategy in shallow waters that did not require the upper portion of the body

383 to get wet. Here, it deployed an active mode of searching for prey, occasionally pawing the

384 waters to flush them out. This suggests that the cat switches from a combination of stationary-

385 and-mobile mode of searching for prey in deeper waters to a more mobile and active mode of

386 hunting in shallow waters in response to the differing needs to conserve energy.

387

388 Contrary to terrestrial foragers, aquatic foragers face additional physiological constraints

389 (Rosen et al. 2007). For example, semi-aquatic foragers such as , muskrat, platypus lose

390 body heat while foraging in water (Grant and Dawson 1978;Sherer and Wunder 1979;MacArthur

391 1984;Carss 1995;Kruuk 2006). Further, predator avoidance strategies of aquatic prey like fish bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

392 (Katzir and Camhi 1993;Harvey and Nakamoto 2013) must be accounted for during a successful

393 hunt. Hence, a successful hunting strategy must compensate for energy loss while optimizing

394 energy gain. For example, our data shows that the cat took measured attempts to hunt in deeper

395 waters, i.e., plunge/dive into water with outstretched forelimbs to catch prey. The ‘attempt to

396 hunt’ in deeper waters was found to be as low as 2.75%. This might be a strategy to maximize

397 chances of prey capture while minimizing energy loss since each attempt to hunt leads to energy

398 loss and too many attempts may alert the prey. The cat was found to be successful 60% of the

399 time it attempted to hunt in deeper waters. We did not record a successful hunt from shallow

400 waters but do not imply that Fishing Cat hunts more successfully in deeper waters. The findings

401 are rather an artefact of the low-sample size of data from shallow water hunting. Irrespective of

402 this, the chances of offsetting the high cost associated with attempting to hunt is variable and

403 thus strategies to conserve energy must evolve.

404

405 Felids are known to mitigate high hunting costs by reducing energy loss in other stages of

406 hunting, such as during pre-kill stages (Williams et al. 2014). The pre-kill stages constitute

407 efforts to detect prey by searching and then approaching/stalking to reduce distance to prey after

408 detection. While the ‘sit-and-wait’ search approach conserves energy, moderate energy might be

409 spent in actively searching for prey. Upon detecting prey, stalking is used selectively to reduce

410 distance to prey and increase chances of a successful hunt and in itself causes low energy loss.

411 The kill stage (attempt to hunt) incurs a high cost in deeper water as against a relatively moderate

412 cost in shallow water. Post-kill stages of fish consumption include subduing prey, processing,

413 consuming and digesting. After subduing prey, some energy is spent in dragging a big fish (1 – 2

414 kg) as was interpreted from a field observation by one of the authors and from online repository bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

415 (Bk Jeong 2017). Smaller prey may be consumed immediately next to the waterbody as was

416 observed in camera traps. However, fish once caught are easy to subdue as against terrestrial

417 prey (Sunquist and Sunquist 2017). Additionally, there is minimal cost of processing hunted fish

418 since scales are mostly excreted out along with scats. More importantly, fish protein is easily

419 digestible despite having high biological value (Vladau et al. 2008;Yilmaz et al. 2018). Thus, in

420 deeper waters, the high cost of hunting is offset by adopting energy-conserving pre-kill stages,

421 economic attempts to hunt and finally by successfully hunting highly-nutritious feed. In shallow

422 waters, more time is spent in actively searching for food which would probably increase the

423 chances of finding one and once a fish is successfully hunted, it could outweigh the energy spent

424 in actively searching for food. This perhaps serves to answer why Fishing Cat prefers fish over

425 rodents, which provide maximum metabolizable energy to small felids in general (Haque and

426 Vijayan 1993; Mukherjee et al. 2004; Hunter 2019). Thus, while the ‘sit-and-wait’

427 (Supplementary Data SD 7) and ‘active foraging’ strategy (Supplementary Data SD 9) from the

428 small cat lineage forms the backbone of Fishing Cat’s hunting strategy, its morphological

429 adaptations to hunt prey in water further enhances the success of this strategy. Such adaptations

430 are common among members of the carnivore guild that exhibit convergent evolution to forage

431 in the ‘semi-aquatic niche’ (Hunter 2019).

432

433 In addition, we recorded scent marking and faecal deposition, which are known

434 marking behaviors in felids (Brown and Bradshaw 2014;Burgos et al. 2018). We also recorded

435 sniffing behavior near resource patches. This is interesting as the strength of the odor in male cat

436 pheromone communicates the presence of high quality resource patches which is an attractant for

437 potential mates (Brown and Bradshaw 2014) whereas sniffing helps in sensing the presence of bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

438 other individuals as well as in gathering information on food resources (Geertsema 1984;Brown

439 and Bradshaw 2014). However, multiple individuals were also recorded to use the same site in

440 the same night in three instances. This could be because of the relatedness of these individuals.

441 While captive studies indicate possible male parental care (Rijsdijk 2011;Castaneda, [Cincinnati

442 Zoo and Botanical Garden, Cincinnati, United States of America], personal communication,

443 [December, 2019]) our data also shows sub-adults hunting in the presence of the mother. Thus,

444 despite being solitary hunters, related individuals could be using the same resource patch,

445 probably because of high density of prey, which is apparent from the use of such wetlands for

446 fishing/pisciculture by humans.

447

448 The strictly nocturnal activity of fishing cat in our study area could be attributed to two

449 reasons: a) the shift in activity pattern with intensifying levels of human presence could have to

450 do with avoidance of humans, much like how it adjusts to daytime hunting in order to co-exist

451 with larger competitors like Tiger and Leopard (Nair 2012) b) its crepuscular and nocturnal

452 activity could also be attributed to activity spikes in fish, its primary prey (Carss 1995; Banerjee

453 2018).

454

455 The major caveat in our study is the post-hoc analysis of opportunistically collected data

456 as opposed to data derived from a structured design with predefined objectives. It is unlikely that

457 the major conclusions of the study will vary, nonetheless, more sampling from various seasons

458 and landscapes will reveal finer details which could have ecological and conservation

459 implications, the latter is particularly relevant in human dominated landscapes. For example, we

460 would expect variations in ‘attempts to hunt’ during cold and warm seasons because heat loss bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

461 might be less of a challenge in the latter. Locals also report greater fish loss during summer

462 months when water levels in ponds decrease and hypoxic conditions prevail. Hence, quantifying

463 ‘attempts to hunt’ and ‘successful hunts’ across seasons could help focus management

464 interventions to minimize negative interactions between fish cultivators and Fishing Cat.

465

466 Finally, prey is central to predator ecology. Analyzing aspects of fish behavior with

467 respect to hunting by Fishing Cat will thus add a vital dimension. For example, anecdotal

468 accounts suggest that the cat taps the surface of the water to attract fish. Moreover, adult Fishing

469 Cat use their paws to feel for fish in muddy waters and in a captive facility, a blind Fishing Cat

470 was seen to feel around for food with his feet before plunging his head into the water to retrieve

471 the food (Buck [Aspinall Foundation, Lympne, United Kingdom], personal communication,

472 [July, 2011])). Additionally, hydrophytic vegetation provides refuge and congregation sites for

473 fish (Romare and Hansson 2003;Sohel and Lindstrom 2015). Predatory frequent

474 hydrophytic vegetation in search of prey (O’Hara 2012;Ren et al. 2019). Therefore, the presence

475 of such vegetation might be an important cue for hunting in both deeper and shallow waters.

476 Future studies which consider avoidance strategies of fish could significantly enhance our

477 knowledge on its hunting strategy.

478

479 Despite its caveats, the study is exemplary in demonstrating how by-catch data from

480 participatory-science initiatives could reveal hitherto unknown aspects of behavioral ecology

481 while also successfully circumventing the problems of equipment theft and need of greater

482 manpower to work in human dominated landscapes. Furthermore, it reveals how Fishing Cat

483 adapts to heavy human presence by shifting its foraging time to a chiefly nocturnal practice. The bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

484 cat could well be taking advantage of human-used fishing gears like bamboo fish-baskets which

485 have unidirectional valves through which fish enters and gets trapped as they were found to

486 perch on beds of dried emergent vegetation formed on banks along which the baskets were

487 deployed in ‘anticipation of prey’. Although we did not record Fishing Cat taking fish from the

488 baskets, locals reported that both Fishing Cat and Otter take the entrapped fish. How piscivorous

489 mammals take advantage of human-used gears requires further exploration as the strategy they

490 deploy gives them access to easy food but also incites negative interactions with fishermen.

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506 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

507

508

509 ACKNOWLEDGEMENT

510 511 The authors would like to acknowledge all participants of the ‘Know Thy Neighbours’

512 program - a participatory science initiative funded by Wildlife Trust of India, Wildlife

513 Conservation Trust and Mohamed Bin Zayed Species Conservation Fund. We would like to

514 extend our gratitude to student volunteers Ryan Rodrigues and Shreya Bhattacharya for initiating

515 the camera trapping exercise and would especially like to thank Subrata Maity, Anil Maity,

516 Bappa Malik, Maidul Islam Khan, Joydeb Pradhan, Sudhin Adhikari, Indrajit Adak, Saswat Pati,

517 Sourav Pati, Kaloshi Behera, Baraju Behera and his team members from Ma Pakshi Suraksha

518 Samity and members of Mahavir Pakshi Suraksha Samity whose enthusiastic participation is

519 especially mention worthy. We would also like to thank Iravatee Majgaonkar, Partha Dey for

520 their vital inputs in developing the manuscript. A special thanks to Dr. Kulbhushansingh

521 Suryawanshi who asked a question, during a discussion over the preliminary findings of the

522 study, which piqued us to re-look at our analysis and discover aspects that excited us.

523

524 SUPPLEMENTARY DATA

525 Supplementary Data SD 1.–––A video showing Fishing Cat fast-walking along waterbody

526 Supplementary Data SD 2.–––A video showing Fishing Cat slow-walking and searching along

527 waterbody

528 Supplementary Data SD 3.–––A video showing Fishing Cat switching between mobile

529 (walking, approaching) and stationary (standing) searching modes along waterbody bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

530 Supplementary Data SD 4.–––A video showing Fishing Cat sniffing and approaching towards

531 edge of waterbody followed by employing stationary searching

532 Supplementary Data SD 5.–––A video showing Fishing Cat approaching waterbody edge

533 Supplementary Data SD 6.–––A video showing Fishing Cat employing mobile stalking along

534 waterbody

535 Supplementary Data SD 7.–––A video showing Fishing Cat employing stationary searching

536 (sitting), followed by stationary stalking along waterbody, followed by attacking (misses the

537 catch), pheromone spraying and taking position again at another spot

538 Supplementary Data SD 8.–––A video showing Fishing Cat subduing and eating small fish

539 along waterbody

540 Supplementary Data SD 9.–––A video showing Fishing Cat employing active search in shallow

541 water

542 Supplementary Data SD 10.–––A video showing Fishing Cat engage in self-grooming along

543 waterbody

544 Supplementary Data SD 11.–––A video showing Fishing Cat swimming out of frame into

545 waterbody

546 Supplementary Data SD 12.–––A video showing Fishing Cat sniffing and depositing faeces

547 along water canal

548 Supplementary Data SD 13.–––A video showing Fishing Cat yawning along waterbody

549 Supplementary Data SD 14.–––A video showing two Fishing Cats seemingly communicating

550 along waterbody

551 Supplementary Data SD 15.–––A video showing two sub-adult Fishing Cats employing active

552 search in shallow water as the mother watches over from some distance bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

553 Supplementary Data SD 16.–––A video showing a male and a female Fishing Cat walking

554 along waterbody together

555

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813 Table 1.–––Weight, prey size and hunting technique in small felids

Serial Species Weight Main Prey Hunting Reference Number Range Technique (in Kg) 1 Black- 1-2.5 rodents, (1)‘Fast- Olbricht and Sliwa1997, footed Cat small birds, hunt’: moves Sunquist and Sunquist 2017, (Felis swiftly in nigripes) ‘bounds’ to IUCN/SSC Cat Specialist flush out prey Group from cover (http://www.catsg.org/) and attacks surprised prey; (2)‘Slow- hunt’: moves steadily, ‘winding’ through grass, moving head from side-to- side and ears in all directions to detect prey, stalks close and attacks in a final rush or jump; (3)‘Sit-down’ hunt: ‘sits with drawn up bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

front paws’ by dens for upto an hour, sometimes with closed eyes yet constantly moving ears and attacks in a final rush consisting of one or more ‘fast jumps out of the crouch’ 2 Pallas’ Cat 2-5 Small (1)‘stalking’: IUCN/SSC Cat Specialist (Otocolobus lagomorphs creeps slowly, Group manul) and rodents stealthily (http://www.catsg.org/), around cover to locate, Ross et al. 2019 move close to and pounce on prey (2)‘moving and flushing’: walks quickly through long grass and undergrowth to flush rodents, small birds, grasshoppers and pounces in attack (3)‘waiting in ambush’: waits outside active burrow for prey to emerge 3 Leopard 1.6-8 Rodents, Actively Sunquist and Sunquist 2017, Cat lizards, searches and IUCN/SSC Cat Specialist bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

(Prionailurus , ‘freezes’, Group bengalensis) small birds, crouches and (http://www.catsg.org/) insects ‘darts forward’ staying low on detecting prey 4 Geoffroy’s 3-6 Rodents Upon prey Branch 1995, IUCN/SSC Cat Cat detection Specialist Group (Leopardus hides behind (http://www.catsg.org/) geoffroyi) cover and closes in by crouching and sneaking from cover to cover and waits in ambush for prey, next it inches closer in a ‘low jerky posture’ and ‘springs forward’ at striking distance 5 Andean 4-6 Small Actively Sunquist and Sunquist 2017, mountain mammals, searches IUCN/SSC Cat Specialist Cat birds, lizards among and Group (Leopardus under rocks (http://www.catsg.org/) jacobita) and stalks once detected 6 European 3-8 Rodents, stalks by Corbett 1979, IUCN/SSC Cat Wildcat lagomorphs lying in wait Specialist Group (Felis near prey (http://www.catsg.org/) silvestris trails or silvestris) burrows and leaps at striking distance 7 Jungle Cat 5-9 Rodents, Stalks and Sunquist and Sunquist 2017, (Felis chaus) birds ambushes Ogurlu et al. followed by a 2010, IUCN/SSC Cat leap or Specialist Group pounce; bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

sometimes (http://www.catsg.org/) digs to retrieve prey , Funny Kitten Videos 2014 hidden in burrows fish Scoops up fish from bank or jumps/dives after fish from bank of water, may also enter shallow water in search of prey 8 Canada 8-12 Hares, Uses ambush Sunquist and Sunquist 2017, Lynx (Lynx squirrels, pounce IUCN/SSC Cat Specialist canadensis) grouse Group (http://www.catsg.org/) 9 African 7-16 Rodents, Uses stalk and Bahaa-el-din et al. 2014, Golden Cat shrews, birds rush upon IUCN/SSC Cat Specialist (Caracal prey detection Group aurata) (http://www.catsg.org/) 10 Caracal 6-18 Birds, Stalks to get Sunquist and Sunquist 2017, (Caracal rodents, close, waits in IUCN/SSC Cat Specialist caracal) hares ambush Group (sometimes (http://www.catsg.org/) for long) and dashes or leaps in attack 11 Serval 7-18 Rodents, Uses ‘high Sunquist and Sunquist 2017, (Leptailurus small birds, pounce’ on serval) lizards, unsuspecting Geertsema 1984, , prey to ‘stun’ IUCN/SSC Cat Specialist frogs, insects them Group (http://www.catsg.org/) 12 6-20 Lagomorphs, Uses a patient Sunquist and Sunquist 2017, (Lynx rufus) rodents sit-and-wait at IUCN/SSC Cat Specialist prey Group trails/burrows; (http://www.catsg.org/) patrols trails looking and listening, bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

leaps in pursuit or crouches and slinks forward on detecting prey (abandons chase after short burst of speed) 13 Eurasian 17-25 Lagomorphs, Uses stalk and Sunquist and Sunquist 2017, Lynx (Lynx rodents, ambush by IUCN/SSC Cat Specialist lynx) birds creeping close Group to prey and (http://www.catsg.org/) jumping (rarely pursues in chase after a miss); waits for prey in ambush by trails

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831 832 833 834 835 836 837 838 839 840 841 Table 2.–––Ethogram of Fishing Cat’s behaviour around resource patches (water body), 842 definitions from Stanton et al. 2015 Behavior Description Supplementary Data Slow Walking Forward locomotion along water body at a normal gait Supplementary with occasional scanning of water body Data SD 2 Fast Walking Forward locomotion along water body at a swift gait, Supplementary faster than slow walking Data SD 1 Approaching Slow and forward locomotion, sometimes in a slightly Supplementary crouched position, directed towards water body with Data SD 5 eyes and ears moving and scanning the water Standing All four legs extended and paws touching ground along Supplementary or within water Data SD 3 Sitting Hind legs flexed and resting on the ground, front legs Supplementary extended, body straight or lying on the ground against Data SD 7 on side of body with head raised, along water Crouching Body close to the ground, whereby all the legs remain Supplementary bent and belly touching or slightly raised above the Data SD 4 ground along water body Mobile Stalking Slow and forward locomotion in a slightly crouched Supplementary position, head low, eyes and ears focused on prey Data SD 6 (movement in water) Stationary Eyes and ears focused while stationary along water-edge Supplementary Stalking Data SD 7

Attempt to Either jumping into water from sitting/standing position, Supplementary hunt/Attacking or, charging towards and plunging front legs first into Data SD 7 water Eating prey Prey is ingested by chewing and swallowing Supplementary Data SD 8 Pheromone spray Spray released backwards against vertical surface or Supplementary object with vertically raised quivering tail while Data SD 7 standing or in a squatting position on the ground, sometimes while walking along water body Defecating Faecal matter deposited on the ground while in a Supplementary squatting position after thoroughly sniffing ground Data SD 12 along water body bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

Self-grooming Paw licked and wiped over head and body scratched Supplementary using hind leg along water body Data SD 10 Sniffing Air inhaled through nose with movement of whiskers Supplementary Data SD 4 Swimming Body propelled in water Supplementary Data SD 11 Yawning Mouth widely opened while inhaling and closed while Supplementary exhaling Data SD 13 843 844 845 Table 3.–––Proportion of time spent in performing each state as observed in deeper water, 846 shallow water and combined by Fishing Cat

Proportion of time spent (%) {(Time spent in performing one state/Time spent in performing all states) X 100}

State Deeper Water (n=180) Shallow water (n=18) Overall (n=198)

Mobile/Active 32.79 (Mobile) 96.32 (Active) 35.38 Searching Stationary Searching 51.65 ___ 49.75

Approaching 5.15 ___ 4.73

Stalking 4.05 3.68 4.29

Sniffing 4.63 ___ 4.25

Self-grooming 1.38 ___ 1.25

Swimming 0.38 ___ 0.35

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860 Table 4.–––Frequency of occurrence of each event as observed in Fishing Cat

Event Frequency of Occurrence

Attempt to hunt 11

Eating prey 3

Pheromone spray 9

Defecating 1

Yawning 1

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876 Fig. 1.––– Map showing study area sites (circled) across West Bengal and . Howrah, 877 Paradip and Chilika are marked with Black, grey and white circles respectively

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892 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

893 Fig. 2.–––Proportion of time spent in performing each state (Mobile Searching, Approaching, 894 Stationary Searching, Stalking, Sniffing, Self-Grooming, Swimming) by Fishing Cat, where 895 error bars represent Standard Error (SE)

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911 Fig. 3.–––Frequency of occurrence of each state (Attempt to hunt, Eating prey, Pheromone 912 Spray, Defecating, Yawning) by Fishing Cat

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928 Fig. 4a.–––Graph showing activity pattern of Fishing Cat in Howrah

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946 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

947 Fig. 4b.–––Graph showing activity pattern of Fishing Cat in Paradip

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966 Fig. 4c.–––Graph showing activity pattern of Fishing Cat in Chilika

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984 bioRxiv preprint doi: https://doi.org/10.1101/2020.04.24.058925; this version posted April 25, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission.

985 Fig. 4d.–––Graph showing general activity pattern (Howrah, Paradip and Chilika combined) of 986 Fishing Cat outside Protected Area (n=98)

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