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The Company Canids Confront: Resource Partitioning in Sympatric Carnivores in an Arid Ecosystem Kadambari Devarajan Post Graduate Program in Wildlife Biology and Conservation, National Center for Biological Sciences, Wildlife Conservation


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The Company Canids Confront: Resource Partitioning in Sympatric Carnivores in an Arid Ecosystem

Kadambari Devarajan

Post Graduate Program in Wildlife Biology and Conservation, National Center for Biological Sciences, Wildlife Conservation Society - India Program, Bangalore. M.Sc. Thesis Defense Guide: Dr. Abi Tamim Vanak (ATREE) Co-guide: Dr. Vishwesha Guttal (CES, IISc)

Kadambari Devarajan (NCBS) November 18, 2015 1 / 43

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Outline

Introduction Goal Prior Research Study Area Study Taxa Methodology Analysis Results Conclusions

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Introduction

Predators important for ecosystem function Multiple predators in a landscape Size-mediated interactions

[Levi & Wilmers, 2012] Kadambari Devarajan (NCBS) November 18, 2015 3 / 43

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Introduction: Carnivore interactions

Lethal interactions

Intra-guild predation Intra-guild competitive killing

Sub-lethal interactions

“Landscape of fear”

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Introduction: Species-scapes

“... a spatial plane of species interactions that combines with resources and habitat structure to drive species distributions”

[Fisher et al., 2012] Kadambari Devarajan (NCBS) November 18, 2015 5 / 43

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Goal

Identify how sympatric carnivores partition resources: space, time, habitat, and diet

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Study Area

http://bannigrassland.klink.co.in/images/Banni%20Location_2.jpg Kadambari Devarajan (NCBS) November 18, 2015 7 / 43

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Habitats

Image source: Pankaj Joshi Kadambari Devarajan (NCBS) November 18, 2015 8 / 43

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Taxa

Indian fox image: Abi Vanak Kadambari Devarajan (NCBS) November 18, 2015 9 / 43

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Other meso- and large carnivores present

Indian wolf - very rare Jungle cat, caracal, desert cat Striped hyena

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Prior Research

Bat-eared fox, cape fox, & black-backed jackals: South Africa

[Kamler et al., 2013]

Indian fox & domestic dog: India

[Vanak & Gompper, 2009a; Vanak & Gompper, 2009b; Vanak & Gompper, 2010]

Domestic dog & golden jackal: India

[Jhala & Aiyadurai, 2006]

Wolf - Coyote - Red/Swift fox: Canada, North America

[Voigt & Earle, 1983; Gese et al., 1996; Smith et al., 2003]

Red & Arctic fox: Canada, North America, Eurasia

[Hersteinsson & Macdonald, 1992] Kadambari Devarajan (NCBS) November 18, 2015 11 / 43

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Research Gaps

Mostly 2-3 species at a time Significant dietary niche separation Human-subsidized carnivores - rarely considered Species-scapes ignored

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Methodology: Overview

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Methodology: Design

Grid-based sampling approach: Occupancy framework

Nested design Single season, multiple species

Imperfect detection:

Spatial replicates → track plots Temporal replicates → camera traps

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Methodology: Grids

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Methodology: Sampling

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Data Collection: Effort

Camera trapping: All canids 74 grids * 4 camera traps * 4 days

≈ 300 cameras * 4 nights → 1200 camera trap

nights

≈ 6500 videos - 30 seconds each

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Data collection: Effort

Photographic capture-recapture: Dogs

17/50 villages in Banni - stratified random sampling based on village size (no. of households)

Kadambari Devarajan (NCBS) November 18, 2015 18 / 43

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Data collection: Covariates measured

On ground:

Vertical density Ground cover Vegetation type Presence of other canid species Food availability - burrow count, indirect signs of prey Anthropogenic influences - dung pat count, lopping

Kadambari Devarajan (NCBS) November 18, 2015 19 / 43

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Data collection: Covariates measured

Remotely-sensed/GIS:

Proximity to human habitation Proximity to road Proximity to water source Vegetation type

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Scat collection

Desert fox & Indian fox - only at active den sites Golden jackal - recorded defecating in front of camera

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Analysis: Camera Trap Videos

Desert fox in Banni Kadambari Devarajan (NCBS) November 18, 2015 22 / 43

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Analysis: Camera Trap Videos

Desert fox in Banni Kadambari Devarajan (NCBS) November 18, 2015 23 / 43

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Analysis: Occupancy

Assumption: Occupancy of larger canids not affected by smaller ones Model selection: Best model from multiple models for each species R package: unmarked Species maps: QGIS

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Results

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Very little dietary partitioning

0.00 0.25 0.50 0.75 0.00 0.25 0.50 0.75 0.00 0.25 0.50 0.75 Desert Fox Indian Fox Jackal rodent hare hedgehog bird stl reptile coleopteran isopteran arachnid

  • rthopteran

hymenopteran

  • donate

crustacean suaeda prosopis grass_root grass_leaf wood_piece zizyphus cucurbit

Diet Composition Frequency of Occurrence

Vertebrates Invertebrates Plants

Kadambari Devarajan (NCBS) November 18, 2015 26 / 43

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Wild canids crepuscular + nocturnal

5 10 5 10 5 10 5 10 Desert Fox Dog Golden Jackal Indian Fox 00h 03h 06h 09h 12h 18h 21h

Time Active

  • No. of Encounters

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Naïve occupancy

From camera trap data: Species Sites Present Total Sites % Indian Fox 61 675 9 Golden Jackal 392 675 58 Desert Fox 81 675 12 Dog 74 675 11

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DF & Dog - Complete separation

  • Kadambari Devarajan (NCBS)

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IF & GJ - Complete overlap

  • Kadambari Devarajan (NCBS)

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IF & DF - Near complete separation

  • Kadambari Devarajan (NCBS)

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GJ & Dog - Near complete separation

  • Kadambari Devarajan (NCBS)

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Spatial partitioning: All canids

  • Kadambari Devarajan (NCBS)

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Spatial partitioning: Conceptual diagram

Desert Fox Indian Fox Jackal Dog Village Kadambari Devarajan (NCBS) November 18, 2015 34 / 43

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Species interactions: Occupancy analysis

From β values estimated from best model: Species IF DF Jackal Dog IF

  • ve

+ve

  • ve

DF

  • ve
  • ve

Jackal

  • ve

Dog

  • ve

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Conclusions

Very little dietary partitioning

Especially among foxes Minor differences in plant matter

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Conclusions

Wild canids primarily crepuscular + nocturnal

Dogs diurnal No canids active - 2 to 4 PM

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Conclusions

Desert fox - spatial partitioning with other canids Dog - spatial partitioning with other canids Indian fox & golden jackal - spatial overlap

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Acknowledgements

  • Dr. Abi Tamim Vanak
  • Dr. Vishwesha Guttal
  • Dr. Ajith Kumar

Chandni Gurusrikar

  • Dr. Pankaj Joshi, Dr. Suhel Quader, Chandrima Home, Kulbushan S. Suryawanshi, Anil Gohil, Girish Punjabi,

Ovee Thorat, N. Lakshminarayanan, Chetan Misher, Chintan Sheth, Ameya Gode, Jayashree Ratnam, Tarun Nair, Priyanka Runwal, Dharmaveer Shetty, Jayashree Ratnam, Megha Bhai, Rasul Bhai, Mutthalib Bhai, Kabul Bhai, Salim Mama

  • Dr. Prabhu Ramachandran

My parents and sister Kadambari Devarajan (NCBS) November 18, 2015 39 / 43

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References

Aiyadurai, A. and Y.V. Jhala. (2006) Foraging and habitat use by Golden Jackal (Canis aureus) in the Bhal Region, Gujarat, India. J. Bombay Nat. Hist.Soc., 103(1): 5-12. Donadio, E. & Buskirk, S.W. (2006) Diet, Morphology, and Interspecific killing in Carnivora. American Naturalist. Vol.167, No.4. Fisher, J.T., Anholt, B., Bradbury, S., Wheatley, M., & Volpe, J.P . (2012) Spatial segregation of sympatric marten and fishers: the influence of landscapes and species-scapes. Ecography, 35: 001-009. Gese, E.M., Stotts, T.E., & Grothe, S. (1996) Interactions between coyotes and red foxes in Yellowstone National Park,

  • Wyoming. J. Mammalogy. 77(2):377-382.

Gompper, M.E. & Vanak, A. T. (2008). Subsidized predators, landscapes of fear, and disarticulated carnivore communities. Animal Conservation 11: 13-14. Hersteinsson, P . & Macdonald, D.W. (1992) Interspecific competition and the geographical distribution of red and Arctic foxes Vulpes vulpes and Alopex lagopus. Oikos, 64:505-515. Jhala, Y. V. & Moehlman, P .D. (2004) 6.2. Golden Jackal Canis aureus Linnaeus, 1758. In: Sillero-Zubiri, C., Hoffmann, M., & Macdonald, D. W. (eds): Canids: Foxes, Wolves, Jackals and Dogs. Status Survey and Conservation Action Plan. IUCN-SSC Canid Specia list Group. Kamler, J. F., Stenkewitz, U., Klare, U., Jacobsen, N.F ., & Macdonald, D.W. 2012. Resource partitioning among cape foxes, bat-eared foxes, and black-backed jackals in South Africa. Journal of Wildlife Management 76:1241-1253. Kamler, J. F., Stenkewitz, U., & Macdonald, D.W. 2013. Lethal and sublethal effects of black-backed jackals on cape foxes and bat-eared foxes. Journal of Mammalogy 94:295- 306. Kadambari Devarajan (NCBS) November 18, 2015 40 / 43

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References

Roemer, G.W., Gompper, M.E., & van Valkenburgh, B. (2009) The ecological role of the mammalian mesocarnivore. BioScience, Vol.59, No.2. Sarmento, P .B., et al. (2010) "Habitat selection and abundance of common genets Genetta genetta using camera capture-mark-recapture data." European Journal of Wildlife Research 56.1: 59-66. Scheinin, S., Yom-tov, Y., Motro, U., & Geffen,E. (2005) Behavioural responses of red foxes to an increase in the presence

  • f golden jackals: a field experiment. Animal Behavior, 71: 577-584.

Sillero-Zubiri, C., Hoffman, M., & Macdonald, D.W. (2004) (Technical Editors) Canids: Foxes, Wolves, Jackals, and Dogs.

  • Gen. Tech. Rep. Status Survey and Conservation Action Plan, IUCN/SSC Canid Specialist Group.

Smith, D.W., Peterson, R.O., & Houston, D.B. (2003) Yellowstone after wolves. BioScience. 53(4):330-340. Levi, Taal, & Wilmers, Christopher. Wolves-coyotes-foxes: a cascade among carnivores. Ecology 93.4 (2012): 921-929. Vanak, A.T., Irfan-Ullah, M. & Peterson, T. (2008). Gap analysis of Indian fox conservation using ecological niche

  • modeling. Journal of the Bombay Natural History Society 105: 49-54.

Kadambari Devarajan (NCBS) November 18, 2015 41 / 43

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References

Vanak, A.T. & Gompper, M.E. (2009). Dogs emph(Canis familiaris) as carnivores: Their role and function in intraguild

  • competition. Mammal Review 39: 265-283 - Included in 2012 Mammal Review Virtual Issue.

Vanak, A. T., Thaker, M & Gompper, M. E. (2009). Experimental examination of behavioural interactions between free-ranging wild and domestic canids. Behavioural Ecology and Sociobiology 64:279-287. Vanak, A.T. & Gompper, M.E. (2009). Dietary niche separation between sympatric free-ranging domestic dogs and Indian foxes in central India. Journal of Mammalogy 90:1058-1065. Vanak, A.T. & Gompper, M.E. (2010). Interference competition at the landscape level: the effect of free-ranging dogs on a native mesocarnivore. Journal of Applied Ecology 47: 1225:1232. Vanak, A. T., Fortin, D., Thaker, M., Owen, C, Lehmann, M, Greatwood, S. & Slotow, R. (2013). Moving to stay in place: behavioural mechanisms of co-existence in an African large carnivore guild. Ecology. Voigt, D.R. & Earle, B.D. (1983) Avoidance of coyotes by red fox families. J. Wildlife Mgmt. Vol. 47, No. 3, pp. 852-857. Zielinski, W.J. & Kucera, T.E., technical editors. (1995) American marten, fisher, lynx, and wolverine: survey methods for their detection. Gen. Tech. Rep. PSW-GTR-157. Albany, CA: Pacific Southwest Research Station, Forest Service, U.S. Department of Agriculture: 163 p. Kadambari Devarajan (NCBS) November 18, 2015 42 / 43

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“Kutch nahi dekha, to kuch nahi dekha!” - The Gujarat Tourism Slogan

THANK YOU!

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