1 Camera trapping for animal monitoring and management: a review of applications Don E. Swann and Nic Perkins

Abstract Introduction Camera traps are being used throughout the world Cameras that record images of wild animals when to address a wide range of issues in wildlife man- humans are not present have a long history in ecol- agement and to address both research and man- ogy, but their use dramatically increased with the agement questions that cannot be easily answered introduction of commercial infrared-triggered with other methods. In addition to detecting rare cameras in the early 1990s. Today, the term ‘camera species and providing answers to practical man- trap’ typically refers to cameras units that are trig- agement questions, camera traps have a potentially gered by the movement of an animal within a large role in assessing global changes in biodiver- detection area, although the term also can describe sity of mammals. The quality of camera traps is cameras set to take photos at set time intervals. continuing to improve, and field and analytical Nearly all camera traps used in current wildlife techniques are also moving rapidly forward. This applications are small (the size of a shoebox or paper reviews the current state of camera trapping smaller), consist of only one piece, shoot digital still in wildlife ecology with a focus on new and or video images, and are passively triggered using emerging applications in management and moni- an infrared light source. Nevertheless, a dazzling toring. Recent papers, including many in this array of commercial camera traps and optional fea- volume, indicate that camera traps have the poten- tures are available (Rovero et al. 2013; Swann et al. tial to be a powerful new tool in areas of animal 2004, 2011; ), and ecology where they have not previously been these can be further modified by researchers. widely used, such as estimation abundance, sam- Camera traps have been applied to nearly every pling of small animals, and establishing conserva- aspect of vertebrate ecology, including to study tion priorities based on regional monitoring. In nest ecology, research activity patterns and behav- addition, the use of camera traps by citizen scien- iour, document rare species or events, and estimate tists and environmental educators continues to state variables such as species richness, occupancy, grow and become more integrated with more tra- abundance (Cutler and Swann 1999; Kucera and ditional scientific studies. Barrett 2011). Data recorded by camera traps

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typically consist of an image (e.g. Plate 1.1), series Chapter 10). Kucera and Barrett (2011) list several of images, or video of an individual or group of recent examples, including a new species of striped animals within the area of detection covered by the rabbit (Nesolagus timminsi) in South-east Asia (Sur- camera trap, as well as other information such as ridge et al. 1999), a range extension for the Sulawesi the date, time, and location of the photograph. palm civet (Macrogalidia musschenbroekii) (Lee et al. Because most individuals in the image can be iden- 2003) and the documentation of the wolverine tified to species, the trap thus records the presence (Gulo gulo) in California for the first time since 1922 of that species at that place and time. Other infor- (Moriarty et al. 2009). Camera traps also have been mation, such as behavioural data (e.g. Meek et al. used to reliably estimate, for the first time, the 2012) or events such as predation or feeding (e.g. abundance of the tiger (Panthera tigris; Karanth and Zimmerman et al. 2011) can also be recorded. In Nichols 1998) and other species where individuals some cases individual animals can be identified can be readily recognised based on their spot pat- either through tags previously affixed by research- tern. After many years of debate and poor informa- ers or by unique natural marks. Some data gath- tion on the number of species of mammals present ered by physical trapping techniques, such as in natural areas, camera traps are now producing reproductive condition and genetic data, cannot reliable estimates of species richness and other usually be obtained by camera traps, but camera community measures (Tobler et al. 2008; O’Brien et trap data are often combined with other techniques al. 2011) that are not based on poor-quality observa- such as radio-telemetry (e.g. Larrucea et al. 2007) tional data or generalised range maps (e.g. New- and genetics (e.g. Janečka et al. 2011). mark 1995; Parks and Harcourt 2002). Despite the limits of the data that can be gath- As demonstrated at the First International ered by camera traps, experience during the past Camera Trapping Colloquium in Wildlife Manage- two decades indicates why they are powerful tools ment and Research, new developments in camera for addressing conservation of populations of trapping are arriving at a staggering pace. Camera native species, especially mammals. First, camera traps themselves are rapidly evolving, becoming traps provide basic knowledge of the distribution faster, cheaper, more resilient, and more versatile of mammals (their presence in a certain place), (Meek 2011). Researchers are developing new tech- which is essential for conservation on both the niques for deploying them, including using verti- local and regional scale, but often previously lack- cal mounts (Swann et al. 2004; Welbourne, Chapter ing for the many species that are nocturnal, avoid 20). More researchers are aware that detectability is humans, and seek cover. Second, they are relatively always < 1 in camera studies (Kéry 2011) and are inexpensive, which means they can be deployed becoming more adept and sophisticated at analys- very efficiently to increase sample sizes over wide ing camera trap data using occupancy and other areas (De Bondi et al. 2010). And third, camera traps methods (O’Connell and Bailey 2011). They are rap- are relatively non-invasive and safe for both idly improving methods for managing data, humans and animals. From a practical point of including extracting data directly from photos, view, it is obviously much easier to sample popula- eliminating the need for data entry, and even tions of very large mammals with camera traps developing methods for using pattern recognition than with live traps. to identify animals automatically (Falzon et al., As a result, camera trapping has been truly sig- Chapter 28). And of course, they are improving nificant for wildlife management and conservation methods for sharing data through the internet and throughout the world. Camera traps have docu- cell phone applications. mented species that are new to science, or occur in What is truly exciting about camera trapping in areas where they were thought to be locally extinct the modern era is that the explosive adoption of or not previously known to exist (e.g. Sangay et al., this technology, in synergy with improvements in

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camera trap quality, wildlife data analysis, and animals) and the characteristics of animals that information management, is resulting in novel can be determined from videos taken at the camera applications in wildlife conservation around the sites, including its size and speed. world. It is a testament to the rapid advances in One of the complications of using the REM has camera trapping that several important develop- to do with the difficulty of accurately estimating ments in camera trapping have occurred since the the area of the detection zone. Although this prob- very recent publication of a major book on the sub- lem has been addressed using distance sampling ject, Camera Traps in Animal Ecology (O’Connell and techniques by Rowcliffe et al. (2011), papers pre- Bailey 2011). The goal of this paper is to provide a sented in this volume (e.g. Welbourne, Chapter short summary of some of these applications and 20) suggest that another approach may be to explore their potential for creating ground-break- mount cameras vertically (facing down at the ing developments in the coming years. ground), so as to more precisely control the area of detection. This approach may not be appropri- ate for larger animals in tropical areas where a Estimating animal abundance and density large amount of vegetation is present, but might Camera traps are often used to provide an index of work in less vegetated areas and for smaller mam- abundance (also called relative abundance), such mals in most habitats. as the number of photos of a species per trap night. However, indices typically provide biased esti- mates of abundance (Anderson 2001), which in Studying small animals camera trapping is primarily due to spatial varia- Another interesting new direction for camera trap- bility and detectability (O’Brien 2011). Many users ping is in studies of smaller animals, including of camera traps assume that the number of photo- mammals such as insectivores, rodents, and small graphs per unit time is an accurate reflection of the marsupials, as well as birds and herpetofauna (e.g. number of individual animals present. However, Welbourne, Chapter 20). Camera traps have been this assumption may not be valid because many used in small mammal research for some time, but factors may influence the number of photographs, typically to determine temporal patterns (e.g. Pear- including attraction to the camera trap, trap shy- son 1959). Recent work on small mammals with ness, use of or type of attractant, weather, ability of more accurate new commercial traps, particularly the camera trap to detect an animal when present, in Australia with ReconyxTM brand traps that pick and others. Several recent studies (see Karanth et up smaller heat signatures (e.g. Meek et al. 2012; al. 2011) have followed Karanth (1995), who used Weerakoon et al., Chapter 29), allow researchers to camera traps and individual natural markings to estimate variables such as abundance (where indi- estimate tiger abundance and density using cap- vidual animals are marked or have identifiable pat- ture–recapture models. However, this approach terns), species richness, and occupancy that does not work for species without features that previously required the use of live traps. The work allow them to be individually recognised. The by Rowcliffe et al. (2008, 2011) applies to estimating recent work by Marcus Rowcliffe and colleaques density of small mammal abundance, which has (Rowcliffe et al. 2008, 2011) is thus truly important the potential to replace live trapping for many for camera trap researchers, as it presents an oppor- studies, especially if combined with vertical camera tunity to estimate density by modelling the under- mounts on a network grid of small mammal sen- lying process of the encounter between the camera sors at random locations with no bait. A paper pre- trap and the animal. Their random encounter sented at the First International Camera Trapping model (REM) relies on characteristics of the camera Colloquium in Wildlife Management and Research trap (the distance and angle with which it detects by Falzon et al. (Chapter 28) presented a method for

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automated species identification that allows for can further developed with the use of species rec- removal of the consistent background image in ognition software and non-infrared triggered cam- photos and use of visual algorithms to identify eras that run continuously. individual species. Although this approach is still in early stages of development, it is expected to grow and have important implications for monitor- Applied management ing not only abundance and density using the While large conservation issues, such as preserv- REM, but also species richness and occupancy. ing the biological diversity of our planet, require a Further, because infrared triggered cameras may large scale vision (Wilson and Peter 1988), most always have imperfect detectability, it seems con- actual conservation happens locally. Every day, ceivable that in the future constant photo or video throughout the world, wildlife managers strive to surveillance of a series of unbaited, random plots reduce the impacts of introduced predators, would produce the potential for higher detection develop structures to prevent threatened species rates and very accurate estimation of small from being killed by cars, create artificial water mammal populations, as well as populations of resources to replace lost natural sources, build herpetofauna and ground birds. gates on caves to protect bats, and engage in many Finally, camera traps have typically been used other types of hands-on conservation efforts. in studies of mammals and birds, and only occa- The basic type of data collected by camera traps sionally used in studies of reptiles and amphibians. – the image or images of an animal in a certain Dustin Welbourne’s presentation at the Colloquium place at a certain time – has been very useful for (Welbourne, Chapter 20) is interesting not only addressing local wildlife management questions because it focuses on herpetofauna, but is poten- for some time, and continues to be very useful. tially ground-breaking in its approach to inventory Recent published papers around the world expand and monitoring of this taxonomic group. Even the uses of camera traps to topics as diverse as more than small mammals, reptiles and amphibi- assessing the value of artificial water sources ans are very difficult to detect due to their small (Krausman et al. 2006); comparing different types of size and (as ectotherms) lack of a heat signature. forestry practices on mammal communities (Same- Many species are rare, cryptic, and active only jima et al. 2012); assessing effects of human presence during specific windows of heat and humidity. on large carnivore populations (Muhly et al. 2011); Herpetologists have traditionally used a variety of assessing effects of predator control on prey species methods to estimate population parameters, populations (Salo et al. 2010); evaluating the poten- including observations and times searches (Visual tial for expansion of the wild boar population in Encounter Surveys, or VES), pitfall traps, hoop Switzerland (Wu et al. 2011); and many others. Case traps, and other methods. Welbourne’s work sug- studies at the colloquium that used camera traps for gests that a more efficient and less invasive applied management applications included approach may be to use camera traps, mounted addressing the effectiveness of different types of vertically over plates that can be naturally warmed baits for decreasing introduced carnivores while to a different temperature than the surrounding preserving native species (Antos and Yuen, Chapter substrate, in conjunction with drift fences. It 2; Moseby and Read, Chapter 14; Bengsen, Chapter remains to be seen whether this technique can be 31); quantifying the use of water holes by intro- modified to sample the range of terrestrial reptiles duced camels and their interactions with water and amphibians with different types of movement availability and native animals (Ninti One Ltd and habitat use (e.g. sand-dwelling, rock-dwelling, 2013); whether and how native mammals use glide aquatic and aboreal). In addition, it may be possible poles and other structures to successfully cross to eliminate drift fences if capture rates are high roadways (Taylor and Goldingay, Chapter 22); and enough. As with small mammals, this technique identifying the predators of the New Zealand kea

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(Nestor notabilis, B. Barrett pers. comm.). In addition, coming these issues. Thus the greatest current camera traps are being used in some truly innova- challenge may be to organise researchers to deploy tive management applications, such as to identify cameras in a systematic way and share the data so Tasmanian devils (Sarcophilus harrisii) with devil the information can be most useful for monitoring facial tumour disease and track the spread of this trends over time and establishing conservation pri- disease (Thalmann et al., Chapter 3). orities. Because the focus is on entire communities, The main trend in using camera traps in applied such an approach requires a randomised study management studies is that camera traps are rap- design with unbaited camera traps. On a local level, idly becoming more affordable and reliable. Many an increasing number of such programs in national wildlife managers, especially in developing coun- parks and other reserves are beginning to monitor tries, have lacked effective methods for determin- communities in this way. ing whether their conservation actions are indeed Even more exciting is the development of pro- effective. The potential for obtaining larger num- grams that monitor the status of communities on bers of camera traps make it easier not only to cap- larger geographic scales. A primary example of ture data that may provide insight into a particular this is the Terrestrial Ecology Assessment and management issue, but also to implement studies Monitoring (TEAM) network (Ahumada et al. 2011; with a higher sample size, thus providing results Jansen et al., Chapter 24), which, in an international that allow for greater inference. partnership at 16 sites across the globe, collects data useful for conservation of tropical forest mam- mals. TEAM has developed a detailed, standard Monitoring of animal communities at local, protocol and data storage and sharing methods regional and international scales that allow for comparisons of species richness, Knowledge of the number of species occurring in a diversity, and evenness among sites. In addition, particular place, either locally as in a national park, programs like the Wildlife Picture Index (WPI) or regionally or even globally, is essential for man- (O’Brien et al. 2010; Townsend et al., Chapter 5) aging and conserving biodiversity, but the poor combine camera traps with occupancy analysis state of this knowledge remains one of the greatest and generalised additive models to allow conser- hindrances to conservation (O’Brien et al. 2011). vationists to monitor trends in biodiversity over Because funding for conserving biodiversity are time. The success of approaches such as TEAM and always less than what is needed, establishing con- WPI suggest that similar standardised approaches servation priorities is one of the most vexing prob- can accomplish important conservation work lems in conservation biology (Isaac et al. 2007). across a variety of regional and national scales, and Three-quarters of all species-based conservation deserve to be implemented more widely. projects are specifically aimed at charismatic meg- afauna (Leader-Williams and Dublin 2000). While part of the problem is the nature of the public’s Citizen science and education attention, the lack of information on many species Two final developments in applications of camera is also an issue. traps in monitoring and management are the grow- There are many challenges to tracking of the ing use of camera traps in education and the estab- status of different taxonomic groups, ranging from lishment of citizen science networks that involve the sheer number of species to the complexities of non-scientists in collecting important scientific their life cycles. The challenge of monitoring ani- data (e.g. Griffiths and Lewis, Chapter 9; Thomas, mals typically detected by camera traps, particu- Chapter 8 volume). There is considerable overlap larly medium and large mammals, is that they are between these two areas. In a recent major event at often rare, nocturnal, and avoid humans. However, Saguaro National Park in Arizona, the 2011 BioBlitz, camera traps have proven to be very useful in over- students learned to use and deploy camera traps in

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Fig. 1.1. Camera trapping as education: students at Saguaro National Park, USA, set a camera trap as part of 2011 BioBlitz.

an effort to both document mammals and learn found by making a Google search of Flickr and more about their habits and conservation needs, camera traps, for example) includes eMammal and the park posted most of the photos gathered (; ). Photos Wildlife/Animals/Archives/2012/Camera-Traps- from camera traps are being used the world over to and-Conservation.aspx> ), which is being developed excite people about the wildlife around them that as an international archive of wildlife photos by the usually remains unseen. These images are often Smithsonian Institute. At the same time, some cau- quite powerful and have great potential for pro- tion is in order. Camera traps are a neutral way to moting conservation among citizens who might detect animals, and can be used just as easily as a otherwise not be interested. tool to detect and kill wildlife as to detect and con- The primary user of camera traps in many coun- serve them. As researchers share information, it is tries, and particularly the United States, is by hunt- important to recognise that the potential downsides ers. In the USA, the value of hunter-collected data in of sharing the secrets of wildlife. the documentation of rare and threatened cannot be underestimated. Where we live in Arizona, USA, for example, camera traps set by hunters have pro- Conclusion vided some of the best recent records of rare tropi- What makes camera trapping unique among the cal cats such as jaguars and ocelots. Networks of many methods used by wildlife ecologists is that it camera traps maintained by volunteers are proving is, in a sense, both a science and an art. It is a science to be valuable for conservation in some areas (e.g. in that camera traps can produce data that are Erb et al. 2012). The effort to tap into the larger pool useful for addressing questions about our world of existing camera trap data (which can be easily works. But the same process produces images that

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can be ‘appreciated primarily for their beauty or De Bondi N, White JG, Stevens M, Cooke R (2010) emotional power’, which is how the Oxford Diction- A comparison of the effectiveness of camera ary () defines art. As Dustin Welbourne trial small-mammal communities. Wildlife recently wrote, ‘the raw data collected with cameras Research 37(6), 456–465. and recorders can be experienced. It is visceral and Erb PL, McShea WJ, Guralnick RP (2012) Anthro- tangible to our senses, and even seasoned research- pogenic influences on macro-level mammal ers are often excited to “‘see’” what has been occupancy in the Appalachian Trail Corridor. detected’ (Welbourne 2012b). It is important for sci- PLoS ONE 7(8), e42574. entists and managers that the art function of photos Isaac NJB, Turvey ST, Collen B, Waterman C, Baillie produced by camera traps does not become a sub- JEM (2007) Mammals on the EDGE: conserva- stitute for the science function; that is, no matter tion priorities based on threat and phylogeny how excellent and interesting a camera trap images PLoS ONE 2(3), e296. are, they cannot replace the requirement of having Janečka JE, Munkhtsog B, Jackson RM, Naranbaatar clear objectives, strong study design, and efficient G, Mallon DP, Murphy WJ (2011) Comparison of and accurate methods for managing photo data. noninvasive genetic and camera-trapping tech- At the same time, the art value of camera traps niques for surveying snow leopards Journal of has a place in conservation biology that is signifi- Mammalogy 92, 771–783. cant and not likely to go away soon. It is this Karanth KU (1995) Estimating tiger Panthera tigris dynamic between science and art that would seem populations from camera-trap data using cap- to ensure that the value and creative uses of camera ture-recapture models. Biological Conservation traps are likely to continue to grow, as today’s 71, 333–338. camera traps merge with newer technologies in Karanth KU, Nichols JD (1998) Estimation of tiger ways that may rapidly change the definition of the densities in India using photographic captures term. The challenge for conservation biologists is and recaptures. Ecology 79(8), 2852–2862. to harness this creativity with sound science in Karanth KU, Nichols JD, Kumar NS (2011) Estimat- ways that will help the Earth’s many magnificent ing tiger abundance from camera trap data: field creatures survive until the next generation. surveys and analytical issues. In AF O’Connell, JD Nichols and KU Karanth (Eds) Camera Traps in Animal Ecology. Pp. 97–118. (Springer Publish- References ing, Tokyo). Ahumada JA, Silva CEF, Gajapersad K, Hallam C, Kéry M (2011) Species richness and community Hurtado J, Martin E, McWilliam A, Mugerwa B, dynamics: a conceptual framework. In AF O’Brien T, Rovero F, Sheil D, Spironello WR, O’Connell, JD Nichols and KU Karanth (Eds) Winarni N, Andelman SJ (2011) Community Camera Traps in Animal Ecology. Pp. 207–232 structure and diversity of tropical forest mam- (Springer Publishing, Tokyo). mals: data from a global camera trap network. Krausman PR, Rosenstock SS, Cain JW, III(2006) Philosophical Transactions of the Royal Society of Developed waters for wildlife: science, percep- London. Series B, Biological Sciences 366, tion values, and controversy. Wildlife Society Bul- 2703–2711. letin 34, 563–569. Anderson DR (2001) The need to get the basics right Kucera TE, Barrett RH (2011) A history of camera in wildlife field studies. Wildlife Society Bulletin trapping. In AF O’Connell, JD Nichols and KU 29, 1294–1297. Karanth (Eds) Camera Traps in Animal Ecology. Cutler TL, Swann DE (1999) Using remote photog- Pp. 9–26. (Springer Publishing, Tokyo). raphy in wildlife ecology: a review. Wildlife Soci- Larrucea ES, Brussard PF, Jaeger MM, Barrett RG ety Bulletin 27, 571–581. (2007) Cameras, coyotes, and the assumption of

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