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Received: 2 May 2018    Revised: 18 September 2018    Accepted: 9 October 2018

DOI: 10.1111/aje.12573

SPECIAL SECTION : CAMERA TRAPPING IN AFRICA

Are camera traps fit for purpose? A rigorous, reproducible and

realistic test of camera trap performance

Peter Apps  | John Weldon McNutt

Botswana Predator Conservation Trust,

Maun, Botswana Abstract
Camera traps are now widely used to produce the data that inform critical, large‐
Correspondence
Peter Apps, Botswana Predator scale conservation strategies, but insufficient attention has been paid to evaluating
Conservation Trust, Private Bag X13, Maun, or validating their performance. A realistic, reproducible and rigorous test of camera
Botswana.
peterjapps@gmail.com trap performance has been developed, using a real animal target moving in different
directions and at different distances from sets of up to six cameras. The test clearly
Funding information
Wild Entrust International; Tusk Trust; shows differences in performance between camera trap brands and models that af‐
Private donors; Self funded fect whether they are fit for research purposes, and the effects on camera perfor‐
mance of the small differences in mounting height and angle that are likely to arise
when camera traps are used in the field.
Résumé
Les pièges photographiques sont aujourd’hui largement utilisés pour produire des
données qui informent des stratégies de conservation critiques et à grande échelle,
mais l’on accorde trop peu d’attention à l’évaluation ou à la validation de leurs perfor‐
mances. Un examen réaliste, répétable et rigoureux des performances des pièges
photographiques a été mis au point, en utilisant une véritable cible animale qui se
déplace dans différentes directions et à différentes distances d’ensembles qui
comptent jusqu’à six appareils photo. Le test montre clairement des différences de
performances selon la marque et le modèle d’appareil, qui influencent le fait qu’ils
soient, ou non, adaptés aux besoins d’une recherche, et les effets sur les perfor‐
mances d’un appareil de petites différences dans sa hauteur et de l’angle de vue, qui
sont susceptibles d’apparaître lorsque l’on utilise des pièges photographiques sur le
terrain.

KEYWORDS
camera trap, capture‐recapture, fit for purpose, occupancy, trail camera, validation

1 | I NTRO D U C TI O N
The use of camera traps in field biology has exploded since about 2002
(Meek & Pittet, 2012; Wearn & Glover‐Kapfer, 2017), and they are now
widely used to generate data that inform critical conservation man‐
agement decisions at ecosystem and landscape scale (Swanson et al.,
2015; TEAM, 2011). Since camera trapping results inform such import‐
ant decisions, there needs to be a rigorous way of evaluating the fitness
for purpose of different camera trap brands and models, verifying the
performance of individual cameras, measuring the effect on perfor‐
mance of how camera traps are set up, and validating camera trap‐
ping methods, so that results and conclusions are robust and reliable.
Testing cameras before they are deployed to the field is recommended
by Wellington, Bottom, Merrill, and Litvaitis (2014), and reviews of
camera trapping (e.g. Burton et al., 2015) have stressed the importance
of camera performance to the collection of quality data that generate

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© 2018 John Wiley & Sons Ltd Afr J Ecol. 2018;56:710–720.
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results that are fit for purpose. Nonetheless, although there has been
some consideration of camera traps’ limitations (Burton et al., 2015;
Meek, Ballard, & Fleming, 2015), comparison between camera models
has been “rudimentary” (Meek et al., 2015), and the performance of
some cameras is disturbingly poor and disturbingly variable between
units (Ballard, Meek, Doak, Fleming, & Sparkes, 2014; Hamel et al.,
2013; Hughson, Darby, & Dungan, 2010; Lyra‐Jorge, Ciocheti, Pivello,
& Meirelles, 2008; Meek et al., 2015; Newey et al., 2015; Pease,
Nielsen, & Holzmueller, 2016; Swann, Hass, Dalton, & Wolf, 2004).
There is no accepted protocol for testing camera traps that is realistic
enough to predict their performance in the field, and most investiga‐
tors simply assume that camera traps are fit for purpose, and then use a
proliferation of increasingly elaborate models to process their data (e.g.
review O’Brien, 2011), which may not improve the quality of results if
the model’s assumptions are violated (Burton et al., 2015).
Camera traps cannot always be set up optimally—they may need
to be up trees to avoid being stolen by humans (Meek, Ballard, &
Falzon, 2016) or high on a pole to avoid being chewed by hyaenas
(Apps, Claase, Yexley, & McNutt, 2017), and if they are fixed to trees
or other natural supports, their heights and angles relative to local
topography will always vary. The effect of large differences in height
and angle has been investigated (Meek et al., 2016; Nichols, Glen,
Garvey, & Ross, 2017; Smith & Coulson, 2012; Swann et al., 2004),
but not the effects of the smaller differences imposed by the prac‐
ticalities of fieldwork. There are calls for the fine details of camera
placement (height, distance to target, angle, etc.) to be included
in published accounts of camera trap studies (Burton et al., 2015;
Meek et al., 2014). While this allows the set up to be repeated with
the same make and model of camera trap, it has limited practical
value in reproducing the camera traps’ detection performance with
other makes and models because the impact of mounting on camera
performance is known only in very broad terms (Meek et al., 2016;
Nichols et al., 2017; Smith & Coulson, 2012; Swann et al., 2004). For
data to be comparable between studies, we need a way of reproduc‐
ing camera performance rather than repeating camera set‐ups, and
that calls for data that directly compare the performance of different
brands and models, and the impact on their performance of different
mounting heights and angles and operating settings. In addition to
the large differences in height and settings that have been investi‐
gated so far, we need data on the effect of the small differences that
arise in practice when camera traps are set up in the field.
In addition to a test to validate the performance of camera
traps and the results they generate, there is a need for a stan‐
dardised, realistic, quantitative performance test that directly
compares camera trap brands and models and detects their
strengths and weaknesses. With camera trap prices differing by a
factor of ten, such a test will save researchers from wasting their
money on cameras that are not fit for purpose, or spending more
than they need to get the job done. While it is straightforward to
test camera traps by walking in front of them (Seccombe, 2017;
see also TrailcamPro https://www.trailcampro.com/pages/trail-
camera-shootout), a walking human is a lot taller than nearly all
mammals and has easily detectable areas of warm, bare skin. Some
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cameras that do well with humans as targets are a bitter disap‐
pointment to field biologists who are trying to detect and record
the presence of animals and their behaviour in the wild. Also, un‐
like the humans in Internet review walk tests, wild animals do not
always walk neatly across the field of view at right angles to the
camera’s direction of aim, but Glen, Cockburn, Nichols, Ekanayake,
and Warburton (2013) had screens in front of the cameras that
constrained their test targets to walk across the fields of view, and
Rowcliffe, Carbone, Jansen, Kays, and Kranstauber (2011) consid‐
ered only the distance from the camera and angular displacement
from the direction of aim, not the direction of the animals’ move‐
ments. A test target that more closely resembles the animals that
biologists are interested in, and that moves around like they do,
would improve how well the tests predict performance in the field.
Field tests with wild animals as targets (e.g. Hughson et al., 2010)
avoid the problem of the target and test conditions being unrealistic,
but suffer from the equally serious problem that the only record
of the cameras’ performance is the images that they capture—the
true positives. The false negatives of animals that were not captured
are unknown unknowns that cannot be counted, and without the
denominator of the number of potential image captures, we can‐
not tell whether a camera captured all, most, or hardly any of the
animals that passed in front of it. Where camera traps have been
tested against independent detections such as spoor, continuous
video or time‐lapse images, the cameras have been shown to be
remarkably poor at capturing medium‐sized carnivores and smaller
animals (Damm, Grand, & Barnett, 2010; Glen et al., 2013; Hamel et
al., 2013; Lyra‐Jorge et al., 2008; Pirie, Thomas, & Fellowes, 2016),
and if camera traps are catching only 30%–50% of potential targets,
then correspondingly more resources are required than with cam‐
eras that catch everything. If cameras catch only half of the pass‐
ing animals, they have to be deployed for twice as long to capture
sufficient images for analysis, and the longer the trapping goes on,
the more the assumptions of closure that underpin many models
are likely to be violated. Validating camera performance requires a
known number of opportunities for the cameras to capture images.
Tests in the field suffer from the further disadvantage that they
cannot be rigorously repeated or reproduced; which animals happen
to pass through the camera’s detection zone varies with time and
from site to site, even within a given habitat with a given suite of
target species. This limits comparisons between cameras to side‐by‐
side shootouts. In the field, data yield rate in terms of images per
unit time is low; depending on species, it may take days or weeks
to capture a single image, and even when cameras are left out for
weeks to generate useful sample sizes (Hamel et al., 2013; Hughson
et al., 2010; Nichols et al., 2017; O’Connor et al., 2017; Rowcliffe
et al., 2011), there is no systematic coverage of their fields of view
or the directions in which animals move. Cameras in the field also
capture a large number of images without the targets of interest in
them, which wastes time during data processing. A practical test of
camera performance needs to generate useful quantities of data in
minutes rather than weeks, with systematic, equal coverage of op‐
erating variables, and without large numbers of extraneous images.
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Rovero, Zimmermann, Berzi, and Meek (2013) called for stan‐
dardised camera trap testing, and there is a need for a test of camera
performance that uses a realistic target, can be reproduced at will, and
quickly generates rigorously comparable data with a known expecta‐
tion of the number of images that will be captured. To realistically sim‐
ulate the movement of wild animals in the field, the target needs to be
at different distances from the camera, moving in different directions
with respect to the camera’s direction of aim, and in different places
within its field of view. Using a trained domestic dog as a real animal
target provides a test that meets these criteria, and that is straightfor‐
ward, cheap and quick to carry out. Here, how the test is conducted is
described and examples of the performance comparisons and valida‐
tion data that it generates are given. Full reports on the performance of
each model of camera trap will be published elsewhere.
2 |  M E TH O DS
A test arena was marked out on an area of short grass, partly shaded
by large trees (Figure 1a). Distances were measured by surveyor’s tape.
At the centre of one end of the arena, a 50 × 50 mm steel square tube
was set vertically into rammed gravel, and sets of up to six test cameras
were mounted using their tripod screw sockets to a 40 × 40 mm angle
iron bar bolted to the pole either vertically or horizontally. Camera angle
from the horizontal (angle of tilt and dip) could be varied continuously
by tilting the cameras relative to the horizontal mounting bar (Figure 1b)
or changing the angle between the vertical mounting bar and the pole
(Figure 1c). Mounting points were at 20 cm vertical increments, with the
exact height of the camera sensors determined by the sensors’ heights
relative to the tripod sockets. Heights were measured to 0.5 cm with a
steel tape, vertical and horizontal were determined with spirit levels, and
angles were measured by a plumb line against a protractor (Figure 1c).
Cameras faced approximately south, and all tests were carried out in the
early morning or evening when shade temperatures were below 26°C.
At the edges of the arena, pairs of stakes 50 cm apart and 2.5, 5,
7.5, 10, 12.5 and 15 m from the cameras had thin cotton rope strung
between them just off the ground to mark pathways for the target
to walk along. One set of paths ran at right angles to the cameras’
direction of aim (Figure 1d) and the other set diagonally across the
field of view and down its centre line (Figure 1e).
F I G U R E 1   (a) The camera trap test
arena; image from a Cuddeback C camera
trap mounted 52 cm off the ground,
upright and level (dip and tilt both 0°)
with Toffee the target dog walking left
to right 7.5 m from the camera. (b) Six
Reconyx Hyperfire camera traps tilted at
2° increments on the horizontal mounting
bar. (c) Six Reconyx Hyperfire camera
traps mounted at 20 cm height increments
and dipped 6°. (d) Layout of the tangential
tracks. (e) Layout of the radial tracks
APPS and McNUTT
A large, mixed breed dog, 68 cm tall at the shoulder (about the
same as a leopard, Panthera pardus, or African wild dog, Lycaon pic‐
tus, two species that we use camera traps to investigate; Figure 1)
was used for all the tests considered here. He wore a light fabric
vest with a matt black spot about 16 cm across on it so that image
processing software could be used to measure his position in the
images (see below). For each test sequence, his handler (PA) left
him standing or sitting at the beginning of a track, walked along the
track between the ropes, and turned and called him, he walked at a
uniform speed along the track and was rewarded with petting and
food treats. The handler then walked back along the same track
and repeated the recall. Tangential tracks were walked in order
from closest to farthest from the camera, and radial tracks in a
clockwise pattern starting from the track along the centre, except
when Cuddebacks were tested when the radial tracks were walked
counterclockwise. Each series was walked once (Cuddeback C
range), twice (Bushnell Aggressor, Reconyx Hyperfire dip and tilt
tangential) or three times (Reconyx Hyperfire radial, Cuddeback
lure simulation and tilt) for each test. The dog is PA’s household
pet who has been trained using rewards, and ethics were compli‐
ant with research permit 8/36/4 XXXVIII (14) from the Botswana
Department of Wildlife and National Parks.
The results from three camera trap models are presented
here: Cuddeback C, Reconyx Hyperfire PC800 and Bushnell
Aggressor (Serial numbers B170511159, B170511162, B170511190,
B170515066, B170515120, B170515137). All the cameras were set
to their highest sensitivity and shortest time between triggers and
recorded one still image per trigger. They all had fresh batteries that
showed at least half charge at the beginning and end of each test.
2.1 | Data analysis
The target dog’s position on radial walks was fixed relative to dis‐
tance markers, the edge of the field of view and the known positions
of the tracks. His position on tangential walks was measured from
the X co‐ordinate of his position in each image and the known dis‐
tance of the tracks in front of the camera. X co‐ordinates were meas‐
ured by locating the spot on his jacket with the open‐source free
software ImageJ (https://imagej.nih.gov/ij/). The colour images from
the cameras were converted to 8‐bit monochrome and then to black
and white using the threshold function, and the spot was located
using the roundness and area criteria of the Analyze Particles com‐
mand. Since each camera model’s images have their own idiosyncra‐
sies, the threshold and particle analysis settings were adjusted to
suit in each case. X‐coordinates in pixels were calibrated to distances
on the ground by measuring the distance between the edges of the
field of view at the 15‐m track.
3 |  R E S U LT S
Each set of tests generates a large body of detailed, precise data that
capture the performance parameters, deficiencies and idiosyncrasies
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of the camera traps. Much of the data are best presented graphically.
Because there is insufficient space here for a full set of test results,
I will illustrate the utility of the tests using selected examples from
three brands of camera. Full reports on the performance of each
model of camera trap will be published elsewhere.
3.1 | Directional bias
Controlling the direction in which the target moves generates tests
of whether cameras preferentially detect targets moving in certain
directions. When the target dog walked tangentially across the field
of view at right angles to the direction of aim (Figure 1d) of cameras
mounted at optimum height, all the camera traps considered here
detected him out to at least 15 m, but the maximum range was re‐
duced along the radial tracks, and when he walked towards the cam‐
eras along the centre line, he was not detected until he was 7.5 m
from the Bushnells, 5 m from the Cuddebacks and 2.5 m from the
Hyperfires. This caused a marked reduction in the area of the detec‐
tion zone compared to when animals are walking across the field of
view (Figure 2, Table 1).
As an example of the results from standard tests being used to
predict the performance of different brands and models for spe‐
cific applications, the radial track layout (Figure 1e) simulates ani‐
mals approaching a scent lure placed 5 m in front of a camera, and
a camera that is suitable for monitoring reactions to the lure will
capture an image, or start a video while the animal is approaching
from any direction. Although every unit of each of the three camera
models considered here captured an image for every replicate on
every radial track, only one model, the Bushnell Aggressor, is suit‐
able for monitoring a lure. The other two would not detect animals
approaching the lure in a sector about 45° wide on the opposite side
to the camera (Figure 2).
3.2 | Effect of camera settings
As an example of a test of the effects of changing camera settings,
Cuddeback Cs have a sliding shutter that blanks off the edges of the
movement detector with the aim of having the target centred on the
images. Tests were run in triplicate with the slider up or down, and
an image was captured for every replicate on every track in both
directions. Due to the short trigger delay, the first image was usually
of the dog’s head as he entered the field of view, independent of the
position of the mask, except when he was moving from right to left
at 10 or 12.5 m from the camera (Figure 3), when raising the mask
captured his whole body on the first images. Overall, masking the
edges of the detector increased the number of full‐body images of
the target at 5 and 7.5 m from the camera, but not at other distances
(Figure 3).
3.3 | Mounting height and angle
The effects on image capture of mounting height and the angles at
which cameras were leaned forward or to the side were tested. Dip
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714       APPS and McNUTT

–6 m at which an animal (but not a human) target was detected, and in‐
creased dip decreased both minimum and maximum range for an
–4 m animal and maximum range for a human. The range interval over
which images could be captured got narrower, and the detection
–2 m area shrank proportionally, as cameras were mounted higher and
cuddeback dipped more, and there was a restricted subset of heights and an‐
gles that allowed detections over the full range of distances (Table 2;
Supporting Information Tables S1–S3).
2m Tilt is the angle at which a camera leans to the side (Figure 1b).
It did not affect range, but with different cameras, it has markedly
4m different effects on where the target is when images are captured.
Cuddeback Cs are robust to tilt up to at least 10°. Tilted Bushnell
6m
Aggressors capture images only after the target has penetrated
–6 m deeper into the field of view on the side away from which the camera
was tilted (corresponding to an animal walking uphill past an upright
–4 m camera). Reconyx Hyperfires are particularly sensitive; when they
are tilted, 10° images are captured only after the target has crossed
–2 m the centre line (Figure 4).
hyperfire

3.4 | The importance of realistic targets

2m
A human is bigger, taller and narrower than a leopard or its sur‐
rogate, and much easier for a camera trap to detect; and a human
4m
target generated misleadingly optimistic results for camera per‐
formance. For example, a Reconyx Hyperfire mounted straight and
6m
level with its sensor 1.2 m above the ground detected a human out
–6 m to 15 m, but the dog only at 5 m (Supporting Information Table
S2) even though the whole arena from 5 to 15 m and beyond was
–4 m in the field of view (Apps, 2018). At short range, cameras that are
mounted higher missed the animal target but detected the human.
–2 m Only a real animal target revealed dead zones where a leopard‐
bushnell sized target could walk across the field of view in both directions
without being detected, even though he was within the maximum
detection range and a human was detected every time. For exam‐
2m ple, Reconyx Hyperfires mounted upright at 80 cm had a dead zone
with no dog detections 7.5–10 m in front of them, even though the
4m
dog was detected moving in both directions at both 5 and 12.5 m
(Table 3), and at 100 cm mounted upright, and 120 cm dipped 2°,
6m
the dead zone was at 7.5 m (Supporting Information Table S2).
2.5 m 5m 7.5 m 10 m 12.5 m 15 m

F I G U R E 2   The locations at which images are captured depend

on the direction that an animal is moving relative to the camera, and
this affects the size of the area in which targets are recorded. The
pale grey triangle is the field of view, the area between the heavy
dotted lines is where a target moving tangentially was captured, the
area inside the heavy solid line is where the target moving radially
was captured (see also Table 1), and the darker grey triangle is the
sector where a target approaching a lure 5 m in front of the camera
would not be detected
is the angle at which cameras are tipped forward to bring the fields of
view and detection lower and closer to the camera (Figure 1c, Apps
& McNutt, 2018). Increased height increased the closest distance
4 | D I S CU S S I O N
The camera trap test proposed here meets the need for a realistic
simulation of field conditions by having a real animal as a target
moving in different directions at different distances from cameras
mounted at increments of height and angle that might be imposed
by field conditions. The track layout and camera mounting can
be reproduced at will and quickly generate rigorously compara‐
ble data with a known expectation of the number of images that
should be captured. In contrast with Meek et al.’s (2015) statement
that “Studies to generate (camera trap) calibration data are time
consuming, expensive and difficult,” this test takes about 15 min
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TA B L E 1   The areas covered by each
camera trap's field of view out to 15 m,
and the areas of the part of the field of
view in which targets moving tangentially
Cuddeback
and radially relative to the camera will be
captured as images Hyperfire
Bushnell
to run a sequence, and 30 min for a set of duplicates. During each
15‐min test replicate, the target dog walks across the field of view
twelve times—equivalent to weeks or months of data from the
wild (e.g. Di Bitetti Paviolo, & Angelo, 2014; O’Connor et al., 2017)
and data are collected equally for different distances, directions,
heights and angles. Having controlled, known target movements
is critical to finding detection dead zones; there is no way of find‐
ing them under field conditions because the most parsimonious
explanation for a lack of images in a particular part of the field of
view is that no animals moved through it. Precise and repeatable
aiming of the cameras is assured by mounting them all on the same
bars, which are bolted together; once the main pole has been ac‐
curately positioned, there is no need for time‐consuming aiming of
individual cameras.
Obviously, the camera traps’ performance with wild animals will
not be exactly the same as with dogs, but they will be a lot more
similar than with a human target. In particular, detection dead zones
for animals are not likely to be detected with a human as a target
(Table 3). To participate, a dog needs only the most basic of pet obe‐
dience in the form of a recall from a wait. Dogs of different sizes can
stand in for different wild species, possibly with the tracks closer
to the cameras for small breeds and species. A variety of camera
trapping scenarios, such as monitoring responses to roadside re‐
pellents, can be simulated by laying out the tracks appropriately
F I G U R E 3   Locations of the target
when the first image on each traverse
was captured by a Cuddeback C camera
trap with the detector mask raised
(circles) or lowered (triangles), and the
number of images showing the whole
target with the detector window slider
raised or lowered at each distance
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Areas (m2)
Field of view Moving tangentially Moving radially
90 59 15
90 55 6
87 65 22
(P. J. Apps, unpublished data), and there is no particular reason to
limit the targets to dogs; opportunistic trials during the dog tests
showed that food treats will induce domestic cats to walk across
the field of view.
Camera traps that are used to record behaviour (e.g. Vogt,
Zimmermann, Kölliker, & Breitenmoser, 2014; Allen, Gunther, &
Wilmers, 2017; Apps et al., 2017) must arguably work better, and
be set up more carefully, than those for population estimates,
because one missed record of, for example, an animal scent mark‐
ing can confound the interpretation of later records of behaviour
at the site, and there is no prospect of using modelling on large
data sets to fill in the gaps. Behaviour will nearly always be re‐
corded on video rather than by still images, and although using
still image trigger delay as a surrogate for video start time is con‐
venient for screening tests, video trigger delays are always longer
than for stills and rigorous validation of video performance must
obviously be run with videos. For example, the Cuddeback Cs
have very fast still image triggers but their video trigger delays
of 3–4 s are much too slow to be useful. To maximise the length
of time that the target is in shot, it needs to be detected as soon
as possible after it enters the field of view, and the effects of tilt
on penetration into the field of view before the camera triggers
(Figure 4) show that this needs cameras to be set up perpendicu‐
lar to the ground surface. If the aim is to video animals reacting to
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TA B L E 2   The ranges of distances (m)

Dip
over which camera traps mounted at
Height (cm) 0° 2° 4° 6° 8° 10° different heights and angles captured
images of a leopard‐sized dog
Cuddeback C
36 2.5–15 2.5–15 2.5–12.5 2.5–2.5 No images No images
56 2.5–15 2.5–15 2.5–15 2.5–10 2.5–2.5 2.5–2.5
76 5–15 5–15 2.5–15 2.5–10 2.5–5 2.5–2.5
96 7.5–15 5–15 5–15 2.5–12.5 2.5–7.5 2.5–5
116 10–15 7.5–15 5–15 5–12.5 5–10 5–7.5
Hyperfire
20 2.5–15 2.5 2.5 No images No images No images
40 2.5–15 2.5–15 2.5 2.5 2.5 No images
60 2.5–15 2.5–15 2.5–15 2.5–5 2.5 2.5
80 2.5–15 2.5–15 2.5–15 2.5–10 2.5–5 2.5–5
100 5–15 5–15 5–12.5 2.5–5 2.5–5 5
120 5–15 5–15 5–10 5–10 2.5–5 2.5–5
Bushnell aggressor
22 2.5–15 2.5– 2.5–5 2.5 2.5 No images
12.5
42 2.5–15 2.5–15 2.5–10 2.5–5 2.5 2.5
62 2.5–15 2.5–15 2.5–12.5 2.5–12.5 2.5–10 2.5–5
82 2.5–15 2.5–15 2.5–15 2.5–10 2.5–7.5 2.5–5
102 5–15 2.5–15 2.5–12.5 2.5–12.5 2.5–10 2.5–7.5
122 5–15 5–15 5–15 5–12.5 2.5–10 2.5–7.5

Note. Cells with bold type have detections over the full range.

lure or repellent placed close to the camera, a camera that does
not detect animals moving towards or away from it (Figure 2) is
unfit for purpose, and the tests with radial tracks can quickly
eliminate certain camera trap models from consideration.
Comparisons between results are valid only if the test is re‐
producible and repeatable. Comparing single units of different
brands and models in a side‐by‐side shootout is straightforward
(Apps, 2018) but needs the cameras to be available at the same
time. Comparisons on the basis of results generated at different
times depend on the test being reproducible, and the three mod‐
els considered here were tested at different times over a period of
9 months. The Cuddeback and Bushnell cameras are what Newey
et al. (2015) refer to as “recreational” in terms of technical specifi‐
cations (and price), and the Hyperfire PC800 is designated as a re‐
search camera trap. Clearly, from the results presented here, these
categories do not correspond to performance.
Testing cameras in sets of six provide strict repeatability in
that each of them has exactly the same opportunities to capture
images; the only difference between them is the parameter under
test: height, dip or tilt. The most likely cause of differences in re‐
sults between replicates would have been the target dog walking at
different speeds, but in practice, his speed was so consistent that
detections on tangential replicates agreed in 95.5% of walks for
the Ultrafires (Supporting Information Table S2), and 98.9% for the
Bushnells (Supporting Information Table S3). With the Cuddebacks
(Supporting Information Table S1), the points from replicates over‐
lapped on plots of target position (Figures 3 and 4), allowing the
number of replicates to be reduced from three to two or one. To re‐
produce the arena layout and camera mounting is a straightforward
application of measurements of lengths and angles, but the repro‐
ducibility between different dogs needs to be investigated if the test
is to be transferable.
Because missed images were recognised as such, and target dis‐
tances and camera mounting were varied systematically, the opti‐
mum height and angle for maximum detections were very quickly
and clearly recognised. The results do not fully support the stan‐
dard advice to mount camera traps at the target’s shoulder height.
The Cuddeback achieved its full range of detection distances only
when it was 20 cm below shoulder height, and although the Reconyx
and the Bushnell achieved detections over the full range at shoul‐
der height, mounting them 20 cm lower provided the same range
of detection, and with a 20 cm buffer above and below. A Reconyx
Hyperfire mounted only 12 cm above shoulder height had a de‐
tection dead zone 7.5–10 m in front of the camera. Differences of
20 cm or less in height, and 2° in dip, which are well within the range
of differences that might be imposed by uneven terrain or when
APPS and McNUTT |
      717

–6
cuddeback
–4

distance from centre (m)

–2

6
0 2.5 5 7.5 10 12.5 15
distance from camera (m)

–6
hyperfire
–4
distance from centre (m)

–2

6
0 2.5 5 7.5 10 12.5 15
distance in front of camera (m)

–6
bushnell
–4
distance from cetre (m)

F I G U R E 4   Shifts in target position –2

when the first image on each traverse
was captured, due to tilting camera traps 0
to the side. Circles are for the target
moving left to right (top to bottom in this 2

orientation) and squares for the target

4
moving right to left. Filled symbols are
captures with the camera upright, empty
6
symbols with the camera tilted 10° to the
0 2.5 5 7.5 10 12.5 15
side. Cameras are mounted at the centre
of the left‐hand edge distance in front of camera (m)

mounting cameras on awkwardly shaped or angled trees, for exam‐
ple, have measurable effects on performance that may jeopardise
the comparability of data between cameras. Occupancy models can
accommodate camera failure, but they require the detection at one
site to be extrapolated to others (MacKenzie et al., 2002), and these
results show that this requirement may not be met unless camera
mounting is more repeatable than it is reported to be. In addition to
copying the variation in camera angles imposed by mounting them
on natural objects, dip and tilt also simulate cameras aimed up or
across a slope, and the results show that for maximum detections
and comparable data, cameras must be angled to match the slope
(see figure 5 in Apps & McNutt, 2018). The results show clearly that
mounting cameras high enough to be out of the reach of animals
will compromise detections for all except the tallest species, unless
the cameras are dipped, and then, maximum range is compromised.
The effects of height and dip are to be expected given the geometry
of the camera traps’ passive infrared detectors’ fields of view, but
the differences in results between models, especially the minimum
range for the Cuddeback Cs and the dead zones for Hyperfires, point
to differences in detector field of view that need to be taken into ac‐
count when setting up cameras; the bottom edge of the Cuddebacks’
detection zone is well above the bottom edge of the image, making
them more vulnerable to being mounted too high or dipped too little
and there is a gap between the two detection bands of a Hyperfire.
 |
718       APPS and McNUTT

TA B L E 3   Number of images of a
Replicate 1 Replicate 2
human and a dog captured on each track
Track (m in L‐R R‐L L‐R R‐L in each direction by a Reconyx Hyperfire
front of camera trap mounted upright 80 cm off
camera) Human Dog Human Dog Human Dog Human Dog the ground
2.5 1 2 1 1 1 1 1 1
5 2 2 1 2 1 1 1 2
7.5 2 0 2 0 2 0 2 0
10 2 1 2 0 2 0 2 0
12.5 3 2 2 2 3 2 3 1
15 3 2 3 2 3 2 3 1

For all three models, their sensitivity to height and dip could make
camera trap detections selective for target height, which might lead
to spurious negative correlations in species occurrence. Mismatches
between the detector and image fields of view, and detection dead
zones within the field of view (Tables 2 and 3), which can be de‐
tected only with known animal movements, mean that even though
the whole of a vehicle track, for example, may be in the field of view,
only one side of it will be in a detection zone. Although missed de‐
tections can be accommodated by occupancy models that are robust
to camera failure (MacKenzie et al., 2002), an improperly mounted
camera will create a gap in a capture‐recapture grid, violating the as‐
sumption that all individuals have a chance of being recorded (Royle,
Nichols, Karanth, & Gopalaswamy, 2009; Tobler & Powell, 2013). A
camera that misses its only chance to photograph a rare animal will
fail to record the presence of that species, which might influence the
conservation status of an area, or lead to a failure to recognise the
new or continued presence of an invasive species.
The controlled tests detected a bias against animals that are mov‐
ing radially with respect to the cameras that violates the requirement
of random encounter models that detections are random with re‐
spect to animal movement (Lucas, Moorcroft, Freeman, Rowcliffe, &
Jones, 2015; Rowcliffe, Field, Turvey, & Carbone, 2008). In addition,
these controlled tests have shown that the area over which camera
traps detect targets and capture their images, which is a parame‐
ter in random encounter estimates (Cusack, Swanson, et al., 2015),
varies by a factor of up to 15 depending on which direction the ani‐
mals are moving relative to the camera. Results from controlled tests
with targets moving in different directions will allow more accurate
estimates of detection area than assumptions based on the angle
of the field of view or detections with targets moving in only one
direction. The smaller detection area for animals that are not walking
tangentially across the field of view is very likely to contribute to the
lower image capture rates of randomly placed cameras compared
to those on game trails (e.g. Di Bitett et al., 2014; Cusack, Dickman,
et al., 2015), and when cameras are aimed across roads, they will
preferentially capture images of animals walking along them rather
than across them (Di Bitetti et al., 2014). If detections are biased by
direction of approach, then data models based on random camera
siting also require the cameras’ directions of aim to be randomly or
uniformly distributed.
The flat, level, grassed test arena and the cool temperatures
under which the tests were run provided optimum conditions for
camera trapping, but obviously do not cover all of the conditions
such as forest, bare sand, rock or snow where camera traps might be
deployed. The test is straightforward enough to be adapted easily
to different conditions, and different‐sized dogs could be used as
surrogates for different target species to validate the cameras and
set up for particular projects.
5 | CO N C LU S I O N S
Rigorously establishing the performance parameters of camera traps
by standard, controlled, validated tests like those described here will
yield several advantages. The suitability of particular makes and
models for particular tasks can be quickly established without using
up valuable field time or wasting funds on unsuitable cameras. If the
results and conclusions that are derived from camera trap data are
to be robust and reliable, data models must be robust to the vagar‐
ies of practical camera trapping, and these tests show aspects of
camera traps’ performance that models need to be robust to, and
how cameras need to be operated in order to minimise the extent
to which unavoidable assumptions are violated. The test results
provide empirical values for model parameters that are currently
estimated by further modelling, and inform best practice in cam‐
era choice and camera set‐up. Empirical test results can be added
to the study design and data modelling that are the current focus
of critiques of camera trapping methods (Tobler & Powell, 2013).
Implementing tests like those described here as a routine compo‐
nent of camera trapping programmes will enhance the quantity and
quality of data, the robustness and reliability of results, and the va‐
lidity of conclusions.
AC K N OW L E D G E M E N T S
I am very grateful for loans of cameras from Tau Consultants
(Christiaan Winterbach and Rob Thomson), WildCru (Dominik
Bauer) and Cheetah Conservation Botswana (Leanne Van
der Weyde). The research was carried out under Botswana’s
Department of Wildlife and National Parks, Ministry of
APPS and McNUTT
Environment, Natural Resources, Conservation and Tourism re‐
search permit number 8/36/4 XXXVIII (14) for which I thank the
government of Botswana. The Botswana Predator Conservation
Trust is funded through Wild Entrust International, Tusk Trust, and
private donors.
ORCID
Peter Apps  http://orcid.org/0000-0002-0129-9468
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S U P P O R T I N G I N FO R M AT I O N
Additional supporting information may be found online in the
Supporting Information section at the end of the article.   
How to cite this article: Apps P, McNutt JW. Are camera
traps fit for purpose? A rigorous, reproducible and realistic
test of camera trap performance. Afr J Ecol. 2018;56:710–
720. https://doi.org/10.1111/aje.12573