CSIRO PUBLISHING

Pacific Conservation Biology, 2016, 22, 48–60

http://dx.doi.org/10.1071/PC15030

Camera traps in the canopy: surveying wildlife

at tree hollow entrances

Nigel Cotsell A,B,C and Karl Vernes B

A
Coffs Harbour City Council, Biodiversity Unit, 2 Castle Street, Coffs Harbour,
NSW 2450, Australia.
B
University of New England, Ecosystem Management, Armidale, NSW 2351, Australia.
C
Corresponding author. Email: nigelcotsell@gmail.com

Abstract. This is the first comprehensive camera trap study to examine hollow usage by wildlife in the canopy of trees.
Eighty cameras directed at tree hollows were deployed across eight sites in nine species of eucalypt in north-east New
South Wales. In total, 38 species (including 21 birds, 9 mammals and 8 reptiles) were recorded at hollow entrances over a
three-month period. There was a significant difference between wildlife hollow usage associated with site disturbance and
tree growth stage (ANOSIM, P . 0.05); however, there was no significant difference associated with tree hollow diameter
(ANOSIM, P . 0.05). The level of anthropogenic disturbance at each site, including vegetation modification of the
understorey, was a significant predictor of species presence. Despite the limitations of using camera traps in the canopy of
trees this study demonstrates the potential to garner useful insights into the ecology and behaviour of arboreal wildlife.

Received 24 September 2015, accepted 9 February 2016, published online 15 March 2016

Introduction knowledge the most comprehensive of these is by Gregory et al.

Our knowledge of arboreal wildlife that use tree hollows is poor
and is generally confined to species that are generally easy to
detect using more traditional methods such as trapping and
spotlighting (Smith et al. 1989; Lindenmayer et al. 1999).
However, there are smaller more cryptic species, or others that
inhabit the high strata or closed forest, that are less likely to be
detected. While our knowledge of the effectiveness of camera
traps at detecting arboreal species is in its infancy, there are now a
range of studies that suggest this technique can provide
significant insights into this poorly explored ecological realm
(Gregory et al. 2014; Harley et al. 2014; Meek et al. 2014a;
Taylor and Goldingay 2014). Information can be collected for
long continuous periods, providing basic data on presence/
absence, behaviour, predation events, hollow use and breeding,
with the potential to collect information on arboreal species far
more efficiently than other survey methods (Gregory et al. 2014).
While there has been a proliferation of ground-based surveys
using camera traps to target medium to large terrestrial mam-
mals (Tobler et al. 2008; Claridge et al. 2010; De Bondi et al.
2010), only a small fraction of these studies have used camera
traps to detect arboreal species and still fewer have incorporated
hollows or hollow usage as part of their survey design. The
techniques employed by these studies are varied and unique but
ultimately rely on mounting camera traps at height in the
arboreal strata. The logistical and safety issue of working at
height obviously provides a unique set of challenges that has
hindered research in this area.
Despite these limitations there have been a range of studies
undertaken using camera traps in the canopy. To the authors’
(2014) on wildlife usage of tree-crossing points in Peruvian
primary rainforests over a six-month period. Other examples
include Cerbo and Biancardi (2012), who used cameras
mounted on trees and used food lures to record small and
arboreal mammals in Italy; Oliveira-Santos et al. (2008) under-
took a study of Atlantic forest arboreal mammals of Brazil using
raised wooden platforms; Otani (2001) examined fig consump-
tion by primates in tree canopies of Japan; and Hirakawa (2005)
studied the efficacy of artificial bat attractants by mounting
camera traps on tree branches. Cockle et al. (2011) monitored
cavity-nesting birds in the Atlantic forests of Argentina using
15-m telescoping poles mounted with video cameras directed at
hollow entrances.
There are even fewer studies in Australia that have adopted this
technique. Davis et al. (2013) examined the relationship between
wildlife and hollow-bearing trees in the urban landscapes of
Sydney; and Harley et al. (2014) developed camera-trapping
protocols to determine site occupancy of Leadbeater’s possum
(Gymnobelideus leadbeateri) by mounting cameras vertically on
trees and using baits as lures. However, cameras were mounted
only 4 m above the ground. Taylor and Goldingay (2012)
examined gliding mammals with camera traps but only within
the context of using wooden poles as ‘stepping stones’ across a
major road.
Camera traps may also provide other opportunities to collect
both qualitative and quantitative data on a range of arboreal
species. Although there are very few examples in the literature
of studies incorporating camera traps at tree hollow entrances,
this technique can answer a range of important ecological

Journal compilation Ó CSIRO 2016 www.publish.csiro.au/journals/pcb

Surveying wildlife at tree hollow entrances
questions. These include, but are not limited to, interspecific and
intraspecific competition, predation, resource selection, niche
partitioning, activity periods, population density estimates and
assemblages of species using hollows.
To better understand the behavioural responses of arboreal
vertebrates, further research is required regarding the interac-
tions between competitors and predators and the role this plays
in cavity choice. While the height of hollows above ground level
has been suggested as a mechanism to avoid predators (Cockle
et al. 2011; Goldingay 2011), no study has specifically exam-
ined this hypothesis. Ambrose (1982) observed that, despite an
abundance of hollows, regular disputes over certain hollows
were noted and that the preferences for hollows needs to be
better understood. Thomson et al. (2008) suggest that targeted
surveys of hollow occupancy by species dependent on hollows
will assist in an understanding of competition for hollows.
Understanding behavioural interactions of cavity-using species
will better inform management, particularly where there is a
scarcity of hollows and populations are threatened (Goldingay
2009).
Camera traps provide the opportunity to better understand
the usage of tree hollows by vertebrate fauna and their behaviour
at hollow entrances. Therefore, the aim of this study was to
compare faunal usage with site characteristics and tree attribute
information including hollows, DBH (diameter at breast height),
age and species. We approached this by deploying a large
number of camera traps in tree canopies at selected locations.
Methods
Study area
The study area encompasses the rural, semirural and forested
landscapes of the Coffs Harbour Local Government Area (LGA)
on the eastern seaboard of New South Wales, Australia (Fig. 1).
The LGA has an area of 117 300 ha, of which 88 964 ha is
vegetated (Office of Environment and Heritage 2012a).
The Coffs Harbour Meteorological office (30.318S,
153.128E) records a mean annual rainfall of 1707 mm (mm), a
mean maximum temperature of 238C and mean minimum
temperature of 148C (Bureau of Meteorology 2014). Overall,
the climate is considered subtropical.
The Coffs Harbour LGA is one of the few places along
eastern Australia where the Great Dividing Range and con-
nected Great Escarpment link to the coastal plain. The area’s
unique topography and geography provide for a diversity of
ecosystems that have representatives of tropical, subtropical and
temperate flora and fauna.
This study was undertaken on the coastal plains of the Coffs
Harbour LGA. The coastal plains are a low-relief landscape with
little variation in altitude. The plains gradually rise from sea
level to around 50 m elevation. Slopes are predominantly flat to
undulating (0–58), though there are some small hills with steeper
slopes. The coastal plain is characterised by alluvial sediments
derived from floodplain deposits and older weathered metasedi-
ments of the Coramba formation (Milford 1999). The Coffs
Harbour LGA coastline stretches 80 km and features a series of
prominent rocky headlands and beaches.
The Coffs Harbour LGA is characterised by a variety of soil
moisture gradients, geologies and topography that support 79
Pacific Conservation Biology 49
different vegetation communities (Office of Environment and
Heritage 2012a). These communities have been grouped into
broader vegetation classes and vegetation formations in accor-
dance with Keith (2006). Nearly 50% of the vegetation in the
LGA is wet sclerophyll forest, almost 20% is dry sclerophyll
forest, 12% is rainforest and nearly 6% is coastal vegetation
communities. Other communities include plantations, exotics
and regenerating pioneers that cover almost 10% of the vegetated
area in the LGA.
Site selection
Mapping of High Value Arboreal Habitats (hereafter HVAH)
(Fisher et al. 2014) on the coastal plain of the Coffs Harbour
LGA was used as a surrogate to identify broad locations of
habitat containing hollow-bearing eucalypt trees. ‘Eucalypts’
for the purpose of this study included the tree genera Eucalyptus,
Angophora, Corymbia, Lophostemon and Syncarpia. The
coastal plain was selected for this study because of the already
depleted areas of HVAH and the increasing development
pressures being placed on this resource.
Eight geographic sites were selected on the coastal flood-
plain on the basis of the number of camera traps available for the
study (n ¼ 80), which allowed for 10 cameras at each site. The
study included eight different vegetation types selected from a
possible 15 eucalypt-based coastal vegetation communities
known from the LGA (Office of Environment and Heritage
2012b). However, due to a paucity of HVAH on the coastal
floodplain, four sites had to be selected from Coast and Escarp-
ment Blackbutt Dry Forest (see Table 1). Blackbutt (Eucalyptus
pilularis) is recognised as the dominant tree species for this
vegetation type (Office of Environment and Heritage 2012b).
The eight sites were allocated a descriptive name based on the
closest road name or geographical feature (Fig. 1).
All selected sites with hollow-bearing trees were arbitrarily
allocated a disturbance category of 1 to 5. High-quality sites
(Category 1), were considered remote from human habitation
and had a complex midstorey and understorey of native plants.
Poor sites (Category 5) were regularly mown, had high human
visitation and little to no understorey (see Table 1).
Sites were also grouped to determine the number of each tree
species used in the study. Faunal species richness was compared
with the number of trees for each species used in the study.
Faunal species richness was based on one record of each species
per location per period.
Camera trap specifications and deployment
For ease of use and cost, camera trap selection included three
Scoutguard digital cameras (SG550, SG560, SG565F-8M).
Cameras were set to normal sensitivity and were programmed to
record two successive still images (1 s apart) upon each ‘trigger
event’. The delay period was set at an interval of 10 s following
each trigger event. Image quality was set to the highest resolu-
tion of 8 Mb per image. For the purposes of this study, an
‘independent picture’ was considered valid after 60 min.
Camera selection was based on a trade-off between unsettling
wildlife with an incandescent white flash but improving the
chance of species identification with the aid of sharp high-
resolution colour images (Glen et al. 2013; Meek et al. 2014b).
50 Pacific Conservation Biology N. Cotsell and K. Vernes

8 - Barcoongere way
COFFS HARBOUR
LOCAL GOVERNMENT AREA

NEW SOUTH WALES

7 - Corindi cemetery

6 - Orara East State Forest

5 - Old Bucca Road

4 - Moonee Caravan Park

3 - Swans road

N
2 - Lawson crescent
SCALE@A4 1 : 135000 GRID NTH
1 - Showground
0 0.5 1 2 3 4 5 6 7

Kilometres

Fig. 1. Camera trap locations within the Coffs Harbour LGA.

Surveying wildlife at tree hollow entrances Pacific Conservation Biology 51

Table 1. Vegetation type and disturbance category for eight selected sites in the Coffs Harbour LGA

Site no. Location Dominant species Vegetation type (OEH 2012b) Disturbance
category (1–5)

1. Show Ground Blackbutt (Eucalyptus pilularis) Coast and Escarpment Blackbutt Dry Forest 5
(CH_DOF01)
2. Lawson Crescent Bastard tallowwood (Eucalyptus Sandstone Bloodwood – Needlebark Stringybark Heathy 4
planchoniana) (CH_DOF02)
3. Swans Road Brushbox Escarpment and Lowland Bangalow – Carabeen – Black 1
(Lophostemon confertus) Booyong Palm Gully Rainforest (CH_RF11)
4. Moonee Reserve Forest red gum Lowlands Swamp Box – Paperbark – Red Gum Dry 4
(Eucalyptus tereticornis) Forest (CH_DOF06)
5. Old Bucca Road Pink bloodwood Foothills Turpentine – Grey Gum – Ironbark Moist 2
(Eucalyptus intermedia) Shrubby Forest (CH_WSF17)
6. Orara East State Forest Blackbutt (Eucalyptus pilularis) Coast and Escarpment Blackbutt Dry Forest 2
(CH_DOF01)
7. Corindi Cemetery Blackbutt (Eucalyptus pilularis) Coast and Escarpment Blackbutt Dry Forest 3
(CH_DOF01)
8. Barcoongere Way Red mahogany (Eucalyptus Coast and Escarpment Blackbutt Dry Forest 1
resinifera subsp. hemilampra) (CH_DOF01)

Camera deployment A one-way analysis of similarity (ANOSIM) (Clarke and

Camera traps were deployed across eight sites (10 cameras at
each site) between September and December 2013 (see Fig. 1).
No single tree had more than two cameras installed.
At each site, all eucalypt trees were considered within a 500-m
transect without bias towards a particular tree species. Tree
selection was based on five criteria:
(1) whether a tree had one or more visible hollows,
(2) whether tree hollows had suitable camera mounting
opportunities,
(3) whether there were opportunities to manoeuvre an elevated
work platform within 20 m of the base of the tree,
(4) whether hollows exceeded the maximum height of the
elevated work platform of 25 m, and
(5) whether the diameter of a hollow was greater than 10 cm (cm).
An elevated work platform with a maximum reach of 25 m
was used to erect the cameras. Depending on the location of the
hollow, cameras were attached to trees using either a ‘Gorillapod’
wrapped around a branch or mounted directly on to the trunk or
branch of a tree using a small plastic mono-pod and fastened
using Tek screws. No food lures or attractants were used in this
study.
Statistical analysis
Automatic data organisation, storage and analysis of camera trap
pictures were after Sanderson and Harris (2013). The program
DataAnalyze 3.2 was used to compare individual camera traps
and GroupAnalyze 2.0 was used to compare across all sites.
To better understand whether species composition varied
significantly across the trap sites and geographic sites, several
tests were run. By using sites as the sample and species as the
variable, data were analysed with PRIMER 6.0. From the Bray–
Curtis similarity measure a resemblance matrix was calculated
for each individual site. The outputs associated with the
similarity values were employed to generate non-metric multi-
dimensional scaling (MDS) plots, which were graphed.
Green 1988) Global R statistic (Clarke and Gorley 2006) was
used to test whether there was a significant difference between
sites. The factors tested included hollow diameter, growth form
and site disturbance. Hollow diameter was classified as follows:
small, #10 cm; medium, 10–40 cm; large, 40–100 cm; very
large, $100 cm. Growth form measures included early mature
(1), mature (2), over-mature (3), and late mature (4). A distur-
bance category was allocated according to the quality of a site’s
mid and lower vegetated understorey and ranged from little to no
observable disturbance (1), minimal disturbance (2), moderate
disturbance (3), high disturbance (4), highly modified including
mown grass (5) (see Table 1).
ANOSIM R is a process that uses the Bray–Curtis resem-
blance matrix to statistically test whether there is a significant
difference between two or more groups (Clarke and Gorley
2006). The ANOSIM R statistic calculates the difference of
mean ranks between groups and within groups. R-values close to
zero indicate no separation between levels of the factor being
tested. Values closer to 1.0 signify an increased amount of
separation between sites; however, if sample sizes are large, low
R-values can still be significant.
Ethics and licence requirements
All field work undertaken as part of this study was in accordance
with relevant national guidelines and was approved by the
University of New England Animal Ethics Committee under
approval AEC13-098. The research component was also
undertaken under a New South Wales National Parks and
Wildlife Scientific Licence SL101168 and Forests NSW Forest
Permit for Research SPPR0027.
Results
Camera trap effort
Cameras were deployed for an 80-day period with a total camera
trap effort of 1158 days. Of the 80 camera traps deployed,
wildlife data were retrieved from 63 cameras. Data were unable
52 Pacific Conservation Biology N. Cotsell and K. Vernes

40

30

Degrees (°C)
20

10

0
1 6 11 16 21 26 31 36 41 46 51 56 61 66 71 76
Days

Minimum temperature (°C)

Maximum temperature (°C)
Linear (Minimum temperature (°C))

100

80
Percent (%)

60

40

20

0
1 6 11 16 21 26 31 36 41 46 51 56 61 66 71 76
Days
9 a.m. relative humidity (%)
3 p.m. relative humidity (%)
Linear (3 p.m. relative humidity (%))

70

60
Rainfall (mm)

50

40

30

20

10

0
1 6 11 16 21 26 31 36 41 46 51 56 61 66 71 76 81
Days

Fig. 2. Meteorological data collected over the course of the camera trap study (September–December 2013). (a) Daily minimum and maximum temperatures,
(b) relative humidity, and (c) daily rainfall.

to be sourced from 12 cameras linked to either problems with the
camera or secure digital card (SD card). Data from five cameras
were also discarded because no wildlife images were recorded,
generally as a result of the SD cards lasting less than a day
following constant background stimuli of leaves and branches,
which resulted in false triggers. Given the exposed windy envi-
ronment into which the camera traps were deployed, it is not
surprising that several cameras and SD cards failed. Also notable
was the large range in the number of days that each of the 63 SD
cards lasted as a result of false trigger events. Limited access to
the elevated work platform only allowed for deployment and
retrieval of cameras, which meant that cameras could not be
checked at intervals to determine their functionality.
Daily weather records over the duration of the camera
trap study
Weather data were sourced for the months of September (part),
October, November and December (part) 2013. Daily minimum
and maximum temperatures remained relatively stable over the
course of the camera trap survey (Fig. 2a). Relative humidity
(Fig. 2b) showed a gradual increase over the course of the field
study; however, there was a significant increase in humidity
Surveying wildlife at tree hollow entrances Pacific Conservation Biology 53

30

Independent pictures
25

20

15

10

0
eet
possum
r
ed lorikeet
Dollarbird
Squirrel glider
htail possum
okuburra
ider
inus
ake
ng
thrush
Noisy minor
g parrot
Galah
gtail possum
ake
duck
er
nded snake
l
ed skink
isher
neyeater
phascogale
k
flying fox
let-nightjar
Grey faintail
rbird
ink
lizard
Little corella
ter dragon
oo
yeater
d glider
Rufous fantai
Lace monito

Grey goshaw
ed treecreep
Pied currawo

black cockat
Bar-sided sk
Rainbow lorik

Brown tree sn

Green tree sn
Feathertail gl
Brown antech

Regent bowe
Sacred kingf
Pacific black

Pink-tongued
Grey shrike-

Lewin’s hone
Yellow-bellie
Australian kin

Dark bar-sid
Brush-tailed

Blue-faced ho
Laughing ko

Grey-headed

Australian wa
Scaly-breast

Australian ow
Stephen’s ba

Brush-tailed
Mountain brus

Common rin

White-throat

Yellow-tailed
Species

Fig. 3. Fauna species ranked by number of independent pictures across all sites in Coffs Harbour LGA
(September–December 2013).

50 haematodus) (26.6%) and the lace monitor (Varanus varius)

R 2 ⫽ 0.9699
45 (8.4%).
40
35 Species accumulation
Species

30 By Day 6, 20 of the 38 recorded species (53%) had been captured

25 by motion-detection cameras. Fig. 4 shows a logarithmic trend
20 line showing the rate at which new species were added with
15 continued camera trapping effort (Thompson and Withers
10 2003). The two most abundant species across all sites were
5 recorded first, while two of the most threatened arboreal
0 mammals, the yellow-bellied glider (Petaurus australis) and the
0 20 40 60 80 100 120 140
brush-tailed phascogale (Phascogale tapoatafa), were recorded
Days
late in the survey (Table 2). These two species were recorded
No. Log. (No.) once each by different cameras at different sites.
Fig. 4. Species accumulation of fauna detected at tree hollow entrances in Species richness
Coffs Harbour LGA (September–December 2013).
There was disproportional usage of blackbutt (Eucalyptus
pilularis) (n ¼ 20 species) and pink bloodwood (Corymbia
intermedia) (n ¼ 19 species) at the 5% significance level. Both
over the last 25 days, probably associated with increased rain- turpentine (Syncarpia glomulifera) (n ¼ 2) and forest red gum
fall. There was a total rainfall of 374 mm, with 302 mm falling in (Eucalyptus tereticornis) (n ¼ 1) were fauna-poor in compa-
the last 22 days (Fig. 2c). rison, although the tree sample size was less than five for both
(Fig. 5).
Data analysis
The total number of pictures processed in the analysis was 2118. Activity patterns
The number of sequential pictures (of the same animal at the Data from each individual camera trap were grouped to determine
same location) was 1474. This was a result of the default camera the activity patterns of the most frequently encountered birds,
trap setting to record two pictures when activated. Of these reptiles and mammals. These groupings best discriminated
pictures, 644 were independent and so were used in the analysis. between diurnal and nocturnal species as only one nocturnal bird
was recorded as part of this study, the Australian owlet-nightjar
Species ranking and frequency (Aegotheles cristatus).
Thirty-eight species were recorded: 21 birds, 9 mammals and 8
reptiles (Fig. 3). Across all sites, the three most frequently Diurnal birds
encountered species included the brush-tailed possum (Tricho- Activity patterns for most diurnal birds peaked between
surus vulpecula) (27.8%), the rainbow lorikeet (Trichoglossus 07:00 and 10:00 hours and then slowly declined until there
54 Pacific Conservation Biology N. Cotsell and K. Vernes

Table 2. Vertebrate fauna species accumulation over 80 days during a camera trap study in the Coffs Harbour LGA
(September–December 2013)

Day Cumulative total Species Common name

1 2 Trichoglossus haematodus Rainbow lorikeet

Trichosurus vulpecula Common brush-tailed possum
2 4 Eurystomus orientalis Dollarbird
Trichoglossus chlorolepidotus Scaly-breasted lorikeet
3 9 Accipiter novaehollandiae Grey goshawk
Alisterus scapularis Australian king parrot
Calyptorhynchus funereus Yellow-tailed black cockatoo
Dendrelaphis punctulatus Green tree snake
Varanus varius Lace monitor
4 14 Strepera graculina Pied currawong
Colluricincla harmonica Grey shrike-thrush
Anas superciliosa Pacific black duck
Eolophus roseicapillus Galah
Aegotheles cristatus Australian owlet-nightjar
5 16 Entomyzon cyanotis Blue-faced honeyeater
Dacelo novaeguineae Laughing kookaburra
6 20 Antechinus stuartii Brown antechinus
Manorina melanocephala Noisy miner
Pseudocheirus peregrinus Common ringtail possum
Cormobates leucophaea White-throated treecreeper
7 21 Cacatua sanguinea Little corella
8 22 Trichosurus caninus Mountain brushtail possum
10 23 Todiramphus sanctus Sacred kingfisher
11 25 Acrobates pygmaeus Feathertail glider
Eulamprus martini Dark bar-sided skink
12 26 Petaurus norfolcensis Squirrel glider
15 27 Cyclodomorphus gerrardii Pink-tongued skink
16 29 Rhipidura fuliginosa Grey faintail
Hoplocephalus stephensii Stephen’s banded snake
17 30 Eulamprus tenuis Bar-sided skink
29 32 Rhipidura rufifrons Rufous fantail
Meliphaga lewinii Lewin’s honeyeater
31 33 Boiga irregularis Brown tree snake
41 34 Petaurus australis Yellow-bellied glider
49 36 Pteropus poliocephalus Grey-headed flying fox
Phascogale tapoatafa Brush-tailed phascogale
57 37 Intellagama lesueurii Australian water dragon
77 38 Sericulus chrysocephalus Regent bowerbird

was virtually no activity between 20:00 and 21:00 hours. The
most active bird was the grey shrike-thrush (Colluricincla
harmonica), which was active from 06:00–07:00 until 21:00–
22:00 hours (see Fig. 6a).
Reptiles
The three most common species of reptile had a core activity
peak between 08:00–09:00 and 14:00–15:00 hours, with a spike
at 10:00–11:00 hours for all three species. Activity periods for
reptiles were most likely correlated with the hottest parts of the
day. There were no records of reptiles before 07:00 hours;
however, both the lace monitor (Varanus varius) and Stephens’
banded snake (Hoplocephalus stephensii) were recorded as
active between 19:00 and 20:00 hours. Core activity periods
for V. varius was between 08:00–09:00 and 11:00–12:00 hours,
with very little activity outside these times (see Fig. 6b).
Arboreal mammals
Activity periods for arboreal mammals were more variable
between species across a 24-h period when compared with
both birds and reptiles (see Fig. 6c). There was very little
arboreal mammal activity until 21:00–22:00 hours, which
continued through until 06:00–0700 hours for most species.
Both the brown antechinus (Antechinus stuartii) and feather-
tail glider (Acrobatus pygmaeus) had minor periods of diurnal
activity but avoided the hottest part of the day between 12:00
and 14:00 hours. The squirrel glider (Petaurus norfolcensis)
did not commence any nocturnal activity until 20:00–21:00
hours, with maximum activity recorded between 02:00 and
03:00 hours. Common brush-tailed possums (Trichosurus
vulpecula) were active during the day and night with peak
periods between 05:00 and 07:00 and between 21:00 and 22:00
hours.
Surveying wildlife at tree hollow entrances
Eucalyptus tereticornis
Syncarpia glomulifera
Eucalyptus resinifera subsp. hemilampra
Eucalypt species
Angophora costata
Lophostemon confertus
Eucalyptus planchoniana
Eucalyptus microcorys
Corymbia intermedia
Eucalyptus pilularis
0 5
Total number of individuals
Fig. 5. Faunal species richness at tree hollow entrances compared with eucalypt species across 63
camera trap sites in the Coffs Harbour LGA.
Commonly detected species
An analysis of the percentage of each species at each site is
provided below for the following commonly recorded species.
Other species could have been included in the results but the
differences at each site were not significant.
Common brush-tailed possum (Trichosurus vulpecula)
This was the highest ranked species by number of independent
pictures (n ¼ 176) and the most abundantly recorded species at
Corindi Cemetary (69.3%), Orara East State Forest (50.9%) and
Old Bucca Road (44.3%). The species was also relatively
common at Lawson Crescent (34.7%), the Showground
(21.4%) and Moonee Reserve (14.1%). The brush-tailed possum
was entirely absent from Barcoongere Way and Swans Road,
which were the lowest-disturbance-category sites (see Table 1).
Rainbow lorikeet (Trichoglossus haematodus)
These large aggressive lorikeets dominated by number and
species percentage at the Showground (46.5% of total number),
Moonee Reserve (46.0%) and Lawson Crescent (42.9%) sites.
By contrast, these birds were totally absent from Barcoongere
Way, Corindi Cemetary, Orara East State Forest and Swans
Road and in very low numbers at Old Bucca Road (1.4%).
The abundance of rainbow lorikeets in areas of high habitat
modification in urban and periurban environments was highly
significant. Perhaps the most interesting result was the total
absence of the species from the least disturbed sites. Their absence
correlated strongly with the presence of lace monitors. Following
removal of camera traps it was noted that several lorikeets had
either eggs or chicks within the subject hollow and both the female
and male were being regularly recorded by the camera trap. It was
also observed that the predatory laughing kookaburra and pied
currawong and brush-tailed possum were occasional visitors to
the hollows that contained eggs and young of rainbow lorikeets.
Lace monitor (Varanus varius)
Lace monitors were common in all bushland areas and were
the third most frequently observed species across all sites
Pacific Conservation Biology 55
1
2
5
6
7
7
7
19
20
10 15 20 25
Frequency
Fauna species richness
overall. They were largely absent from sites that were highly
disturbed or contained high levels of human occupation, including
frequent domestic dog activity. As a total percentage of each
species, monitors were plentiful in bushland areas at Old Bucca
Road (38.57%) and Orara East State Forest (25.5%). They were
moderately common at Corindi Cemetery (8.1%), Swans Road
(8.0%) and Barcoongere Way (5.8%). However, monitors were
absent or recorded at very low frequencies at sites with distur-
bance categories of 4 or 5 (see Table 1), including Moonee
Reserve (0.6%), Showground (0%) and Lawson Crescent (0%).
Despite no feeding events being captured at hollow
entrances, the camera trap results show that lace monitors were
actively engaged in searching hollows. Sequences of camera
trap footage revealed monitors’ dexterity in being able to access
hollows on the end of branches, climb inside and then exit head
first. While no predation events were captured at the entrance of
hollows, it is likely that such events occurred within the hollow.
Possible targets are likely to have been bird eggs, chicks, or adult
birds or mammals of varying age classes.
Observations of the size and markings of monitors confirmed
that the same hollow was being visited by the same individual on
multiple occasions or was frequented by up to four separate
individuals over the duration of the field study. In some cases,
monitors were captured visiting the same hollow every day for
up to five consecutive days and also basked in the same tree at
the same location over extended periods. Two separate indivi-
duals were recorded residing in the same tree hollow over
several nights.
Lace monitor activity in trees was solely diurnal, reaching a
peak between 9:00 and 11:00 hours. This behaviour was often
repeated by the same individual over several days. Lace monitors
were captured at the highest-set cameras (25 m). Activities
beyond this height were not recorded.
Dollarbird (Eurystomus orientalis)
The migratory dollarbird was the fifth highest ranked species
by number of independent pictures (n ¼ 40). One camera
recorded a particularly long photographic sequence of a nesting
pair of birds, and two chicks were present at the time of camera
56 Pacific Conservation Biology N. Cotsell and K. Vernes

0.7 Resemblance: S17 Bray Curtis similarity

0.6
0.5 3D Stress: 0.01

0.4
0.3
0.2
0.1
0
00:00-01:00
01:00-02:00
02:00-03:00
03:00-04:00
04:00-05:00
05:00-06:00
06:00-07:00
07:00-08:00
08:00-09:00
09:00-10:00
10:00-11:00
11:00-12:00
12:00-13:00
13:00-14:00
14:00-15:00
15:00-16:00
16:00-17:00
17:00-18:00
18:00-19:00
19:00-20:00
20:00-21:00
21:00-22:00
22:00-23:00
23:00-24:00
CC5b
SG2b
SG6
M5a BW5b
OE4a
LC3b
M1a
M3
M5b
M6
OBR4b
OBR2a
OBR2b
OBR1b
BW4b
OBR6
SG1b
CC4a
LC3a
LC4b
SG4
M1b
LC6a
OBR1a
SG2a
SG5a
M7
OBR3b
LC1a
OE4b
LC5
OE5
SG5b
OBR4a
CC3a
SR2b
BW6
OE6
OE2b
CC2b
M4a
CC2a
M4b
OBR3a
OE1b
OE7a
OE1a
CC1a
BW4a
BW3a
OE2a
BW2a
SR1a
SR3
SG1a
LC2
BW2b
CC3b
BW5a
SR2a
SR1b

Hours

Rainbow lorikeet Scaly-breasted lorikeet

SR5a
Pied currawong Dollarbird
Laughing kookaburra Grey-shrike thrush

0.7
Fig. 7. Bray–Curtis similarity showing the clustering of sites (cameras)
0.6 and outliers.
0.5
Frequency

0.4
retrieval. As for the rainbow lorikeet, the hollow was visited by
0.3
laughing kookaburras, pied currawongs and common brush-
0.2
tailed possums at intervals sometimes weeks apart. These
0.1
predators were possibly attracted by the noise of the chicks or
0 the hollows were within their home range and part of a regular
00:00-01:00
01:00-02:00
02:00-03:00
03:00-04:00
04:00-05:00
05:00-06:00
06:00-07:00
07:00-08:00
08:00-09:00
09:00-10:00
10:00-11:00
11:00-12:00
12:00-13:00
13:00-14:00
14:00-15:00
15:00-16:00
16:00-17:00
17:00-18:00
18:00-19:00
19:00-20:00
20:00-21:00
21:00-22:00
22:00-23:00
23:00-24:00

inspection route. Other than their head, these predators were too
large to pass through the hollow entrance to reach the eggs or
chicks.
Hours
Squirrel glider (Petaurus norfolcensis)
Brown tree Lace monitor Stephen’s banded The squirrel glider was the sixth most common species
snake snake ranked by frequency of independent pictures (n ¼ 22) and was
recorded at four sites with a disturbance category of #3 (see
Table 1). The species was particularly abundant at Barcoongere
0.6 Way (30.8%) as a percentage of all species.
0.5
Patterns in usage of hollows by fauna
0.4
Results for the Bray–Curtis similarity resemblance in Fig. 7
0.3 show that species at individual hollows were similar except at
0.2
two outliers: BW5b (red mahogany, hollow diameter 16 cm, site
Barcoongere Way) and SR5a (brushbox, hollow diameter
0.1 12 cm, site Swans Road). These two hollows were classed as
0 outliers because they attracted a single species only, which was
00:00-01:00
01:00-02:00
02:00-03:00
03:00-04:00
04:00-05:00
05:00-06:00
06:00-07:00
07:00-08:00
08:00-09:00
09:00-10:00
10:00-11:00
11:00-12:00
12:00-13:00
13:00-14:00
14:00-15:00
15:00-16:00
16:00-17:00
17:00-18:00
18:00-19:00
19:00-20:00
20:00-21:00
21:00-22:00
22:00-23:00
23:00-24:00

not recorded by any other camera trap. They included the brown
tree snake (Boiga irregularis) at BW5b and a regent bowerbird
(Sericulus chrysocephalus) at SR5a.
A second MDS test generated a Bray–Curtis similarity resem-
Hours blance using disturbance, growth form and hollow diameter as
factors. There was little separation in the first two factors but there
Common brushtail possum Mountain brushtail possum was a relatively strong separation between species based on the
Squirrel glider Brown antechinus
Feathertail glider
level of disturbance.
The 2D stress test separated out Category 1 sites, based on
Fig. 6. (a) Activity patterns across all geographic areas for the six most
their site quality; that is, they were classified as having good
common diurnal birds over 80 days (b) Activity periods across all sites for ecological condition in that they showed low levels of overall
the three most common arboreal reptiles after 80 days (c) Activity periods disturbance and had an understorey and mid-stratum that was
across all sites for the five most common arboreal mammals after 80 days. intact and were represented by a diversity of plant species. Both
Surveying wildlife at tree hollow entrances Pacific Conservation Biology 57

Resemblance: S17 Bray Curtis similarity

2D Stress: 0.09 Disturbance
1
2
3
4 5 1 1
5 4 4
4 55 2
4 2
4 5423
4 1 1 5
5 4 2 2
5 444
2 22 2 1
3 54
2 3 2 2
2 1
3 1
4 1 1
3 2 1
4 33 2
2
1 1

Fig. 8. MDS plot showing ecological separation based on five levels of disturbance for 63 camera
traps located on the coastal floodplain of the Coffs Harbour LGA.

Barcoongere Way and Swans Road sites fell into this category
(see Fig. 8).
A one-way analysis of similarities (ANOSIM) was undertaken
on the same factors of disturbance, growth form and hollow
diameter to test whether there was a significant difference
between two or more sites. The results were statistically signifi-
cant for disturbance (ANOSIM Global R ¼ 0.27, P ¼ 0.001),
growth form (Global R ¼ 0.11, P ¼ 0.003), but not hollow dia-
meter (Global R ¼ 0.004, P ¼ 0.40). While the P-values suggest a
significant result for disturbance and growth form, the low R
values indicate that there was not a high level of separation
between the factors.
Incidental observations
Arboreal termite nest
A camera trap located at the Barcoongere Way site was
mounted 12 m above ground on a pink bloodwood (Corymbia
intermedia) directed at a ‘mud hollow’ instead of a natural tree
hollow. All other tree hollow options in the vicinity had been
exhausted. Despite the SD Card lasting only 34 days, the camera
recorded some of the most cryptic arboreal mammals including
the feathertail glider, brown antechinus, squirrel glider and
brush-tailed phascogale. The phascogale record was the only
one obtained over the duration of the study. For each of these
species, the sequence of photographs show each species taking
an active interest in the termite mound, often climbing over the
mound. The mud hollow was also visited by three individual
lace monitors, two of which entered the cavity and appeared to
actively excavate the mound to source termites.
European and native bees
Although bees were not the target of this study, up to 10% of
hollows inspected were found to contain the European honey
bee (Apis mellifera) – the presence of bees excluded these
hollows from the study. Bee hives were often near to small
native bees, with the introduced bees occupying large cavities,
generally .10 cm in diameter, while the smaller native bees
appeared to prefer small fissures or cracks in the branches of
eucalypts.
Discussion
The detection of wildlife at tree hollow entrances is a relatively
new technique. This study has demonstrated that, with the
appropriate camera trap accessories and some prior planning,
it is possible to survey tree hollow entrances for vertebrate
arboreal wildlife.
Some refinement of camera deployment methods may be
required to minimise the 15% failure rate of camera traps
experienced in this study. Working from an Elevated Work
Platform in an environment that was often windy and rainy
exacerbated by a confined work space and limited time created
less than optimal deployment conditions. Like Gregory et al.
(2014), we found that cameras were often triggered by non-
target motion (e.g. leaves and branches), which rapidly filled
memory cards well before camera retrieval after 80 days.
Mistakes made at set-up also had to be carried through until
retrieval as quality checks of the data were not possible due to
the costs associated with operating the Elevated Work Platform.
More expensive camera traps that use remote or continuous
download of data via Bluetooth or wireless would provide a
better alternative and allow checks to monitor camera functionality
and data quality.
Eucalypt species versus wildlife hollow preferences
The overall percentage of each eucalypt species used in the
camera trap study is broadly reflective of the overall composi-
tion of hollow-bearing trees remaining on the coastal floodplain
of the Coffs Harbour LGA. Both blackbutt and pink bloodwood
constitute a significant proportion of remaining hollow-bearing
trees on the coastal floodplain (authors’ obs.). The number of
fauna species active at hollow entrances of these two eucalypts
may simply be reflective of the frequency at which these two
58 Pacific Conservation Biology
tree species occur. For this study, hollow selection was inde-
pendent of tree species selection. It was not the aim of this study
to determine whether arboreal wildlife had a preference for
hollows of a specific eucalypt tree species although this would
make an informative future study.
Some of the individual camera trap set-ups recorded parti-
cularly long sequences of breeding events, multiple species
visits or unusual behaviours. Comparison across sites and
preferential use by wildlife of the nine eucalypt species is also
discussed, as are some of the interactions between individual
wildlife species and site quality.
Lace monitor (Varanus varius)
The frequency of visits to hollows and the abundance of lace
monitors suggest that large arboreal monitor lizards may have a
significant impact on hollow-dependent arboreal fauna, parti-
cularly those species that depend on hollows for at least a part of
their lifecycle. The absence of monitors in urban and periurban
environments is likely to account for the high abundance of
rainbow lorikeets at these sites. Whether this is a result of habitat
modification, interactions with humans, domestic dogs, or a
combination of all three, is a subject for further research.
There is growing evidence that monitor lizards may be
released from predation pressure as a result of targeted manage-
ment programs focussed on mammalian predators including
wild dogs, foxes and cats (Olsson et al. 2005; Sutherland et al.
2011; Allen et al. 2013; Anson et al. 2013; Jessop et al. 2013).
Given the results of this study, it is reasonable to speculate that
wild dog, and potentially fox, control may be indirectly impact-
ing on native arboreal wildlife by releasing lace monitors from
predation pressure in bushland ecosystems. This in turn may be
negatively impacting on native bird and arboreal mammal
populations.
Conversely, the absence of lace monitors from sites with higher
levels of anthropogenic disturbance resulted in significantly high
densities of rainbow and scaly-breasted lorikeets. Long sequences
of camera trap data suggest that both rainbow lorikeets and
dollarbirds select small (generally ,10 cm diameter) hollows to
exclude predators. Alternatively, only birds that select hollow
entrances ,10 cm diameter are successful in fledging their young.
Tree hollows containing eggs and young were visited by a range of
avian and mammalian predators that were ultimately too large to
take advantage of the food resources contained within.
Threatened species
The yellow-bellied glider is listed as vulnerable under both the
Environmental Protection and Biodiversity Conservation Act
1999 and New South Wales Threatened Species Conservation Act
1995 (TSC Act). The brush-tailed phascogale is listed as vulnerable
under the TSC Act. The rarity of these two species was a notable
feature of this camera trap study, with both species being recorded
only once by separate cameras. On the basis of records in the Atlas
of Living Australia, both of these species are considered to be
relatively rare within the study area, although better detection rates
may have been achieved using more traditional methods such
as spotlighting and ‘stag watching’. Potentially, the use of cameras
trained on targeted baits in high-quality habitat may have achieved
higher detection rates for both species.
N. Cotsell and K. Vernes
The squirrel glider, a TSC Act Schedule 2 species, was far
more common, occurring at four sites (disturbance ranking of 3
or less), suggesting that this species prefers sites that have
minimal human disturbance.
A camera at Barcoongere Way also recorded the presence of
Stephens’ banded snake, a species listed as vulnerable under the
TSC Act. The species’ presence was notable for its occurrence in
dry sclerophyll forest as it is traditionally thought to inhabit
mesic forests (Fitzgerald et al. 2002). The snake was recorded
spending long periods basking in the sun in a vertical-facing
hollow during the hottest part of the day.
The level of predation pressure that lace monitors are
imposing on threatened arboreal fauna such as yellow-bellied
gliders, squirrel gliders, brush-tailed phascogales and Stephen’s
banded snake should be further investigated.
Behavioural responses
Probably the most surprising and opportunistic result of this
study was the level of wildlife activity at an active termitarium.
During camera deployment at Barcoongere Way, the absence of
any suitable tree hollow resulted in a camera trap being sited on
the entrance of an arboreal ‘mud hollow’, or termitarium, 12 m
from the ground. Given the size of the entrance, the hollow was
probably excavated by a bird, most likely the laughing kooka-
burra, which is known to nest in arboreal termitaria (Hindwood
1959). The SD Card of this camera lasted just 34 days but
recorded four mammals, including two threatened species. Most
visiting fauna were observed to be actively engaged with the
termite nest, possibly for the food resources it held. Additionally,
three individual lace monitors spent long periods investigating the
termitarium, climbing inside and excavating the entrance to
source termites. Monitor lizards are also known to lay their eggs
in termite nests and seal the entrance (Carter 1991).
Conclusions
There are limitations to undertaking camera trap studies in the
high canopy of trees; however, the wealth of data that can be
obtained can add substantially to our understanding of arboreal
wildlife. There are obvious limitations including cost and access
to appropriate sites and the problems associated with ‘false
triggers’ as a result of branch and leaf movement. However, with
appropriate hollow selection and astute camera placement this
study has shown that it is possible to record long series of
behavioural information that, until recently, was otherwise
unavailable.
A closer examination of isolated old trees and high-value
arboreal habitat on the coastal floodplain and their importance to
wildlife seems to be of critical importance if we are to maintain
and enhance terrestrial landscapes that have high-quality habitat
for arboreal species.
Similarly, the functional role of large trees in the landscape
matrix is very poorly understood. Given the recent emphasis and
investment in protecting threatened species nationally, the spatial
relationship and the importance of old trees needs to be more
closely examined. A strong policy and legislative response is
required, but this will not transpire unless there is a better
understanding of the dynamics of tree hollows and their impor-
tance to wildlife across the broader landscape.
Surveying wildlife at tree hollow entrances
There are ongoing human demands being placed on the
eucalypt forests of the Australian landscape. What is abundantly
clear is that future targeted research and long-term monitoring
programs of old, isolated trees and HVAH are essential if we are
to understand and interpret the legacies of past forest manage-
ment practices. This information will give us a better under-
standing of how current land uses are impacting on Australia’s
native arboreal fauna.
Suggested future research initiatives using camera traps
 Whether lace monitors are exerting significant predation
pressure on threatened arboreal wildlife and if their presence/
absence is influencing the ecological guild
 The role and function of arboreal termitaria to vertebrate
arboreal wildlife
 Arboreal species preferences for hollows in different
eucalypts
 The importance of isolated old trees and HVAH on the coastal
floodplain to arboreal vertebrate wildlife with consideration
of the spatial context of these trees in the landscape matrix.
Supplementary material
A selection of arboreal wildlife images captured at tree hollow
entrances by camera traps is available on the Journal’s website.
Acknowledgements
We thank the Coffs Harbour offices of National Parks and Wildlife and
Forests NSW for allowing us to undertake research on publicly owned land.
Appreciation is also extended to Coffs Harbour City Council for allowing
access to its elevated work platform, and to Council’s arborists, Mark Baker
and Brett Prendagast, who provided valuable assistance in organising and
operating the heavy machinery. The GIS section of Council also provided
support with the preparation of spatial layers and maps. Helpful reviewer
comments were provided by Ross Goldingay and an anonymous reviewer.
We thank Skye Ravenscroft and Donella Andersen for editorial comment.
Dave Scotts assisted with identification of some of the more cryptic species.
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