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\begin{document}

\title{Conventional wisdom on roosting behaviour of Australian flying foxes - a
critical review, and evaluation using new data}



\author[1]{Tamika Lunn}%
\author[2]{Peggy Eby}%
\author[1]{Remy Brooks}%
\author[1]{Hamish McCallum}%
\author[3]{Raina Plowright}%
\author[4]{Maureen Kessler}%
\author[1]{Alison Peel}%
\affil[1]{Griffith University}%
\affil[2]{University of New South Wales}%
\affil[3]{Montana State University}%
\affil[4]{Montana State University System}%


\vspace{-1em}



  
  \date{June 10, 2024}


\begingroup
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\selectlanguage{english}
\begin{abstract}
1. Fruit bats (Family: Pteropodidae) are animals of great ecological and
economic importance, yet their populations are threatened by ongoing
habitat loss and human persecution. A lack of ecological knowledge for
the vast majority of Pteropodid bat species presents additional
challenges for their conservation and management. 2. In Australia,
populations of flying-fox species (Genus: Pteropus) are declining and
management approaches are highly contentious. Australian flying-fox
roosts are exposed to management regimes involving habitat modification,
either through human-wildlife conflict management policies, or
vegetation restoration programs. Details on the fine-scale roosting
ecology of flying-foxes are not sufficiently known to provide
evidence-based guidance for these regimes and the impact on flying-foxes
of these habitat modifications is poorly understood. 3. We seek to
identify and test commonly held understandings about the roosting
ecology of Australian flying-foxes to inform practical recommendations
and guide and refine management practices at flying-fox roosts. 4. We
identify 31 statements relevant to understanding of flying-fox roosting
structure, and synthesise these in the context of existing literature.
We then contribute contemporary data on the fine-scale roosting
structure of flying-fox species in south-eastern Queensland and
north-eastern New South Wales, presenting a 13-month dataset from 2,522
spatially referenced roost trees across eight sites. 5. We show evidence
of sympatry and indirect competition between species, including spatial
segregation of black and grey-headed flying-foxes within roosts and
seasonal displacement of both species by little red flying-foxes. We
demonstrate roost-specific annual trends in occupancy and abundance and
provide updated demographic information including the spatial and
temporal distributions of males and females within roosts. 6. Insights
from our systematic and quantitative study will be important to guide
evidence-based recommendations on restoration and management and will be
crucial for the implementation of priority recovery actions for the
preservation of these species into the future.%
\end{abstract}%



\sloppy


\subsection*{Conventional wisdom on roosting behaviour of Australian
flying foxes -- a critical review, and evaluation using new
data}

{\label{354287}}

Tamika J. Lunn\textsuperscript{1*}, Peggy Eby\textsuperscript{1,2}, Remy
Brooks\textsuperscript{1}, Hamish McCallum\textsuperscript{1}, Raina K.
Plowright\textsuperscript{3}, Maureen K. Kessler\textsuperscript{4},
Alison J. Peel\textsuperscript{1}

\emph{\textbf{1} Environmental Futures Research Institute, Griffith
University, 130 Kessels Road, Nathan, AUS 4111}

\textbf{2} \emph{School of Biological Earth and Environmental Sciences,
University of New South Wales, Sydney, AUS}

\emph{\textbf{3} Department of Microbiology and Immunology, Montana
State University, Bozeman, USA}

\emph{\textbf{4} Department of Ecology, Montana State University,
Bozeman, USA}

* Correspondence author: tamika.lunn@griffithuni.edu.au

\subsubsection*{}

{\label{abstract}}

\subsection*{Abstract}

{\label{427984}}

\begin{enumerate}
\tightlist
\item
  Fruit bats (Family: Pteropodidae) are animals of great ecological and
  economic importance, yet their populations are threatened by ongoing
  habitat loss and human persecution. A lack of ecological knowledge for
  the vast majority of Pteropodid bat species presents additional
  challenges for their conservation and management.
\item
  In Australia, populations of flying-fox species
  (Genus:\emph{Pteropus}) are declining and management approaches are
  highly contentious. Australian flying-fox roosts are exposed to
  management regimes involving habitat modification, either through
  human-wildlife conflict management policies, or vegetation restoration
  programs. Details on the fine-scale roosting ecology of flying-foxes
  are not sufficiently known to provide evidence-based guidance for
  these regimes and the impact on flying-foxes of these habitat
  modifications is poorly understood.
\item
  We seek to identify and test commonly held understandings about the
  roosting ecology of Australian flying-foxes to inform practical
  recommendations and guide and refine management practices at
  flying-fox roosts.
\item
  We identify 31 statements relevant to understanding of flying-fox
  roosting structure, and synthesise these in the context of existing
  literature. We then contribute contemporary data on the fine-scale
  roosting structure of flying-fox species in south-eastern Queensland
  and north-eastern New South Wales, presenting a 13-month dataset from
  2,522 spatially referenced roost trees across eight sites.
\item
  We show evidence of sympatry and indirect competition between species,
  including spatial segregation of black and grey-headed flying-foxes
  within roosts and seasonal displacement of both species by little red
  flying-foxes. We demonstrate roost-specific annual trends in occupancy
  and abundance and provide updated demographic information including
  the spatial and temporal distributions of males and females within
  roosts.
\item
  Insights from our systematic and quantitative study will be important
  to guide evidence-based recommendations on restoration and management
  and will be crucial for the implementation of priority recovery
  actions for the preservation of these species into the future.
\end{enumerate}

\subsubsection*{Keywords : Pteropodidae; fruit bat; camp; habitat;
management;
conservation}

{\label{518416}}

\subsubsection*{}

{\label{introduction}}

\subsection*{Introduction}

{\label{770874}}

Fruit bats (Family: Pteropodidae) are animals of extraordinary
ecological and economic importance (Fujita \& Tuttle 1991). As long
distance seed dispersers and pollinators, fruit bats play a crucial role
in the maintenance and regeneration of forest ecosystems
(Shilton\emph{et al.} 1999; Hodgkison \emph{et al.} 2003; Oleksy, Racey
\& Jones 2015). Moreover, fruit bats are responsible for the propagation
of at least 289 plant species, 186 of which have economic value, making
fruit bats important contributors to the sustainability of human
livelihoods (Fujita \& Tuttle 1991). Despite their importance, many
fruit bat species are in severe decline. Half are listed as near
threatened to extinct according to the IUCN (88 of the 177 species with
sufficient data) (IUCN 2020), with human persecution and habitat loss
identified as two of the largest threats imposed on these species
(Jenkins \emph{et al.} 2007; Acharya, Bumrungsri \& Racey 2011;
Andrianaivoarivelo\emph{et al.} 2011; IUCN 2020). While measures have
been taken in some countries to reverse this trend -- including
increased legislative protection (Eby \& Lunney 2002b; Thiriet 2010;
Aziz \emph{et al.} 2016) and community awareness campaigns (Carroll \&
Feistner 1996; Trewhella \emph{et al.} 2005; Anthony, Tatayah \& De
Chazal 2018) -- conservation and management efforts for the majority of
these species remain hindered by an enduring absence of ecological
knowledge (Fujita \& Tuttle 1991; Mickleburgh, Hutson \& Racey 2002) and
ongoing conflict with humans (Aziz \emph{et al.} 2016; Currey \emph{et
al.} 2018).

These same conservation challenges persist for Australian flying-foxes
(Genus: \emph{Pteropus} ) despite improved levels of protection.
Indiscriminate and widespread persecution and killing of flying-foxes
were persistent until the \textasciitilde{}1990's (Ratcliffe 1931;
Fujita \& Tuttle 1991; Hall 2002). Species listed as threatened are now
afforded national protection under the Environment Protection and
Biodiversity Conservation Act 1999 (EPBC Act) (Department of Agriculture
Water and the Environment 1999) and other species are protected from
harm under state-level native species legislations (Department of
Environment and Primary Industries State Government of Victoria 1988;
Queensland Government 1992; New South Wales Government 2016). However,
loss and degradation of roosting habitat continues to pose a substantial
threat, and management of these species must additionally balance
conservation outcomes with negative public perception and human-wildlife
conflict (e.g. BBC News Australia 2017; Kohut 2017; Welle 2021).

A major challenge for these species is that policies for conservation
and conflict management are often in direct contrast. The
identification, management and protection of roosting habitat are listed
as priority recovery actions for the Vulnerable grey-headed flying-fox
(\emph{Pteropus poliocephalus} ) and Endangered spectacled flying-fox
(\emph{P. conspicillatus} ) (Commonwealth of Australia 2017a). Yet in
direct contrast, roost management policies and guidelines that aim to
reduce human-wildlife conflict often promote removal of roost trees to
create perimeter buffers between the roost and private properties, which
can exceed 50 meters in some cases (State of NSW and Office of
Environment and Heritage 2018). In more extreme cases, flying-fox roost
management permits can be granted to disturb, drive away or destroy
flying-fox roosts entirely (Mo \emph{et al.} 2020a; Mo \emph{et
al.}2020b).

Management challenges in Australia are being further compounded by an
emerging and accelerating trend of urbanisation of flying-fox roost
sites, and fragmentation of roost populations (Williams \emph{et
al.}2006; Tait \emph{et al.} 2014). Roost structures are transitioning
from large roosts that are seasonally occupied by nomadic individuals
into smaller, continuously occupied roosts in urban areas (Van der
Ree\emph{et al.} 2006; Eby et al in review). This fragmentation, or
fissioning, of roost populations has been attributed to environmental
change, both land clearing of winter flowering native species in
south-eastern Australia (Eby \emph{et al.} 1999) and the concurrent
increase in availability of exotic winter food resources in urban areas
(Parry-Jones \& Augee 2001; Williams \emph{et al.} 2006). As a
consequence, increasing numbers of roosts have formed near residential
housing, particularly in metropolitan areas like Sydney, the Gold Coast
and Brisbane, despite overall population declines (Tait \emph{et
al.}2014). These urban roosts often develop into sites of ongoing
conflict with neighbours (Commonwealth of Australia 2017b) and there has
been growing demand to reduce the impact of roosts on local communities
through active management of flying-fox camps (Currey \emph{et al.}
2018). Similar changes with fragmentation and urbanisation have been
observed elsewhere (Hahn \emph{et al.} 2014a; Hahn \emph{et al.} 2014b;
Peel \emph{et al.}2017) suggesting that this occurrence is likely
representative of other systems across the range of Pteropodids.

A second major challenge for management of these species is that
systematically informed, baseline ecological knowledge is limited, so
the impact and effectiveness of efforts to contribute to either
conservation (roost restoration) or conflict (roost modification) goals
are unknown. Roosting requirements of these species are not well
understood (Commonwealth of Australia 2017a) beyond broad scale trends
in roosting patterns (e.g. Tidemann \emph{et al.} 1999; Vardon \&
Tidemann 1999), migration (Eby 1991; Eby \emph{et al.} 1999) and studies
on sociality and behaviour (Nelson 1965b; Welbergen 2005; Klose \emph{et
al.} 2009). Detailed (fine-scale) spatio-temporal patterns in animal
density and tree-use remain unquantified (Commonwealth of Australia
2017a), and knowledge on historical usage patterns (e.g. Ratcliffe 1931;
Nelson 1965b; Tidemann\emph{et al.} 1999; Vardon \& Tidemann 1999) may
be inconsistent with current usage patterns. This lack of detailed
information is of particular concern, as current conservation strategies
that aim to identify, protect and restore important roosting habitat,
and practices for managing conflict, are necessarily founded on
observations that may not fully reflect the habitat requirements of the
animals. In this context, the number of flying-fox roosts exposed to
programs of vegetation modification is increasing rapidly in Australia,
yet the potential impact of modifications to roosting habitat on
flying-foxes is largely unknown. More information is needed to provide
baseline ecological data in this time of rapid ecological change, and to
guide and support vegetation management practices and decision-making
criteria to provide a realistic representation of the roosting habitat
needs and preferences of flying-foxes. Systematic and comprehensive
examination of multiple species in Australia may also help identify
whether generalities exist among Pteropodids, and guide understanding in
systems where more limited data and resources are available.

In this paper, we seek to identify and evaluate commonly held
understandings about the roosting ecology of Australian flying-foxes,
focusing on species on the Australian mainland. We first review `grey
literature' (management, recovery and restoration plans or reports
published by state government and local groups) to identify commonly
held understandings concerning flying-fox roosting structure. We then
review the existing empirical literature, to critically evaluate the
extent of empirical support for these statements and highlight gaps in
empirical evidence. Lastly, we utilise high resolution spatial mapping
techniques and monthly field observations to systematically and
quantitatively document spatial and intra-annual temporal patterns in
flying-fox roost and tree use in south-east Queensland and north-east
New South Wales. This approach allows us to highlight where quantitative
information on flying-fox roosting has been missing, and where updated
information may be required. Our new dataset is the first to capture
fine-scale spatial and temporal dynamics of flying-fox roost use in a
structured, repeatable design, and provides baseline information in a
time of rapid ecological change. Such systematic and quantitative study
will be important for informing evidence-based recommendations to guide
vegetation modification practices and improve roost management
strategies for flying-fox conservation. This will be crucial for
implementation of effective habitat restoration projects, to
successfully balance the management of these threatened, contentious and
urbanising wildlife, and to guide comparable approaches in other
Pteropodid species across their range.

\subsection*{}

{\label{943361}}

\subsection*{Methods}

{\label{943361}}

\emph{Review of grey literature}

Flying-fox management is generally undertaken in line with site-specific
roost management plans (e.g. EcoLogical 2014; Scenic Rim Regional
Council 2015; Council of Ipswich 2016), which are adopted by local
government councils based on their state's flying-fox camp management
policy (e.g. Queensland: SEQ Catchments (2012), State of Queensland
Department of Environment and Science (2020); and New South Wales: State
of NSW and Office of Environment and Heritage (2018)). We focused on
statements made in state-level documents, as these are the primary
resource for individual roost plans. We identified common
statements/understandings across these documents, with particular
emphasis on those that pertain to 1) routine vegetation management
activities (weed removal and trimming under-storey vegetation); 2)
creation of buffers (either by clearing/trimming canopy trees, or
disturbing animals at the roost boundary); and 3) restoration
interventions.

\emph{Review of existing empirical support}

We conducted a systematic literature search of peer-reviewed published
literature using ISI's Web of Knowledge (July
27\textsuperscript{th}2020). Keywords were chosen to target studies
evaluating the within- and between-roost structure of Australian
flying-foxes (Table S1). This included any studies relevant to 1) the
physical structure of roosts (e.g. area, tree structure, tree/roost
selection), 2) the social structure of roosts (e.g. demographic and
species structuring), 3) roosting behaviour (e.g territoriality and
fidelity of individuals), 4) movement and migration relating to
occupancy and abundance of roosts, and 5) roost microclimate. In
addition to the literature search, reference lists and relevant studies
already known to the authors were also screened to identify potentially
relevant studies not captured by our initial search. We also included
empirical support from key unpublished sources (e.g. theses).

\emph{Empirical data collection}

We collected data on roosting structure at eight sites in south-east
Queensland and north-east New South Wales (Figure 1). These sites were
chosen to represent a gradient of habitats utilised by flying-foxes,
ranging from metropolitan areas of Brisbane and the Gold Coast, to
roosts in peri-urban and rural areas (Figure 1, Table 1). All sites were
previously documented as having a continuous population of grey-headed
or black flying-foxes. Little red flying-foxes visited some roost sites
intermittently, however no roost sites occurred within the distribution
of spectacled flying-foxes (National Flying-Fox Monitoring Program
2017).

We mapped the spatial arrangement of all overstory, canopy and midstory
trees in a grid network of 10 stratified random subplots (20 x 20 meters
each) per roost site. Trees were mapped and tagged using tree survey
methods described in the ``Ausplots Forest Monitoring Network, Large
Tree Survey Protocol'' (Wood \emph{et al.} 2015). To evaluate
spatio-temporal patterns in roosting, we revisited all tagged trees and
scored the extent of species occupancy using the following tree
abundance index: 0= zero bats; 1= 1-5 bats; 2=6-10 bats; 3=11-20 bats;
4=21-50 bats; 5=51-100 bats, 6=101-200 bats, 7= \textgreater{}200 bats.
For a subset of trees (N=60 per site, consistent through time) absolute
counts and minimum/maximum roosting heights of each species were taken.
Overall roost perimeter (perimeter of area occupied) was mapped with GPS
(accurate to 10 meters) immediately after the tree survey to estimate
perimeter length and roost area. Total abundance at each roost was also
estimated with a census count of bats where feasible (i.e. where total
abundance was predicted to be \textless{}5,000 individuals), or by
counting bats as they emerged in the evening from their roosts
(``fly-out''), as per recommendations in Westcott \emph{et al.} (2011).
If these counts could not be conducted, population counts from local
councils (conducted within \textasciitilde{}a week of the bat surveys)
were used, as total abundance of roosts are generally stable over short
timeframes (Nelson 1965b). Because roost estimates become more
unreliable with increasing abundance we also converted the total
estimated abundance into an index estimate for use in analyses, as per
values used by the National Flying-Fox Monitoring Program (2017). Census
index categories were as follows: 1 = 1-499 bats; 2 = 500-2,499 bats; 3
= 2,500 - 4,999 bats; 4 = 5,000 - 9,999 bats; 5 = 10,000 - 15,999 bats;
6 = 16,000 - 49,999 bats; and 7 = \textgreater{} 50,000 bats. Roosting
surveys were repeated once a month for 13 months (August 2018 - August
2019). More detailed methods of empirical data collection can be found
in Appendix S1 in the Supporting Information.

\emph{Statistical analyses}

The main statistical comparisons tested with our empirical data were: 1)
whether frequency of occupation is greater for subplots in `core' areas
of the roost compared with subplots in irregularly occupied `peripheral'
areas (defined by occupation greater than or less than 80\% of surveys
respectively (Appendix S1); 2) whether bat occupation decreases with
distance from the roost centre (per species); 3) whether bat species
segregate in vertical space; and 4) whether dominant individuals occupy
the centre of roosts, and subdominant individuals the outer area (per
species). We also provide qualitative comparisons of 5) seasonal
patterns of abundance and occupancy per species; and 6) whether bat
species segregate in horizontal space.

We utilised generalized additive models for all statistical comparisons
to allow for nonlinearity, with random effects modelled with smooth
functions. Roost site and subplot were modelled using a standard random
effects smoothing function. Session was modelled using a cyclic cubic
regression spline in cases where seasonality in the time series was
evident (all comparisons except those involving the proportion of male
black, male grey-headed and combined male bats per tree), otherwise
session was modelled with a standard random effects smoothing function.
We accounted for non-independence (nesting) of random effects by
including an autoregressive model for errors in the model (Yang \emph{et
al.} 2012; Laurinec 2017). For the comparisons involving evaluation of
species, models were run separately for each species owing to
differences in seasonality of occupation (and so, differences in the fit
of cyclic cubic regression splines). Error distribution for comparisons
were specified according to data type and extent of zero-inflation (as
per Crawley 2013). We fit the models and performed checks of
standardised residuals in R (Version 4.0.2), using the `mgcv' package
(functions `gamm' and `gam.check') (as per Wood 2017). See Appendix S1
in the Supporting Information for more detailed information on modelling
decisions and a summary table of comparisons. Summarised data and
annotated R code are available on GitHub at: \textless{}
https://github.com/TamikaLunn/FF-roost-ecology \textgreater{}.

\subsubsection*{}

{\label{results}}

\subsection*{Results}

{\label{395246}}

From our review of management, recovery and restoration documents
published by state government, we highlighted 31 commonly held
understandings relevant to flying-fox roosting structure (Table 2). From
our systematic search for empirical literature we generated a total of
79 search results. Of these, 52 were removed through screening (10 being
outside the Australian mainland, 4 on non-Pteropus species, and 38
focused on topics other than roost structure). An additional 18
published studies and 4 honours/PhD theses were included from citations
and the author's reference collections, giving 49 included studies in
total (Appendix S1). Lastly, we generated an empirical dataset
consisting of 13 monthly repeat measures from 2,522 trees across eight
roost sites. Roost sites contained 118-474 measured and tagged trees
each, with an average of 2 (sparsely structured) to 75 (densely
structured) trees per 20x20 meter subplot. Tree roosting height and
count was recorded for 9,056 trees out of 32,206 repeat measures. (Note
that our total repeat measures were less than 32,786 owing to cases of
tree removal through the duration of the survey.) We report model
outputs of main interest in the main text, but see Appendix S2 in the
Supporting Information for full model output.

Below, and in Table 2, we synthesise how commonly held understandings
compare with existing literature and new data from our study.

\emph{Use of area}

\emph{{``Some areas of permanent camps are more consistently occupied
('core areas') than others''}}

This understanding was generally reported by previous studies, with none
contradicting it (Table 2). Consistent with these other studies (e.g.
Nelson 1965b; Welbergen 2005) we observed some areas of roosts to be
more consistently occupied than others (Figure 2). Occupancy of subplots
ranged between 100\% (30 subplots) to under 10\% (15 subplots) across
surveys when bats were present in roosts.

\emph{{```Core areas' are more densely occupied than `peripheral
areas'''}}

Existing empirical data broadly supported this statement (Table 2). In
our study, peripheral areas (those occupied less than 80\% of the time)
generally were less densely occupied than core areas, though density
varied substantially across roost site, subplot and session (all
contributed substantially as random effects). Here, lower density refers
to both a lower number of bats per subplot in peripheral subplots
(-0.581 \selectlanguage{ngerman}± 0.177, p= 0.001, Figure 3), and a lower proportion of occupied
trees (-0.222 ± 0.078, p= 0.005,

Appendix S2). Within subplots, we also note that some trees were more
consistently used than others, including trees that were occupied in
100\% of surveys where bats were present at the roost (Appendix S1). The
number of bats per tree in irregularly occupied trees (occupied less
than 80\% of the time) were typically lower than for regularly occupied
trees (-0.606 ± 0.034, p\textless{} 0.001).

We observed negative relationships between bat occupation metrics and
distance from the roost centre, including in the number of bats per
occupied subplot (-1.639 ± 0.016, p\textless{} 0.001, Figure 4) and
proportion of occupied trees per subplot (-0.315 ± 0.034, p\textless{}
0.001, Appendix S2). This decline with distance from the centre of
subplot was largely driven by little red flying-foxes (Figure 4). Roost
site, subplot and session also all contributed substantially as random
effects (Appendix S2).

\emph{{``Roost area fluctuates with total abundance''}}

Studies have previously reported changes to total roosting area, but
none to date have formally quantified the relationship between area and
abundance (Table 2). From our data, we observed substantial fluctuations
in total roost area within some roost sites across monthly surveys, and
overall, a positive relationship with total roost abundance. The extent
of variation was variable across roosts, however (Figure 5). We note
that relationships between total abundance and area were likely masked
in many roosts by the large span of population values in some index
categories (e.g. index 6 spans 16,000 - 49,999 bats). It is probable
that data of finer resolution may have detected this relationship more
strongly for roosts in this size range, but are not available in this
dataset.

\emph{Spatial segregation of species}

Results from our new dataset included systematic recording of the three
species that occur in south-east Queensland -- \emph{P. alecto}
,\emph{P. poliocephalus} and \emph{P. scapulatus} . The majority of
observations were made of \emph{P. alecto} , and \emph{P. poliocephalus}
, which occupy this region continuously through the year. \emph{P.
scapulatus} was found irregularly at some roosts, which is consistent
with the seasonal migration patterns of this species (Nelson 1965a).
Fine-scale spatial overlap between species was evaluated during surveys
when multiple species were present (N=73, 70.2\% of surveys). Black and
grey-headed flying-foxes co-occurred in 65 surveys (62.5\%), black and
little red flying-foxes co-occurred in 17 surveys (16.3\%), and
grey-headed and little red flying-foxes co-occurred in 9 surveys
(8.7\%). We observed roost-dependent support for spatial segregation of
species.

\emph{{``Species share roosts sites, but segregate spatially within''}}

Observations from previous studies commonly report co-occupation of
roosts by multiple species, with anecdotal observations of inconsistent
overlap or separation within and between trees (Table 2). We observed
some horizontal spatial segregation of species, with species showing
preference for discrete areas in roosts. In the `Lismore' roost, for
example, black flying-foxes were commonly distributed toward the eastern
part of the roost and grey-headed flying-foxes in the western part of
the roost (Appendix S3). Likewise, in the `Clunes' roost, black
flying-foxes were commonly observed toward the north-eastern part of the
roost and grey-headed flying-foxes in the south-western part of the
roost (Appendix S3). Of 659 occupied subplots across the survey period,
only 34.1\% (225, binomial confidence interval: 0.31-0.38) showed
co-occupation by two different species (Figure 6A). Co-occupation of
individual trees by two different species was also relatively low --
across surveys where two species were present, 4.6\%-7.9\% of occupied
trees were co-occupied by two species, versus 92.1-95.4\% that were
occupied by only one species (Figure 6B). Only six trees were ever
observed to occupy all three species at once.

\emph{{``Large influxes of species into roosts (especially little red
flying-foxes) can displace other species''}}

Only one previous study had reported displacement by species, reporting
an anecdotal observation of black and grey-headed flying-foxes being
displaced by little red flying-foxes (Table 2). Our quantitative data
document changing distribution of regular species occupants in response
to (`invading') irregular species occupants, supporting this prior
observation. Little red flying-foxes, in particular, were observed to
displace black and grey-headed flying-foxes from their usual roosting
locations (most notably at the `Redcliffe' roost: Appendix S3). Black
and grey-headed flying-foxes tended to co-occur in roosts without too
much impact on each other (Appendix S3).

\emph{{``Species roost at different heights''}}

Previously only one study had formally documented differences in
roosting height between species (Table 2). This included record of black
flying-foxes and grey-headed flying-foxes only, and did not provide
measures of absolute height (rather, roosting in different quadrants of
trees) (Welbergen 2005). From our new dataset, we observed segregation
of species by roosting height, with black flying-foxes typically showing
the highest roosting heights (average maximum height with interquartile
range: 18.0, 14.6-21.0; average minimum height with interquartile range:
14.3, 11.3-17.2), followed by grey-headed (maximum: 15.1, 11.2-18.9;
minimum: 12.6, 8.8-16.2), then little red flying-foxes (when present)
(maximum: 11.4, 9.2-13.6; minimum: 8.8, 7.1-10.4) (Figure 7). Note,
however, that topographical variation within roosts was not taken into
consideration in measures of height. Differences in heights presented
here reflect a relative difference in roosting heights from the ground
within trees, but may not reflect true, realised height relative of the
canopy.

\emph{Demographic/social structure}

\emph{{``The majority of roost trees are occupied by mixed groups of
adults, with territories comprised of a single male and one or more
females and their dependent young''}}

We commonly observed roost trees to be occupied by mixed groups of
sexes, with a single tree occupied by one or more males, and one or more
females and their dependent young. This is inconsistent with general
knowledge based on historical studies like Nelson (1965a) and Nelson
(1965b), but consistent with more contemporary observations (Table 2).
We also observed cases where trees were occupied by entirely male
individuals (consistent with reports of `bachelor male' trees in Markus
(2002)). We would note here that a single tree may contain multiple male
territories (Markus 2002; Connell 2003) and the survey methods did not
allow inference on the composition of individual territories, only
individual trees. The proportion of males per tree appeared to follow
seasonal patterns that was mostly consistent between black and
grey-headed flying-foxes within roosts (Appendix S2). Some roosts
(`Toowoomba', `Avondale', `Lismore') showed an increase in the
proportion of males per tree after parturition in September/October,
while other roosts (`Sunnybank', `Canungra') decreased immediately after
this time. We also did not observe complete segregation of sexes at any
time of the year, in contrast to Nelson (1965b) who noted complete
segregation between September until early December, and March to April.

\emph{{``Dominant individuals (defined as reproducing males and females)
occupy the centre of roosts and subdominant individuals (defined as
non-reproducing males and females) the outer area''}}

Prior studies reported inconsistent spatial patterns in flying-fox
occupation (Table 2). From our new dataset, we observed that the
proportion of males per tree increased with distance from the roost
centre (0.15 ± 0.039, p\textless{}0.001), though this effect was
relatively small and variable across roosts and species (Figure 8). If
we assume that only dominant males share their territory with females
and their young (Markus 2002; Welbergen 2005), a lower proportion of
males in trees closer to the centre of roosts may indicate that dominant
individuals occupy the centre of some roosts and subdominant individuals
the outer area. The small effect sizes observed would suggest that there
is no clear spatial structure to reproductive groupings or dominance
groupings. This can be seen also in maps showing male composition by
tree relative to the roost perimeter, given in Appendix S4 in the
Supporting Information.

\emph{Roost abundance/occupancy}

\emph{{``Individual roosts have distinguishable seasonal patterns of
abundance and occupation.'' \& ``Intra- and inter-annual variations in
abundance can be extreme'' \&``Roost abundance peaks in March''}}

Prior studies reported inconsistent patterns in occupancy and abundance
(Table 2). In out dataset, seasonal patterns in abundance and density
were roost specific (Figure 9). Some roosts showed patterns consistent
with the general notion that total roost abundance peaks towards March
(Nelson 1965b; State of NSW and Office of Environment and Heritage 2018)
(e.g. `Redcliffe', `Canungra' and `Clunes'). Others showed no
considerable fluctuation in abundance (`Burleigh') or peaks at other
times (`Toowoomba', `Sunnybank', `Avondale', `Lismore') (Figure 9). The
latter cases potentially highlight that population dynamics are more
strongly driven by local dynamics in these roosts (e.g. food
availability) (Parry-Jones \& Augee 1992; Eby \emph{et al.} 1999;
Parry\selectlanguage{english}-Jones \& Augee 2001; Giles \emph{et al.} 2016), than reproductive
cycles as described in Nelson (1965b). Little red flying-foxes showed
seasonal trends in occupancy and density, peaking in February-March
(Appendix S2). Seasonal trends in grey-headed and black flying-fox
numbers were less consistent between roost sites (Appendix S2).

\subsubsection*{}

{\label{discussion}}

\subsection*{Discussion}

{\label{182303}}

The success of efforts to conserve Pteropodid bats across their
distribution relies on effective population and habitat management.
Pivotal to this is a baseline understanding of species ecology and
behaviour, which is currently lacking for the majority of these species
(Fujita \& Tuttle 1991; Mickleburgh, Hutson \& Racey 2002). Here we
provide a synthesis on all existing literature, as well as an
unprecedented empirical dataset, to meet that need for Australian
species of \emph{Pteropus} . We highlight that many existing beliefs on
which conservation and management decisions are based, are unsupported
or outdated, and suggest that management plans should be updated to
incorporate improved knowledge. Most importantly, we highlight that a
one-size-fits-all approach to roost management will be inappropriate,
given the extent of variation between sites even within a regional area.
Roost management guidelines need to be changed to promote a more
tailored approach that requires preliminary data acquisition before
management plans are formulated and approved.

\emph{Existing understanding not supported}

\emph{{``Individual roosts have distinguishable seasonal patterns of
abundance and occupation''}}

All roost sites in our empirical dataset were occupied continuously
throughout the year by adults and juveniles/sub-adults of both sexes.
This type of roost occupation has been noted from 1981 onwards
(Puddicombe 1981; Parry-Jones 1985) and has become common in recent
decades (e.g. Aston 1987; Eby 1991; Larsen \emph{et al.} 2002; Van der
Ree \emph{et al.} 2006). This pattern of occupation contrasts to the
`summer' and `winter' pattern described historically by Nelson (1965a)
and Nelson (1965b) and cited in the \emph{Flying-fox Roost Management
Guideline} for Queensland, where `summer roosts' of reproducing
individuals would form between \textasciitilde{}September/October and
April/May, and `winter roosts' of dispersed animals would form between
April/May and September (Ratcliffe 1931; Nelson 1965a; Nelson 1965b;
Parry-Jones \& Augee 1991; Parry-Jones \& Augee 1992). For these roost
types, overwintering animals at summer roosts were rare, and when
present, were documented as being predominantly juveniles or lone adult
males (Nelson 1965b).

While seasonally occupied colonies are still observed (e.g.
Klose\emph{et al.} 2009), an increasing number of roosts are now
consistently occupied year around, particularly in urban areas
(Parry\selectlanguage{english}-Jones \& Augee 2001; Tait \emph{et al.} 2014). The cyclic
patterns of summer aggregation and winter dispersal were originally
thought to reflect social drivers and availability of resources
(Parry-Jones \& Augee 1992). Specifically, territory formation (from
January) and conception (from \textasciitilde{}March) (\emph{P.
poliocephalus} and \emph{P. alecto} ) (Welbergen 2005) coupled with
abundant flowering of native flora in these months (Nelson 1965a), were
understood to drive and support aggregative living in summer/autumn,
while decreased food availability and the cessation of mating from
\textasciitilde{}May triggered the animals to disperse and adopt a
less-gregarious living style in winter (Parry-Jones \& Augee 1992). This
ecology has changed in more recent decades, where continuous
availability of exotic foods in urban areas has reduced the need for
migratory behaviours and allows aggregate groups to remain year round
(Parry\selectlanguage{english}-Jones \& Augee 2001; Williams \emph{et al.} 2006).

Policy documents containing only historical information on flying-fox
occupation patterns (including the most recent \emph{Flying-fox Roost
Management Guideline} for Queensland: State of Queensland Department of
Environment and Science (2020)) are of concern, as recommendations based
on historical usage patterns may be inconsistent with current usage
patterns, particularly in urban areas where occupation patterns have
changed the most (Larsen \emph{et al.} 2002; Tait \emph{et al.} 2014),
and where human-bat conflict is often the highest (Kung \emph{et
al.}2015). Roost monitoring prior to management actions should encompass
every season, and not assume that bats will disperse in winter.
Similarly, contemporary overwintering roosts commonly contain
individuals from all age and sex groups and may be consistently utilised
through time (Larsen \emph{et al.} 2002; Tait \emph{et al.} 2014).

\emph{{``The majority of roost trees are occupied by mixed groups of
adults, with territories comprised of a single male and one or more
females and their dependent young'' \& ``Dominant individuals (defined
as reproducing males and females) occupy the centre of roosts and
subdominant individuals (defined as non-reproducing males and females)
the outer area''}}

These historic perspectives also describe complete separation of males
and females between September until early December (the period
immediately before parturition, during lactation, and before conception)
and again post March (after conception) (Nelson 1965a; Nelson 1965b).
During these times, animals were historically noted to segregate by tree
or height, such that all social contacts were between individuals of the
same sex. However, these observations contrast with more recent
observations of flying-fox social groupings (Puddicombe 1981; McWilliam
1984; Eby \emph{et al.} 1999; Welbergen 2005), and observations from
this study. In contemporary roosts, mixed-sex groups are commonly
present all year around, such that males and females co-occur in the
roost and within trees year around.

Historically, during the times that males and females co-occurred within
roosts, four types of social groupings were noted: 1) guard groups on
the outsides of roosts, 2) family groups of one male, one female and one
young, 3) other adult groups including polygamous males, 4) groups of
juveniles. More recent observations, and results from this study
suggest, however, that there is no clear spatial structure in the
distribution of the sexes within the roost. Puddicombe (1981) notes that
reproductive groups (mixed groups of males, females and their young)
were uniformly distributed through the camp and present in peripheral
areas (McWilliam 1984). Additionally, in this study we observed randomly
distributed groups of mixed males and females, and groups of males. This
potentially reflects the change in occupancy patterns in flying-fox
roosts, where aggregative living was historically believed to be driven
by strong social drivers (i.e. mating), whereas aggregative living in
contemporary roosts is thought to be driven by continuous resource
availability in the urban environment (Parry\selectlanguage{english}-Jones \& Augee 2001;
Williams \emph{et al.} 2006). The observations will have implications
for current management plans. Specifically, in support of current
guidelines, managers should avoid management actions during times of the
year when females are in late stages of gestation and have dependant
young that cannot fly on their own (as per Commonwealth of Australia
2015; Department of Environment and Science 2020a). Importantly (and in
contrast to current guidelines), actions scheduled within this time
should note that restricting work to edges of roosts will likely not
circumvent disturbances to gestating females and dependant young

\emph{Existing understanding supported, but conditional on roost site
and local conditions}

{}\emph{{``Roost abundance peaks in March'' \& ``Intra- and inter-annual
variations in abundance can be extreme''}}

March was identified in some management documents as being the time for
peak abundance in flying-fox roosts (e.g. State of Queensland Department
of Environment and Science 2020). However, studies on \emph{P.
poliocephalus and P. alecto} identify a typical pattern of increasing
abundance from September-October (when females give birth) until a peak
in January-February (when the season's young are able to fly
independently) (Nelson 1965b; Eby 1991; Eby \& Palmer 1991; Parry\selectlanguage{english}-Jones
\& Augee 2001). Roost sizes then decrease during March-April (the period
of mating) to low winter counts in continuously occupied/overwintering
roosts, or zero winter counts in seasonally occupied summer roosts
(Nelson 1965b; Eby 1991; Eby \& Palmer 1991). These studies note that
cyclical patterns of occupation are driven by reproductive factors (i.e.
timing of birth and independent flight), but highlight that irregular,
local dynamics of food availability can superimpose variability into
these patterns of abundance (Parry-Jones \& Augee 1992). Indeed, many
studies note high intra- and inter-annual variability in abundance.
Parry\selectlanguage{english}-Jones and Augee (2001), for example, note that animals from their
study roost appeared to migrate away and decrease in abundance in
response to a blossoming event, presumably to move to a roost in closer
proximity to the blossoming.

In our study, some roosts showed patterns consistent with a total roost
abundance peak towards March (e.g. `Redcliffe', `Canungra' and
`Clunes'). Others showed either no considerable fluctuation in abundance
(`Burleigh') or peaks at other times (`Toowoomba', `Sunnybank',
`Avondale', `Lismore'). Drivers of peaks were variable between roosts.
For the `Redcliffe' roost, seasonal migration of little red flying-foxes
from \textasciitilde{}January 2019 contributed to a peak in abundance
around March (see species abundance plots in

Appendix S2). For the `Lismore' roost, a blossoming event in winter 2018
triggered an influx of nomadic bats into the population, driving the
peak observed in August 2018. Dynamics observed in other roosts were
likely the result of local dynamics of food availability.

We note also that estimates of abundance from our study were much
smaller than those of historical estimates. Ratcliffe (1931) describes
`small' roosts as \textasciitilde{}5,000-10,000 animals, `medium' as
10,000-50,000, and `large' as anything over this size. Ratcliffe (1931)
also report roosts in northern Queensland with bats `into the millions'
(Red River) and `exceeded a quarter of a million, possibly considerably'
(Burnett River). Likewise, Lunney and Moon (1997) report historical
observations of flying-foxes in the Richmond Valley (1870's) as into the
millions. The maximum roost site observed in this current study was
\textasciitilde{} 95,000, recorded at the Lismore roost in August 2018
in response to a local eucalyptus flowering event. Roost sizes of
\textless{}5,000 were more common for the roost sites surveyed and,
extending from the sizes in Ratcliffe (1931), may constitute a new
category of `very small'. Local management areas should expect that
local conditions can change substantially and rapidly for flying-fox
populations, resulting in population changes outside of times predicted
by demographic driven dynamics alone. An understanding of the timing and
productivity of flower resources within the feeding range of roosts is
likely to be of greater importance to forecasting and interpreting large
population fluctuations than are reproductive considerations.

\emph{Existing understanding supported}

\emph{{``Some areas of permanent camps are more consistently occupied
('core areas') than others'' \& ``\,`Core areas' are more densely
occupied than `peripheral areas''' \& ``Roost area fluctuates with total
abundance''}}

Variability in the usage and occupation of areas within roosts have been
highlighted in management documents (e.g. SEQ Catchments 2012). This
includes more persistent usage of `core' areas, higher occupation of
`core' areas, and variability in the roost perimeter (reflecting
expansion and contraction from the core area). All existing literature
(to our knowledge) and the new data from our study support these
understandings. We would note however, the distinction between a
`core/peripheral' roost area and a `central/edge' roost area. We defined
the core area based on consistency of occupation, not spatial location.
Areas identified to be `core' were not necessarily in the centre of the
roost (see location of roost centroid relative to the roost perimeter
and surveyed subplots, in Appendix S3). This distinction has not
necessarily been made in literature and management plans to date but has
important implications for the interpretation of `core' roosting areas,
and management recommendations specific for `core/central' or
`peripheral/edge' areas. For example, it cannot be assumed that buffer
creation via vegetation removal from the roost edge will not affect a
`core' area of bat roosting, and so will not have a substantial impact
of flying-foxes. Management activities should be prescribed for specific
zones in roosts, based on prior monitoring of the roost, and recognising
the ecological importance of different areas (Pallin 2000; Ku-ring-gai
Council 2018). In addition, prior monitoring of core/peripheral roosting
areas will be important to inform the location and potential
effectiveness of buffer creation. Given the potential for roost area to
fluctuate with abundance, creation of buffers via vegetation removal may
reduce the area of normal roost habitat available, and result in an
expansion into new areas when flying-fox numbers increase (as noted in
Currey \emph{et al.} 2018). The prescription of buffers should be
planned with care to avoid unintended outcomes during periods of high
population abundance.

\emph{{``Species share roosts sites, but segregate spatially within'' \&
``Large influxes of species into roosts (especially little red
flying-foxes) can displace other species'' \& ``Species roost at
different heights''}}

The range of black flying-foxes underwent a phase of rapid southern
expansion in the late 1990s and early 2000s, increasing the area of
overlap with grey-headed flying-foxes (Roberts \emph{et al.} 2012a). As
the two species co-occupy roosts where their distributions overlap, this
process has substantially increased the number of roosts occupied by
both species, and thereby increased the pertinence of understanding the
structure of mixed species roosts. There has been relatively little
formal documentation of species overlap and segregation within roosts.
Ratcliffe (1932) noted that sections of roosts were occupied by
different species - specifically, that little red flying-foxes and black
flying-foxes occupied different areas. Some horizontal separation has
also been noted by Nelson (1965b) and Klose \emph{et al.} (2009), and
notes of displacement by little red flying-fox have been described in
Birt and Markus (1999). We contribute quantitative, spatial information
on the extent and overlap little red flying-fox, black flying-fox and
grey-headed flying-fox, extending on the predominantly anecdotal
observations underlying management documents to date. Findings from our
data support common understandings of flying-fox roost structure:
species commonly showed preferences for discrete areas of roosts, and
even more commonly, preference for occupation of separate trees. We also
observed segregation of species by roosting height, with black
flying-fox foxes showing the highest roosting, followed by grey-headed
flying-foxes and little red flying-foxes. These findings flag the
importance of species monitoring of roost sites prior to management
interventions. It cannot be assumed, for example, that species occupy
areas of the roost uniformly, and management actions need to consider
areas that may be more or less important to vulnerable species, like the
grey-headed flying-fox. These results also give interesting insights
into understanding disease transmission dynamics within roosts, relating
to the extent of mixing of primary host species (e.g. black flying-foxes
for Hendra virus) and other species presumed to be incidental hosts
(e.g. grey-headed and little red flying-foxes).

\emph{Final comments and implications for roost management}

State-level management guidelines, including the \emph{Flying-fox Camp
Management Policy} (State of NSW and Office of Environment and Heritage
2018) and the \emph{Flying-fox Roost Management Guideline} (State of
Queensland Department of Environment and Science 2020) outline several
camp-based management approaches that involve the modification or
removal of vegetation within roost sites. `Routine camp management
actions' include the removal of tree branches or whole trees, weed
removal, trimming of understorey vegetation, and minor habitat
augmentation. The aim of such actions are often to encourage roosting in
alternative areas of the roost (e.g. Geolink 2010; EcoLogical 2014), or
to increase the sustainability of existing roosting habitat for
flying-foxes (e.g. Ku-ring-gai Council 2018). These actions are
considered to be low impact activities (Department of Environment and
Science 2020b) and do not require referral under the EPCB act
(Commonwealth of Australia 2015), however these actions may considerably
alter the structure of roost vegetation and decrease the suitability of
a roost as habitat (Ku-ring-gai Council 2018). For example, the removal
of mature weed vines in the canopy and midstory, as well as the clearing
of understory, can reduce the structural complexity of roost vegetation.
This may have immediate and direct effects on roosting flying-foxes, and
may accidently cause bats to disperse or adjust use of roost trees in
ways contradictory to conflict management. This may also have long-term,
indirect implications for the ability of flying-foxes to survive extreme
weather events, by altering roost macroclimate and removing physical
refuge needed at times of extreme heat (Welbergen \emph{et al.} 2008).

Individual and council-level roost management plans developed by local
governments under the guidance of these policies, commonly utilise these
vegetation management measures (e.g. EcoLogical 2014; Logan City Council
2015; Sunshine Coast Regional Council 2016; Ku-ring-gai Council 2018),
though the long-term implications for flying-foxes of vegetation works
are rarely noted (with the exception of Ku-ring-gai Council 2018). We
recommend that vegetation removal should not be considered low impact by
default. Routine management actions should follow a mosaic pattern
(State of NSW and Department of Planning Industry and Environment 2019),
or target weeding on a weed-by-weed case basis (Ku-ring-gai Council
2018), and seek to maintain refuges in the mid- and lower storeys at all
times. Special care not to disturb bats should be taken in identified
core areas of the roost.

\subsubsection*{}

{\label{conclusion}}

\subsection*{Conclusion}

{\label{519309}}

This study takes a thorough, multifaceted approach to better understand
the ecology of flying-fox roost use and structure in Australia. We build
upon broad-scale knowledge of historic roosting occupancy and abundance
patterns, and provide updated baseline information on roosting structure
in urban and peri-urban roosts by providing fine-scale spatial, and
temporal data on roost and tree use. Specifically, we demonstrate high
variation in patterns of occupancy and abundance between roosts sites,
and provide updated demographic information including the spatial and
temporal distributions of males and females within roosts. We also show
evidence of sympatry and indirect competition between species, including
spatial segregation of black and grey-headed flying-foxes within roosts,
and seasonal displacement of both species by little red flying-foxes.
The outcomes of this research will be of immediate, practical benefit to
management and conservation of flying-fox roosts in Australia, and meets
research needs specifically identified in the draft Recovery Plan for
the Vulnerable grey-headed flying-fox. The level of spatial and temporal
detail provided in our empirical study will be important in designing
management plans that are sensitive to flying-fox habitat needs, and in
identifying and protecting important habitat areas within roosts that
are reflective of current movements and preferences. Most importantly,
we highlight that a one-size-fits-all approach to roost management will
be inappropriate, given the extent of variation between sites even
within a regional area. Fine-scale information on roost tree preferences
will also improve understanding of the potential impacts of existing
conflict management strategies involving vegetation removal, including
buffer creation, and can guide vegetation removal efforts to heed these
habitat requirements. This information is timely, and much needed in
advance of the recently announced Environmental Trust grants program for
flying-fox habitat restoration, and in the face of continued and
increasing urbanisation of flying-foxes in Australia.

\subsubsection*{}

{\label{acknowledgements}}

\subsection*{Acknowledgements}

{\label{303242}}

We would like to thank Beccy Abbot, Kirk Silas, Devin Jones, Liam
Chirio, Rachel Smethurst and Cara Parsons for their assistance in the
field. Fieldwork for this work was supported by a Paddy Pallin Research
Grant and a `Grants in Need' science grant awarded to TJL, sponsored by
the Paddy Pallin Foundation, The Royal Zoological Society of NSW and The
Foundation for National Parks and Wildlife, and the National Science
Foundation (a Coupled Dynamics of Natural and Human Systems grant
DEB1716698 and a DARPA PREEMPT program Cooperative Agreement
\#D18AC00031). TJL was supported by an Endeavour Postgraduate Leadership
Award and a Research Training Program scholarship sponsored by the
Australian Government, AJP was supported by an ARC DECRA fellowship
(DE190100710) and a Queensland Government Accelerate Postdoctoral
Research Fellowship, RB was supported by a Griffith University Honours
Allowance and an EFRI Honours Thesis Write-Up Scholarship, and RKP was
supported by USDA National Institute of Food and Agriculture (Hatch
project 1015891). This research was conducted under a Griffith
University Animal Research Authority permit (DEB-1716698), a Scientific
Purposes Permit from the Queensland Department of Environment and
Heritage Protection (WISP17455716), a permit to Take, Use, Keep or
Interfere with Cultural or Natural Resources (Scientific Purpose) from
the Department of National Parks, Sport and Racing (WITK18590417), a
Scientific Licence from the New South Wales Parks and Wildlife Service
(SL101800) and general and products liability protection permit (GRI 18
GPL), and with permission to undertake research on council and private
land. The content of the information does not necessarily reflect the
position or the policy of the U.S. government, and no official
endorsement should be inferred. We acknowledge the Danggan Balun, Kabi
Kabi, Turrbal, Widjabul Wia-bal, Yugambeh and Yuggera Ugarapul people,
who are the Traditional Custodians of the land upon which this work was
conducted.

\textbf{Data Accessibility Statement}

Summarised data and annotated R code will be made available on GitHub
at: \textless{} https://github.com/TamikaLunn/FF-roost-ecology
\textgreater{}. Summarised data will also be made available on Dryad
after publication.

\textbf{Competing Interests Statement}

The authors have no competing interests to declare.

\textbf{Author Contributions}

TJL and PE conceived and designed the research; TJL acquired funding for
the research and led project administration; TJL and RB collected and
curated the data; TJL, AJP and HM analysed and visualised the data; AJP,
HM, RKP and PE provided supervision; TJL drafted the manuscript, and all
authors participated in review and editing of drafts.

\subsubsection*{}

{\label{references}}

\subsection*{References}

{\label{335648}}

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\par\null

\textbf{Tables}

Table 1: Information on roost sites included in the study. Contemporary
roost types (features greyed) are characterised by: being a new
overwintering site (defined as having either formed since 2007 or
changed to an overwintering site since 2007), having a high number of
neighbouring roosts within 45 km, having a high proportion of
surrounding urban land-cover, and by being in close proximity to urban
land-cover. Foraging radii enclose 45 km from roost study sites (as per
Giles \emph{et al.} 2018).\selectlanguage{english}
\begin{longtable}[]{@{}lllllll@{}}
\toprule
Roost site & Type & Year of formation/new overwintering & Number of
neighbouring roosts within foraging radius & Area of urban land-use
within foraging radius (km\textsuperscript{2}) & Distance to nearest
urban edge (km) & Number of tagged trees inside subplots (number of
trees per km\textsuperscript{2})\tabularnewline
\midrule
\endhead
& \emph{Within 45 km foraging radius} & \emph{Within 45 km foraging
radius} & \emph{Within 45 km foraging radius} & \emph{Within 45 km
foraging radius} & \emph{Within 45 km foraging radius} &\tabularnewline
Toowoomba & & 2009 & 9 & 135.2 & 0 & 118 (29.5)\tabularnewline
Redcliffe & & 2003 & 41 & 1447.3 & 0 & 286 (71.5)\tabularnewline
Sunnybank & Contemporary & 2009 & 67 & 1650.0 & 0 & 226
(56.5)\tabularnewline
Avondale & Contemporary & 2008 & 68 & 1650.0 & 0 & 268
(67)\tabularnewline
Burleigh & Contemporary & 2013 & 50 & 1601.6 & 0 & 327
(81.8)\tabularnewline
Canungra & & 1996 & 59 & 1601.6 & 11.5 & 474 (118.5)\tabularnewline
Clunes & & 2014 & 40 & 20.8 & 14.4 & 349 (87.3)\tabularnewline
Lismore & & 2007 & 37 & 20.8 & 0.4 & 474 (118.5)\tabularnewline
\bottomrule
\end{longtable}

Table 2: Common understandings in state-level management documents. Note
that additional evidence from our 13-month empirical study only
addresses questions that require less than one year of data (i.e.
intra-annual patterns in roost structure). Statements not addressed with
our empirical data are coloured grey. An extended version of this table
with details on study results is provided in

Appendix S2 in the Supporting Information.\selectlanguage{english}
\begin{longtable}[]{@{}llllll@{}}
\toprule
\begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Understandings}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Referenced by}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Empirical evidence}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Empirical evidence}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Additional evidence (this study)}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Additional evidence (this study)}\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Support}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Contradict}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Support}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Contradict}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Use of area:\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Some areas of permanent camps are more consistently occupied ('core
areas') than others\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); EcoLogical (2014)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005); Richards (2002); Nelson (1965b)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Figure 2\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
`Core areas' are more densely occupied than `peripheral areas'\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Nelson (1965b); Welbergen (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Figure 3; Figure 4; Appendix S2\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Roost area fluctuates with total abundance\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); EcoLogical (2014)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005); Pallin (2000); Larsen \emph{et al.} (2002)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Figure 5\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Flying-foxes adjust the location of `core areas' through time\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Hall (2002) \& Pallin (2000)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Areas outside of the 'core area' are used by more transient
animals\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Spatial segregation of species:}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Species share roosts sites, but segregate spatially within\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Commonwealth of Australia (2017a)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005); Ratcliffe (1932); Parsons \emph{et al.} (2010); Nelson
(1965b); Klose \emph{et al.} (2009)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Parsons \emph{et al.} (2010); Markus (2002)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Figure 6; Appendix S3\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Large influxes of species into roosts (especially little red
flying-foxes) can displace other species\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Birt and Markus (1999)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Appendix S3\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Species roost at different heights\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Geolink (2010)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005); Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Figure 7\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Indirect competition favours black flying-foxes over grey-headed
flying-foxes\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Commonwealth of Australia (2017a); EcoLogical (2014)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Ratcliffe (1931)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Markus (2002); Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Demographic/social structure:}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
The majority of roost trees are occupied by mixed groups of adults, with
territories comprised of a single male and one or more females and their
dependent young\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); State of Queensland Department of Environment and
Science (2020)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005); Puddicombe (1981); Nelson (1965b); Nelson (1965a);
Markus and Blackshaw (2002); Markus (2002); Eby \emph{et al.} (1999);
McWilliam (1984); Connell (2003)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005); Nelson (1965b); Nelson (1965a)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Figure 8\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Dominant individuals (defined as reproducing males and females) occupy
the centre of roosts and subdominant individuals (defined as
non-reproducing males and females) the outer area\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
State of Queensland Department of Environment and Science (2020)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Nelson (1965b); Welbergen (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Puddicombe (1981); Markus and Blackshaw (2002)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Figure 8; Appendix S4\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Individuals at the periphery of groups act as `guards'\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
State of Queensland Department of Environment and Science (2020)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Nelson (1965b); Klose \emph{et al.} (2009)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Juveniles wean and leave their mothers from January and form groups on
the edge of their existing roost or at another site\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
State of Queensland Department of Environment and Science (2020)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005); Nelson (1965b); Nelson (1965a); Markus and Blackshaw
(2002); Eby \emph{et al.} (1999); Connell (2003)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
The roosting positions of individual males are highly consistent and
animals return to the same branch of a tree over many weeks or
months\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005); Markus and Blackshaw (2002); Markus (2002)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tidemann and Nelson (2004); Roberts \emph{et al.} (2012b); Parsons,
Robson and Shilton (2011)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Roost abundance/occupancy:}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Individual roosts have distinguishable seasonal patterns of abundance
and occupation.\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\emph{Abundance:} Commonwealth of Australia (2017a); \emph{Occupation:}
State of Queensland Department of Environment and Science (2020)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\emph{Abundance} Westcott \emph{et al.} (2018); Welbergen (2005); Tait
\emph{et al.} (2014); Parry-Jones and Augee (2001); Parry-Jones and
Augee (1992); Nelson (1965b); Nelson (1965a); Meade \emph{et al.} (2019)
\emph{Occupation} Welbergen (2005); Vardon and Tidemann (1999);
Parry-Jones and Augee (1992); Parry-Jones (1985); Nelson (1965b) \&
Nelson (1965a); Nelson (1965b) \& Nelson (1965a); Klose \emph{et al.}
(2009); Puddicombe (1981); Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\emph{Abundance} Shilton \emph{et al.} (2008); Richards (2002); Roberts
(2005) \emph{Occupation} Van der Ree \emph{et al.} (2006); Richards
(2002); Puddicombe (1981); Shilton \emph{et al.} (2008)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Figure 9\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Intra- and inter-annual variations in abundance can be extreme\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Commonwealth of Australia (2017a)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Westcott and McKeown (2004); Tait \emph{et al.} (2014); Welbergen
(2008); Welbergen (2005); Vardon and Tidemann (1999); Ratcliffe (1931);
Ratcliffe (1932); Eby (1991); Eby and Palmer (1991); Van der Ree
\emph{et al.} (2006); Eby and Lunney (2002a); Roberts \emph{et al.}
(2012a); Richards (2002); Parry-Jones and Augee (2001); Parry-Jones and
Augee (1992); Pallin (2000); Meade \emph{et al.} (2019); Loughland
(1998); Giles \emph{et al.} (2016); Forsyth, Scroggie and
McDonald-Madden (2006); Eby \emph{et al.} (1999); Lunney and Moon
(1997)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Figure 9\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Roost abundance peaks in March\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
State of Queensland Department of Environment and Science (2020)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Van der Ree \emph{et al.} (2006); Tait \emph{et al.} (2014); Meade
\emph{et al.} (2019); Eby (1991); Eby and Palmer (1991); Nelson
(1965a)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Westcott \emph{et al.} (2018); Welbergen (2005); Vardon and Tidemann
(1999); Vardon \emph{et al.} (2001); Roberts \emph{et al.} (2012a);
Richards (2002);Parry-Jones and Augee (2001); Parry-Jones and Augee
(1992); Pallin (2000) (citing personal communication with M. Beck);
Nelson (1965b); Nelson (1965a)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Figure 9; Appendix S2\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Consistent (inter-annual) patterns in abundance and use are more
commonly observed in roosts located in 1) extensive areas of rainforest,
and 2) urban areas\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
SEQ Catchments (2012); Commonwealth of Australia (2017a)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\emph{Extensive rainforest} Parry-Jones (1985) \emph{Urban areas} Tait
\emph{et al.} (2014); Welbergen (2005); Van der Ree \emph{et al.}
(2006); Richards (2002); Williams \emph{et al.} (2006); Parry-Jones and
Augee (2001); Parry-Jones and Augee (1992)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Habitat preferences:}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Stager and Hall (1983)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
The habitat patch must be at least 1ha in size but be large enough to
accommodate and sustain large numbers of flying-foxes. For a small roost
(10,000 bats) the area needed is approximately 3ha and for a large roost
(50,000) the area needed is 10ha\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); State of NSW and Department of Planning Industry
and Environment (2019) as per State of NSW and Office of Environment and
Heritage (2018); EcoLogical (2014)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Pallin (2000); Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Flying-foxes prefer complex vegetation structure (upper, mid- and
understorey layers)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); State of NSW and Department of Planning Industry
and Environment (2019) as per State of NSW and Office of Environment and
Heritage (2018)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Pallin (2000) (citing report by Buchanan)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Flying-foxes prefer dense vegetation\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Flying-foxes prefer a dense understory\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Flying-foxes prefer a closed canopy at least 3-5m high\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); State of NSW and Department of Planning Industry
and Environment (2019) as per State of NSW and Office of Environment and
Heritage (2018); EcoLogical (2014)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Tidemann \emph{et al.} (1999); Tidemann (1999); Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
The structure of roost-wide vegetation is more important than the
characteristics of individual roost trees (e.g. species, canopy
cover)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Palmer and Woinarski (1999); Pallin (2000); Vardon \emph{et al.} (2001);
Tidemann \emph{et al.} (1999); Vardon and Tidemann (1999); Hall and
Richards (2000); Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Flying-foxes prefer level topography (\textless{}5\selectlanguage{ngerman}° incline)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); State of NSW and Department of Planning Industry
and Environment (2019) as per State of NSW and Office of Environment and
Heritage (2018)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Flying-foxes prefer to roost within 50 km of the coastline or at an
elevation \textless{}65 m above sea level\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); State of NSW and Department of Planning Industry
and Environment (2019) as per State of NSW and Office of Environment and
Heritage (2018)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Hall and Richards (2000); Roberts (2005)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Ratcliffe (1931); Ratcliffe (1932)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Roost macroclimate:}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
The mid-storey vegetation within roosts is critical for maintaining a
cool, humid and sheltered environment that is stable against the outside
environment\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); State of NSW and Department of Planning Industry
and Environment (2019) as per State of NSW and Office of Environment and
Heritage (2018)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Loughland (1998)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Snoyman and Brown (2010)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\textbf{Negative impacts from flying-foxes:}\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Impacts sustained over several years of flying-fox occupancy can lead to
damage and/or death of individual roost trees\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); State of Queensland Department of Environment and
Science (2020)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Welbergen (2005); Richards (2002); Pallin (2000); McWilliam (1984); Hall
(2002)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Some tree species are more resilient to damage by flying-fox roosting
than others\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
In small remnant patches, the process of opening the canopy (from tree
damage by roosting) will increase the impact of invasive weeds\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); State of Queensland Department of Environment and
Science (2020)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Pallin (2000); McWilliam (1984); Hall (2002)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Where sufficient roosting space is available, flying-foxes shift their
roosting areas, which lessens their damage to vegetation over time\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
SEQ Catchments (2012); EcoLogical (2014)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Pallin (2000); Hall (2002)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

\par\null

\textbf{Figure Legends}

Figure 1: Map of roost sites included in the study. Grey shading
indicates urban land cover of dense human habitation (as per Schneider,
Friedl \& Potere 2009) and grey circles are locations of flying-fox
roosts. Circles show 45km foraging radii from roost study sites (as per
Giles \emph{et al.} 2018). GIS land-cover data was downloaded from
Natural Earth (2020) and flying-fox roost locations obtained from the
National Flying-Fox Monitoring Program (2017).

Figure 2: Occupancy of subplots across survey period, for surveys when
at least one bat was present. A) shows the total number of bats per
subplot, and B) shows the proportion of surveys the subplot was
occupied. Facets/colour indicates separate roost sites. ``C'' indicates
roosts that have features of contemporary roost types (see Table 1).
Note that construction works at the `Avondale' roost during the survey
caused the bats to shift their roosting location, such that only one
subplot was utilised thereafter.

Figure 3: Occupancy of subplots in `core' and `peripheral' areas, shown
by average total number of bats per occupied subplot across the survey
period. Data is filtered to show numbers of bats when subplots were
occupied (i.e. unoccupied subplots are removed). `Core' subplots were
identified as those occupied in at least 80\% of surveys (when bats
present at the roost), and `peripheral' subplots as those occupied less
than 80\% of the time. A) Shows areas split by roost site (facet and
colour), and B) shows all roosts combined. Area displayed in subplot has
been cropped to remove extreme outliers. ``C'' indicates roosts that
have features of contemporary roost types (see Table 1).

Figure 4: Distance from roost centre and occupancy of bats, shown by the
average total number of bats per occupied subplot during the survey
period. Data is filtered to show numbers of bats when trees are occupied
(i.e. unoccupied subplots are removed). Roost centre is calculated for
each survey as the centroid of the roost area at the time of the survey.
Distance from the centre is calculated as the mean distance of trees in
each subplot from this centroid, scaled by the maximum observed distance
value per session. A) shows values per species (line type) split by
roost (facets); and B) shows species and roost combined. Trend line is
by loess fit (local polynomial regression fit) with standard error bands
(grey shading). ``C'' indicates roosts that have features of
contemporary roost types (see Table 1).

Figure 5: Relationship between total roost abundance (x axis) and total
roost area (y axis) for each roost site. A) shows relationship split by
roost (facets) and B) shows relationship with roosts combined. Trend
line is by loess fit (local polynomial regression fit) with standard
error bands (grey shading). Note that trend lines could not be fitted
for all sites and are omitted. ``C'' indicates roosts that have features
of contemporary roost types (see Table 1).

Figure 6: Co-occupation of subplots (A) and individual trees (B) by
species. Total subplots/total trees observed are shown in text labels
and include subplots/trees across sessions where every bat species in
the species comparison were present. (e.g. for the black and grey-headed
flying-fox comparison, only sessions where both black and grey-headed
flying-fox were present were included in the subplot/tree tally). `BFF'
refers to black-flying-fox, `GHFF' grey-headed flying-fox and `LRFF'
little red flying-fox. Confidence intervals are binomial, calculated
with a Wilson test.

Figure 7: Difference in roosting height per species, over time. Fill
shows average roosting height range per species (minimum height to
maximum height). Fill boundaries (minimum and maximum curves) are by
loess fit (local polynomial regression fit). A) shows relationship split
by roost (facets) and B) shows relationship with roosts combined. In A),
dashed line represents the average canopy height per site; for roost
sites where species occupy distinctly different areas (`Clunes' and
`Lismore'), canopy height is split by areas the species predominantly
occupy. ``C'' indicates roosts that have features of contemporary roost
types (see Table 1). Note that height data are taken from the tree
subset only (up to N=60 per roost site), and that trend lines could not
be fitted for all site by species combinations and are omitted.

Figure 8: Proportion of male bats per occupied tree versus distance of
tree from the roost centre, scaled by the maximum distance value
observed per session. A) shows values per species (row facet) split by
roost (column facet); B) shows combined species value pooled by roost.
Trend line is by loess fit (local polynomial regression fit) with
standard error bands (grey shading). `BFF' refers to black-flying-fox,
`GHFF' grey-headed flying-fox and `LRFF' little red flying-fox. ``C''
indicates roosts that have features of contemporary roost types (see
Table 1).

Figure 9: Different scales of bat abundance measures through time. A)
shows total roost abundance; B) shows total roost area; C) shows the
proportion of occupied trees per subplot; D) shows the total abundance
of occupied subplots. Total roost abundance is measured by an index
score of abundance: 1 = 1-499 bats; 2 = 500-2,499 bats; 3 = 2,500 -
4,999 bats; 4 = 5,000 - 9,999 bats; 5 = 10,000 - 15,999 bats; 6 = 16,000
- 49,999 bats; and 7 = \textgreater{} 50,000 bats.

\par\null

\subsection*{Figures}

{\label{595705}}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Lunn-et-al-Figure-1/Lunn-et-al-Figure-1}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Lunn-et-al-Figure-2/Lunn-et-al-Figure-2}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Lunn-et-al-Figure-3/Lunn-et-al-Figure-3}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Lunn-et-al-Figure-4/Lunn-et-al-Figure-4}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Lunn-et-al-Figure-5/Lunn-et-al-Figure-5}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Lunn-et-al-Figure-6/Lunn-et-al-Figure-6}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Lunn-et-al-Figure-7/Lunn-et-al-Figure-7}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Lunn-et-al-Figure-8/Lunn-et-al-Figure-8}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/Lunn-et-al-Figure-9/Lunn-et-al-Figure-9}

\end{center}
\end{figure}

\selectlanguage{english}
\FloatBarrier
\end{document}

