Saving species beyond the protected area fence: Threats must be managed across multiple land tenure types to secure Australia's endangered species

Abstract

The main effort to secure threatened species globally is to set aside land and sea for their conservation via governance arrangements such as protected areas. But not even the biggest protected area estate will cover enough area to halt most species declines. Consequently, there is a need for assessments of how species habitats are distributed across the tenure landscape, to guide policy and conservation opportunities. Using Australia as a case study, we assess the relationship between land tenure coverage and the distributions of nationally listed threatened species. We discover that on average, nearly half (48%) of Australian threatened species' distributions occur on privately owned (freehold) lands, despite this tenure covering only 29% of the continent. In contrast, leasehold lands, which cover 38% of Australia, overlap with only 6% of species' distributions while protected area lands (which cover 20%) have an average of 35% of species' distributions. We found the majority (75%; n = 1199) of species occur across multiple land tenures, and those species that are confined to a single tenure were mostly on freehold lands (13%; n = 201) and protected areas (9%; n = 139). Our findings display the opportunity to reverse the current trend of species decline with increased coordination of threat management across land tenures. We quantify the overlap of threatened species with land tenure across Australia. On average, half of threatened species' distributions occur on freehold lands and three‐quarter of the species occur across multiple land tenures.

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CONTRIBUTED PAPER
Saving species beyond the protected area fence: Threats
must be managed across multiple land tenure types to
secure Australia's endangered species
Stephen G. Kearney
1
| Josie Carwardine
2
| April E. Reside
3
|
Vanessa M. Adams
4
| Rebecca Nelson
5
| Anthea Coggan
2
|
Rebecca Spindler
6
| James E. M. Watson
1
1
School of Earth and Environmental
Sciences, University of Queensland,
Brisbane, Queensland, Australia
2
CSIRO Land and Water, Brisbane,
Queensland, Australia
3
School of Biological Science, University
of Queensland, Brisbane, Queensland,
Australia
4
School of Technology, Environments and
Design, University of Tasmania, Hobart,
Tasmania, Australia
5
University of Melbourne Law School,
Melbourne, Victoria, Australia
6
Bush Heritage Australia, Melbourne,
Victoria, Australia
Correspondence
Stephen G. Kearney, School of Earth and
Environmental Sciences, Steele Building,
University of Queensland, St Lucia, QLD
4072, Australia.
Email: stephen.kearney@uq.edu.au
Abstract
The main effort to secure threatened species globally is to set aside land and
sea for their conservation via governance arrangements such as protected
areas. But not even the biggest protected area estate will cover enough area to
halt most species declines. Consequently, there is a need for assessments of
how species habitats are distributed across the tenure landscape, to guide pol-
icy and conservation opportunities. Using Australia as a case study, we assess
the relationship between land tenure coverage and the distributions of nation-
ally listed threatened species. We discover that on average, nearly half (48%) of
Australian threatened species' distributions occur on privately owned (free-
hold) lands, despite this tenure covering only 29% of the continent. In contrast,
leasehold lands, which cover 38% of Australia, overlap with only 6% of species'
distributions while protected area lands (which cover 20%) have an average of
35% of species' distributions. We found the majority (75%; n=1199) of species
occur across multiple land tenures, and those species that are confined to a sin-
gle tenure were mostly on freehold lands (13%; n=201) and protected areas
(9%; n=139). Our findings display the opportunity to reverse the current
trend of species decline with increased coordination of threat management
across land tenures.
KEYWORDS
conservation, extinction, habitat destruction, invasive species, national park, threat
management, threatened species
1|INTRODUCTION
Human activities are causing a decline in native species
at a rate and scale unprecedented in recent history
(Butchart et al., 2010; Maxwell et al., 2016). In response,
the global community is working together to halt this
decline (United Nations, 1992), an effort that is recog-
nized as one of the greatest challenges for humanity
Received: 20 April 2021 Revised: 28 October 2021 Accepted: 9 December 2021
DOI: 10.1111/csp2.617
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided
the original work is properly cited.
© 2022 The Authors. Conservation Science and Practice published by Wiley Periodicals LLC on behalf of Society for Conservation Biology.
Conservation Science and Practice. 2022;4:e617. wileyonlinelibrary.com/journal/csp2 1of15
https://doi.org/10.1111/csp2.617

(IPBES, 2019). To date, considerable and increasing
action is being conducted across Earth (Tittensor
et al., 2014), the cornerstone of which has been the desig-
nation of protected areas (Watson et al., 2014), which
now cover 15% of the global land surface (Maxwell
et al., 2020). However, collective conservation efforts
have so far had limited success in slowing the rate of
decline in these threatened species (Maxwell et al., 2020).
While there are localized success stories of efforts to
arrest decline of a limited number of threatened species
(Bolam et al., 2021; Butchart et al., 2006), numerous
examples highlight site-based conservation efforts failing
to halt the decline of many species (Craigie et al., 2010;
Wayne et al., 2017; Woinarski et al., 2017).
The vital importance of the 15% of terrestrial earth
within the protected area estate for threatened species is
well established (Watson et al., 2014; Woodley
et al., 2019); and while there are global discussions
around the fact that land area under protection should be
far larger (Dinerstein et al., 2017; Wilson, 2016), at pre-
sent the management of the other 85% of Earth will likely
have far greater implications for threatened species out-
comes (Maron et al., 2018; Watson et al., 2021). The non-
protected landscape is a complex matrix of ownership
and use regimes, with some areas consistent with, or at
least not impeding, positive conservation outcomes
(e.g., some military lands; Stein et al., 2008) while others
are far less compatible (e.g., freehold land under inten-
sive agriculture; Dudley & Alexander, 2017). Historic and
current ownership and use regimes have significant
implications for the legal and policy approaches taken for
threatened species conservation and threat management
(Fischer & Lindenmayer, 2007). Hence, mapping and
understanding the tenure landscape is critical for guiding
policy, understanding collaborative pathways to effective
management, and maximizing conservation opportuni-
ties to respond to the species extinction crisis.
Land tenure is the legal regime under which land is
owned, accessed, and used (FAO, 2002). In some regions,
it has been found that land tenure impacts environmen-
tal outcomes (Robinson et al., 2014; Woinarski
et al., 2013) and tenure can influence (or at least is corre-
lated with) threatening processes to biodiversity
(Evans, 2016; Fensham et al., 1998). This is consistent
with the foundational concept of common law systems
that owning land carries the right to exploit the natural
resources on that land, though that right is regulated via
legislation (Bates, 2019). Important mechanisms available
to manage the processes threatening species vary across
land tenure, with specific laws only applicable on some
land tenure types and jurisdiction (e.g., land clearing
laws in Queensland, Australia; Hamman, 2019). Other
legal mechanisms theoretically apply regardless of land
tenure type but relate to activities that are more likely to
occur on private land (e.g., liability for pollution depen-
dent on the degree of environmental harm caused;
Bates, 2019). Furthermore, it is increasingly established
that achieving effective environmental management
often relies on coordination across the tenure landscape
(McCune & Morrison, 2020). Therefore, understanding
the distribution of threatened species across an entire
tenure landscape may provide critical insights into the
law and policy options as well as opportunities for land-
holder contributions to halt the on-going decline of
threatened species.
Here, using Australia as a case study, we integrate a
current land tenure map with distribution maps of
nationally listed threatened species to assess how these
species are distributed across the entire tenure landscape.
Australia is one of only 17 megadiverse countries
(Mittermeier et al., 1997) and is a continent in the grips
of an extinction crisis, having already lost 10% of the ter-
restrial mammal fauna (Woinarski et al., 2015) and with
an additional 1797 species at high risk of extinction in
the near-term (Commonwealth of Australia, 2017a,
2017b). The decline of the continent's threatened species
is driven by an array of threatening processes, including
habitat loss, inappropriate fire regimes and invasive spe-
cies (Kearney et al., 2019). Although our knowledge of
the distribution of species threatened with extinction
across parts of the Australian landscape has improved in
recent years (e.g., protected areas: Watson et al., 2011; cit-
ies and urban areas: Ives et al., 2016; Soanes &
Lentini, 2019; Indigenous owned and managed lands:
Renwick et al., 2017), a comprehensive understanding of
how tenure overlays with threatened species is lacking. A
quantification of species across different tenures is
needed to formulate appropriate conservation actions
that can be implemented in line with relevant legal
frameworks; and therefore, critical if Australia is to
improve its response to the extinction crisis. We explore
this concept with a particular focus on the variation
across taxonomic groups, range size and extinction risk
categories as well as the importance of single and multi-
tenure approaches to conservation efforts.
2|METHODS
2.1 |Land tenure
In Australia, land tenure is the legal regime within which
land is owned and occupied (Walker & Dale, 2013). Ten-
ure defines the responsibilities or required actions of
landholders, as well as circumscribing the range of activi-
ties or opportunities available to landholders from the
2of15 KEARNEY ET AL.

land (Ostrom & Schlager, 1996). Tenure legally mandates
the permissible uses of land, constraints to such uses and
specific property rights, as well as influencing the objec-
tives of management (Schlager & Ostrom, 1992; van
Etten, 2013). For present purposes, there are two high
level categories of land tenure in Australia: freehold and
Crown land (Geoscience Australia, 2018). Freehold land
is that which is owned by the landholder in perpetuity.
Crown land includes numerous land tenure types
(e.g., leasehold; reserves; unallocated Crown land) owned
by the state (the Crown) and leased or licensed to allow
particular land uses (Australian Trade and Investment
Commission, 2020). Relevant legislative acts govern the
use of Crown land for each state and territory. For exam-
ple, leasehold or pastoral lands acts govern the use of pas-
toral leasehold lands (e.g., New South Wales: Crown
Land Management Act 2016) while environmental legis-
lation govern the uses and management of the conserva-
tion estates (e.g., Queensland: Nature Conservation
Act 1992).
At the time of undertaking this analysis (November
2018), there was no single comprehensive and up-to-date
map of land tenure in Australia. As such, we created a
tenure map by combining multiple datasets (Figure 1a).
The basis of this map was Public Sector Mapping
Agency's (PSMA) CadLite and Land Tenure datasets
(PSMA Australia, 2018) which, while the most up-to-
date, are not comprehensive as they contain areas of no
data across parts of Western Australia, New South Wales,
and Victoria. Where this spatial dataset was incomplete,
we filled those data gaps with information from Geosci-
ence Australia's 1993 Land Tenure map (Geoscience
Australia, 1993), the only comprehensive tenure dataset
for Australia that we are aware of. Finally, we
incorporated protected area spatial data from the Collab-
orative Australian Protected Area Database (CAPAD;
Commonwealth of Australia, 2017c). CAPAD has incom-
plete coverage of privately protected areas (PPAs;
Fitzsimons, 2015; Ivanova & Cook, 2020); as such, PPAs
that are in place in some jurisdictions are missed, which
could influence the results of our analysis. However, as
noted by Fitzsimons (2015), CAPAD contains a signifi-
cant majority of the total area within PPAs including
those of major Australian nongovernment organizations
(e.g., Bush Heritage Australia, Australian Wildlife Con-
servancy), so the PPAs not contained within CAPAD
likely only account for a very small proportion of the
Australian landmass and therefore threatened species
distributions.
The PSMA dataset contains 23 standardized land ten-
ure types (one freehold and 22 Crown land), the Geosci-
ence Australia land tenure dataset has 16 land tenure
types (two freehold and 14 Crown land) and the CAPAD
dataset has seven protected area management categories.
For this analysis, we combined tenure categories within
and across data sets to produce seven consistent and
mutually exclusive tenure categories relevant to our
analysis. These were: (1) Freehold; (2) Leasehold; (3)
FIGURE 1 (a) The Australian land tenure map produced for this analysis and (b) threatened species distribution density across
Australia
KEARNEY ET AL.3of15

Protected area; (4) Forestry; (5) Defense; (6) Unallocated
Crown land; and (7) Other tenures (Figure 1a; Table 1).
Categories (2)–(7) are all Crown land, and are differenti-
ated according to whether a private party controls them
under a lease (primarily pastoral leases; Category 2), or a
state entity for a particular purpose (Categories 3, 4, 5, and
7) or for no designated purpose (Category 6). The category
“Other tenures”include the land tenure types from the
PSMA and Geoscience Australia datasets that did not fit
into any of the other categories (e.g., Other reserve; Water
reserve; Mining reserve; Stock route; Transportation
reserve; Other infrastructure reserve). These tenures make
up 2.3% of the continent.
Land tenure in Australia can be complicated with
multiple overlapping tenure arrangements. Often one
parcel of land will have a number of tenure interests
associated with it (Maguire, 2009). Notably native title
can be exclusive or sit alongside other tenures (with the
exception of freehold land that extinguishes native title).
The Native Title Act 1993 (Cth) provides a statutory pro-
cess to protect the rights of native title holders (Native
Title Act 1993, s. 3). Further, minerals and petroleum
belong to the Crown regardless of tenure and mineral
rights are governed under individual state and territory
acts (e.g., Northern Territory: Mineral Titles Act 2010).
Thus, for our analysis, we focus on the primary tenure,
rather than any multiple overlapping titles which could
also affect which conservation actions can take place as
well as who to engage with (Adams & Moon, 2013).
The focus on primary tenure was particularly promi-
nent in our treatment of protected areas (tenure Category
3). Although some protected areas have underlying ten-
ure categories, such as freehold lands leased back to gov-
ernment for protection (e.g., some parts of Kakadu
National Park; Director of National Parks, 2016) and con-
servation covenants over private land (Fitzsimons, 2015),
we were primarily interested in the protected area designa-
tion. As such, if an area was within the official protected
area network of Australia (as recognized in CAPAD), the
protected area superseded the underlying tenure type
because to be named a protected area, the primary goal
must be nature conservation (Dudley, 2008). For these
areas, the tenure underlying the protected area designation
were not considered in this analysis. It is also important to
note that there are many legal mechanisms that pursue
nature conservation goals on certain lands, which are not
included in the CAPAD database. For example, legal land
use instruments are used to designate areas of environmen-
tal significance (Nelson, 2019) or conservation-oriented
legal agreements between municipal councils and land-
owners (Harwood et al., 2016). These differ between states,
are not aggregated in centralized databases even at the state
level and lie outside the scope of the present analysis.
Further details on the land tenure categories reported
on here and the corresponding PSMA and Geoscience
Australia tenure categories and CAPAD protected manage-
ment categories are provided in Supplementary Table 1.
2.2 |Species data
We focus on terrestrial and freshwater species and subspe-
cies (hereafter, taxa) listed as threatened (Vulnerable,
Endangered and Critically Endangered) under Australia's
Environment Protection and Biodiversity Conservation Act
1999 (EPBC Act). This includes taxa from vertebrate
(Classes: Mammalia,Aves,Amphibia,Reptilia,Chondri-
chthyes,Actinopterygii,Sarcopterygii), invertebrate (Classes:
Insecta,Bivalvia,Malacostraca,Gastropod,Arachnid,
Onychophora,Remipedia,Oligochaeta) and plant (Classes:
Bryopsida,Cycadatae,Liliopsida,Lycopodopiatae,Mag-
noliopsida,Pinatae,Polypodiatae,Psilotatae,Streptophyta)
TABLE 1 The seven land tenure categories used in this analysis.
Tenure category name Details
Percent of Australian
terrestrial area
Freehold Lands owned privately and in perpetuity 28.7
Leasehold Lands owned by the state (the Crown) and leased for various uses 38.2
Protected area Lands recognized, dedicated, and managed primarily for the conservation
of nature (IUCN Categories: I–VI)
19.7
Forestry Lands that are used or reserved for timber harvest and production
(multiple use forest; state forest; timber reserve; other forestry reserve)
1.4
Defense Lands reserved for use by the armed forces. 0.1
Unallocated Crown land Vacant, unallocated, unreserved, or other Crown land 9.6
Other tenure Lands that do not fall within the other categories used in this analysis
(e.g., Other reserve; Water reserve; Mining reserve; Stock route;
Transportation reserve; Other infrastructure reserve)
2.3
4of15 KEARNEY ET AL.

taxonomic groups. At the time of writing, 1797 (1342 plant,
455 animal) taxa were listedasthreatenedunderthe
EPBC Act.
Distribution data for these taxa are made available by
the Australian Government through the Species of
National Environmental Significance (SNES) database
(Commonwealth of Australia, 2018). Taxa distribution
data are classified into three categories based on the
degree of certainty that a taxon is present: “known to
occur”;“likely to occur”and “may occur.”Following pre-
vious studies (Ives et al., 2016; Renwick et al., 2017; Ward
et al., 2019; Watson et al., 2011), we included only the
two categories of higher certainty (known and likely to
occur) and combined them into a single category for this
analysis (Figure 1b).
2.3 |Analysis
We quantified the overlap of each threatened taxon with
each tenure category by intersecting taxa distribution
maps with the Australian land tenure map. In the spatial
overlay analysis, 1602 taxa were included. Spatial ana-
lyses were undertaken in ArcMap 10.6 (ESRI, 2011). We
explored the variation across taxonomic groups, range
sizes and extinction risk categories. Although, we
acknowledge that taxa range size and extinction risk are
related, as range size thresholds can be used to assign
extinction risk category. However, it is still valuable to
explore both factors as not all small range taxa have
higher extinction risks (e.g., Lancelin Island skink
[Ctenotus lancelini]; SNES known and likely distribution
0.2 km
2
; EPBC Act status: Vulnerable) and not all
higher extinction risk taxa have small ranges (e.g., plains-
wanderer [Pedionomus torquatus]; SNES known and
likely distribution: >175,000 km
2
; EPBC Act status: Criti-
cally Endangered). We also quantify the number of ten-
ures that a taxon is distributed across to understand the
importance of single and multi-tenure approaches to
threatened species conservation.
3|RESULTS
3.1 |Threatened taxa distribution across
each tenure category
The three dominant tenure types across Australia were
leasehold (38%), freehold (29%), and protected areas
(20%) (Table 1), although their importance for threatened
taxa varied considerably (Figure 2). The greatest propor-
tion of Australian threatened taxa distributions occurred
on freehold land, accounting for an average of nearly half
of all threatened taxa distributions (average and median
taxa overlap: 48%; Table 2, Figure 3). Protected areas
facilitated an average of 35% of taxa distributions
(median: 24%; Table 2, Figure 3), while leasehold lands
contained an average of just 6% of threatened taxa
FIGURE 2 The distribution (%) of threatened taxa within each taxonomic group across each broad land tenure category in Australia.
Each vertical bar represents one Environment Protection and Biodiversity Conservation Act listed taxa. (amphibians: n=29; fish: n=37;
reptiles: n=45; invertebrates: n=54; birds: n=76; mammals: n=99; plants: n=1262)
KEARNEY ET AL.5of15

distributions (median: 0%; Table 2). In contrast, a similar
proportion of threatened taxa distributions (5.9%)
occurred on forestry reserves while only occupying 1.4%
of the Australian landmass (Table 2).
3.2 |Taxonomic groups across tenures
The majority of taxonomic groups had the greatest
overlap with freehold lands (plants 51%, reptiles: 49%,
invertebrates 47%, birds: 41%, fish: 37%; Figure 3;
Supplementary Table 3), while protected areas covered
the greatest proportion of average taxa distributions for
amphibians (49%) and mammals (48%) (Figure 3, Supple-
mentary Table 3).
Mammals, reptiles, and birds had the greatest overlap
with leasehold lands (15.6, 15.5, and 14.8%, respectively),
while amphibians and fish have the least (2.6 and 3.5%,
respectively). Amphibian and invertebrates have the
greatest overlap with forestry reserves (13.6 and 10.9%,
respectively), while reptiles and birds had the least (1.9
and 2.7%; Figure 3, Supplementary Table 3).
TABLE 2 The average and median species distribution overlap with each land tenure category (n=1602). The number of species in
each overlap category is also shown.
Freehold
Protected
areas Leasehold Forestry Defense
Unallocated
land
Other
tenure
Average (%) species overlap with
tenure
48.0 35.0 5.8 5.9 0.3 2.4 2.6
Median (%) species overlap with
tenure
48.0 23.9 0.0 0.0 0.0 0.0 0.4
The number of species (and % of total) that fall into the range overlap categories for each tenure
>95% 201 (13%) 139 (8%) 14 (1%) 6 (<1%) 1 (<1%) 5 (<1%) 0 (0%)
50–94.9% 576 (36%) 331 (21%) 58 (4%) 35 (2%) 0 (0%) 12 (<1%) 10 (<1%)
5–49.9% 559 (35%) 783 (49%) 166 (10%) 341 (21%) 24 (2%) 117 (7%) 182 (11%)
<5% 266 (17%) 349 (22%) 1364 (85%) 1220 (76%) 1577 (98%) 1468 (92%) 1410 (88%)
FIGURE 3 The average distribution overlap (%) of each species group (taxonomic group, extinction risk category, and range size
category) with each land tenure. (Amphibians: n=29; fish: n=37; reptiles: n=45; invertebrates: n=54; birds: n=76; mammals: n=99;
plants: n=1262; Vulnerable: n=729; Endangered: n=660; Critically Endangered: n=213; >10,000 km
2
:n=205; 1000–10,000 km
2
:
n=364; 100–1000 km
2
:n=566; 10–100 km
2
:n=349; <10 km
2
:n=118)
6of15 KEARNEY ET AL.

3.3 |Land tenure and extinction risk
category
On freehold lands, average taxa overlap decreased with
decreasing extinction risk, with Critically Endangered and
Endangered taxa having over half of their distributions
(54 and 53%, respectively) and Vulnerable taxa having 42%
on this tenure type (Figure 3, Supplementary Table 3). Vul-
nerable taxa had the greatest overlap with almost all other
tenure types, including protected areas, which accounted for
an average of 38% of these taxa distributions (EN: 32%; CR:
33%) and leasehold lands, accounting for 9% of their distribu-
tion (CR: 3% and EN: 4%; Figure 3, Supplementary Table 3).
3.4 |Land tenure and range size
Freehold lands accounted for the greatest proportion of dis-
tributions for taxa within each range size category except
for narrowly distributed species (<10 km
2
distribution;
Figure 3). These narrowly distributed taxa had, on average,
44.9% within protected areas, followed closely by 44.5% on
freehold lands. The average taxa distribution overlap with
protected areas decreased with increasing range size
(<10 km
2
: 44.9%; >10,000 km
2
: 25.0%; Supplementary
Table 3), while average taxa overlap with leasehold lands
increased with increasing range size (<10 km
2
:2.2%;
>10,000 km
2
: 18%; Supplementary Table 3).
3.5 |Single- and multi-tenure species
The majority of Australian threatened taxa occurred on
multiple land tenure categories (Figure 4a). When tenure
categories making up less than 5% of a taxon's distribution
were disregarded, 75% of taxa (n=1199) occurred across
two or more tenure categories, with 38% (n=609) occur-
ring across three or more (Figure 4a). One-quarter of taxa
(n=366, 22.8%) had the vast majority (>95%) of their
range on a single tenure type. Over half of these (n=201,
12.5% of all taxa) occurred almost solely on freehold lands,
while 139 taxa (8.7% of all taxa) occurred almost solely on
protected lands and 14 (1% of all taxa) on leasehold lands
(Table 2). Of the combinations (pairs) of tenure categories,
two-thirds (64%) of species (n=1024) had >5% of their
distributions on both protected areas and freehold lands
(Figure 4b). The combination of tenure categories that had
the next highest taxa count was forestry and freehold
lands, with 21% of species (n=344) having >5% of their
distributions on both tenure types (Figure 4b). The overlap
of each EPBC Act listed threatened taxa with each land
tenure category included in this analysis is provided in
Supplementary Table 4.
4|DISCUSSION
Our study revealed that freehold lands and protected
areas covered the largest proportion of species distribu-
tions, with freehold lands being particularly important
for those species at greatest risk of extinction
(i.e., Endangered and Critically Endangered) and protec-
ted areas for narrow-range species. The majority of spe-
cies (75%) occurred across multiple land tenures,
highlighting the critical need for multi-tenure efforts in
Australia's response to the species extinction crisis. This
effort in its nature needs to be inclusive and account for
the values and preferences of local land managers,
FIGURE 4 (a) The number of taxa that are distributed across one or more land tenure types, when tenures making up <5% of their
distribution are removed; and (b) the number of species that had ≥5% of their distribution overlapping with each land tenure (colored cells)
and the paired combination of each land tenure (intersecting cells)
KEARNEY ET AL.7of15

landowners and custodians, creating environmental co-
benefits and job opportunities. Supporting Indigenous
leadership in decision-making and culturally appropriate
management will be critically important for species and
places with Indigenous interests (Ens et al., 2016;
Moorcroft et al., 2012), which overlap with all of the land
tenure categories considered in this analysis (Garnett
et al., 2018; Renwick et al., 2017).
4.1 |Conservation on freehold land has
significant benefits for Australia's
biodiversity
Our analysis showed how important freehold land man-
agement is when it comes to abating species extinction in
Australia. Freehold land accounts for the greatest propor-
tion of threatened taxa distributions (48%) and a majority
of the distributions of those species at greatest risk of
extinction (Endangered: 54%; Critically Endangered
53%). It is not surprising that many threatened species
are known or are likely to exist on freehold land, given
that these lands have seen the greatest impact of some
processes that cause species endangerment in the first
place (e.g., land clearing and other forms of habitat loss
and degradation, Evans, 2016). However, our analysis
and a recent analysis by Ivanova and Cook (2020) high-
light the great urgency and enormous potential of achiev-
ing positive biodiversity outcomes on privately owned
lands if Australia employs different strategies to land-
scape conservation.
Building on opportunities for farmers and other land-
holders and custodians to contribute to cross-tenure and
landscape-wide threat management therefore has signifi-
cant benefits for Australia's threatened species. There are
a multitude of instruments and mechanisms available to
governments and nongovernment entities to achieve pos-
itive threatened species outcomes across the tenure land-
scape, particularly in relation to habitat retention, a
critical action needed for many (86%) Australian threat-
ened species (Kearney et al., 2020). These include govern-
ment, private and Indigenous protected areas (Archibald
et al., 2020), economic incentives (e.g., biodiversity stew-
ardship: Burns et al., 2016; carbon farming: Evans, 2018),
disincentives (e.g., land-clearing legislation: Evans, 2016),
educational programs (Byron et al., 2014) as well as
income diversification and influencing reduced demand
for products that drive habitat loss and degradation
(e.g., beef: McAlpine et al., 2009; Selinske et al., 2020).
Often it is a mix of these interventions that generates the
most cost-effective outcome, informed by understanding
public and private benefits and costs (see Pannell (2008)
for more on this balance). The many mechanisms and
instruments available for retaining habitat quality and
quantity need to be activated across a spectrum of ten-
ures to overcome this prevailing threat (Ward
et al., 2019).
4.2 |Securing the fate of range-restricted
species
Our analysis shows that one in seven (n=227)
Australian threatened taxa occur almost entirely (>95%
of distribution) on a single non-protected tenure type,
with the majority of these (n=201) occurring solely on
freehold lands. The pygmy blue-tongue lizard (Tiliqua
adelaidensis; Endangered), endemic to South Australia,
occurs only on freehold lands in a highly fragmented
landscape dominated by agriculture (predominantly
sheep grazing). For this species, mitigating the impacts of
the identified threats from intensive agriculture, pollu-
tion, and urban development (Fenner et al., 2018) on pri-
vate lands is now vital to its persistence. Furthermore,
the 2012 recovery plan for the species noted that all
known habitat is critical to the survival of the species
(Duffy et al., 2012, p. 10), highlighting that the recogni-
tion and effective management of species' critical habitats
is vital to the persistence of this freehold-endemic species.
Likewise, the shapely zieria (Zieria formosa, Endan-
gered), a plant species endemic to south-eastern New
South Wales, occurs only on privately owned lands.
Known only from a single population across three rural
residential properties (Threatened Species Scientific
Committee, 2016), the species persistence now relies on
actions to ensure agriculture, pollution, invasive species,
and fire are effectively managed on these properties.
On leasehold lands, the Carpentarian rock-rat,
Aywalirroomoo (Zyzomys palatalis; Endangered), is
known only from a single pastoral property in the North-
ern Territory (Puckey et al., 2003). Management of fire
regimes is the primary action needed, along with man-
agement of introduced herbivores and predators
(i.e., feral cats) and mitigation of the impacts of climate
change (e.g., through translocation; Threatened Species
Scientific Committee, 2019). The persistence of these
(and the other >200) species depends on effective, coordi-
nated threat management and conservation efforts on
non-protected lands. These species and their critical habi-
tats must be the focus of specific planning that works
with all relevant stakeholders to ensure their extinctions
are prevented. At present, most species do not have ade-
quate recovery plans (McDonald et al., 2015) to ensure
their persistence, particularly on non-protected lands.
Greater effort to tailor strategies to other land tenures,
especially private lands, is required.
8of15 KEARNEY ET AL.

While working with existing private landholders will
be critical, another important component of securing the
fates of threatened species, particularly range restricted
ones, on private land is through the direct acquisition
and protection of this land. Nongovernment organiza-
tions have become involved in proactively securing
important habitat on non-protected tenures and
safeguarding it for conservation. This has been a success-
ful mechanism for securing habitat for species like the
night parrot (Pezoporus occidentalis; Endangered) in east-
ern Australia. After the night parrot's rediscovery, Bush
Heritage Australia purchased 56,000 ha of land that was
previously under pastoral lease (Bush Heritage
Australia, 2020). The reserve has since been declared
Australia's first Special Wildlife Reserve, providing
National Park (IUCN category II) level protection of the
species habitat on private land (Bush Heritage
Australia, 2020). However, this level of protection for pri-
vate land is currently only available in Queensland, dem-
onstrating the importance of more widespread legal
support for innovative conservation mechanisms.
4.3 |Coordination across multiple
tenures is needed to recover the majority
of species
Coordination and collaboration of efforts within and
across tenure types is vital to halt Australia's biodiversity
loss as three-quarters of species occur across multiple ten-
ures. This requires integrating multiple tenures into
regional planning initiatives up front, to understand which
policies are appropriate where (Karimi & Adams, 2019).
Regional natural resource management groups play a key
role in the coordinated and collaborative management of
biodiversity across Australia (Youl et al., 2006), and the
private land conservation sector is growing
(Fitzsimons, 2015). However, critical deficiencies remain
in the size and coordination of these efforts to ensure their
potential benefits can be fully realized (Robins, 2018).
Although coordinated and broadscale cross-tenure man-
agement of some threats occurs in some regions of
Australia (e.g., European red fox; Saunders et al., 2010;
Fleming et al., 2014), many are not. For example, feral
cats, a nationally recognized key threatening process, lack
a coordinated large-scale management and monitoring ini-
tiative (Garrard et al., 2020). Similarly, despite the 2006
and 2016 Threat Abatement Plan for chytrid fungus call-
ing for national coordination of management efforts
(Commonwealth of Australia, 2006, 2016), such an
approach is still lacking (Skerratt et al., 2016).
Many Australian species are also highly mobile
(Runge et al., 2015) and use different parts of the
landscape depending on lifecycle stages or the need to
follow ephemeral resources (Griffioen & Clarke, 2002;
Kerezsy et al., 2013). Species such as regent honeyeater
(Anthochaera phrygia; Critically Endangered), swift par-
rot (Lathamus discolor; Critically Endangered) and gray-
headed flying-fox (Pteropus poliocephalus; Vulnerable)
have very large distributions and are highly dependent
on seasonal and shifting resources (Franklin et al., 1989;
Roberts et al., 2012; Saunders et al., 2007). These species
are in decline due to acute threatening processes, particu-
larly habitat destruction due to logging, urban develop-
ment, and agriculture, operating in parts of their range
that are periodically critical for the species survival.
Multi-tenure actions are now vital for these and similar
species to secure them from extinction.
Recognizing the multi-tenure distribution of many
EPBC-listed species also points to the need for legal
reform to improve the implementation of recovery plans
and threat abatement plans (EPBC Act sections
188, 270A) across a broad range of tenure types. At pre-
sent, both recovery plans and threat abatement plans
bind only the Commonwealth (EPBC Act sections 139(1)
(b), 268) and lack requirements of implementation out-
side Commonwealth land (section EPBC Act 269(2)), a
subset of Crown land. As a result, threatening process
can occur across many tenure types where Common-
wealth approvals or actions are not involved.
While the findings of our analysis highlight the broad
trend of the importance of freehold lands and protected
areas for many species, the reality is that the inter-
section between threatened species and land tenure
across Australia is highly diverse, with many unique
combinations of land tenures being important for species.
These combinations present unique opportunities and
challenges to coordinating conservation efforts across
Australia. As highlighted by our analysis, the most
species-rich tenure combination is freehold lands and
protected areas, with over 1000 taxa having >5% of their
distribution on both tenures. Approaches to landscape-
scale and cross-tenure conservation, such as Biosphere
Reserves and Conservation Management Networks, pro-
vide a means of coordinating conservation efforts across
public and private lands (Fitzsimons et al., 2013;
Wyborn, 2011). Concerningly, land tenure combinations
including forestry (e.g., freehold and forestry; forestry
and protected areas) are important for the conservation
of hundreds of Australian threatened taxa. At present,
forestry remains a troublesome land tenure in terms of
threatened species conservation efforts in Australia
(Lindenmayer & Burnett, 2021), especially as forestry is a
major threatening process to many endangered species
(Kearney et al., 2019; Ward et al., 2021), making cross-
tenure conservation efforts difficult (Webb et al., 2018).
KEARNEY ET AL.9of15

Managing threats that impact imperiled species beyond
protected area boundaries can drive a multitude of co-
benefits such as increased agricultural profitability (Rees
et al., 2020) and carbon sequestration (Carwardine
et al., 2015; Evans et al., 2015). A growing body of evidence
suggests that many threats to biodiversity are having
increasingly negative impacts on other land uses as well as
human livelihoods and well-being. For example, widespread
land clearing in many areas has had significant impact on
local weather and climate in Australia (Andrich &
Imberger, 2013; Pitman et al., 2004) and elsewhere
(McAlpine et al., 2018; Salazar et al., 2016), impacting agri-
cultural productivity and water sources for human settle-
ments. Similarly, widespread land clearing across large parts
of Australia has resulted in major salinity issues, rendering
these areas inhospitable for agriculture and biodiversity
alike (Murray-Darling Basin Authority, 2015). Furthermore,
the management of numerous invasive species has obvious
benefits for agricultural activities and other ecosystem ser-
vices(Pejchar&Mooney,2009),asinvasivespeciesin
Australia cost billions in economic losses and control mea-
sures (Hoffmann & Broadhurst, 2016). The European rabbit
(Oryctolagus cuniculus), an invasive species most commonly
listed as impacting threatened species in Australia (Kearney
et al., 2019), has an estimated economic impact on
Australian agriculture in excess of AUD$200 million annu-
ally (Gong et al., 2009). Similarly, lantana (Lantana camara)
impacts nearly 100 Australian threatened species (Kearney
et al., 2019) and costs the livestock industry over AUD$100
million in lost productivity and management expenses
annually (Grice et al., 2014). Obviously, those threats to bio-
diversity that have already been identified to impact human
livelihoods should not be the sole focus of management
interventions; however, it does provide useful insights into
threat management opportunities and co-benefits.
Conservation success for threatened species requires
going beyond active threat management (e.g., invasive spe-
cies control; habitat restoration), to ensure that the manage-
ment and use of habitat on non-protected lands becomes
more compatible with the persistence of threatened species
(and maintaining common species, preventing their decline
into threatened status). Technology and innovation will
have a role to play in this (Sayer & Cassman, 2013), as will
landscape-scale planning and a shift toward biodiversity-
compatible and regenerative agriculture (Fischer
et al., 2008; White, 2020) and multifunctional landscapes
(Kremen & Merenlender, 2018). In addition, many threat-
ened species are unable to tolerate negative human distur-
bances, so intact areas free from incompatible human
pressures are critical to their survival (Di Marco et al., 2018;
Nori et al., 2018). Ambition and leadership are urgently
needed to establish institutions that signal the public desire
for long-term sustainable production and biodiversity
conservation (Leclère et al., 2020; Springmann
et al., 2018). Fundamental to this is mapping the core
habitats of threatened species, monitoring their trajecto-
ries, and making this information available to prioritize
the most important locations for these efforts. The Key
Biodiversity Area (KBA) process provides a useful frame-
work within which to achieve this (Langhammer
et al., 2017). KBAs are sites that contribute significantly
to the global persistence of biodiversity, including threat-
ened species (IUCN, 2016). The KBA criteria are globally
standardized, consistent, repeatable, quantitative, and
transparent and provide a tenure-blind approach to iden-
tifying significant sites for threatened species. The recog-
nition of a site that is important for a threatened species
as a KBA can help inform land management practices
and provides a clearly delineated area that is priority for
the management of threatening processes.
5|CONCLUSION
Understanding the distribution of threatened species
across the human landscape is critical to informing how
best to respond to species decline. We assessed this in the
context of Australian land tenure, to help inform the
response to the extinction crisis by combining threatened
species distributions maps with land tenure spatial data.
Our research points to the need for strengthening
Australia's multi-tenure approach to the conservation of
threatened species. Working with landholders and custo-
dians across all tenures is critical for better threatened
species outcomes, as coordinated actions by land man-
agers are essential to the persistence of many species.
Key to achieving this will be mapping out the key habitat
areas and gaining agreement and resources for the criti-
cal actions needed to secure threatened species from
extinction, as well as ensuring that the benefits of this
management to the broader environment and its people,
are realized. Furthermore, threatened species survival
hinges on a step change in both the laws that manage
conservation outcomes across tenure types and the
opportunities available for landholders to contribute to
threat management and positive threatened species con-
servation outcomes on their lands.
ACKNOWLEDGMENTS
The authors thank Scott Atkinson for assistance in pre-
paring spatial datasets and Matt Hayward, Richard
Schuster, and an anonymous reviewer for helpful com-
ments that improved our manuscript.
CONFLICT OF INTEREST
The authors declare no conflicts of interest.
10 of 15 KEARNEY ET AL.

AUTHOR CONTRIBUTIONS
Stephen G. Kearney, Josie Carwardine, Vanessa
M. Adams, and James E. M. Watson conceived the idea
and designed the research. James E. M. Watson acquired
the data. Stephen G. Kearney analyzed and interpreted
the data, with assistance from all authors. Stephen
G. Kearney, Josie Carwardine, and James E. M. Watson
drafted the manuscript with input from all authors. All
authors contributed to critically editing and revising the
manuscript.
DATA AVAILABILITY STATEMENT
This study used a combination of commercial, restricted
access, and freely available datasets. The species' distribu-
tion data was provided by the Australian Government's
Department of the Environment via a Data License Deed.
A publically available (and lower resolution) version of
this dataset is available from the URL link in the refer-
ence list. The PSMA CadLite and Land Tenure data are
commercial products which require a paid subscription
to access and use. The Geoscience Australia land tenure
dataset and the Collaborative Australian Protected Areas
Database are freely available via the relevant URL links
in the reference list.
ETHICS STATEMENT
No ethical approval was required for this study.
ORCID
Stephen G. Kearney https://orcid.org/0000-0002-0026-
970X
Josie Carwardine https://orcid.org/0000-0002-5153-
4622
April E. Reside https://orcid.org/0000-0002-0760-9527
Vanessa M. Adams https://orcid.org/0000-0002-3509-
7901
Rebecca Nelson https://orcid.org/0000-0002-7891-0296
Anthea Coggan https://orcid.org/0000-0003-2625-9435
Rebecca Spindler https://orcid.org/0000-0001-6309-
0435
James E. M. Watson https://orcid.org/0000-0003-4942-
1984
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How to cite this article: Kearney, S. G.,
Carwardine, J., Reside, A. E., Adams, V. M.,
Nelson, R., Coggan, A., Spindler, R., & Watson, J.
E. M. (2022). Saving species beyond the protected
area fence: Threats must be managed across
multiple land tenure types to secure Australia's
endangered species. Conservation Science and
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