Bioregionalization Approaches For Conservation: Methods, Biases, and Their Implications For Australian Biodiversity

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Biodiversity and Conservation

https://doi.org/10.1007/s10531-019-01913-6

REVIEW PAPER

Bioregionalization approaches for conservation: methods,


biases, and their implications for Australian biodiversity

Cristian S. Montalvo‑Mancheno1   · Stefania Ondei1 · Barry W. Brook1,2 ·


Jessie C. Buettel1,2

Received: 14 February 2019 / Revised: 13 October 2019 / Accepted: 21 November 2019


© Springer Nature B.V. 2019

Abstract
Biogeographic classification schemes have been developed to prioritize biodiversity con-
servation efforts at large scales, but their efficacy remains understudied. Here we develop
a systematic map of the literature on bioregional planning, based on a case study of the
Interim Biogeographic Regionalization for Australia (IBRA), to identify where and how
such schemes have been used in scientific research. We identified 67 relevant studies,
finding that the majority investigated biodiversity exclusively within a single bioregion
(65.7%), with 18 of these studies splitting the targeted bioregion based on administrative
boundaries. Most used inferential techniques (74.6%) or pattern-based measures (68.7%),
and few studies (9%) both considered biodiversity across multiple bioregions and com-
pared findings between bioregions. Species were investigated ten times more frequently
than ecosystems attributes, with mammals and birds monopolizing scientists’ attention.
These findings show that our knowledge of biodiversity at bioregional scales is patchy,
even for well-studied taxa, and that we have a limited understanding of the synthetic rela-
tionship between biodiversity and IBRA bioregions (which are demarcated according to
other biophysical factors). This creates a barrier for systematic conservation planning,
which requires unbiased information on the spatial attributes of biodiversity, and therefore
this knowledge deficit warrants more attention.

Keywords  Bioregionalization · Terrestrial biodiversity · Australia · Interim Biogeographic


Regionalization for Australia (IBRA) · Systematic map

Communicated by Dirk Sven Schmeller.

Electronic supplementary material  The online version of this article (https​://doi.org/10.1007/s1053​


1-019-01913​-6) contains supplementary material, which is available to authorized users.

* Cristian S. Montalvo‑Mancheno
cristian.montalvomancheno@utas.edu.au
1
School of Natural Sciences, College of Sciences and Engineering, University of Tasmania, Private
Bag 55, Hobart, TAS 7001, Australia
2
ARC Centre of Excellence for Australian Biodiversity and Heritage (CABAH), University
of Tasmania, Private Bag 55, Hobart, TAS 7001, Australia

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Biodiversity and Conservation

Introduction

Biogeographically-based conservation schemes have played an important role in guiding


conservation efforts at large spatial scales (Jepson and Whittaker 2002; Whittaker et  al.
2005). They have been used to identify areas where biodiversity (e.g., endemic species
or habitat) are highly threatened (Hoekstra et al. 2005; Mittermeier et al. 2004), evaluate
conservation priorities (Long et al. 1996; Olson and Dinerstein 1998), attract conservation
investment to specific regions (Sloan et al. 2014) and to guide environmental policy and
biodiversity research (Smith et  al. 2018). Yet, biodiversity continues to decline, even in
regions prioritized as having high conservation value (Butchart et al. 2010; Rodrigues et al.
2014).
Despite constituting important planning tools, conservation schemes based on biogeo-
graphic principles have been criticized on conceptual, methodological and implementation
grounds. Although these schemes have incorporated systematic conservation principles
(i.e., vulnerability and irreplaceability) to various degrees (Brooks et al. 2006), they typi-
cally fail to incorporate complementarity, to optimize the representation of all mapped taxa
(Humphries 2001; Mace et al. 2000). Setting priorities using major habitat boundaries or
species of one taxon (even if speciose, like birds) may fail to capture diversity in other
taxa and biodiversity levels (Brummitt and Lughadha 2003). Using species and endemism
as the levels of specificity for analyses can be problematic, because of the possible vari-
ance in patterns of richness at different taxonomic levels and the not necessarily positive
relationship between species richness and endemism (Whittaker et  al. 2005). It has been
shown that the unequal size of these schemes’ planning units has skewed prioritization
results towards planning units of small size, because there was no correction for the non-
linearity of the species-area relationship (Ovadia 2003). Further, the large size of these
schemes’ planning units has been argued to be impractical for implementation of conserva-
tion actions because of the need to move back-and-forth between planning scales before
these schemes can be effectively executed on ground (Humphries 2001; Mace et al. 2000).
Despite this criticism, which prompted the response of some of the original biogeographic
conservation advocates (Brooks et  al. 2006; Myers and Mittermeier 2003), research on
how the patterns and processes of biodiversity relates to the spatial units of biogeographi-
cally based conservation schemes remains an understudied topic.
Globally, there are marked biases on where, what and how biodiversity is researched
(e.g., Clark and May 2002; Fardila et  al. 2017; Martín-López et  al. 2009). The study of
single species has, to date, been the main research focus in scientific fields related to biodi-
versity conservation (Carmel et al. 2013; Fazey et al. 2005). Research on mammal and bird
species is still the dominant trend across biological sub-disciplines (Hecnar 2009; Velasco
et al. 2015), with data skewed toward the Palearctic and Nearctic biogeographic realms and
focused on developed countries in those regions (Collen et  al. 2008; Ondei et  al. 2018).
Observational and experimental research is most common in ecological research, with an
increase in problem-solving studies and the use of secondary sources of data for modeling
and big data analysis over the last three decades (Carmel et al. 2013), often at the expense
of field-based research (Ríos-Saldaña et al. 2018). Evidence syntheses of the literature on
conservation science have also found disparities in the relative effort given to different
methodological, geographic or biodiversity foci (e.g., Fardila et al. 2017; Roe et al. 2014).
The inconsistency in biodiversity research is systemic; therefore, it is reasonable to expect
that biodiversity research based on the spatial units of biogeographic conservation schemes
will reflect similar patterns.

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Biodiversity and Conservation

Systematic reviews and syntheses are important for evaluating strength of evidence in
conservation biology and environmental science (James et  al. 2016; Pullin and Knight
2009). Although the gold standard of evidence synthesis is to produce an aggregate quan-
titative measure of an intervention’s impact on a study system (e.g., systematic review,
meta-analysis), this is not always feasible, due to the lack of suitable empirical data or the
scope of the question of interest (Collaboration for Environmental Evidence 2018). ‘Sys-
tematic mapping’, developed as a response to this limitation, seeks to describe the nature of
a research field in terms of distribution and abundance of available evidence, but also as a
means to identify sub-sets of studies suitable for systematic review in one or more areas of
the systematic map (Gough et al. 2012; James et al. 2016). In this study, our overall objec-
tive was to create a systematic map that charts the factors underpinning our understanding
of biodiversity research within the context of bioregionalized conservation schemes. To do
this, we considered how bioregions have been studied for conservation, what dimensions
of biodiversity have been investigated, and how research effort has been distributed within
and across the spatial units of a biogeographically based conservation framework.
To meet these objectives, we chose to focus on the development of research on pat-
terns and processes of Australia’s terrestrial biodiversity, based on the spatial units (i.e.,
bioregions) of the Interim Biogeographic Regionalization for Australia (IBRA) framework.
This provides an ideal case study, because Australia is the only megadiverse country (Mit-
termeier et al. 1997) where a biogeographical approach has been used explicitly to prior-
itize conservation actions on the ground. Implementation of the IBRA framework has been
instrumental in curbing the inherent bias of Australia’s national reserve system towards
areas of low agricultural productivity (Barr et  al. 2016). Nevertheless, conservation of
large intact landscapes in Australia has decreased (Watson et al. 2009), and it is recognized
that IBRA bioregions and threatened species with relatively large ranges are not uniformly
represented (Taylor 2017; Taylor et al. 2014) or adequately protected (Watson et al. 2011).
The mixed success of the IBRA framework at conserving biodiversity has, like its global
counterparts, been attributed to an inadequate use of systematic conservation planning
principles (Craigie et al. 2015) and inherent methodological limitations, though this has yet
to be quantitavely assessed.

Methods

Search, selection, classification and summary of the literature

To identify the relevant literature, we searched three databases: Web of Science, Scopus,
and Google Scholar. In the first two, the following keyword string was used ([*region* OR
biogeograph* OR “IBRA”] AND [“Australia” OR “Australian” OR “Australia’s”] AND
[“species” OR *divers* OR conserv* OR communit* OR assemblage* OR ecosystem* OR
guild* OR tax*] NOT [*water* OR mari* OR aqua* OR sea OR ocean*] NOT [“New
Zealand”]). In Google Scholar, a simpler keyword string was used instead ([“Interim Bio-
geographic Regionali Australia”]), retrieving the first 50 records. We constrained the date
range to commence from 1995 (the year IBRA was first officially released; Thackway and
Cresswell 1995) through to 2017. Duplicates were removed; and the reference lists of key
biodiversity-focused studies using IBRA bioregions were cross-checked to ensure all rel-
evant studies were captured.

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Biodiversity and Conservation

To identify relevant studies, we implemented a stepwise screening process. This started


with the application of a broad filter (based on a set of exclusion criteria), followed by
use of a text mining technique (n-gram analysis) to prioritize a list of potentially relevant
papers that was made available for a final, in-depth manual screening (O’Mara-Eves et al.
2015). A detailed description of the screening phase is provided in Online Resource 1.
The ‘Preferred Reporting Items for Systematic Reviews and Meta-Analysis’ protocol
(Moher et al. 2009) was followed, to record the results of our search and screening process
(Fig.  1). EndNote X8.2 (Clarivate Analytics 2018) was used to manage references; and

Fig. 1  Identification and selection of relevant studies used in our systematic mapping of biodiversity studies
of Australian biogeographic units, based on application of the PRISMA protocol (Moher et al. 2009). Note:
WoS, GS and IBRA respectively stand for Web of Science, Google Scholar and the Interim Biogeographic
Regionalization for Australia

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Biodiversity and Conservation

three packages in Program R program (R Core Team 2018)—tm (Feinerer et  al. 2008),
RWeka (Hornik et al. 2009) and SnowballC (Bouchet-Valat 2014)—were used to construct
structured data from the text.
To characterize research on biodiversity for all relevant studies, we extracted a range of
attributes, including: IBRA information (e.g., number of IBRA bioregions studied), spa-
tial location, study design (e.g., type of analysis, data source, biodiversity response), focal
entity (flora, fauna, and ecosystem), and various biodiversity attributes (Online Resource
2). We also recorded information on the number of species and ecosystem services investi-
gated. We used the shapefile of IBRA version 7 (Department of the Environment 2012) to
spatially summarize the number of times bioregions and dimensions of biodiversity were
reported. To represent this accurately, the four IBRA bioregions that lie outside of main-
land Australia and Tasmania (i.e., Coral Sea, Indian Tropical Islands, Pacific Subtropical
Islands, and Sub-Antarctic Islands) were omitted, and polygons extending across two or
more administrative units were split based on state and territory boundaries, excluding
Australia Capital and Jervis Bay territories due to their small scale and absence from rel-
evant studies. This resulted in 32 bioregion polygons being artificially split into 71 dis-
crete spatial units—hereafter referred as state-split discrete bioregions—and 53 bioregion
polygons located exclusively within a state or territory—hereafter referred as full-extent
discrete bioregions (Fig. 2).
In this systematic map, the development of evidence matrices was informed by the most
salient themes covered in the biodiversity research literature (e.g., de los Ríos et al. 2018;
Martín-López et al. 2009) and systematic analyses on the nature of scientific fields applied
to biodiversity conservation (e.g., Cronin et al. 2014; Fardila et al. 2017). To provide an
overview of the existing evidence on biodiversity research at the IBRA scale, we compared
counts within and across all categories/groupings and calculated descriptive statistics for
both the number of species investigated, and the number of times bioregions and dimen-
sions of biodiversity were reported in relevant studies. We used ArcGIS 10.5.1 (ESRI
2017) to generate spatial data, and three R packages to analyze, summarize and visualize
data: dplyr (Wickham et al. 2017), ggplot2 (Wickham 2016) and expss (Demin 2018).

Quality assurance of evidence synthesis process

To minimize possible sources of bias and error, we started by structuring our overarch-
ing evidence-synthesis question to contain the population (P) and outcome (O) elements—
often referred as ‘PO’ question type (Collaboration for Environmental Evidence 2018;
James et al. 2016). We used an iterative-keyword-screening process to identify the search
strategy that best captured our studies of interest across three databases, and a stepwise
process with pre-defined exclusion criteria to ensure our work is reproducible and transpar-
ent (Online Resource 1). The accuracy of the text-mining technique was estimated based
on selective manual validation, and both the interpretation of our exclusion criteria and the
reliability of the classification of relevant studies were independently examined (Online
Resource 3). Due to the breadth of our topic, an explicit critical appraisal of study valid-
ity was impractical; this is justifiable given that it was unlikely to influence the collation,
description and mapping process (Collaboration for Environmental Evidence 2018). We
instead focused on removing duplicate publications of the same data to avoid double count-
ing (Frampton et al. 2017), given that frequency of study attribute categories were pivotal
to the reliability of our evidence synthesis.

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Fig. 2  Extent (expressed in percentage) of bioregions within administrative units (states and territories),
and number of times discrete bioregions were reported in biodiversity studies. IBRA Interim Biogeographic
Regionalization for Australia

Results

Of the 15,190 references uncovered during the identification phase (Fig. 1), 67 studies met
all eligibility criteria. The number of biodiversity-focused studies on IBRA bioregions
published annually has increased over time (Online Resource 4), with more than half (58%)
using bioregions based on the IBRA 6.1 revision, published in 2004 (Online Resource
5). The majority (65.7%) of relevant studies undertook biodiversity research inside only
a single bioregion. For 18 of the studies, the reported bioregion was artificially split by
administrative boundaries, with a fifth (19.4%) investigating biodiversity from bioregions
extending across two or more administrative units. Five studies considered all bioregions
found across the entire Australian continent (mainland and Tasmania), and in two studies,
Tasmania was excluded.
The frequency of relevant studies differed among bioregions, and within bioregion
when its area extends across two or more administrative units (Fig.  2). Nine of the 124

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Biodiversity and Conservation

discrete spatial units into which IBRA bioregions were split were reported anywhere
between 12 and 17 times (Online Resource 6). The largest number of times any of the 53
full-extent discrete bioregions (i.e., IBRAs found exclusively within one state or territory)
was reported ranged from 6 to 17, in Tasmania and Queensland respectively (Table 1). Yet,
in New South Wales and Victoria, a state-split discrete bioregion (i.e., IBRAs extending
across administrative units) was more frequently reported than any full-extent bioregion
found within their boundaries.
Nearly half (47.8%) of relevant studies were based exclusively on new data, but only
two provided spatial information to locate sample sites (Table 2). Inferential analysis was
used ~ 3.5 times more frequently than predictive or descriptive analyses (combined). Meas-
ures of species distribution and/or diversity were most common (68.7%), and 6/23 studies
that considered biodiversity across multiple bioregions compared findings between biore-
gions (9% of all relevant studies).
There were seven relevant studies that investigated biodiversity at the ecosystem level
(Online Resource 7), all of which focused on one bioregion each or two bioregions (n = 6

Table 1  Number of full-extent and state-split discrete bioregions within Australia’s administrative units (n),
and total, mean, standard deviation (SD), minimum (Min.) and maximum (Max.) number of times full-
extent and state-split discrete bioregions were reported

Administrative unit With all relevant studies


Type of discrete bioregion n Total Mean Median SD Min. Max.

New South Wales


 Full-extent 3 31 10.3 10.0 0.6 10 11
 State-split 15 162 10.8 10.0 1.26 9 14
Northern territory
 Full-extent 12 101 8.4 8.0 1.4 8 13
 State-split 13 107 8.2 8.0 0.6 8 10
Queensland
 Full-extent 6 65 10.8 10.0 3.4 8 17
 State-split 12 106 8.9 8.0 2.0 7 14
South Australia
 Full-extent 4 34 8.5 8.5 0.6 8 9
 State-split 13 102 7.9 8.0 0.4 7 8
Tasmania
 Full-extent 8 48 6.0 6.0 0.0 6 6
 State-split 1 6 6.0 6.0 – 6 6
Victoria
 Full-extent 2 15 7.5 7.5 0.7 7 8
 State-split 9 70 7.8 7.0 1.4 7 11
Western Australia
 Full-extent 18 185 10.3 10.0 0.6 10 12
 State-split 8 80 10.0 10.0 0.0 10 10
Australia
 Full-extent 53 479 9.0 9.0 2.1 6 17
 State-split 71 633 8.9 8.0 1.7 6 14

– Not applicable

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Table 2  Number of studies by categories of study design attributes (n), and mean, median, standard devia-
tion (SD), minimum (Min.) and maximum (Max.) number of IBRA bioregions reported by categories of
study design attributes

Attribute With all relevant studies


Categories n Mean Median SD Min. Max.

Data source
 Primary data 32 1.7 1.0 3.0 1 18
 Secondary data 27 27.4 8.0 34.8 1 85
 Primary and secondary data 8 1.6 1.5 0.7 1 3
Geographic analytical scope
 General 65 12.4 1.0 25.7 1 85
 Localized 2 1.0 1.0 0.0 1 1
Type of analysis
 Inferential 50 9.7 1.0 22.4 1 85
 Predictive 8 32.8 2.0 43.3 1 85
 Descriptive 6 9.5 5.5 10.4 1 26
 Mechanistic 3 1.3 1.0 0.6 1 2
Biodiversity response
 Distribution and/or diversity 46 12.6 1.0 26.4 1 85
 Genetics 9 5.2 1.0 11.9 1 37
 Population change 6 16.2 4.5 30.0 1 77
 Species trait 3 26.7 2.0 43.6 1 77
 Mechanistic relationship 3 1.3 1.0 0.6 1 2
Type of Study
 One bioregion 44 – – – – –
 >  1 bioregion and no compare 17 43.3 26.0 35.4 2 85
 > 1 bioregion and compare 6 4.3 2.5 3.3 2 9

– Not applicable. Description of categories for each attribute can be found in Online Resource 2

and 1 papers, respectively). In six studies, the research topic was ecosystem functioning
(Online Resource 8). The remaining study evaluated three provisioning and five regulatory
ecosystem services provided by forests in the Wet Tropic bioregion, with this being the
only bioregion reported twice.
Of the 60 papers that investigated living organisms, species assemblages were used four
and six times more frequently than species communities and single species, respectively
(Online Resource 8). Fauna was the predominant and exclusive target across species stud-
ies (76.7%), and in only four studies, flora was investigated in combination with fauna. Out
of the 55 species studies with enough information to classify species at higher taxonomic
rank, 86 observations spread across 16 taxa were identified and examined. Mammals and
birds were commonly studied, followed by reptiles and vascular plants (Online Resource
9). Vertebrates, which constituted 67.4% of taxonomic records (Fig.  3a), were studied
more frequently than plants (Fig. 3b) or invertebrates (Online Resource 10) in all discrete
bioregions.
Five of the 86 observations lacked enough information to identify how many species
were studied at higher taxonomic rank, affecting four taxa (i.e., Aves, Arachnida, and Mal-
acostraca only once, and Mammalia twice). Although the average number of species varied

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Biodiversity and Conservation

Fig. 3  Number of times a vertebrates and b plants were investigated in discrete bioregions, based on rel-
evant studies that had sufficient information to classify fauna and flora into higher taxonomic groups
(n = 55). IBRA Interim Biogeographic Regionalization for Australia

greatly between taxa—ranging from 1 to 692 species of non-vascular plants and insects,
respectively—the standard deviation in the number of species that were studied was hun-
dred or more for mammals, reptiles, snails/slugs, insects, and vascular plants (Online
Resource 11).
For the nine most-reported discrete bioregions (i.e., 12–17 relevant studies; Online
Resource 6), similar frequency patterns are apparent for the categories of study design
attributes (Table 3) and taxa (Fig. 4). The only striking difference is that research on biodi-
versity in the most-reported discrete bioregions was largely based on secondary data, and
most of these considered biodiversity across multiple IBRA bioregions without comparing
findings between bioregional spatial units (Table 3).

Discussion

Biogeographic classification schemes have been developed to prioritize conservation


efforts, and have been repeatedly refined in some countries (e.g., IBRA is now up to its
fourth official release [version 7]). However, the factors underpinning our understanding
of biodiversity research within the context of such schemes has yet to be studied. Through
a detailed look (evidence synthesis) at the scientific use of the IBRA framework, we found
disparities in the relative effort given to different geographic, methodological and biodiver-
sity foci, and a lack of integrative work spanning multiple bioregions.

Geographical bias

Research effort is distributed unevenly both among IBRA bioregions, and within biore-
gion when its area extends across two or more administrative units (Fig. 2). This suggests

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Biodiversity and Conservation

Table 3  Administrative unit, extent of discrete bioregion within administrative unit (expressed in percent-
age), total number of times most-reported discrete bioregions were reported, and the percentage of times
the categories for each of the study design attributes were used in most-reported discrete bioregions

Attribute Most-reported discrete bioregions


Categories CYP MAC MUL NAN NET NSS PIL RIV WET

General information
 Administrative unit QLD NT QLD NSW NSW NSW WA NSW QLD
 Extent (%) 100 100 73.9 76.7 95.2 93.5 100 72.5 100
 Times represented 12 13 14 12 12 14 12 12 17
Data source
 Primary data 16.7 15.4 21.4 25.0 16.7 35.7 16.7 8.3 29.4
 Secondary data 66.6 84.6 71.5 75.0 83.3 57.2 83.3 83.4 53.0
 Primary and secondary data 16.7 – 7.1 – – 7.1 – 8.3 17.6
Geographic analytical scope
 Localized – 7.7 – – – – – – –
 General 100 92.3 100 100 100 100 100 100 100
Type of analysis
 Inferential 58.4 76.9 64.4 66.7 66.7 71.5 66.7 58.4 52.9
 Predictive 33.3 23.1 21.4 25.0 25.0 21.4 25.0 25.0 47.1
 Descriptive – – 7.1 8.3 8.3 7.1 8.3 8.3 –
 Mechanistic 8.3 – 7.1 – – – – 8.3 –
Biodiversity response
 Distribution and/or diversity 75.1 76.9 71.6 66.8 66.8 85.8 58.3 66.8 64.6
 Genetics – 7.7 7.1 8.3 8.3 – 25.0 8.3 5.9
 Population change 8.3 7.7 7.1 16.6 16.6 7.1 8.3 8.3 17.6
 Species trait 8.3 7.7 7.1 8.3 8.3 7.1 8.3 8.3 11.8
 Mechanistic relationship 8.3 – 7.1 – – – – 8.3 –
Type of study
 One bioregion 16.7 38.5 35.7 8.3 8.3 28.6 16.7 8.3 52.9
  > 1 bioregion and no compare 66.6 61.5 64.3 83.4 83.4 64.3 83.3 83.4 41.2
  > 1 bioregion and compare 16.7 – – 8.3 8.3 7.1 – 8.3 5.9

– Not applicable. Most-reported refers to those discrete bioregions that were reported ≥ 12 times. CYP,
MAC, MUL, NAN, NET, NSS, PIL, RIV and WET respectively corresponds to Cape York Peninsula,
MacDonnell Ranges, Mulga Lands, Nandewar, New England Tablelands, New South Wales South Western
Slopes, Pilbara, Riverina, and Wet Tropics. NSW, NT, QLD, and WA respectively stand for New South
Wales, Northern Territory, Queensland, and Western Australia. Description of categories for each attribute
can be found in Online Resource 2

that Australia’s state and territory boundaries are important determinants of the geographic
extent of biodiversity research, taking priority over the integrity of IBRA bioregions as
unit of analysis. Similar patterns have been observed, at a global scale, in efforts devoted
to different countries, again likely driven by historical and political/administrative circum-
stances (Ahrends et al. 2011; Meyer et al. 2015). Although the planning units of the IBRA
framework (i.e., bioregions) are independent from political boundaries, their demarcation
and description were achieved by aggregating environmental information (e.g., Tasmania
nature conservation regions, Queensland biogeographic regions) provided independently

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Fig. 4  Number of times each taxon was represented in the most-reported discrete bioregions. Note: Most-
reported refers to discrete bioregions that were reported ≥ 12 times. CYP, MAC, MUL, NAN, NET, NSS,
PIL, RIV and WET respectively stand for: Cape York Peninsula, MacDonnell Ranges, Mulga Lands,
Nandewar, New England Tablelands, New South Wales South Western Slopes, Pilbara, Riverina and Wet
Tropics

by states and territories (Thackway and Cresswell 1995). Further, the planning and man-
agement of natural resources in Queensland, for example, is driven by a hierarchical clas-
sification approach that merged the IBRA bioregions extending outside the state with
those lying exclusively within (Wilson et  al. 2002), which might have inadvertently pro-
moted those bioregions as landscape features that are also, or can be confined to, political
boundaries.

Methodological preferences

There are marked preferences on how biodiversity research within the context of the IBRA
framework have been designed in the last two decades (Table  2). Unlike global trends,
where a decline in fieldwork-based studies have been observed (Carmel et al. 2013; Ríos-
Saldaña et  al. 2018), we found that researchers largely used field data to carry out their
analyses at a bioregional scale. This means that biases in biodiversity research may also
stem from factors limiting the digitalization and access of first-hand collected data (Meyer
et al. 2015). The large number of studies carried out at a relatively small scale (i.e., 65.7%
of relevant studies undertook biodiversity research inside only a single IBRA bioregion)
could amount to a considerable scientific output if their data were properly stored in an
open-access form (Hampton et al. 2013). This potential contribution is, however, hampered

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by the lack of geographic information on sample sites (e.g., 97% of relevant studies uncov-
ered in this evidence synthesis), limiting accountability and replicability, and thus increas-
ing the challenge in incorporating these data into national- or global-scale analyses.
The preference for pattern-based measures of biodiversity was expected, as these can be
easily interpreted and readily used to investigate different spatial arrangements (alpha, beta
and gamma diversity) of biodiversity (Colwell 2009) and are less sensitive to differences in
survey design (Magurran et al. 2010). However, as a consequence of their community ecol-
ogy focus, they provide less accurate estimates of autecological processes (e.g., population
dynamics, species’ dispersal), compared with more complex response measurements on
individual species, or the directed study of species interactions (Fardila et  al. 2017). We
also found a higher proportion of inferential research compared to those that focused on the
likelihood and magnitude of environmental changes or their underlying mechanisms (pre-
dictive and mechanistic approaches). Further, the lack of comparison between spatial units
of analysis—IBRA bioregions in this review—is a limiting factor for systematic conserva-
tion planning (de los Ríos et al. 2018).

Biases in biodiversity foci

Our Australian findings reinforce the view that there is a bias towards species as the level
of biological organization, as recorded in the global literature (Fazey et al. 2005; Velasco
et al. 2015). As such, the emphasis on species occurrence and community/diversity met-
rics—60% of relevant studies investigating living organisms at IBRA scale conducted
research at assemblage level—is at odds with the claim that species interactions and eco-
system processes enjoy a major emphasis in ecological research (Caliman et  al. 2010;
Nobis and Wohlgemuth 2004). However, research on single species is also rarely under-
taken within the context of the IBRA framework, contrary to what has been found in other
reviews (e.g., Fardila et al. 2017; Fazey et al. 2005). Further, our bioregion findings align
with previous literature reviews that have pointed out both the overrepresentation of mam-
mals and birds (Clark and May 2002; Martín-López et al. 2009) and the lack of research
on more speciose taxa (e.g., insects) or relatively highly threatened ones (e.g., amphibians),
despite a slight increase in research effort towards poorly represented taxa in recent years
(Di Marco et al. 2017).
The predominant focus of research on mammals and birds might also be linked to these
taxa being perceived as umbrella or flagship species (de los Ríos et al. 2018; Hecnar 2009)
which are conspicuous and more vulnerable to extinction in Australia (Johnson 2006; Loe-
hle and Eschenbach 2012). Emphasis on highly visible and charismatic taxa is not new in
the literature (Clark and May 2002; Ford et al. 2017; Martín-López et al. 2009). While such
a focus can serve to attract public support to conservation, it also risks diverting conserva-
tion resources away from less alluring but nevertheless threatened species (Seddon et al.
2005) with important ecological roles (Gascon et al. 2015; Lavelle et al. 2006), whose pro-
tection might require less investment for a greater conservation impact (Walsh et al. 2013).

Implications for systematic conservation planning

The design of environmental policies has been linked to both knowledge on biodiversity
and bioregionalized conservation schemes (Martín-López et al. 2009; Smith et al. 2018).
As such, disparities in the relative effort given to where, what and how biodiversity has
been investigated might undermine systematic conservation planning at large scales, due

13
Biodiversity and Conservation

to assessments of biodiversity status, as well as past and future trends (and their drivers),
being based on misleading baselines (Magurran et  al. 2010; Mihoub et  al. 2017). This
could, in turn, translate into conservation policies, targets and actions (de los Ríos et  al.
2018; Martín-López et al. 2009) that fail to achieve their intended goals (Di Marco et al.
2017; Pyšek et al. 2008). For example, incomplete knowledge on distributions and habitat
use has been identified as a crucial factor influencing the listing of species as threatened
and the development of plans for their recovery in Australia (Walsh et al. 2013).
To deal with biodiversity knowledge gaps, which limit our understanding of the rela-
tionship between biodiversity—ranging from genes to ecosystems—conservationists have
proposed three main solutions: (1) greater effort directed towards poorly represented taxa
in the scientific literature (Bonnet et al. 2002; Clark and May 2002); (2) development and
maintenance of biodiversity databases, preferably open-source, relational in structure, and
based on clear standards (Hampton et  al. 2013); and (3) a more equitable allocation of
funding for biodiversity research for meeting the above goals (Andelman et al. 2004; Clark
and May 2002). Improvement has been reported for the first two points, as poorly rep-
resented taxa (e.g., insects) are increasingly salient as study organisms (Di Marco et  al.
2017), and open-source, long-term databases continue to increase in number and quality
(Ondei et  al. 2018). Yet, biodiversity research largely depends on priorities set by fund-
ing organisms (Ahrends et al. 2011; Stroud et al. 2014). All this suggests that incomplete
knowledge on biodiversity will be the norm for quite some time; therefore, conservationists
and managers working at macroecological scales should consider the implications of, and
account for gaps and biases in, research effort when designing policy instruments aimed to
systematically conserve biodiversity. Our evidence synthesis is an important first step in
that direction, as it provides an overview of where, what and how terrestrial biodiversity
has been researched for almost a quarter of a century of systematic conservation planning
at bioregional scale in Australia.

Acknowledgements  This study was supported by Australian Research Council Laureate Fellowship (Grant
Number FL160100101).

Funding This work was funded by Australian Research Council Laureate Fellowship (Grant Number
FL160100101).

Compliance with ethical standards 


Conflict of interest  All authors declare that they have no conflict of interest.

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