Austral Entomology (2016) 55, 1–17

Review
Biosystematics and conservation biology: critical scientific disciplines for
the management of insect biological diversity
Michael F Braby1*† and Matthew R Williams2
1
Department of Land Resource Management, PO Box 496, Palmerston, NT 0831, Australia.
2
Department of Parks and Wildlife, Bentley Delivery Centre, Locked Bag 104, Perth, WA 6983, Australia.

Abstract Biosystematics and conservation biology are critical scientific disciplines that underpin the management of
biological diversity. This is because biosystematics provides two basic elements that are fundamental to
conservation management: the circumscription of species and the spatial distribution of species. These
elements in turn allow conservation biologists to determine the components of biodiversity, such as local
species richness (α-diversity), composition and community structure, patterns of spatial turnover and hetero-
geneity (β-diversity), levels of endemism, and location of ‘biodiversity hotspots’. This information ultimately
provides a framework for systematic conservation planning for the management of biological diversity and
natural resources. In this review, drawing on examples of Australian diurnal Lepidoptera (butterflies and
day-flying moths), we discuss three areas of conservation biology that are crucial for insect biodiversity
conservation: (1) inventory and estimation of faunal richness; (2) monitoring for conservation management
and the selection and use of bioindicators; and (3) assessment of conservation status and recovery of threatened
species. We then explore the capacity of biosystematics to complement and enhance these programmes. Major
challenges for biosystematics are to catalogue and map the Earth’s known species, to discover and describe
new or as-yet-unknown species, to reconstruct the evolutionary history or tree of life and to incorporate
phylogenetic diversity (taxonomic distinctiveness) as a component of biodiversity into conservation planning
and practical nature conservation. The first two tasks, which need to be completed relatively urgently in an era
of biodiversity crisis and a limited and declining pool of taxonomic expertise, are required in order to optimise
conservation effort of the world’s biodiversity. It is recommended that to overcome the taxonomic impediment
for insect conservation taxonomic attention should focus on a limited set of ‘priority’ taxa, and the rate at
which new species are discovered and described needs to be accelerated (by at least an order of magnitude).
An agenda for future research in biosystematics and conservation biology is proposed as a guideline for
biodiversity conservation for Australian entomology.
Key words biological indicator, butterfly conservation, flagship taxon, invertebrate conservation, umbrella taxon.

I NTRODUCT IO N Vane-Wright 2013). Biosystematics is thus a major scientific

Biological systematics or biosystematics is the study of the
diversification of organisms, both past and present, and the
relationships among those organisms through time. It
includes the subdisciplines of taxonomy and systematics
(Probert 2010). Taxonomy includes the tasks of species dis-
covery and recognition, identification, diagnosis, comparison,
classification and naming (Vane-Wright 2008, 2013),
whereas systematics seeks to explore relationships between
species and higher taxonomic units (such as genera and fami-
lies) in an evolutionary context and provide the framework
into which species are classified (e.g. Wiens 2007;
*michael.braby@anu.edu.au
†
Present address: Research School of Biology, The Australian National
University, Canberra, ACT 0200, Australia.
© 2015 Australian Entomological Society
discipline that underpins many areas of biological science –
as May (1990, p. 130) succinctly stated: ‘Without taxonomy
to give shape to the bricks, and systematics to tell us how
to put them together, the house of biological science is a
meaningless jumble’.
Conservation biology, on the other hand, has a major focus
to ensure that biodiversity values or assets (from genes through
species to ecosystems or biotypes) are managed sustainably
and the ecological and evolutionary processes that support that
diversity are maintained (Frankel & Soulé 1981; Wilson
1992). The core areas of scientific endeavour in conservation
biology include inventory, monitoring, recovery of threatened
species, restoration ecology and invasive species management.
However, until relatively recently, biosystematics and conser-
vation biology have remained somewhat disconnected from
one another (May 1990; Vane-Wright et al. 1991; Williams
et al. 1991; Faith 1994, 2002; Humphries et al. 1995;
doi:10.1111/aen.12158
2 M F Braby and M R Williams
Vane-Wright 1996, 2008; Wilson 2004; Braby 2010b;
Cranston 2010; Harvey et al. 2011; Braby et al. 2012).
In essence, conservation management relies on two basic
elements provided by biosystematics: (1) the circumscription
of species and (2) the spatial distribution of species. Estimates
of these taxonomic elements allow conservation biologists to
determine the components of biodiversity, such as local
species richness (α-diversity), composition and community
structure, patterns of spatial turnover and heterogeneity
(β-diversity), levels of endemism, and location of ‘biodiversity
hotspots’, information which ultimately provides a framework
for systematic conservation planning of biological diversity
and natural resource management (e.g. May 1990, 1994;
Humphries et al. 1995; Vane-Wright 1996; Crozier 1997;
Myers et al. 2000; Sarkar et al. 2006; Margules & Sarkar
2007; Boero 2010). In other words, conservation biology is
reliant on taxonomy and, increasingly, on systematics; and
management of biodiversity such as threatened species
requires knowledge from both disciplines to be effective
(Fig. 1). Biosystematics and conservation biology are thus
critical scientific disciplines for the management of biological
diversity because they provide the units or currency on which
almost all knowledge of biodiversity is based: its makeup, its
spatio-temporal distribution, its ecological role/function, the
key threatening processes impacting upon it and the manage-
ment actions and policies required to maintain it. In short, they
provide an understanding of our natural heritage and provide
the scientific information needed by government agencies and
non-government organisations for setting conservation priori-
ties and implementing practical biodiversity conservation
(Nielsen & West 1994).
Yet, many conservation biologists and ecologists simply
regard biosystematics as a service provider for taxonomic
names and species identification, subservient to the appar-
ently more important tasks of experimental design, quantita-
tive sampling and analysis (New 1996b). This perception
© 2015 Australian Entomological Society
Fig. 1. Conceptual model of the interre-
lationships between the disciplines of
biosystematics (taxonomy and systemat-
ics) and conservation biology in the man-
agement of threatened species as a
component of biological diversity.
persists despite the fact that taxonomy and systematics are
hypothesis-driven sciences in which species delimitation and
relationships among species are tested and refined as new
evidence accumulates (Yeates et al. 2011). Conversely, many
contemporary taxonomists are reluctant to integrate a stand-
ard biosystematics research agenda with practical conserva-
tion outcomes (Harvey et al. 2011).
In this review of biosystematics and conservation biology,
we discuss three areas in which the science of conservation
biology is critical for the management of insect biodiversity:
(1) inventory and estimation of faunal richness; (2) monitoring
for conservation management and the selection and use of
bioindicators; and (3) conservation status evaluation and
recovery of threatened species. We then explore the capacity of
biosystematics to complement and underpin these three pro-
grammes and deliver outcomes for insect biodiversity conser-
vation in the context of the biodiversity crisis and limited pool
of taxonomic specialists, and conclude by offering suggestions
for future research.
We draw primarily on examples from the Australian diurnal
Lepidoptera (butterflies and day-flying moths) to illustrate our
points from an entomological perspective. The rationale for this
is that diurnal Lepidoptera in Australia over the past 25 years
has played a major role in conservation biology – as surrogates
for fostering conservation of biodiversity associated, or
co-existing, with specific ecological communities or habitats
(New 1997), including native vegetation remnants in frag-
mented landscapes (Braby & Edwards 2006; Collier et al.
2006; Williams 2008, 2009, 2011); as biological indicator taxa
to monitor the quality and health of natural environments
(Lomov et al. 2006); and as ‘flagship’ taxa in promoting aware-
ness of conservation need for insects generally or focusing
attention on particular threatened species and their habitats in
need of protection (New 1991a, 1992, 1995, 1996a, 2011; Dunn
et al. 1994; New et al. 1995; Sands & New 2002; New & Sands
2004).
 Biosystematics and conservation biology 3

I NVE NTORY review). Roger Kitching (Kitching 1993a,b; Kitching et al.

Inventory is the recording (surveying, sampling, sorting, iden-
tifying, cataloguing, databasing and documenting) of species
(or other components of biodiversity) at various spatial scales:
local (site, landscape), regional (bioregion, biome, continent) or
global (Stork 1995; Stork et al. 1996; Samways 2005). When
the recorded occurrences of species from many sites or loca-
tions are collated they can be mapped to estimate the geographi-
cal range of each species. This information can then be used for
spatial comparison to identify areas of high conservation value
based on patterns of richness and endemism. That is, the objec-
tive of inventory is to document both the composition and
spatial distribution of biodiversity, to identify the important
components of this diversity (i.e. values or assets), and to
prioritise areas for protection and conservation management.
The complete inventory or enumeration of all insect taxa
from a site or landscape is rarely, if ever, possible because of
the large diversity, taxonomic impediment and constraints on
time, finances and personnel (Ehrlich 1992, 2003; Kitching
1993a,b; Kitching et al. 2001; Spector 2006). Therefore, in
inventory studies, typically a single taxon or a relatively small
suite of taxa is selected and used as surrogates (‘umbrella’ taxa
or biodiversity indicators) to estimate broad spatial patterns of
biodiversity (Samways 2005, 2007). These selected taxa are
assumed to represent the diversity of other organisms and the
relative biodiversity of the sampled sites. The majority of
studies, however, caution against using single taxa and recom-
mend a suite of taxa be used for such purposes (see Cranston
& Trueman 1997; McGeoch 2007; Gerlach et al. 2013 for
2000, 2001; Ashton et al. 2011; Kitching & Ashton 2013) has
developed this concept further by the recognition of a ‘predic-
tor set’ − a subset of insect/invertebrate taxa that captures
the spatial patterns characteristic of the entire insect/
invertebrate assemblage. Another approach is to employ
species-occupancy modelling to identify putative biodiversity
indicator species, which are then used to predict species rich-
ness of taxonomic groups, such as been done for butterflies
(Mac Nally & Fleishman 2002; Fleishman et al. 2005).
The selection of priority taxa for documenting patterns of
biodiversity is open to considerable debate (New 1993,
1996b). Building on the framework of New (1993), Cranston
and Trueman (1997), McGeoch (1998), Andersen (1999),
Spector (2006) and Kitching and Ashton (2013), the Northern
Territory Department of Land Resource Management
(NTDLRM) recently proposed several criteria for the selection
of candidate groups of insects and other invertebrates for
inventory (Table 1). Three of the criteria in this list were con-
sidered to be the most crucial, namely: (1) good taxonomic
knowledge of the group; (2) a proven record as informative
bioindicators, with context elsewhere; and (3) performing a
critical role in ecological processes. Of particular importance,
it was considered desirable to capture the major ecological
functions performed by a suite of insects/invertebrates at dif-
ferent levels (herbivory, predation, pollination, nectarivory,
decomposition, seed dispersal etc.) so that the ecosystem ser-
vices (see Raffaelli 2006) were complementary. The list in
Table 1 demonstrates the importance of both biosystematics
and ecology in this process for biodiversity conservation.

Table 1 Criteria for the selection of terrestrial invertebrate groups for inventory adopted by the Northern Territory Department of Land
Resource Management (Division of Flora and Fauna). Criteria are organised under four key areas: taxonomic, ecological, practical and
social

Taxonomic
†Good taxonomic knowledge. Taxonomic coverage relatively complete and accessible so that samples can be accurately identified with aid of
published keys/reference collections
Diversity appropriate with sufficient species richness
Range of geographical distributions with endemic components present, to allow comparisons of assemblages at different spatial scales
Scientific value (e.g. evolutionary or taxonomically distinct)

Ecological
†Informative as bioindicators, with context elsewhere. Preferably show a range of responses to environmental/ecological change or are indicative of
biodiversity patterns of other organisms
†Critical role in ecological processes (e.g. keystone, ecosystem engineer).
Natural history and general biology well understood
Temporal distribution (seasonality) predictable among years
Ecological amplitude with sufficient habitat/spatial heterogeneity

Practical
Sampling can be incorporated into pre-existing survey programmes using standardised, quantitative methods
Ease of sampling in space and time, with sufficient detectability and representation
Samples easily collected, sorted, stored, accessioned and curated
Historic spatial data available (specimens, literature)

Social
Resonance with the general public
Indigenous cultural value

Criteria marked with a dagger symbol (†) are considered to be the most crucial.
© 2015 Australian Entomological Society
4 M F Braby and M R Williams
Another important consideration in inventory is seasonality
or the temporal component of species richness (Hill 1999;
Ashton et al. 2011; Ribeiro et al. 2015). Most insects show
pronounced seasonal patterns in appearance and abundance, in
both temperate and tropical latitudes. It is therefore important
that not only the temporal distribution of priority taxa be
predictable among years (Table 1), but that surveys are under-
taken at an appropriate time of year to optimise the time when
the taxon is likely to be present and sampled.
Species discovery curves
In inventories, usually only a fraction of the total species
present is sampled, and it is often desirable to know the total
number of species by estimating the richness of the unsampled
fraction. This is particularly important when patterns of
species richness are compared among sites or regions to
inform conservation management. A commonly used method
of estimating the adequacy of inventory is through species
accumulation curves and species discovery curves. Species
accumulation curves estimate the expected species richness of
a site or landscape and provide an estimate of how well a given
area has been surveyed (Colwell & Coddington 1994). In
contrast, species discovery curves are used to estimate the total
number of extant species within the group and provide an
estimate of how well a particular taxonomic group has been
sampled and documented regionally or globally (Solow &
Smith 2005; Bebber et al. 2007; Joppa et al. 2011a).
For species discovery curves, the discovery process involves
the stages of field collection of samples, curation of material
and recognition of new taxa, followed by taxonomic descrip-
tion and formal publication of the new name (Bebber et al.
2010). The usual approach with species discovery curves is to
extrapolate from this temporal pattern of publication of known
species, by statistical modelling of the cumulative discovery
record with the expectation that when the inventory
approaches completion the rate of new species discoveries will
diminish as they become harder to find.
Thus, there are two quantities that need to be determined in
species discovery curves: the number of known species at a
given point in time and the estimated number of species that
remain to be discovered (i.e. the number of as-yet-unknown or
missing species). The estimated total number of species of the
group is thus the sum of these two numbers. However, such
analyses do not take into account potential problems associated
with issues of nomenclature, synonymy, species delimitation
and ranks below species level (e.g. subspecies), not to mention
the species concept adopted. Joppa et al. (2011a) recommended
incorporating aspects of the taxonomic effort or taxonomic
efficiency into the model to account for some of these issues.
Several authors have also discussed problems with the species
concept used, noting that the phylogenetic species concept not
only results in taxonomic inflation but also has the most limited
utility in practical biodiversity conservation (Agapow et al.
2004; Isaac et al. 2004; Frankham et al. 2012).
Compared with most other insects, butterflies have long held
a prominent place in scientific and popular cultures with several
© 2015 Australian Entomological Society
centuries of systematic research. Indeed, Kristensen et al.
(2007) concluded that the global inventory of butterflies was
probably close to completion, with nearly 20 000 described
species, and that comparatively few species remained to be
discovered/named compared with the micro- and macro-moths.
Given the widespread use of butterflies in conservation biology
(see Sands & New 2002; New & Sands 2004; New 2011 for
review inAustralia), we ask, in the following example, how well
is the butterfly fauna ofAustralia known taxonomically and how
close is the taxonomic inventory to completion at the regional
scale? The important point here is that if taxonomic resolution
is poor, then their effectiveness as an umbrella group (New
1997) to identify biodiversity values for conservation manage-
ment is likely to be compromised.
The species discovery curve of the Australia butterfly fauna
(Braby 2010b) suggests the inventory for this group is almost
complete (Figs 2,3). The rate of taxonomic description of new
species over the past 240 years shows a high level of activity
during the 19th century, reaching a peak around 1870–1910,
followed by a steady decline during the 20th century (Fig. 2).
However, closer analysis of the past 110 years shows an
increasing rate in the number of species recognised (Fig. 3).
During the first 70 years of the 20th century there was an
exponential increase in the rate of taxonomic knowledge; from
1970 to 2000 the rate steadied and increased linearly; and
during the last decade there is a hint that the trajectory is
starting to level off towards an asymptote, as would be
expected as the taxonomic inventory nears completion. New
species have been added as a result of three processes:
undescribed species being discovered and named, especially
from remote areas of the country; species being recorded from
Australia for the first time but were previously known else-
where (e.g. from south-east Asia or mainland New Guinea);
and resolution of taxonomic uncertainty, such as species com-
plexes (e.g. subspecies proving to be specifically distinct on
closer examination). This trend of increasing taxonomic
knowledge parallels the more general trend in the inventory of
butterflies worldwide in which the rate of new species being
recognised has increased over the past 50 years (Kristensen
et al. 2007). Similar patterns have been documented for many
other taxa worldwide (Joppa et al. 2011b).
Statistical modelling of the 1903–2010 data (Fig. 3) predicts
an asymptote of around 472 butterfly species in Australia, based
on projections from past taxonomic research effort. Extrapolat-
ing from the temporal pattern of the species discovery curve
suggests that in 2010, about 91% of the total estimated fauna
had been documented. In other words, the estimated number of
as-yet-undescribed species is 42 (9%). By analogy with other
taxa, many of these ‘missing’ species (some of which may be
already known in museum collections but yet to be described)
are likely to be rare and local, with small geographic ranges,
cryptic, and concentrated in biodiversity hotspots (Joppa et al.
2011a; Scheffers et al. 2012). Bebber et al. (2007) questioned
the reliability of estimating the total number of extant species in
taxonomic groups based on plotting cumulative frequency
curves. They identified two constraints on being able to accu-
rately predict total species number: the completeness of the
 Biosystematics and conservation biology 5

Figs 2,3. Species discovery curve for

Australian butterflies showing: (2) the
rate of taxonomic description of recog-
nised species (black) and subspecies
(cross-hatched) with an Australian-type
locality, in 5-year intervals (after Moulds
1999); and (3) the relationship between
the cumulative number of species recog-
nised over time according to major publi-
cations during the past 100 years (after
Braby 2010b). Fitted curve is a general-
ised logistic function S = 329 + 144 /
(1 + e98.3-0.0493*t) (r2 = 0.9975), where S is
480 3 472
the number of species, and t is the year
(1880–2090). The quantity γ is the 460
‘knowledge gap’ of the estimated number γ = 42
Number of species

of missing species that remain to be dis- 440

covered, and the quantity λ is the time
estimated to complete the taxonomic 420
inventory. Modelling the rate of species
400
accumulation based on taxonomic
research effort over the past 100 years
380
suggests that about 91% of the Australian
λ = 80
butterfly fauna has been catalogued so far: 360 2090
the model predicts an estimated total
number of species of 472 (γ = 42 species), 340
which will be reached by the year 2090
(λ = 80 years) if it is assumed that the 320
1880 1900 1920 1940 1960 1980 2000 2020 2040 2060 2080 2100
discovery effort continues at similar
levels to the past century. Year

current inventory, and variations in the discovery process (i.e.
taxonomic and publication method, access to unexplored areas,
and the species concept adopted). They found that unless the
inventory was approaching completion (>90%), extrapolations
from existing data yielded very large errors. Because the taxo-
nomic inventory of the Australian butterfly fauna is close to
completion, we concluded that the upper estimate of around 472
species is a reasonable estimate. However, should a different
species concept be adopted with greater emphasis on DNA
barcodes/molecular markers to reveal cryptic species (Adams
et al. 2014), estimates for the Australian butterfly fauna are
likely to be substantially higher.
MONITORING
Monitoring is the process of measuring temporal changes in
species abundances, species richness or species assemblages
(or other components of biodiversity, such as geographic range
size/boundaries) (Field et al. 2007). It is used to gauge how a
site or location might be changing over time from a predeter-
mined standard (baseline), usually in relation to environmental
features (e.g. threats, habitat restoration) (Stork et al. 1996;
Samways 2005). Importantly, monitoring should be integrated
adaptively to improve land management, for instance, to assess
the effects of management actions to mitigate a particular
threatening process or to measure the recovery of damaged
ecosystems through ecological restoration (Nichols &
Williams 2006; Lindenmayer & Gibbons 2012; Lindenmayer
et al. 2012, 2013). It provides a strategic framework for facili-
tating management options, and is really the only way to
effectively assess the success of conservation efforts and return
on investment in natural resource management and biodiver-
sity conservation. In essence, monitoring provides the early
warning system for detrimental environmental change.
Despite being one of the world’s mega-diverse nations, Aus-
tralia has a poor record of monitoring its biodiversity, and an
appalling record of loss/decline of that biodiversity (Lindenmayer
& Gibbons 2012; Lindenmayer et al. 2012). Terrestrial insects in
Australia have rarely been incorporated into biodiversity moni-
toring programmes, and this has arisen largely because of a lack of
taxonomic knowledge (Kitching 1993a; Ponder & Lunney 1999),
a traditional ‘vertebrate wildlife culture’ among government con-
servation agencies that have little familiarity with insects/
invertebrates, and a general lack of taxonomic expertise and
facilities within those agencies (Kitching 1993a; Andersen &
Majer 2004; Andersen et al. 2004; Samways 2007; New & Yen
2013; Yen & New 2013). Taxonomic knowledge, which is
© 2015 Australian Entomological Society
6 M F Braby and M R Williams
perhaps the greatest impediment to the science of insect conser-
vation (Kitching 1993a; New 1996b; Gerlach et al. 2013), has two
dimensions: availability of taxonomic expertise, and the level of
knowledge according to access to published literature and/or
scientific reference collections. These two elements need to be
considered because groups that may be poorly known taxonomi-
cally, with a large proportion of undescribed species, can still be
useful for monitoring by parataxonomists using a morphospecies
approach (Cranston & Hillman 1992; Beattie & Oliver 1994;
Oliver & Beattie 1996a,b; Farr et al. 2011). However, this infor-
mal taxonomic approach is only possible for local studies in
which specialists with well-sorted reference collections of
vouchered material are available – Australian ants (Andersen &
Majer 2004; Andersen et al. 2004) and macromoths (Kitching
et al. 2000; Ashton et al. 2011; Kitching & Ashton 2013) being
excellent cases in point. The morphospecies approach is less
useful when data from a range of studies are integrated into larger,
regional or global data sets (Novotny & Miller 2014) because
morphospecies do not serve as a convenient unambiguous inter-
national label to facilitate communication among scientists that
taxonomic names provide.
In monitoring, as for inventory, typically a limited taxon set
is selected and used as a biological indicator (Andersen 1999;
Nakamura et al. 2003, 2007; Andersen & Majer 2004).
However, the prioritised taxa are often different to those
selected for inventory because different criteria are required to
fulfil the objective(s) of monitoring. The key criteria include
good taxonomic knowledge, informativeness as bioindicators,
predictability in both space and time with sufficiently high
detectability, and the ability to be incorporated into a sampling
regime using a standardised, quantitative method that is rela-
tively simple and cost-effective (and, preferably, one useable
by non-specialists).
A key part of monitoring is detectability, which is the prob-
ability that a species present at a site is actually detected.
Detectability is a crucial aspect that needs to be considered in
designing monitoring protocols because it allows for correc-
tion between the relative value and true value to be estimated
according to the sampling unit adopted (see Gibson & New
2007; Kéry & Plattner 2007; Williams 2008, 2009; Bried &
Pellet 2012; Pellet et al. 2012; Ribeiro et al. 2015 for moni-
toring programmes of butterflies and day-flying moths).
Detectability may vary among observers, as well as in space
(e.g. between sites) and time (e.g. between seasons), and
recent statistical methods employing occupancy modelling
enable the probability of detection to be quantified
(MacKenzie et al. 2002; MacKenzie & Royle 2005).
MacKenzie and Royle (2005) recommended that when detec-
tion probability is high (>0.5) sampling units should be sur-
veyed a minimum of three times to avoid false absences (i.e.
species were present but were not detected).
Bioindication
Bioindication is a critical element of monitoring and it has
several components. For terrestrial insects, McGeoch (1998,
© 2015 Australian Entomological Society
2007) recognised three different categories of biological indi-
cators: environmental, ecological and biodiversity depending
upon the goals, objectives and questions proposed for the
monitoring programme. Environmental and ecological indica-
tors are the categories most frequently used in monitoring
(Gerlach et al. 2013): the former respond predictably to a
change in environmental state or conditions (e.g. pollutants,
pesticides, acidification), whereas the latter are sensitive to the
impacts of environmental stress (e.g. habitat disturbance, frag-
mentation, climate change) (McGeoch 1998, 2007). Monitor-
ing for these effects may be undertaken in two complementary
ways: measuring the temporal changes in relative abundance
of ‘characteristic’ indicator species highly specific to a particu-
lar habitat or ecological state over time, or measuring the
spatial changes in relative abundance of ‘detector’ indicator
species across two or more habitats or ecological states over
time (McGeoch et al. 2002; McGeoch 2007).
The selection (and testing) of terrestrial insect bioindicators
is difficult, and few rigorous bioindication systems have been
developed for these organisms (McGeoch et al. 2002;
McGeoch 2007). Dufrêne and Legendre (1997) developed a
quantitative indicator value (IndVal) based on habitat specific-
ity (relative abundance among sites) and fidelity (occupancy or
frequency of occurrence among sites) to identify potential
indicator species, and this method has been used to discriminate
among beetle and ant assemblages for future monitoring of
rainforest restoration in south-eastern Queensland (Leach et al.
2013). In contrast, in an extensive study of a wide range of
invertebrates associated with jarrah (Eucalyptus marginata)
forests of south-west Western Australia for timber production,
Farr et al. (2011) found none that served as effective indicators
for monitoring the recovery of harvested (disturbed) forest
because of substantial species turnover among sites and years.
Globally, the taxa used most frequently as bioindicators
include Coleoptera (especially ground beetles), Hymenoptera
(especially ants) and Lepidoptera (especially butterflies) (see
McGeoch 2007 for review). Gerlach et al. (2013) recom-
mended that for the various structural attributes of the envi-
ronment, different suites of terrestrial invertebrates should be
used as bioindicators – soil (isopods, mites), ground layer
(landsnails, millipedes, spiders, ground beetles, ants) and
foliage (theridiid spiders, orthopterans, leaf beetles, butterflies
and some diurnal moths, ants). In Australia, terrestrial insects
or other invertebrate groups targeted, or proposed, for moni-
toring include spiders, spring-tails (Collembola), dragonflies,
grasshoppers, some families of beetles, ants, butterflies and
some groups of moths (see Kitching 1993a; New 1996b, 1998;
Andersen et al. 2001; Andersen et al. 2004; Lomov et al.
2006; Michaels 2007; Kitching & Ashton 2013; Leach et al.
2013 for review).
Butterflies and other diurnal Lepidoptera have been identi-
fied for use as bioindicators globally (e.g. Ehrlich 1992, 2003;
Warren et al. 2001; Thomas et al. 2004; Vane-Wright 2005,
2008; van Swaay et al. 2006; van Swaay et al. 2008; Merckx
et al. 2013). In the northern hemisphere, particularly in Europe
and North America, this guild has been used extensively as
environmental and ecological indicators (see Gerlach et al.

2013 for review). These insects possess many of the recom-
mended criteria for bioindication, including being taxonomi-
cally well known, easily recognised and readily identified (most
species are day-flying, relatively large and often brightly col-
oured, and regional field guides are usually available); easily
sampled or observed; sampled individuals are abundant; spatial
and temporal distributions are predictable, they exhibit a wide
range of spatial patterns (including narrow-range endemics)
and host-specificities (including ecological specialists); and are
sensitive to a multitude of threatening processes, such as habitat
loss, disturbance (e.g. through inappropriate fire regime), frag-
mentation, degradation (through spread of invasive species),
pollution, pesticides and climate change. Moreover, baseline
biological and distributional data are available for many
species, and the geographical range can be determined to a high
degree of accuracy based on historical distribution records.
Such historical data can also be used to assess changes in extent
of occurrence and area of occupancy in relation to land use
practices or climate change (Warren et al. 2001), indices that
are frequently unobtainable for most other insect groups (New
1991a, 2009; New et al. 1995; Samways 2005).
Monitoring of butterflies in Australia, however, is still very
much in its infancy, with most attention focused on individual
species rather than assemblages. Well-established monitoring
programmes designed to assess changes in relative abundance
of individual threatened species of Australian diurnal
Lepidoptera include Paralucia pyrodiscus (Doubleday, 1847)
(Braby et al. 1999; New 2011), Ornithoptera richmondia
(Gray, [1853]) (Sands 2008; Sands & New 2013) and Synemon
plana Walker, 1854 (Richter et al. 2013). The few ecological
studies that have assessed the composition and species rich-
ness of assemblages (and sometimes seasonal changes in
abundance) through repeated sampling have tended to be short
term (maximum of 2.5 years) (Braby 1995; Hill 1999;
Haywood & Wilson 2002; Collier et al. 2006; Lomov et al.
2006; Ginn et al. 2007; Williams 2009, 2011; Franklin 2011).
An exception is the study by Newland (2006) who compared
the species richness, composition and relative abundance of
butterflies at a hilltop site in northern New South Wales before
and after disturbance. However, only two comparisons were
made, 12 years apart, rather than a series of repeated samples
designed to detect temporal changes over time. National long-
term recording schemes to evaluate broad changes in spatial
distribution and relative abundance of all butterfly species are
currently not in existence in Australia. This contrasts markedly
with schemes such as those in the northern hemisphere pro-
duced by the Mapping European Butterflies and related moni-
toring programmes (van Swaay et al. 2008; Kudrna et al.
2011), and large collaborative initiatives such as the Tropical
Andean Butterfly Diversity Project in South America
(Willmott et al. 2011; Merckx et al. 2013).
THRE ATE NED SPE C IE S
Lists of threatened species and threatened ecological commu-
nities are compiled at various levels of government. In addition
Biosystematics and conservation biology 7
to the international threatened species list prepared by the
International Union for the Conservation of Nature (the IUCN
Red List of Threatened Species) (Rodrigues et al. 2006), Aus-
tralia has a national list as well as individual state and territory
lists (New & Yen 2013). These lists provide a temporal snapshot
of which components of biodiversity are facing extinction
according to their conservation status. Although the conserva-
tion status of relatively few insects has been assessed globally
compared with vertebrates (Rodrigues et al. 2006), the listing
of threatened insects has the advantages of raising the profile of
individual species (especially the more iconic or ‘charismatic’
species) and their conservation needs. Listing also identifies
key threatening processes impacting upon the species that may
also be symptomatic of the wider habitat or ecological commu-
nity to which the species belongs. That is, a threatened species
can act as a ‘flagship’ for promoting the conservation of other
insects and allied organisms and/or their habitats. Moreover, the
co-occurrence of multiple threatened species can be used to
identify and prioritise important areas for conservation through
the compilation and analysis of digital spatial data, both glob-
ally and regionally (Rodrigues et al. 2006).
Disadvantages of using threatened species in biodiversity
conservation arise when lists are inaccurate, threatening pro-
cesses are either not identified or not addressed strategically,
and taxonomic knowledge is poor, all of which may lead to
inefficient or inappropriate expenditure of scarce conservation
resources (Garnett 2013; Lindenmayer et al. 2013). Moreover,
single species-oriented conservation may divert a substantial
proportion of available resources and attention while achiev-
ing little practical outcome or broader environmental benefit
(New 1991b). Ideally, such a ‘fine-filter’ species-level
approach should be complemented with a ‘coarse-filter’ col-
lective approach to protect ecological communities, biotypes
or landscapes (Yen & Butcher 1997; Samways 2007; Merckx
et al. 2013).
Threatened species research and successful conservation
management usually involves six sequential steps: (1) protec-
tive legislation, either at the state/territory and/or national
level; (2) evaluation of conservation status; (3) identification of
threatening processes; (4) preparation of a recovery or action
plan with recommendations giving priority actions; (5) imple-
mentation of recovery plan through a recovery team to miti-
gate threats and achieve practical conservation management
(which may include security and/or habitat restoration, or
captive breeding programme followed by reintroduction or
translocation of populations); and (6) monitoring to gauge
effectiveness of management actions, with capacity to refine
management if required (New & Yen 1995; New et al. 1995;
New 1996a, 2007; Yen & New 2013).
The legislative framework for protection (step 1), which
occurs at state, national and international levels, is necessary
because it provides the legal basis on which conservation
measures can be initiated. In Australia, the insects that have
received most attention for threatened species listing are the
dragonflies and butterflies – groups for which taxonomic/
ecological knowledge is relatively complete (New & Sands
2003).
© 2015 Australian Entomological Society
8 M F Braby and M R Williams
Table 2 Summary of the five criteria (A–E) proposed by the IUCN (IUCN Standards and Petitions Subcommittee 2014) to evaluate
if a taxon belongs in a threatened category (Critically Endangered, Endangered or Vulnerable)

A. Declining population (past, present and/or projected). Decline measured over the longer of 10 years or 3 generations.
B. Small geographic range size, fragmentation, decline or extreme fluctuations. Decline in geographic range in the form of extent of occurrence
(EOO) and/or area of occupancy (AOO), and severely fragmented or small number of locations.
C. Small population size and fragmentation, decline or extreme fluctuations. Low numbers of mature individuals and ongoing decline.
D. Very small population or very restricted distribution. Very low numbers of mature individuals and restricted AOO or small number of locations.
E. Quantitative analysis of extinction risk (e.g. population viability analysis). A high probability of extinction in the wild within defined timeframes.

At least one criterion must be satisfied for a taxon to be evaluated as threatened.

IUCN, International Union for the Conservation of Nature.

Conservation status evaluation (step 2) determines the
extent to which a taxon is threatened and therefore at risk of
extinction under current circumstances. It is usually based on
objective criteria developed by the IUCN Red List guidelines
(Rodrigues et al. 2006; IUCN Standards and Petitions
Subcommittee 2014) for classifying species at high risk of
global extinction (Table 2).
Under the IUCN guidelines, criteria for status evaluation
include an analysis of quantitative measures such as popula-
tion size, changes in abundance or spatial distribution
(Rodrigues et al. 2006). This information does not necessarily
provide a framework for assessing priorities for action, but
when assessed accurately does highlight urgency of conserva-
tion need (IUCN Standards and Petitions Subcommittee
2014). However, most of the IUCN criteria are rather limited
in scope for insects because they require a sound understand-
ing of population trends, which are not available for most
species (Sands & New 2002; New & Sands 2003; Warren et al.
2007). The most frequently used measure for insects is geo-
graphic range size (Criteria B and D) in the form of either
extent of occurrence (EOO) or area of occupancy (AOO),
together with evidence of severe fragmentation and/or decline,
which is typically based on an analysis of critical habitat and
threatening processes. This criterion has been used success-
fully to evaluate the conservation status of several butterflies in
Australia in recent years (Braby & Douglas 2004, 2008;
Eastwood et al. 2008; Braby 2010a). An exception to this is
the Graceful Sun-moth, Synemon gratiosa Westwood, 1877, in
Western Australia, which was assessed against Criterion A by
using standardised sampling methods (transect counts) and
treating the resulting counts as an index of abundance for the
species (IUCN Standards and Petitions Subcommittee 2014).
Changes in population size over a 10-year time frame were
then estimated by determining the effect of projected future
habitat loss (clearing of the moth’s habitat for urbanisation) on
the population index (M.R. Williams unpublished data).
It is noteworthy that the first questions addressed in formally
assessing a species for listing and status evaluation relates to
taxonomy: What is the scientific name of the species? Is the
species conventionally accepted? Is the species taxonomically
distinct? Threatened species conservation invariably assumes
that the taxonomic status or species delimitation is well under-
stood, which frequently is not the case for insects. Indeed, the
lack of taxonomic information on many species is a major
impediment in evaluating conservation status and developing
recovery plans for effective conservation management (New &
© 2015 Australian Entomological Society
Yen 2013). In three of the threatened butterfly cases noted
above (Braby & Douglas 2004, 2008; Eastwood et al. 2008),
the species-level status (and conservation status) were not
resolved until more detailed taxonomic investigations were
undertaken: one species (Candalides noelkeri – Endangered)
was previously considered to be a geographical form of a more
widespread species, and two species (Ogyris halmaturia –
Endangered; Jalmenus eubulus – Vulnerable) were formerly
regarded as subspecies of more widely distributed localised
species. In the case of Synemon gratiosa from south-western
Western Australia, the taxonomic boundary of this species is
clouded by the fact that specimens are difficult to distinguish
morphologically from the closely related S. jcaria R. Felder,
1874. Genetic analysis based on mitochondrial DNA
(mtDNA) has also shown that the two taxa differ by only
1–2%, and subsequent surveys have located populations
that likely represent hybrids between these two species (M.R.
Williams, unpublished data).
Conservation management of
significant populations
The genetics revolution – the explosion in the use of genomic
data and the methods to analyse them (Wolfe & Li 2003) – has
greatly increased the capacity of taxonomy to deliver outcomes
that are directly applicable to conservation issues, including the
recognition of conservation units based on patterns of genetic
variation within species (Moritz 1994, 2002; Waples 1995;
Crandall et al. 2000; Funk et al. 2012). Knowledge of the
genetic structure and diversity of populations within species can
be used to inform conservation management of threatened
species (Fig. 1). This is particularly important because the goals
of conservation are to maintain the adaptiveness and adaptabil-
ity of organisms (Frankel & Soulé 1981; Frankham 2010).
Much of the conservation assessment of Australian butterflies,
for example, has focused on subspecies, many of which are of
doubtful status (Sands & New 2002), with little attention given
to the genetic diversity and structure of populations of threat-
ened species (Collier et al. 2010). Although the current legis-
lative framework in Australia does allow for the conservation
status of subspecies to be assessed separately, there is limited
provision for the management of Evolutionary Significant
Units (ESU) within species (Braby et al. 2012). The Australian
Environmental Protection and Biodiversity Conservation Act
does allow for the recognition of important populations as
‘distinct’, but for widely distributed species across multiple
 Biosystematics and conservation biology 9
land tenures and state boundaries, it may be more practical to
0 100 200
focus conservation efforts on managing ESUs or distinct popu-
Kilometres
lation segments (i.e. adaptive genetic potential) as subset areas
within the overall geographical range without the need to list MACKAY
and protect the species or subspecies as a whole. 4
For example, a recent study investigating the phylogeo-
graphy and population genetics of a threatened butterfly in
ROCKHAMPTON
coastal South Australia, Theclinesthes albocinctus
(Waterhouse, 1903) ‘southern coastal form’ (Eastwood 2006;
Collier et al. 2010), found that populations in a fragmented
landscape were at greater risk of extinction because of their
smaller population size and reduced genetic diversity as a
consequence of ongoing habitat loss and isolation. Manage- Jalmenus eubulus
ment of these isolated, remnant populations may require Jalmenus evagoras BRISBANE

several approaches, including augmentation of existing popu-

lations, restoration of breeding habitats or translocation to
QLD
more suitable sites (Collier et al. 2010). Thus, with appropriate NSW

use, this revolution in conservation genetics and the delinea-

tion of distinct evolutionary units within species has the poten-
5 0 100 200
tial to deliver better outcomes for conservation management of
Kilometres
threatened species, particularly those in anthropogenic modi-
fied landscapes, by providing fundamental information on MACKAY
minimum viable population size, connectivity and the mainte-
nance of genetic diversity (Frankham 2010; Mills 2013).

ROCKHAMPTON
A case study: Jalmenus eubulus and
brigalow woodland
The following example serves to illustrate how biosystematics
incorporating genetic evidence resolved long-standing taxo-
nomic confusion and clarified the conservation status of a
threatened species, the Pale Imperial Hairstreak Jalmenus BRISBANE

eubulus Miskin, 1876. This species is restricted to old growth

brigalow woodland habitats dominated by Acacia
QLD
harpophylla, a highly threatened ecological community NSW
limited to central and southern Queensland and northern New
South Wales (Figs 4–6). For many years J. eubulus was not
6 0 100 200
formally recognised as a species, being treated as a subspecies
Kilometres
of J. evagoras (Donovan, 1805), which occurs widely in
south-eastern Australia. However, recent evidence from MACKAY

▶
ROCKHAMPTON
Figs 4–6. Spatial distribution of Jalmenus eubulus showing: (4)
extent of occurrence (hatched area) based on known sites (open
circles), together with the spatial distribution of its sister species
J. evagoras in Queensland (QLD; closed circles represent point
localities, with thick black line representing geographical bound-
ary); (5) extent of its brigalow habitat (shaded black) before clear-
ing; and (6) after clearing based on extent of remnants in 2003 BRISBANE
(after Eastwood et al. 2008). Taxonomic revision of the
J. evagoras complex revealed that it comprised two species that
are parapatric, with a very narrow range of overlap between QLD
Kroombit Tops and Toowoomba, and provided the foundation for NSW

a more rigorous evaluation of the conservation status of

J. eubulus, which had been grossly underestimated. Under Inter-
national Union for the Conservation of Nature Red List criteria,
the species’ conservation status nationally would be Vulnerable.
© 2015 Australian Entomological Society
10 M F Braby and M R Williams
multiple data sources (wing colour pattern, morphology,
ecology and genetics) revealed fundamental differences
between the two taxa, including fixed differences in the
mitochondrial genomes with absence of matrilineal gene flow,
whereas allozyme data showed significant structure within and
between populations of both species consistent with recent
diversification (Eastwood et al. 2008).
Given that species-level status/delimitation is a hypothesis
that is refined as additional evidence becomes available
(Yeates et al. 2011), one might assume that the conservation
status of a taxonomic entity, whether it is a species, subspecies,
ESU etc., should not be related to its taxonomic categorisation.
However, in the case of J. evagoras complex poor taxonomic
knowledge of J. eubulus had not only led to considerable con-
fusion over its delimitation and identification, but also to poor
understanding of its distribution and ecology, and conse-
quently its conservation status was grossly underestimated.
Prior to the taxon being recognised as a distinct species from
its closely related sister species J. evagoras (Eastwood et al.
2008), its conservation status was considered to be ‘Least
Concern’ (Sands & New 2002).
Clarification of the taxonomy of J. eubulus provided the
basis for a revised, clearer assessment of its spatial distribution
(Fig. 4) and conservation status under the IUCN Red List
criteria. This assessment indicated that it would qualify as a
threatened species nationally (Vulnerable), Vulnerable in the
State of Queensland, and Critically Endangered in New South
Wales (Eastwood et al. 2008). That is, information on its
spatial distribution, critical habitat and threatening processes
indicated that the geographic range of the species had an
estimated AOO of less than 2000 km2; was severely frag-
mented, known from few widely dispersed locations; and the
extent or quality of its habitat, which is poorly conserved and
currently regarded as a threatened ecological community
under Queensland legislation, continued to decline. Analysis
of the pre-cleared and remnant extent of old-growth brigalow
within the known spatial boundary of J. eubulus showed that
only 5.3% of habitat remained uncleared (Figs 5,6). In other
words, almost 95% of habitat or potential habitat of J. eubulus
has been lost because of land-clearing practices since Euro-
pean settlement, most of which had occurred during the past
50 years. Moreover, only a small fraction of this habitat
(c. ∼ 0.5% of the estimated total area of original extent) is
reserved in protected areas, and these reserves face additional
threatening processes associated with edge effects, including
weed invasion and fire, and require conservation management
(Eastwood et al. 2008).
BIOSYSTE MATICS, C AN IT D E LIVER?
It is often argued that we cannot hope to conserve and manage
our biological diversity effectively if we do not know the
makeup of the Earth’s biological constituents (e.g. May 1988,
1994; Wilson 2004; Wheeler et al. 2012). However, knowing
the constituent taxa is of limited use, unless there is basic
understanding of where those taxa occur in relation to land-
© 2015 Australian Entomological Society
scape features and tenure (i.e. their spatial distribution). In
other words, it is equally important to map the distributions of
species as it is to recognise them formally (Wheeler et al.
2012). Systematic monographs and taxonomic revisions (and
the museum collections on which they are based) provide these
two basic elements: the circumscription of species, and the
geographical distribution of those species.
Two major challenges for biosystematics are therefore to:
(1) systematically catalogue and map the Earth’s known
species, and (2) discover, describe and document new species
(the as-yet-unknown or missing species). The former task
includes resolving synonymy of the numerous cases where
species have been described more than once, redescribing and
diagnosing taxa currently placed in synonymy that prove to be
specifically distinct, and resolving other problems of nomen-
clature (e.g. homonyms). Both tasks are needed in order to
optimise conservation effort of the world’s biodiversity
(Scheffers et al. 2012; Costello et al. 2013; Stork & Habel
2014). Other challenges for biosystematics are to: (3) recon-
struct the evolutionary history or tree of life and establish the
phylogenetic relationships among species and (4) incorporate
phylogenetic diversity (taxonomic distinctiveness) as a com-
ponent of biodiversity into conservation planning. The ration-
ale behind the latter task is that phylogenetic diversity captures
the evolutionary legacy or distinctiveness of a taxon or taxo-
nomic group, but so far this has apparently contributed little
towards practical nature conservation (Winter et al. 2013).
Isaac et al. (2007), however, have developed a method for
incorporating both phylogenetic diversity and extinction risk
for threatened species that are both ‘evolutionarily distinct and
globally endangered’ (i.e. EDGE species), which can be used
to generate global priority lists for conservation. Moreover,
there is a sense of urgency to complete these challenges
because of increasing levels of extinction against a background
of declining systematic expertise (Wilson 1987, 1992, 2004;
Dirzo & Raven 2003; Mallet & Willmott 2003; Yeates &
Raven 2004; Oliver & Lee 2010; Probert 2010; Maddison
et al. 2012; Wheeler et al. 2012; Yen & New 2013). Not only
are professional taxonomic specialists across all biota declin-
ing, but this expertise is mismatched against the species rich-
ness of taxa and the geographical location of biodiversity (e.g.
Gaston & May 1992; New 1996b; Godfray 2002; Tautz et al.
2003; Wheeler et al. 2004; Yeates 2009; Probert 2010; Boero
2010; Löbel 2014).
The general view is that the task of describing and naming
all of the world’s species is probably unachievable (New 1993,
1996b; Stork et al. 1996; Wheeler et al. 2012; Costello et al.
2013) and that complete inventories of invertebrates even at
relatively small spatial scales are logistically impossible
(Harvey et al. 2011). Kitching (1993a) estimated that between
50–80% of species among some Australian rainforest canopy
taxa are unnamed, and Yeates et al. (2003) and Austin et al.
(2004) estimated that at least 75% of the Australian Insecta
(c. 205 000 known species) have not been formally described
or are yet to be discovered. Kristensen et al. (2007) estimated
that only about one third of the global Lepidoptera (c. 160 000
species described so far) has been systematically catalogued,

with around 1000 new species being described annually in
recent years. Insect conservation in Australia is thus a formi-
dable task, being carried out by a small number of informed
specialists using incomplete taxonomic and biological knowl-
edge (Samways 2007; New & Yen 2013; New & Samways
2014). Not surprisingly, insects receive low priority on the
conservation agenda in all state agencies, and setting priorities
for insect conservation is difficult and sometimes controver-
sial.
Several solutions have been proposed to overcome this
enormous taxonomic impediment for insect conservation
biology. One proposal is that taxonomic attention should be
diverted towards small geographical regions known to
support high levels of endemism where detailed inventories
may be possible (Samways 2007), or focus on a limited set
of ‘priority’ taxa (Ehrlich 1992, 2003; Kitching 1993a,b;
New 1993; Stork et al. 1996; Samways 2007; Kitching &
Ashton 2013). These proposals may be preferable than
attempting to catalogue the entire biological diversity of the
planet because most species will almost certainly go extinct
before they are even discovered, let alone named and classi-
fied. Ehrlich (1992, 2003) argued that this prioritised taxon
set be used for inventory and monitoring – to design conser-
vation area networks, to evaluate ecosystem health and to
gauge the effectiveness of management and persistence of
ecological and evolutionary processes. Given finite resources
for taxonomic research, which groups should be prioritised
for conservation biology? The answer to this question will
require careful consideration – we suggest at least two of the
essential criteria listed in Table 1 be considered for selection
of taxonomic groups: (1) the taxon is reasonably well known
taxonomically (i.e. total inventory is estimated to be 90%
complete, and/or morphospecies have been circumscribed
through the availability of parataxonomists and well-curated
reference collections); and, (2) the taxon is known to be
informative as bioindicators. Terrestrial insect and other
invertebrate groups in Australia that fit both criteria include
landsnails (Stanisic 1999; Slatyer et al. 2007; Braby et al.
2011), some spiders (Churchill 1997; New 1999), dragonflies
(Brooks 1996; Clausnitzer et al. 2009), dung beetles (Hill
1996; Spector 2006; Nichols & Gardner 2011; Monteith
2015), ground beetles (Grimbacher et al. 2007; Michaels
2007; but see New 1998), butterflies, some diurnal moths,
some nocturnal macromoths (New 1996b, 2011; Kitching
et al. 2000; Lomov et al. 2006; Ashton et al. 2011) and ants
(Andersen & Majer 2004; Andersen et al. 2004).
The second proposal is to accelerate the rate at which new
species are discovered and described, at least by an order of
magnitude (Wiens 2007; Bacher 2012; Wheeler et al. 2012;
Costello et al. 2013; Novotny & Miller 2014). This proposal
includes several initiatives, such as increasing the efficiency of
the publication process (e.g. moving towards early view pub-
lication, online registration through ZooBank (Yeates 2009));
open-access to comprehensive online taxonomic databases and
literature, and development of other information technology
platforms or bioinformatics (e.g. Global Biodiversity Informa-
tion Facility, Integrated Taxonomic Information System, Tree
Biosystematics and conservation biology 11
of Life, All Taxa Biodiversity Initiative); increased resources
for specimen collection and curation, integrated with digital
imaging and DNA barcoding; and, perhaps most importantly,
training of more taxonomists and better coordination between
amateur and professional scientists. Boero (2010) recom-
mended that for the science of taxonomy to survive it must
become a multidisciplinary science, integrating molecular
biology and computational science with traditional morpho-
logical approaches. We suggest that a fourth dimension be
added to raise the profile/relevance of biosystematics – inte-
gration and collaboration with conservation biologists, conser-
vation agencies and relevant community groups (e.g. citizen
science programmes).
The cost in terms of the amount of both time and effort
required to complete the taxonomic inventory (should that be
possible) is debatable, but clearly considerably more invest-
ment is needed than at present (Wheeler et al. 2012; Costello
et al. 2013). Although there has been a global increase in the
number of people describing species (Joppa et al. 2011b;
Bacher 2012; Costello et al. 2013), the professional work-
force of traditional taxonomists has been in a steady state of
decay for several decades, at least in Australian entomology
(particularly those associated with premier natural history
museum collections such as the Australian National Insect
Collection). In our opinion, many of the people who describe
new species or who have their name attached to publications
describing new species, are not necessarily experts nor ‘good’
taxonomists. Many new generation taxonomists have little
or no formal training in the principles of the International
Code of Zoological Nomenclature (ICZN) (International
Commission on Zoological Nomenclature 1999), have limited
understanding of species concepts and the importance of
types, do not prepare formal diagnoses, have limited knowl-
edge of morphology, and typically publish works of single
species rather than the comprehensive revisions of higher taxa
(e.g. genera, families) that require high levels of taxonomic
expertise.
Some proposals to accelerate taxonomic description, such
as web-based taxonomy and/or side-stepping peer review
(Scoble 2004; Maddison et al. 2012), are not without problems
and inevitably will produce poor quality science, resulting in
long-term taxonomic chaos that will eventually need resolu-
tion. Moreover, taxonomists who reveal the existence of
unrecognised taxa on the internet run the risk of having those
candidate taxa described by unscrupulous individuals using
non-refereed, and often in privately published, works (Oliver
& Lee 2010). The rules of the ICZN currently do not regulate
the quality of taxonomic research, just the use of taxonomic
names.
We contend that predictions that the global inventory
(recent estimates vary from c. 10 million species (Chapman
2009) to 3.7–6.1 million species (Hamilton et al. 2010, 2013)
and 6.8 million species (range 5.9–7.8 million) for terrestrial
arthropods (Stork et al. 2015)) can be completed within five
decades by boosting taxonomic capacity (Wheeler et al.
2012; Costello et al. 2013) are overly optimistic. Extrapola-
tions from species discovery curves fail to appreciate that the
© 2015 Australian Entomological Society
12 M F Braby and M R Williams
relationship between the cumulative number of species over
time is non-linear. In well-known groups, a disproportionate
amount of time/effort is required to document the as-yet-
unknown species (Joppa et al. 2011b), largely due to the
diminishing pool of missing species that are harder to ‘catch’,
refinement of species boundaries (hypothesis testing) as new
evidence becomes available, and inevitable synonymy. The
inventory of Australian butterflies, for example, is not
expected to be completed until about the year 2090 (i.e. 75
years from now), despite the fact that the pool of missing
species is estimated to be only around 9% of the total fauna
(Fig. 3). Historically, the taxonomy of Australian butterflies
as a science has been driven by private researchers compris-
ing largely amateur experts using traditional methods; this is
still very much the case today (Moulds 1999; Edwards et al.
2001). Clearly, if the inventory of even such well-known
groups is to be completed within a few decades the rate of
taxonomic resolution needs to change drastically.
FINAL THOU G HT S
The following agenda for Australian entomology is suggested
as a guideline for future research in biosystematics and con-
servation biology:
• Broad agreement of insect/invertebrate taxa to be
prioritised, both for taxonomic focus and for use in
inventory and as bioindicators in monitoring. A sys-
tematic approach such as that adopted by NTDLRM
using objective criteria (Table 1) provides a possible
framework.
• Increased investment in taxonomic research capacity
of prioritised taxa to systematically catalogue known
species and describe new species, including university
training and resources for museum collections,
curation, digitisation of material (including type speci-
mens), DNA sequencing and development of taxo-
nomic databases on the Internet (Scoble 2004; Probert
2010). Considerable progress has already occurred in
increasing the efficiency with the publication process
and open-access online taxonomic databases (e.g.
Austral Entomology, ZooBank, Australian Faunal
Directory/Atlas of Living Australia (ALA)) (see
Yeates & Raven 2004; Yeates 2009).
• Greater attention towards molecular genetic methods,
such as DNA barcode technology and application of
genomic data through next-generation sequencing
together with population genetic studies of species
will assist with rapid identification of field samples,
especially where morphospecies approach is used for
some prioritised taxa, as well as recognition of conser-
vation units within species. This technology should not
be used at the expense of traditional taxonomy, but
rather complement it.
• Complete reconstruction of the evolutionary history or
tree of life, which will form the framework for the
development of reliable guidelines and conceptual
© 2015 Australian Entomological Society
basis as to how phylogenetic diversity can be incorpo-
rated into biodiversity conservation planning compared
with traditional measures, such as species richness,
endemism and threatened species (Winter et al. 2013).
For example, in the NTDLRM, taxa that are members
of a species-poor higher systematic rank (family,
order), or belong to a monotypic genus, score more
highly than those that belong to more species-rich
clades in the prioritisation framework for threatened
species (see also Isaac et al. (2007) for more sophisti-
cated methods for threatened EDGE species).
However, there appears to be a general lack of consid-
eration of how to use phylogenetic diversity as a com-
ponent of biodiversity in land use decision-making and
conservation reserve design at the landscape and
bioregional levels.
• Complete inventory and targeted field surveys of Key
Biodiversity Areas and other small geographical
regions rich in narrow-range endemics in Australia (e.g.
Wet Tropics World Heritage Area, Queensland; central
Arnhem land plateau, Northern Territory; south-west
corner of Western Australia) and relatively unexplored
regions that have been poorly sampled (e.g. Kimberley,
Western Australia, western Cape York Peninsula and
the Gulf of Carpentaria, Queensland). These areas
(especially biodiversity hotspots) are likely to harbour
the majority of unnamed, cryptic, short-range endemic
species. The national BushBlitz programme has been a
major initiative in this direction.
• Development of national databases to document the
spatial distribution of species, and predictive spatial
modelling of biodiversity. The ALA and related activi-
ties, such as the Australian Natural Heritage Assess-
ment Tool, are excellent government-funded initiatives
in the field of bioinformatics and are currently the best
platforms on which to build at the national level.
• Establishment of rigorous, long-term monitoring pro-
grammes through development of national databases
designed to detect changes in geographical range, rela-
tive abundance (occupancy) and species richness or
composition through national and state mapping
schemes.
• Increased capacity-building through greater participa-
tion of citizen science programmes, such as eButterfly
in North America (McFarland et al. 2015) and the
ALA-BowerBird inAustralia (Walker 2015), in collect-
ing spatial and temporal data and indices of relative
abundance of species. Such checklist data (species,
location, date, observer) can now be collected by visual
observations or photographs and then uploaded by
computer, smartphone or tablet. These data can then be
used in conjunction with databased vouchered speci-
mens for evaluation of geographical range, phenology
and conservation status of threatened species. These
initiatives also have the potential to collate and analyse
large spatial data sets and hence inform government
agencies and non-government organisations on

conservation priorities. Networks consisting of such
agencies/organisations, scientists, community groups,
natural history societies and volunteers are likely to be
the future for the conservation management and moni-
toring of insect diversity in Australia (New 2010; Yen &
New 2013).
ACKNOWLE D G E ME NTS
We are most grateful to Alan Andersen, Roger Kitching,
David Yeates and Alan Yen for their thoughts and constructive
comments on the manuscript. Max Moulds kindly provided
permission to reproduce Figure 2.
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Accepted for publication 25 May 2015.
© 2015 Australian Entomological Society