Biocultural aspects of species extinctions
Richard J. Ladle1,2,3 , Fernanda Alves-Martins1,2 , Ana C.M. Malhado1,2,3,
Victoria Reyes-García4,5,6 , Franck Courchamp7, Enrico Di Minin8,9,10 , Uri Roll11,
Ivan Jarić7,12 and Ricardo A. Correia8,9,13
1CIBIO-InBIO, Research Centre in Biodiversity and Genetic Resources, University of Porto, Vairão, Portugal; 2BIOPOLIS
Program in Genomics, Biodiversity and Land Planning, CIBIO, Vairão, Portugal; 3Institute of Biological and Health
Sciences, Federal University of Alagoas, Maceió, Brazil; 4Institució Catalana de Recerca i Estudis Avançats, Barcelona,
Spain; 5Institut de Ciència i Tecnologia Ambientals, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain;
6Departament d’Antropologia Social i Cultural, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain;
7Ecologie Systématique Evolution, Université Paris-Saclay, CNRS, AgroParisTech, Gif-sur-Yvette, France; 8Helsinki Lab
of Interdisciplinary Conservation Science, Department of Geosciences and Geography, University of Helsinki, Helsinki,
Finland; 9Helsinki Institute of Sustainability Science (HELSUS), University of Helsinki, Helsinki, Finland; 10School of Life
Sciences, University of KwaZulu-Natal, Durban, South Africa; 11Mitrani Department of Desert Ecology, The Jacob
Blaustein Institutes for Desert Research, Ben-Gurion University of the Negev, MidreshetBen-Gurion, Israel; 12Biology
Centre of the Czech Academy of Sciences, Institute of Hydrobiology, České Budějovice, Czech Republic and 13Biodiversity
Unit, University of Turku, Turku, Finland
Abstract
Predicting whether a species is likely to go extinct (or not) is one of the fundamental objectives of
conservation biology, and extinction risk classifications have become an essential tool for
conservation policy, planning and research. This sort of prediction is feasible because the
extinction processes follow a familiar pattern of population decline, range collapse and frag-
mentation, and, finally, extirpation of sub-populations through a combination of genetic,
demographic and environmental stochasticity. Though less well understood and rarely quan-
tified, the way in which science and society respond to population decline, extirpation and
species extinction can also have a profound influence, either negative or positive, on whether a
species goes extinct. For example, species that are highly sought after by collectors and hobbyists
can becomemore desirable and valuable as they become rarer, leading to increased demand and
greater incentives for illegal trade – known as the anthropogenic Allee effect. Conversely, species
that are strongly linked to cultural identity are more likely to benefit from sustainable manage-
ment, high public support for conservation actions and fund-raising, and, by extension, may be
partially safeguarded from extinction. More generally, human responses to impending extinc-
tions are extremely complex, are highly dependent on cultural and socioeconomic context, and
have typically been far less studied than the ecological and genetic aspects of extinction. Here, we
identify and discuss biocultural aspects of extinction and outline how recent advances in our
ability to measure and monitor cultural trends with big data are, despite their intrinsic
limitations and biases, providing new opportunities for incorporating biocultural factors into
extinction risk assessment.
Impact statement
Human responses to impending extinctions are complex, are highly dependent on cultural and
socioeconomic context, and have typically been far less studied than the ecological and genetic
aspects of extinction. Specifically, the way in which science and societies respond to population
decline, extirpation and species extinction can also have a profound influence, either positively
or negatively, on whether a species goes extinct. For example, while some rare species suffer
higher extinction risk the rarer they become, some charismatic species benefit from significantly
higher conservation efforts and elevated levels of scientific research. Amore comprehensive and
nuanced understanding of which species will go extinct, and which will be “rescued” by
conservation and stewardship efforts, requires an explicit interdisciplinary, biocultural approach
to extinction that draws on expertise from the natural and social sciences, and dialog with
holders of different knowledge systems and, in particular, with Indigenous Peoples and local
communities. Ultimately, many currently threatened species will only go extinct if society allows
it to occur, through a lack of motivation, knowledge, resources, or local conservation capacity.
Introduction
Extinction is typically viewed as a logical end point of the process of population decline – the
point on the graph where the population size curve meets the x-axis and terminates abruptly and
Cambridge Prisms: Extinction
www.cambridge.org/ext
Review
Cite this article: Ladle RJ, Alves-Martins F,
Malhado ACM, Reyes-García V, Courchamp F,
Di Minin E, Roll U, Jarić I and Correia RA (2023).
Biocultural aspects of species extinctions.
Cambridge Prisms: Extinction, 1, e22, 1–9
https://doi.org/10.1017/ext.2023.20
Received: 09 March 2023
Revised: 14 July 2023
Accepted: 08 August 2023
Keywords:
biocultural; culturally important species;
anthropogenic Allee effect; culturomics
Corresponding author:
Richard J. Ladle;
Email: richardjamesladle@gmail.com
© The Author(s), 2023. Published by Cambridge
University Press. This is an Open Access article,
distributed under the terms of the Creative
Commons Attribution licence (http://
creativecommons.org/licenses/by/4.0), which
permits unrestricted re-use, distribution and
reproduction, provided the original article is
properly cited.
https://doi.org/10.1017/ext.2023.20 Published online by Cambridge University Press
finally (Ladle and Jepson, 2008). Accordingly, the International
Union for the Conservation of Nature (IUCN) defines a species
as extinct if there is no reasonable doubt that the last individual has
died (Hughes et al., 2021). Reasonable doubt in this context is the
lack of evidence in the face of exhaustive surveys or extrapolation
from historical observations (Solow, 2005). The intrinsic uncer-
tainty about extinction led E. O. Wilson to characterize it as “the
most obscure and local of all biological processes” (Wilson, 1992,
p. 255). If scientists were to restrict their analysis to only well-
documented extinctions, there would be a huge risk of underesti-
mating current extinctions (Pimm et al., 2014). Thus, extinctions
are often extrapolated onto unknown species, whose existence is
inferred from species discovery curves, from biodiversity ratios, or
from species–area relationships (Bebber et al., 2007; Chisholm et al.,
2016; Kunin et al., 2018; García‐Robledo et al., 2020).
Early conceptualizations of the extinction process strongly
emphasized the role of population decline and the effects of small
population size on population viability (Caughley, 1994). The
former process is the result of deterministic factors such as habitat
loss, degradation and overexploitation, ultimately leading to small
fragmented populations that are highly susceptible to stochastic
factors (Lande, 1998). Accordingly, extinction risk assessment
schemes such as the IUCN Red List emphasize the rate of popula-
tion decline, distributional range, and the size and fragmentation of
extant populations as important factors that influence long-term
prospects of a species’ survival (https://www.iucnredlist.org/assess
ment/process). Likewise, these important insights led directly to the
development of the concept of minimum viable populations and
sophisticated tools for performing population viability analysis
(Ladle, 2009; Traill et al., 2010; Flather et al., 2011).
In summary, extinction is typically understood to mean the
death of the last individual of a species and is a consequence of
the process of population decline and the negative consequences of
small population size. However, contemporary extinction is almost
never a purely biological process (Ladle and Jepson, 2008). Rather,
contemporary extinction is almost always a consequence of the
interaction of cultural and biological phenomena, that is a biocul-
tural process. Indeed, cultural practices now influence almost every
aspect of the extinction process – howwe perceive it, measure it and
act upon it – and, critically, whether a threatened species actually
goes extinct (or stays extinct), echoing calls for the adoption of
biocultural approaches to conservation (Garibaldi and Turner,
2004; Gavin et al., 2015; Bridgewater and Rotherham, 2019). We
contend that there is a need for a broader conceptualization and
exploration of species extinctions as a biocultural process. Here, we
outline some of the key biocultural aspects of extinction, including
taxonomic change, human impacts on population decline, and how
the behavior of the global conservation movement plays a key role
in contemporary extinction dynamics.
Taxonomy and extinction
Although extinction is sometimes used to describe the extirpation
of populations, geographic variants or subspecies (often called
“local extinction”), the term is more commonly applied to the loss
of all populations of an officially recognized species (Ladle and
Jepson, 2008). Extinction statistics are therefore highly sensitive to
changes in taxonomic practice, especially changes in normative use
of species concepts and species delimitation criteria (Zachos, 2016).
Such changes have become increasingly prevalent, partly as a
consequence of advances in molecular taxonomy, leading to
significant recent increases (and sometimes decreases) in the num-
ber of recognized species in many taxa (Garnett and Christidis,
2017). Every taxonomic decision to “split” a species into two or
more species or to “lump” two or more species into a single species
necessarily has consequences for the extinction risk of each newly
defined species (Mace, 2004). Splitting means that each newly
recognized species will have a smaller population size and/or
geographic range, potentially increasing the threat level. Such
increasing threat may potentially be compensated by the increased
conservation attention afforded to a fully recognized species rather
than a subspecies or regional variant. It could also reduce the
average societal attention given to each newly designated species
(Ladle et al., 2019). Moreover, estimates of the numbers of
unknown species (sometimes referred to as the Linnean Shortfall;
Hortal et al., 2015) are highly sensitive to the number of docu-
mented species, with knock-on effects for estimates of global
extinction rates (Stropp et al., 2022).
Cultural push-factors
The first scientists to study extinction at the end of the nineteenth
century had enormous difficulties attributing the decline and even-
tual loss of a species to human actions (Ladle and Jepson, 2008). For
example, despite clear evidence of overhunting (Bengtson, 1984),
James Orton described his confusion about the causes of the
extinction of the Great Auk (Pinguinus impennis) as follows:
“The upheaval or subsidence of strata, the encroachments of other
animals, and climatal revolutions—by which of these great causes
of extinction now slowly but incessantly at work in the organic
world, the Great Auk departed this life, we cannot say” (Orton,
1869, p. 540). This reluctance to attribute human causes to species
extinction continued well into the twentieth century (Ladle and
Jepson, 2011). In contrast, modern current conceptualizations of
the factors driving population declines foreground the indirect and
direct role of human actions and how they are shaped by socio-
cultural practices and beliefs (Lande, 1998; Díaz et al., 2019).
Although extinction can occur in the absence of human influ-
ence (De Vos et al., 2015), the vast majority of contemporary
extinctions are ultimately or proximately connected to human
action (Ceballos et al., 2015; Díaz et al., 2019, 2015), and are
underpinned by societal values and behaviors. Ultimate causes
include human population growth (McKee et al., 2004), the seem-
ingly universal desire to accumulate surplus capital (McBrien,
2016), the political need for economic growth (Spash and Smith,
2019) and the grinding hardship of rural poverty that forces indi-
viduals into a reliance on natural resource exploitation (Adams
et al., 2004). These factors, in turn, drive the proximate causes of
population decline, the most significant of which are habitat loss,
fragmentation and transformation (Maxwell et al., 2016; Powers
and Jetz, 2019), climate change (Thomas et al., 2004; Cahill et al.,
2013), biological invasions (Clavero andGarcía-Berthou, 2005) and
overexploitation (Bennett et al., 2002; Maxwell et al., 2016).
While habitat loss, climate change and biological invasions can
be considered as by-products of other human activities such as
agriculture, trade, transport and recreation, population decline due
to overexploitation of species is a direct consequence of human
cultural practices and values. This is clearly illustrated by the
anthropogenic Allee effect, the idea that human predisposition to
place an exaggerated economic value on rarity drives the dispro-
portionate exploitation of rare species, causing them to become
rarer and therefore even more desirable (Courchamp et al., 2006;
2 Richard J. Ladle et al.
https://doi.org/10.1017/ext.2023.20 Published online by Cambridge University Press
Palazy et al., 2012a; Tournant et al., 2012). Even when a species
becomes so rare that it cannot, alone, support livelihoods, oppor-
tunistic exploitation, while targeting more common species, will
ensure that population decline continues (Branch et al., 2013). For
example, Chinese bahaba (Bahaba taipingensis) is a highly sought-
after fish for use in traditional Chinesemedicine, but fishers seeking
this species must make their living off other species because only a
few are caught each year (Sadovy and Cheung, 2003). The
anthropogenic Allee effect, an explicitly biocultural model of
extinction risk, is particularly applicable to “collectable” exotic
species (Siriwat et al., 2019) or their products, as in the example
of traditional Chinese medicine highlighted above (but see Mateo-
Martín et al., 2023). More generally, it illustrates the complex ways
in which cultural practices and beliefs can become entangled in the
process of population decline and extinction. In this case, human
perceptions of rarity and economic dynamics interact with the
population trends of the exploited species to create an “extinction
vortex” (Courchamp et al., 2006).
The ultimate and proximate drivers of population declines of
species are typically considered to be insufficient to cause the actual
extinction of a species (Lande, 1998). Instead, populations become so
small and fragmented that they become subject to a range of sto-
chastic processes (genetic drift, demographic and environmental
stochasticity, natural catastrophes) that ultimately lead to the death
of the last individual (extinction) (Lande, 1998). As described above,
in a few cases the increasing rarity valueof these last individuals vastly
inflates their economic value and therefore incentivizes their capture
or elimination (Courchamp et al., 2006; Hall et al., 2008). In species
without economic value and without sufficient human intervention
(see below), remnant populations eventually succumb to one of the
many risk factors associatedwith small populations, as highlighted by
individual animals that become famous for being the “last” of their
species (Nicholls, 2012; Jarić et al., 2023).
Cultural push-back factors
While cultural “push-factors” for extinction are generally well
known and quantified, far less attention has been given to the role
of humans in delaying or preventing extinction (“push-back” fac-
tors). Since the emergence of the global conservation movement in
the late nineteenth century (Soulé, 1985), conservationists have
increasingly monitored threatened populations and, when deemed
necessary, intervened in aiming to halt or slow the extinction
process (Hoffmann et al., 2015; Bolam et al., 2021).
The increasing capacity of the global conservation community
to identify species at risk of extinction and to take action tomitigate
this risk highlights the key role that humans now play in the
extinction process. In other words, species become extinct
(or avoid impending extinction) due to the interplay between
human-mediated biological processes (range collapse, population
decline, small populations) and human capacity to monitor and
successfully intervene in this process (Ladle and Jepson, 2008).
There is a global safety net provided by the conservationmovement
as represented by government bodies and various international and
national non-governmental conservation organizations (NGOs).
In a similar way that extinction threats vary geographically, the
motivation, capacity, resources and effectiveness of conservation
also vary immensely by country and region (Waldron et al., 2013).
Sometimes extinction threats and capacity to deal with those threats
align, but frequently there is a mismatch, with geographic areas
hosting a high frequency of threatened species mainly located in the
“Global South” (Schipper et al., 2008) while the capacity of the
global conservation movement to act is often more concentrated in
the “Global North” (Balmford et al., 2003). While this is generally
true, there are many exceptions, and further research into this area
is needed.
Although conservation capacity is clearly central to the prob-
ability and time-scale of extinction for many species, it is a very
poorly defined concept, hampering measurement and mapping
efforts. The broadest definition of conservation capacity in relation
to species extinctions includes at least three dimensions (Table 1): i)
willingness and motivation to act; ii) knowledge to design effective
interventions; and iii) institutional, technical and economic
resources to implement effective interventions. The interactions
of these three dimensions will largely determine whether a species is
identified as being at risk of extinction, whether efforts are made to
reduce the risk of extinction and whether those efforts are success-
ful in the short and long term.
Cultural willingness and motivation to prevent extinction
It is self-evident that societies vary enormously in their willingness
to allocate resources to conserve different species depending upon
their values and valuation of nature (Díaz et al., 2015). Among the
conservation community, charismatic and culturally iconic species
of vertebrates (mainly mammals and birds) are prioritized for
conservation funding and action over equally threatened but cul-
turally less visible species (Davies et al., 2018; Mammola et al.,
Table 1. Main dimensions of conservation capacity and some of the methods of measurement (see text for details)
Dimension Definition Example methods References
Willingness and
motivation to act
Societies vary in their desire to prevent different species fromgoing extinct,
largely depending on the societies’ cultural values and on the species’
cultural characteristics (public awareness, interest, sentiment, etc.)
Culturomic analysis Millard et al. (2020)
Social surveys Samojlik et al. (2023)
Knowledge The amount of scientific and/or local knowledge of species that is
relevant to their conservation, management and stewardship varies
due to a wide range of cultural and historic factors
Bibliometrics dos Santos et al. (2020)
Expert assessment Pearce-Higgins et al. (2017)
Reports from Indigenous and local
knowledge systems
Ziembicki et al. (2013)
Institutional,
technical and
economic
resources
The capacity of local actors and conservation organizations (NGOs,
governmental bodies, international institutions and private
organizations) to fund and implement successful conservation
interventions varies geographically in relation to complex
socioeconomic, political and historic factors
Desk-based analysis of
institutional capacity
Malhado et al. (2020)
Social surveys Fu and Shumate (2020)
Cambridge Prisms: Extinction 3
https://doi.org/10.1017/ext.2023.20 Published online by Cambridge University Press
2020). A recent analysis of the internet salience of 36,873 vertebrate
taxa revealed that search interest was higher for more threatened
mammal and bird species than it was for fish, reptiles and amphib-
ians (Davies et al., 2018). Similarly, Kim et al. (2014) examined web
search data for 246 threatened species in Korea and found that the
interest for mammals, birds, amphibians and reptiles were 10 times
higher than those for other taxa. This bias toward vertebrates also
has a geographical component, with temperate species receiving
more conservation attention than those in the tropics (Titley et al.,
2017). Although plants generally receive less conservation attention
than vertebrates, they are also strongly influenced by cultural
perceptions (Adamo et al., 2021), with a recent study indicating
that species with attractive flowers received more funding, irre-
spective of extinction risk (Adamo et al., 2022). Similarly, fungi
conservation is significantly biased toward macrofungi since these
are most easily observed and include many edible taxa (Gonçalves
et al., 2021). It should be noted that even charismatic vertebrate
speciesmay still be lacking adequate resources to prevent continued
population decline (Di Minin et al., 2015; Courchamp et al., 2018).
Culturally prominent species that generate high public interest
and positive sentiment are more likely to be the target of conser-
vation actions for two main reasons. Firstly, it is easier to mobilize
support and resources through campaigns and other fund-raising
actions when a species already has a high public profile (Thomas‐
Walters and J Raihani, 2017). Secondly, societal preferences also
extend into scientific research, with researchers across the world
preferentially collecting data on larger, more charismatic taxa
independent of the threat status (Caro, 2007; Troudet et al.,
2017). Conversely, many species receive little to no attention and
are likely to suffer from a process of societal extinction – the decline
of collective attention and memory of an extinct or threatened,
extant species (Jarić et al., 2022). The process of societal extinction
of species is linked to that of biological extinctions, as it is likely to
result in decreased support for conservation action, ultimately
affecting negatively the outcome of such efforts.
Until recently it was challenging to quantify the level of public
awareness, interest and sentiment about threatened species at a
national, regional and global scale because this required the use of
time-consuming and costly social surveys. However, the recent
development of “conservation culturomics” (Di Minin et al.,
2015; Ladle et al., 2016; Correia et al., 2021), the analysis of digital
data generated by people to provide novel insights on human–
nature interactions, allows the evaluation of multiple aspects of
societal preferences for species and higher taxa. For example, Ladle
et al. (2019) found that the salience of bird species on the global
internet was strongly correlated with species that have wide geo-
graphic ranges that overlap with technologically advanced societies,
that are phenotypically conspicuous and, critically, that have direct
interactions with humans (e.g., hunting, pet keeping, etc.). A related
study based on Wikipedia page views for all extant species of birds
found that farmed species and species in the pet bird trade were
particularly prominent over multiple language editions
(Mittermeier et al., 2021). These examples demonstrate how cul-
turomic metrics can be used to capture and quantify different
aspects of human interest in nature at scales that are beyond the
reach of standard social surveys (see also Fink et al., 2020; Falk and
Hagsten, 2022; Johnson et al., 2023).
Big data approaches such as culturomics have enormous poten-
tial but also many limitations related to scale and coverage
(reviewed inCorreia et al., 2021,DiMinin et al., 2021). For example,
many Indigenous Peoples (and many other socially and econom-
ically marginalized groups) have limited access to the global
internet, and understanding their interactions, attitudes and senti-
ment toward local species is also critical for effective conservation,
but frequently ignored in conservation management (Zanotti and
Knowles, 2020). Recognizing Indigenous Peoples and local com-
munities’ rights and agency in conservation management (Reyes-
García et al., 2022) is of critical importance, both because much of
the world’s biodiversity now exists in landscapes and seascapes
traditionally owned, managed, used and/or occupied by Indigenous
Peoples (Garnett et al., 2018) or by local communities (Brondizio
and Le, 2016) and because such a strategymight ultimately improve
conservation outcomes (Büscher and Fletcher, 2019). Rates of
biodiversity decline are slower in such areas than elsewhere, includ-
ing protected areas (Garnett et al., 2018; Fa et al., 2020; O’Bryan
et al., 2021). Reyes-García et al. (2023) recently developed a frame-
work around the concept of culturally important species that could
be used to integrate different nature values into the management of
threatened species. Such species predominated among areas where
Indigenous Peoples live and, critically, include a high proportion of
species that the IUCN classify as Data Deficient. Species in their
study were more likely to be culturally than biologically threatened,
especially those associated with Indigenous Peoples due to the high
levels of cultural loss they have experienced.
Conservation-relevant knowledge of species
It has long been recognized that there are large and persistent
taxonomic biases in which species have been researched and,
consequently, the volume and quality of scientific knowledge
about different species (Clark and May, 2002; Fleming and Bate-
man, 2016). Such variations potentially have a significant impact
on the capacity of societies to prevent species from going extinct.
For many species, to be “saved” from extinction there should be
sufficient biological, ecological and cultural knowledge of the
species and its habitat to support the design and implementation
of appropriate conservation interventions (Murray et al., 2015;
Cooke et al., 2017). For other species, habitat and site-based
conservation may be sufficient to prevent extinction and popula-
tion decline. Moreover, more knowledge of a species or higher
taxon does not always lead to better conservation interventions or
swifter responses when a species is threatened, but, all things being
equal, a well-studied species or group is more likely to be the
subject of effective conservation actions than a poorly known
counterpart. It should be noted that it is not only published
scientific knowledge that is potentially important but also the
practical and contextual knowledge of researchers (and other
stakeholders) about the species in question. The greater the
research effort, the greater number of people with such knowledge
that can be mobilized to facilitate conservation efforts.
The causes of these biases are relatively well understood (Jarić
et al., 2019). For example, scientists tend to study species within the
country where they work due to a combination of funding prior-
ities, cost and convenience. It follows that countries with low
scientific capacity typically have fewer qualified conservation sci-
entists and ecologists and less resources available for research,
leading to geographical biases in conservation research effort
(Meyer et al., 2015). Species also vary in their “researchability” –
any characteristic of the species that potentially increases the costs
of data collection or which impedes or reduces the feasibility of a
research project (da Silva et al., 2020; dos Santos et al., 2020). For
field-based conservation research, this could include characteristics
that make it harder to observe a species, such as small body size,
habitat characteristics and accessibility, nocturnal activity patterns,
4 Richard J. Ladle et al.
https://doi.org/10.1017/ext.2023.20 Published online by Cambridge University Press
elusiveness or cryptic coloration. Researchability could also be
correlated with “conservability,” if the traits that make a species
more challenging to study overlap with those that make it more
difficult to implement conservation interventions. Moreover, when
species are challenging to study they become less desirable targets
for researchers whose chances of career advancement may depend
on their publication record or the completion of a high-level
research dissertation (Caro, 2007).
As with human interest in species (see above), the last decade
has seen great advances in our capacity to quantify taxonomic
biases in research at scale through the analysis of bibliometric
databases such as Scopus, Web of Knowledge or Google Scholar.
For example, a recent regional-scale bibliometric analysis of
Australian birds demonstrated significantly more publications
on species with larger body sizes, larger ranges, higher relative
abundance and presence in urban environments (Yarwood et al.,
2019). A similar study on all extant species ofmammals found that
research volume was strongly associated with the scientific cap-
acity within the range of species, high body mass and whether the
species was non-native, with a very weak effect of conservation
threat status (dos Santos et al., 2020).
Additionally, it should be noted that there can be a mismatch
between the “researchability” of a species, as determined by sci-
entists, and its cultural relevance, as defined by local criteria
(Crane et al., 2016). Reyes-García et al. (2023) found that cultur-
ally important species had a much higher proportion of Data
Deficient species than the full set of IUCN species, most likely
resulting in an underestimation of their biological threat, as
species categorized as Data Deficient by the IUCN seem to be
more threatened than data-sufficient species (Borgelt et al., 2022).
The data gap underscores that cultural considerations remain
disregarded in much current biological research (Bridgewater
and Rotherham, 2019) despite the fact that Indigenous and local
knowledge has long been deemed as essential to setting realistic
and effective biodiversity targets (Berkes et al., 2000; Brondízio
et al., 2021; Reyes-García et al., 2022).
Institutional, technical and economic capacity to intervene
Even when there is good scientific knowledge about a threatened
species and strong public support for conservation action, weak
institutional capacity means that interventions may be poorly
planned and executed or not even implemented. In this context,
institutional capacity normally refers to governmental depart-
ments, conservation NGOs and other civil society groups, and
occasionally private sector organizations. Measuring such capacity
is highly challenging, and there have been very few systematic
analyses of conservation organizations at national, regional or
global scales (Brockington et al., 2018). To our knowledge, there
have not yet been attempts to evaluate institutional conservation
capacity at the level of species or geographic regions (e.g., coun-
tries), though such quantifications could play a major role in
determining the number, type and quality of interventions in the
face of endangerment (Ladle and Jepson, 2008). Such a lack is partly
attributable to the difficulties of collecting data on diverse conser-
vation actors (Malhado et al., 2020), and partly due to the com-
plexity of factors that contribute to institutional capacity, severely
limiting the potential to develop robust metrics.
Over the last decades, there has been an institutionalization of
co-management and bottom-up approaches to conservation
(e.g., Indigenous and Community Conserved Areas, community
monitoring). For example, community-based monitoring is
increasingly proposed to improve scientific understanding of
biodiversity status and trends, or local uses of plants and animals,
among other processes (Danielsen et al., 2021). Understanding
the role of such initiatives in curving extinction processes also
requires monitoring.
How many species have been “saved” from extinction?
Estimating the number of species that would have gone extinct if
conservationists had not intervened is, by definition, exceedingly
challenging, and the fact remains that the number of species
currently threatened with extinction is unprecedented in human
history (Ceballos et al., 2010; Díaz et al., 2019). Well-known
examples of “near extinction” events, such as the black-footed ferret
(Dobson and Lyles, 2000) or the Chatham Island Black Robin (von
Seth et al., 2022), are indisputable. However, the imminent demise
of the rescued population is often less clear-cut and there have been
few large-scale estimations, mainly restricted to birds and mam-
mals (Table 2). For example, Bolam et al. (2021) estimated the
number of species “saved” by canvassing the opinion of experts;
their estimate that bird and mammal extinction rates would have
been 2.9–4.2 times greater without conservation action is almost
certainly an underestimate given that only clear cases were con-
sidered. There are many more situations where, had conservation
not intervened earlier in the extinction process (i.e., before a species
becomes Critically Endangered), a species would arguably have
gone extinct. Moreover, many species have avoided extinction
due to actions aimed at conserving sites, habitats and ecosystems.
This category of “saved” species is even more difficult to quantify
since they include many lesser-known taxa, some of which may be
undescribed (Hortal et al., 2015).
A closely related issue to species saved from extinction is the
number of species that would eventually go extinct without
continued conservation investment. Such “conservation-
dependent” species could justifiably include, among others, spe-
cies whose i) populations are periodically augmented with
captive-bred individuals; ii) populations are not declining due
to continued management efforts such as anti-poaching meas-
ures and surveillance, genetic management, control of invasive
species, supplemental feeding, etc.; and iii) last remaining indi-
viduals exist only in captivity. Taking the latter group of species
as an example, there are 85 species currently classified as Extinct
in the Wild (i.e., only ex-situ populations remain), and some of
these species have persisted in captivity for over 70 years (Smith
et al., 2023). Other forms of conservation dependence are poorly
quantified but potentially represent a significant limitation to
Table 2. Estimated number of bird and mammal species that would have gone
extinct without direct human intervention
Taxon
Number of
species Timeframe References
Birds
16 1994–2004 Butchart et al. (2006)
21 to 32 1993–2020 Bolam et al. (2021)
9 to 18 2010–2020 Bolam et al. (2021)
Mammals (all)
7 to 16 1993–2020 Bolam et al. (2021)
2 to 7 2010–2020 Bolam et al. (2021)
Mammals (Ungulates) 6 1996–2008 Hoffmann et al. (2015)
Cambridge Prisms: Extinction 5
https://doi.org/10.1017/ext.2023.20 Published online by Cambridge University Press
future conservation actions, given the limited resources available
for new initiatives.
Extinction risk forecasting using push and push-back factors
As quantitative assessments of extinction risk, Red Lists are a
crucial knowledge product and underpin much conservation law
and policy (Rodrigues et al., 2006; Hoffmann et al., 2008). Red List
categorizations are used, among other things, to i) support conser-
vation decisions at, and across, multiple governance levels; ii) guide
strategy and investments in species conservation; and iii) inform
progress toward targets of international agreements. Perhaps more
importantly, Red Lists have translated the key conservation value of
avoiding anthropogenic extinctions into a governance tool that has
helped produce global norms governing relations between society,
economy and the nonhuman world (Jepson et al., 2011).
IUCNRedLists assign species to extinction threat categories based
on five quantitative criteria: i) population vulnerability; ii) population
size reduction; iii) geographic range; iv) population size; and v)
population viability analysis. These categories have a population
ecology/life history focus, yet, as argued above, extinction (and its
avoidance) is a biocultural phenomenon. Before and after a taxon is
assigned to a Red List category it is subject to cultural forces that
determine the success (or otherwise) of conservation actions (Ladle
and Jepson, 2008). We would argue that a species well known to
science is, ceteris paribus, less likely to be at risk of extinction com-
paredwith a lesser-known species in the same threat category because
the public and institutions will mobilize more effectively to save
it. Exceptions may include species that are highly sought after as pets,
trophies, food or fashion accessories (Gault et al., 2008; Palazy et al.,
2012b; Leclerc et al., 2015) that may suffer more intense exploitation
than less-desirable species (Courchamp et al., 2006).Moreover, IUCN
species lists do not explicitly include the importance of species for
local cultures (Reyes-García et al., 2023), a factor that could also play a
vital role in the success of any proposed conservation intervention. In
short, IUCN Red List categories currently omit a range of non-
biological factors thatmay be critical in determiningwhether a species
will be “saved” from extinction (or not).
Creating a systematic and comprehensive system of assessing
public interest and local cultural importance of species could, along
with information of scientific knowledge of species, provide inter-
esting complementary information to support and add nuance to
extinction categorizations (Figure 1). Specifically, information
from Indigenous and local knowledge systems or from macroscale
cultural analysis (e.g., culturomics) could potentially be used to i)
identify threatened species assemblages and geographic regions
(or parts of regions) where the potential for rallying support may
be weaker and where greater investment may be required to
improve the conservation status of a taxon (Ladle et al., 2016); ii)
further support the use of Red Lists in business and investment
decisions (Bennun et al., 2018), making material the reputational
risks associated with activities that impact publicly visible globally
threatened species; iii) enhance the quality of conservation actions
by increasing the recognition of Indigenous Peoples and local
communities’ knowledge, values and rights (Reyes-García et al.,
2022); and iv) provide complementary information to support and
prompt innovative actions to reduce extinction risk. For example,
“digital interventions” can raise the public profile of threatened
species and more effectively link communities of interest with
specific taxa.
The above suggestions come with the caveat that cultural evalu-
ation based on big data approaches such as culturomics has many
limitations and biases (Troumbis and Iosifidis, 2020; Correia et al.,
2021), and much research is still needed to develop robust, well-
validated metrics that can be used with confidence for conservation
planning and assessment. Furthermore, as cultural metrics are
eventually integrated into extinction risk assessment, it is inevitable
that different threatened species might benefit or lose out depend-
ing on how conservation organizations choose to use this informa-
tion; for example, deciding on the balance between funding the
conservation of well-known species versus promoting the conser-
vation of species deemed to have little or no cultural importance.
All things being equal, in areas where conservation capacity is
low, i) species are less likely to be “saved” from extinction due to a
lack of scientific knowledge, resources and effective conservation
interventions; ii) threatened species may be less effectively moni-
tored (Fisher et al., 2011) leading to incomplete knowledge of
species distributions/population status and slow or absent conser-
vation responses; and iii) technological interventions such as cap-
tive breeding, reintroductions and translocations are less likely to be
implemented or successful. However, the willingness and capacity
of institutions to act to prevent a species from imminent extinction
are not straightforward to evaluate and partially depend on the
cultural characteristics of the threatened species, with far less effort
expended on the conservation of non-charismatic species. For
example, Bellon (2019) found that species popularity (higher inter-
net salience) had a greater effect than federal priority ranking for
the funding of threatened species by various US federal agencies
under the Endangered Species Act.
Conclusions
Species extinction is a complex phenomenon that involves both
biological and social factors. Understanding and addressing these
factors is crucial for the conservation of biodiversity.Most, if not all,
species currently in danger of imminent extinction are in that state
due to the direct and/or indirect impacts of humans on the envir-
onment. Moreover, whether these species actually go extinct will
largely depend on the willingness to act and the technical capacity
of local, national and international conservation organizations,
along with the support of local communities and other stake-
holders. In contrast to the assessment of biological and ecological
risk factors, our understanding and quantification of the cultural
and political vulnerability of species is at an early stage of develop-
ment. A more comprehensive understanding of which species will
Figure 1. Enhanced extinction assessment by combining the IUCNRed List withmetrics
of scientific knowledge and cultural salience. Species in block Y are more likely to go
extinct compared to block A because, in addition to biological drivers of extinction (e.g.
small population size, small range size, declining populations, etc.), species in this
category have low cultural salience, reducing willingness/motivation to act, and low
scientific knowledge, reducing ability to mount effective conservation interventions
6 Richard J. Ladle et al.
https://doi.org/10.1017/ext.2023.20 Published online by Cambridge University Press
go extinct and which will be “rescued” by conservation and stew-
ardship efforts will require an explicit interdisciplinary, biocultural
approach to extinction that draws on expertise from the social
sciences, and dialog with holders of other knowledge systems
and, in particular, with Indigenous Peoples and local communities.
For many currently threatened species, extinction will only occur if
society allows it to occur, through a lack of motivation, knowledge,
resources, or local conservation capacity.
Open peer review. To view the open peer review materials for this article,
please visit http://doi.org/10.1017/ext.2023.20.
Author contribution. All authors contributed to the conceptualization, writ-
ing the original draft, and reviewing and editing of the draft. Writing was
coordinated by R.J.L.
Financial support. R.J.L., F.A-M. and A.C.C.M. are supported by the
European Union’s Horizon 2020 research and innovation programme (grant
agreement #854248). V.R-G. is supported by the European Research Council
under an ERC Consolidator Grant (FP7–771056-LICCI). This research con-
tributes to the “María de Maeztu Unit of Excellence” (CEX2019–000940-M).
E.D.M. thanks the European Research Council (ERC) for funding under the
European Union’s Horizon 2020 research and innovation program (grant
agreement 802,933). R.A.C. acknowledges funding from the Research Council
of Finland (Grant agreement #348352) and the KONE Foundation (Grant
agreement #202101976).
Competing interest. The authors declare none.
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Cambridge Prisms: Extinction 9
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