Thesis

UNIVERSITY OF COPENHAGEN
FACULTY OF SCIENCE
CENTER FOR MACROECOLOGY, EVOLUTION AND CLIMATE
Impacts of human use on species
Changes in biological traits and populations
Louise Kjær-Hansen 60 ECTS
Thesis submission: 14.10.2016
Academic supervisor Professor Neil Burgess
Co-advisor Professor Nathan Sanders
External advisor Doctor Jonas Geldmann

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Goal 12 Ensure sustainable consumption and production patterns
Goal 14 Conserve and sustainably use the oceans, seas and marine resources for
sustainable development
Goal 15 Protect, restore and promote sustainable use of terrestrial ecosystems,
sustainably manage forests, combat desertification, and halt and reverse land
degradation and halt biodiversity loss
United Nations
(the 2030 agenda for Sustainable Development)

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Target 11 By 2020, at least 17 per cent of terrestrial and inland water, and 10
percent of coastal and marine areas, especially areas of particular importance
for biodiversity and ecosystem services, are conserved through effectively and
equitably managed, ecologically representative and well connected systems of
protected areas and other effective area-based conservation measures, and
integrated into the wider landscapes and seascapes.
Target 12 By 2020 the extinction of known threatened species has been
prevented and their conservation status, particularly those most in decline, has
been improved and sustained.
Convention on Biological Diversity
(Aichi Biodiversity Targets – Strategic Biodiversity Plan 2011-2020)

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ABSTRACT
The way to sustainably manage the use of ecosystems and wild living resources still
remains a highly debated topic. Humans have harvested animals and plants, both in
marine and terrestrial environments, for most of their existence, where hunting and
gathering predates the development of agriculture. However, human use, both direct and
indirect, and the intense pressure on usage of wild living resources, is having profound
effects throughout the natural environment. These issues are recognised in global policy
such as the Sustainable Development Goals and the Aichi Biodiversity Targets, where
actions are taken towards helping nationals to achieve a sustainable Earth.
In this thesis, I have done two things that are relevant to the biological sustainability
debate. First, I reviewed existing literature on anthropogenic trait-selective harvesting.
Changes in biological traits was seen in 68% of the reviewed species, and six studies on
harvested ungulate males saw an average decrease of 10.7% in horn measurements. The
studies in this review indicate that the pressure on certain traits leads to changes in
species morphology and life history.
Second, I examined the change in population abundance over time in relation to utilised
and not utilised vertebrates. Here, the Living Planet Index as developed by WWF was
included to be able to analyse trends for all vertebrate populations in protected areas, the
Afrotropical and Palearctic realms as well as globally. For utilised vertebrates the
population trends showed a decline of c. 25% in the global trend, c. 90% in the
Afrotropical realm and c. 35% in protected areas worldwide.
The impact that trait-selective harvesting has on biological traits and the magnitude of
the decline in utilised populations illustrates a grim outcome for future perspectives.
Therefore, being able to successfully monitor conservation strategies is key in future
recommendations. This includes the need for policies and science to interact at a larger
scale and draw benefits from each other to achieve sustainability in accordance with the
Sustainable Development Goals and Aichi Biodiversity Targets.

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RESUMÉ
Den måde, hvorpå man forvalter brugen af økosystemer og levende vilde dyr og planter
på en bæredygtig måde er stadig et meget debatteret emne. Mennesket har jaget og
høstet dyr og planter i både marine og terrestriske miljøer, langt før at landbruget blev
udviklet. Men den måde mennesket, både direkte og indirekte, lægger pres på
naturressourcer, har dybdegående konsekvenser for miljøet. Disse problematikker er
også afspejlet i den globale politik, hvor der er blevet udviklet bæredygtige mål for
fremtiden, herunder de globale mål for bæredygtig udvikling udarbejdet af de Forenede
Nationer og Aichi bæredygtighedsmålene fra Konventionen om biologisk diversitet.
I denne afhandling har jeg undersøgt to områder der er relevante for debatten
omhandlende biologisk bæredygtighed. Først undersøgte jeg den eksisterende litteratur
omkring, hvorledes jagt på dyr påvirker dyrets morfologi over tid. Ændringer i
biologiske egenskaber blev set i 68% af de gennemgåede arter, og seks undersøgelser på
hovdyr viste for jagtede hanners horn størrelser at det i gennemsnit blev 10,7% mindre.
Undersøgelserne tyder på, at presset på visse træk kan føre til ændringer i artens
morfologi og livshistorie.
Efterfølgende undersøgte jeg ændringer i populations størrelser i forhold til, om
populationen er udnyttet eller ikke udnyttet af mennesker. Hertil brugte jeg, the Living
Planet Index som er udviklet af WWF, for at kunne analysere tendenser inden for
populationers størrelser over tid for hvirveldyr i beskyttede områder - i den Afrotropiske
og Palæarktiske zone og globalt. Resultaterne viste tilbagegang i populations
størrelserne på 25% for den globale tendens, 90% i den Afrotropiske zone og 35% i
beskyttede områder over hele verden.
Dette illustrerer en uønskelig fremtid for vores nulevende arter, og fremtidige
anbefalinger omfatter derfor, behovet for at politikkere og videnskaben interagerer på et
større plan for at opnå bæredygtighed i overensstemmelse med de globale mål for
bæredygtig udvikling og Aichi bæredygtighedsmålene.

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ACKNOWLEDGEMENTS
First and foremost a huge thanks to my supervisor Neil Burgess and my external advisor
Jonas Geldmann for their invaluable support throughout this thesis. The amount of
enthusiasm and interest they have shown for this project has been a great help and
motivation, despite the distance from Cambridge to Copenhagen. I would also like to
thank Nate Sanders and Katarzyna Nowak for their encouragement and helpful inputs.
Furthermore, this thesis would have been very different without collaboration with The
Zoological Society of London, wherefrom the Living Planet Index was obtained. Here, I
would like to especially thank Robin Freeman and Louise McCrae for their
collaboration, R scripts and guidance into the complex world of the LPI data set. My
thanks should also extend to Ben Collen who is an LPI wizard and who provided me
with the diagnostic package used to analyse my findings further. I would also like to
thank the people I met at UNEP-WCMC for welcoming me during my visits. Also many
thanks to Rosamunde Almonde, Mike Hoffmann as well as the IUCN specialist groups
for their input and advice during the process.
Also, a big thank you to my friends and family for a few favours along the way,
especially with R assistance provided by Sophie Kjær-Hansen and Andreas Bock, and
proofreading by Johannes D. Hedegaard, Birgitte Holt Andersen and Johan Kjær-
Hansen. Finally, I would like to thank everyone at CMEC, especially the thesis office
aka “Kagestuen”, where I have spent most of my time during this past year.

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TABLE OF CONTENTS
ABSTRACT ..................................................................................................................................... 4
RESUMÉ.......................................................................................................................................... 5
GENERAL INTRODUCTION .................................................................................................... 10
TIMELINE OF SUSTAINABLE DEVELOPMENT.................................................................................. 10
DEVELOPMENTS IN SUSTAINABILITY............................................................................................... 12
AIM AND APPROACH.............................................................................................................................. 14
PART ONE ................................................................................................................................. 15
INTRODUCTION......................................................................................................................... 16
EVOLUTION............................................................................................................................................... 16
POLICY FRAMING.................................................................................................................................... 18
AIM AND EXPECTATIONS...................................................................................................................... 18
METHODS .................................................................................................................................... 19
RESULTS....................................................................................................................................... 21
EFFECTS OF SELECTIVE HUNTING ..................................................................................................... 28
DISCUSSION ................................................................................................................................ 30
PART TWO................................................................................................................................ 32
INTRODUCTION......................................................................................................................... 33
BACKGROUND ......................................................................................................................................... 33
AIM.............................................................................................................................................................. 34
METHODS .................................................................................................................................... 35
THE LIVING PLANET INDEX ................................................................................................................. 35
METHODOLOGY....................................................................................................................................... 36
RESULTS....................................................................................................................................... 38
GLOBAL TRENDS..................................................................................................................................... 38
REALMS ..................................................................................................................................................... 41
PROTECTED AREAS ................................................................................................................................ 43
DISCUSSION ................................................................................................................................ 44
REALMS ..................................................................................................................................................... 44
PROTECTED AREAS ................................................................................................................................ 46
GLOBAL TRENDS..................................................................................................................................... 47
LIMITATIONS AND STRENGTHS TO THE LPI.................................................................................... 48
GENERAL DISCUSSION............................................................................................................ 50
CONCLUDING REMARKS....................................................................................................................... 53
BIBLIOGRAPHY ......................................................................................................................... 54
APPENDICES ............................................................................................................................... 60

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NOMENCLATURE
Armaments* Species armour such as horns, antlers, and tusks
Bush meat Refers to meat from non-domesticated mammals, reptiles, amphibians and birds hunted for
food in Africa
Ecosystem goods and services The benefits ecosystems provide including food, water, timber, air
purification, soil formation and pollination
Natural Capital The world’s stocks of natural assets which includes geology, soil, air, water and all
living things
Not Utilised* Not utilised populations are considered to be all other populations that do not experience
utilisation, see ‘Utilised’ below
Realm A biogeographic realm or ecozone is the broadest biogeographic division of Earth’s land surface,
based on distributional patterns of terrestrial organisms. They are subdivided into ecoregions which are
classified in biomes or habitat types. There are eight realms according to WWF, including the
Afrotropical realm and Palearctic realm (appendix 2)
Ungulate Members of a diverse clade of primarily large mammals that includes odd-toed ungulates and
even-toed ungulates
Utilised* Utilisation includes recreational-, sport-, legal-, illegal-, trophy-, and commercial hunting,
poaching, native subsistence use, egg-, fur-, legal-, illegal-, and commercial harvest, whaling, culling,
recreational-, commercial-, illegal-, and legal fishing, pet trade
* The definitions may only apply to this thesis for clarification

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ABBREVIATIONS & ACRONYMS
CBD Convention on Biological Diversity
CITES The Convention on International Trade in Endangered Species of Wild Fauna and Flora
CSD Commission on Sustainable Development
IISD International Institute for Sustainable Development
IUCN The International Union for Conservation of Nature and Natural Resources
LPI The Living Planet Index
MDGs Millennium Development Goals
SDGs Sustainable Development Goals
UN United Nations
UNCED The United Nations Conference on Environment and Development
UNEP United Nations Environment Programme
UNESCO The United Nations Educational, Scientific and Cultural Organization
WCED World Commission on Environment and Development
WWF World Wide Fund for Nature (also known as World Wildlife Fund)
ZSL The Zoological Society of London

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GENERAL INTRODUCTION
As this thesis is broadly about the sustainable use of species by humanity and the
challenges in achieving sustainable use, I have framed the thesis in terms of the global
debate on sustainability and how this has evolved over the past 50 years.
TIMELINE OF SUSTAINABLE DEVELOPMENT
Efforts to achieve sustainable use of our planet have been underway since the
Intergovernmental Conference for Rational Use and Conservation of the Biosphere
coordinated by UNESCO which took place in 1968 (UNESCO, 1993). The first United
Nations conference on the Human Environment, also known as the Stockholm
conference, followed in 1972. The conference focused on international environmental
issues and reflected a growing concern for conservation issues worldwide. The
conference led to the formation of the United Nations Environment Programme, or
UNEP, later that year. UNEP’s role is to coordinate global efforts to promote
sustainability (UNEP, 2016). In 1975, the Convention on International Trade in
Endangered Species of Flora and Fauna, or CITES, came into force (IISD, 2012). The
Stockholm conference further led to the collaboration between the International Union
for Conservation of Nature (IUCN), The World Wildlife Fund (WWF) and UNEP,
which resulted in the World Conservation Strategy, where the aim was to identify
priority conservation issues and key policy options to further sustainable development,
stressing the need for a new development strategy (Drexhage & Murphy, 2010).
In 1983, the UN called for a commission to be created by both developing and
developed countries to address and assess the ‘accelerating deterioration of the human
environment and natural resources and the consequences of the deterioration for
economic and social development’ (Drexhage & Murphy, 2010). Four years later the
World Commission on Environment and Development (WCED), published Our
Common Future, or the Brundtland report (WCED, 1987). The principles of sustainable
development were ratified in 1992 at the UN Conference on Environment and
Development, also known as UNCED, in Rio de Janeiro, Brazil, also referred to as the

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Rio Summit or the Earth Summit. Here, the UN General Assembly created the
Convention on Biological Diversity (CBD), and the Commission on Sustainable
Development or CSD (Drexhage & Murphy, 2010).
Recognising that the majority of CITES Parties are parties to the CBD, both entities
have adopted this definition of sustainable use: ‘The use of components of biological
diversity in a way and at a rate that does not lead to the long-term decline of biological
diversity, thereby maintaining its potential to meet the needs and aspirations of present
and future generations’ (CITES, 2016). In 2000, the United Nations Millennium
Development goals, or MDGs, were agreed on by the largest-ever gathering of world
leaders to prevent further environmental degradation by 2015. The World Summit on
Sustainable Development followed in 2002, and was held in Johannesburg, South
Africa. The overall outcome of this was the reporting of some positive outcomes the last
ten years, but mostly failed attempts of implementations at the national and international
levels. Overall trends for sustainable development had actually worsened since the Rio
Summit in 1992. In 2009, an article introducing the nine planetary boundaries was
published in Nature (Rockstrom et al., 2009). The article argues that human activities
have over-stretched the Earth system and propose a framework to stay within the
‘planetary boundaries’ by defining a safe operating space. The nine planetary
boundaries include change in land use, biodiversity loss, atmospheric aerosol loading,
chemical pollution, climate change, ocean acidification, stratospheric ozone depletion,
nitrogen and phosphorous cycle as well as global freshwater use.
Under the CBD, the Nagoya Protocol was adopted in 2010, as a supplementary
agreement to the CBD, including also the Strategic Biodiversity Plan which highlights
the Aichi Biodiversity Targets. The aim of the protocol is the ‘fair and equitable sharing
of benefits arising from the utilisation of genetic resources, thereby contributing to the
conservation and sustainable use of biodiversity’ (CBD, 2010). In 2012, Rio +20 was
held, 20 years after the Rio Summit, introducing the Future We Want (UN, 2012). The
summit committed governments to create a set of Sustainable Development Goals, or
SDGs (IISD, 2012; Griggs et al., 2013), which leads us to the present state of our

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worlds natural capital and how we are still struggling to prevent further exploitation
from happening. The SDGs are by far the most ambitious targets to date, but it is vital
that a change to our way of living and our consumption patterns occurs, if we are to
pass on a healthy Earth to future generations.
DEVELOPMENTS IN SUSTAINABILITY
The way to sustainably manage the use of ecosystems and wild living resources still
remains a highly debated topic (Kareiva & Marvier, 2012). A ‘New Conservation’
debate has emerged and the old ways of doing conservation are being questioned, also
commonly referred to as the ‘parks vs. people’ debate (Miller et al., 2011; Kareiva et
al., 2012; Marvier, 2014). The ‘New Conservation’ is largely focused on the
anthropogenic aspect of nature, how nature can benefit humans and how prioritising
ecosystem goods and services can improve human livelihoods. Whereas, traditional
conservation emphasises the intrinsic value of nature by prioritising biological diversity
and preventing species from going extinct (Minteer & Miller, 2011; Holmes &
Sandbrook, 2016).
Legislation and policies still remain as effective solutions to this issue, by adapting a
strict protected status for highly affected areas and banning all use and trade of the most
critical species (Mace et al., 2008). However, if the areas are the primary source for
rural community’s livelihood, it is an almost impossible task to implement. Enforcing
and restricting all exploitation of wild living resources, may lead to species being
displaced from their territories due to human encroachment and domestic livestock
overtaking areas to compensate for the reduction in the resources from the wild (Hutton
& Leader-Williams, 2003). Thus, a successful conservation strategy leading to both
benefits for rural communities as well as preservation of biological resources, can be
difficult in practice (McShane et al. 2011).
Conversely, there have been some positive efforts to avoid this dilemma. Forms of
sustainable use can include the removal of individuals in the wild where the population
is still reasonably healthy, despite intense exploitation, like moose (Alces alces)

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(Sutherland, 2001). Moreover, the extractive use of animals does not necessarily mean
the individuals have to be permanently removed from the population, parts of the
animal will suffice, as in the case of sheared wool from vicunas (Vicugna vicugna) in
Peru. The local communities as well as the species benefit greatly from this
arrangement (Wheeler & Hoces, 1997). Furthermore, it is argued that profits from
trophy hunting of large mammal species can contribute to conservation if the money is
managed properly and off-take is carried out sustainably (Freeman & Wenzel, 2006;
Edwards et al., 2014; Brink et al., 2016).
To ensure conservation efforts are sustainable and successful, an attempt at identifying
the natural capital is needed, and thereby which strategy should be implemented for the
best outcome when taking into account both use for humans and the revival of the
biological community (Ekins et al., 2003). When priorities have been recognised, the
political aspect can thus be made clear to further the management. It is not the wild
resources that need managing, it is the humans who use it (Ekins et al., 2003).
It is very important that political frameworks are highlighted and complied to, in order
to stop over-exploitation and to sustain the natural capital for generations to come. For
example, the Sustainable Development Goals (SDGs), where the UN have produced a
set of goals on sustainable development to be met by 2030. The agenda means to ensure
sustainable consumption and production patterns, to conserve and sustainably use the
oceans, seas and marine resources and to protect, restore and promote sustainable use of
terrestrial ecosystems and forests and to halt and reverse land degradation and
biodiversity loss (Goal 12, 14 and 15) (IISD, 2012; Griggs et al., 2013). Also, the Aichi
Biodiversity Targets, from the CBD, have created a strategic biodiversity plan to be met
by 2020, where at least 17% of terrestrial and inland water, and 10% of coastal and
marine areas are to be conserved through effectively and equitably managed and well
connected systems of protected areas and that the extinction of known threatened
species have been prevented and their conservation statuses have been improved and
sustained (Target 11 and 12) (CBD, 2010).

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AIM AND APPROACH
The aim of this thesis is to tackle two aspects of sustainable use by surveying a change
in biological traits of species due to human use, and a change in population trends for
used species. This will then be discussed in light of the global sustainability agenda.
The first part of the thesis is a literature review, to survey existing literature on human
use on animals and how this may have affected and impacted species morphology. A
similar attempt at a literature review on this has previously been made, though this was
almost a decade ago (Fenberg & Roy, 2008).
The second part of this thesis will report on data analyses on population trends for used
and not used species from the Living Planet Index developed by the WWF. The data
analyses will include populations in protected areas, the Afrotropical realm and
Palearctic realm as well as globally.

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INTRODUCTION
EVOLUTION
Humans have harvested animals and plants, both in marine and terrestrial environments,
for most of their existence. In fact, hunting and gathering predates the development of
agriculture (Mitchell, 2012). However, human use, both direct and indirect, and the
intense pressure on usage of wild living resources, is having profound effects
throughout the natural environment (Wake & Vredenbrug, 2008). Although the main
causes are still a highly debated topic, Earth’s most recent major extinction episode, the
Quaternary Megafauna Extinction, seems to have been partly driven by humans through
a large kill-off of non-human megafauna species (Martin, 1996; Alroy, 2001; Koch &
Barnosky, 2006; Wroe & Field, 2006; Barnosky, 2008). In light of that episode, some
argue that a sixth mass extinction is under way, which should be taken seriously
(Ceballos & Ehrlich, 2002; Wake & Vredenburg, 2008; Barnosky et al., 2011).
Much of the current concern over another extinction episode involves the loss of species
on a global scale (Hughes et al., 1997). This is partly due to hunting which can be
divided into four types; 1) hunting for food, 2) hunting for certain traits, 3) hunting for
safety and 4) to some extent, population control (McNay, 2002; Festa-Bianchet et al.,
2004; Kaltenborn et al., 2005; Mysterud, 2012).
Trait-selective harvesting, where certain traits in individual’s morphology are preferred
by human hunters, is becoming an increasingly significant factor when considering the
fitness of species and their survival. When trait-selective harvesting is done intensively,
humans could be imposing a non-natural selection pressure working against natural
selection (Allendorf & Hard, 2009). Morphological traits are considered an important
influence on the overall life history of an individual in a given species, where natural
selection optimises reproduction and survival strategy (Bro-Jorgensen, 2014).
Assuming natural selection, at least in some cases, is being overwritten by
anthropogenic selection, an imbalance may ensue on specific life history traits within

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target species (Allendorf et al., 2009). Trait based harvest of wild populations can lead
to various changes in the population. The fishing industry harvests a vast number of fish
every day. They target the largest individuals as value increases with size, leading to the
largest individuals disappearing from the populations because of the constant fishing
pressure (Uusi-Heikkila et al., 2016). This creates a loss of large successful breeders
causing females to mate with smaller males which over time leads to smaller offspring
(Roy et al., 2003; Birkeland & Dayton, 2005). Hermaphroditic species where larger size
classes are primarily one sex, are also threatened as size-selective harvesting leads to a
sexual skew in the population (Hamilton et al., 2007; Fenberg & Roy, 2012).
Similarly, males in terrestrial environments are hunted for their large body size and their
large armaments often carried solely by the males (Hartl et al., 2003; Roy et al., 2003;
Loehr et al., 2006; Hamilton et al., 2007; Schmidt et al., 2007; Crosmary et al., 2013;
Monteith et al., 2013). The hunting of large vertebrates can lead to a female-skewed
adult sex ratio as well as a younger age distribution, influencing on reproduction
strategies, body mass variations and resilience towards survival (Garel et al., 2007;
Bro-Jorgensen, 2014). A Canadian study (Ciuti et al., 2012) showed how harvesting
leads to altered behaviour in elk (Cervus elaphus), where bold elk were more likely to
be killed than shy elk. This meant that the lifespan differed between the two types of
behaviour. Shy elk would stay away from open areas, roads and human settlements.
Whereas, bold elk would venture into open and accessible areas for hunters and were
more curious about their surroundings than the shy elk. This inclination could alter the
behavioural composition of individuals in populations and select for more shy elk.
Furthermore, it has been speculated that changes in morphology could lead to changes
in behaviour. Tusklessness in elephants (Loxodonta africana) has increased in females
from 10.5% to 38.2% between 1969 and 1989 and from 1% to 10% in 1970 to 1993 in
males. This sudden decrease in armaments in a species could potentially impact the
behavioural patterns in individuals (Jachmann et al., 1995).

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Even plants are pressured by anthropogenic trait-selective harvest. Such is the case with
the Snow Lotus (Saussurea laniceps), which saw a significant change in size over time
due to harvesting (Law & Salick, 2005). Hence, the necessity for clarifying the
magnitude of trait-selective harvesting for all widely harvested species.
POLICY FRAMING
If alterations of species morphology are happening on a large scale due to intensive
harvesting strategies based on certain traits, it is crucial for wildlife managers, hunters
and conservationists to be aware of this so they can take relevant precautions to
minimize the effect it might have. Distorting population dynamics by altering or even
depleting certain groups based on body and trophy size, which often correlates with age,
can trigger severe consequences throughout biological communities. Our actions need
to be surveyed so deliberate measures can be taken, if it turns out that the effect our
actions are having on biological levels, is causing species to change, influencing on
their life history.
When formulating conservation efforts, it is therefore an advantage that so many nations
on a global scale have committed themselves to the United Nations Sustainable
Development Goals (SDGs) and the Aichi Biodiversity Targets. The SDGs and Aichi
Targets have multiple goals concerning the sustainable development of our planet.
Having an international political agenda that incorporates all these aspects that are
directly or indirectly linked to sustainable harvest of species, should clear a path for
protecting our planet and its resources.
AIM AND EXPECTATIONS
The aim of this part of the thesis was to conduct a literature review that focused on how
human usage, in terms of hunting, has impacted species morphology over time. The
increase or decrease in morphology and absence of larger individuals was expected to
be correlated with intensity of harvest and duration of study period, with males being
the most targeted sex within mammals.

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METHODS
In order to review existing literature on anthropogenic trait-selective harvesting, the
available literature for peer-reviewed studies was scanned. The aim of the literature
search was to target studies that looked at examples of changes in species morphology
due to harvesting. Changes in behaviour, age or sexual skew were not included due to
limitations of the scope.
With careful consideration, as well as a test run of what search terms could target the
relevant papers related to the question, a comprehensive word list was produced prior to
the literature search (appendix 1). Web of Science was used as the platform for the
search, which took place in December 2015. The search results were refined to include
only articles from evolutionary and biological conservation topics as well as including
only document types that were articles, review papers and book chapters. This narrowed
the results down to 1743 articles (step one in figure 1.1.). These articles were then
sorted into two categories – ‘maybe relevant’ and ‘not relevant’ - based on a screening
of the title and abstract (step two in figure 1.1.). Beforehand, a trial of 120 articles had
been sorted through to better the systematic approach in classifying the articles as
relevant or not. The ‘maybe relevant’ category, comprised of 77 articles, was then
investigated more thoroughly and put into three categories – ‘relevant’, ‘supporting’
literature or ‘not relevant’ literature. All the references from the ‘relevant’ articles were
then looked through to find any further literature that would be important to include in
the search, resulting in 36 articles in the final ‘relevant’ category (step three in figure
1.1.).
Figure 1.1. Visualisation of amount of articles after step one, two, three and four in the literature review.
1743 articles
(step one)
77 articles
(step two)
36 articles
(step three)
20 articles
(step four)

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Next, it was interesting to compare the remaining articles in a table to try to systematise
the studies and present the information in a schematic way. Out of the 36 relevant
articles, 20 were considered suitable to put into a schematic table (step four in figure
1.1.) (table 1.2.).
The rest were deemed unfitting since some were literature reviews, (Jerozolimski &
Peres, 2003; Fenberg & Roy, 2008; Hendry et al., 2008; Caro et al., 2009; Allendorf et
al., 2009), some were replicate studies which would have biased the results (Milner-
Gulland & Mace, 1991; Festa-Bianchet & Pelletier, 2014; Pigeon et al., 2016). Others
had some mentioning of change in species traits over time, but either no mention on
what species exactly and what traits were targeted, or the trait mentioned in the study,
was not related to morphology (Brooks et al., 2007; Archie et al., 2008; Darimont et al.,
2009; Ciuti et al., 2012; Bro-Jorgensen, 2014; Tierney et al., 2014; Benitez-Lopez et
al., 2015). There was also one study based on Silverside fish (Menidia menidia) in a
manipulated setting, and therefore not useful in the schematic table (Conover et al.,
2009).

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RESULTS
The literature review found 36 relevant articles on trait-selective harvesting and what
effects this has on species. However, in order to analyse these references further, a
selection of 20 references were chosen. The 20 references from the literature review
contained 34 study species, where 23 of the study species (or 68%) had seen changes to
either body size, antler, tusk or horn size over time. These are presented in a schematic
way to give an overview of the changes seen (table 1.2).
Table 1.1. List of species from the 36 relevant articles in the
literature review.

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Table1.2.Schematicoverviewofarticlesonanthropogenictrait-harvestingfromrelevantarticlesintheliteraturereview.

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24

25

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Figure 1.2. Overview of results from table 1.2. Top left: distribution of terrestrial and non-terrestrial species. Top right:
distribution of classes. Middle left: distribution of ungulates, marsupials and ‘other’ in mammals. Middle right: distribution
of effect and no-effect in mammals. Bottom left: distribution of targeted traits. Bottom right: distribution of sexes.

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Analysing the 34 study species from the 20 references from the literature review (table
1.2.), it was found that out of the 34 study species, 19 were terrestrial species, seven
were marine or freshwater species and two were plants (figure 1.2.).
The 34 study species were comprised of mammal species (n = 18), gastropod species (n
= 4), fish species (n = 2), one reptile species and one crustacean (figure 1.2.).
The mammals consisted mainly of ungulates (n = 12), seven of the study species were
marsupials and three were categorised as ‘other’ (figure 1.2.) (Jachmann et al., 1995;
Monteith et al., 2013; Ingram et al., 2015).
Thirteen of the study species that were mammals saw a change in either body size,
antler, tusk or horn size over time, nine did not. The change could be seen as attributes
getting either smaller or bigger, and did not necessarily mean that individuals were
getting smaller in size or that armaments were being reduced (figure 1.2.).
Of the 34 study species, 17 were targeted due to their body size, 14 were targeted only
for the trophy and three were targeted for both body size and trophy size (figure 1.2.).
Of the 34 study species, 13 were targeted for being males, 11 were not specified (N/A)
and in 10 of the cases no sex was preferred over the other and therefore both sexes were
considered as targets (figure 1.2.).

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EFFECTS OF SELECTIVE HUNTING
The study setups were only comparable in six of the study species, where six of the 34
study species saw an average decrease of 10.7% in horn measurements. The six study
species were all harvested ungulate males (Garel et al., 2007; Hengeveld & Festa-
Bianchet, 2011; Perez et al., 2011; Crosmary et al., 2013).
Therefore, it can be more meaningful to evaluate each study on its own and note the
changes seen. Thus the studies from table 1.2., where the results have been quantified
will be highlighted here:
• Mouflon (Ovis gmelini) saw a decrease between 3.4% and 38.3% in male horn
measurements (Garel et al., 2007).
• The horn length of male California bighorn sheep (Ovis canadensis
californiana) saw a decline of 3.9% in three to five-year olds in their study
period, whilst older males saw a decline in horn measurements of 3.7%
(Hengeveld & Festa-Bianchet, 2011).
• The average horn length of Iberian wild goat (Capra pyrenaica) was 15.4% less
from 1975 to 1985 (Perez et al., 2011).
• Horns of male aoudads (Ammatragus lervia) aged 7-8 years, decreased in length
by 10.9% over the 11-year study period (Perez et al., 2011).
• Tusklessness in female African elephants (Loxodonta africana) has increased
from 10.5% to 38.2% in the population between 1969 and 1989, where
tusklessness in male African elephants increased from 1% in the 1970s to 10%
in 1993 – both as a result of hunting but also migration due to displacement
(Jachmann et al., 1995).
• From 1974 to 2008 the general horn length of harvested male impalas
(Aepyceros melampus) decreased with a loss of 4%.
• Similarly, harvested male sable antelopes (Hippotragus niger) saw a decrease of
6%. The difference in the two responses for impalas and sable antelopes is
explained by the higher hunting pressure and the higher trophy value on sable
antelopes.

29
• Surprisingly, harvested males of greater kudus (Tragelaphus strepsiceros) saw
an increase in horn length of 14% during the same period (Crosmary et al.,
2013). The expectation was that greater kudus would see a horn reduction
similar to the impala and sable antelopes, however the increase could be
explained by an increasing proportion of older males in the harvest and therefore
not necessarily representing an actual increase in male horn length.
• Further, Monteith et al. (2013) conducted a study on 25 species of native North
American big game and saw a decreasing trend in trophy size for 21 of the
species, whilst four saw a positive trend in size. The change in trophy size
ranged from a 4.93% decrease in Columbia black-tailed deer (Odocoileus
hemionus columbianus) to a 0.18% increase in Canadian moose (Alces alces).
The increase in trophy size for four of the species is explained by less hunting
pressure and conservation efforts where reintroductions have supplied the
population with healthy older males. Further, the hunting pressure on antlers can
be hard to estimate as antlers are shed annually, unlike horns which are
permanent structures and size-correlated with age and genetics.

30
DISCUSSION
Changes in biological traits was seen in 68% of the reviewed species (table 1.2.).
Darimont et al. (2009) saw a change in morphological traits such as body and horn size,
in 94.9% of the estimates. They combined data from 29 species and the results were
comprised of 475 estimates where the average decrease for the changes seen in the
results were 18.3%. Mammals were the most likely class in the terrestrial environment
to be targeted by hunting, where ungulates were the majority. Ungulates are commonly
large bodied and carry armaments, thus good targets in trophy hunting (Coltman et al.,
2003). The review could show a taxonomic bias towards mammals, as there is a
disproportionate amount of research on vertebrates, especially mammals and birds
(Czech et al., 1998; Stein et al., 2002). Further, large-bodied mammals are often
selected as ‘flagship’ species to increase awareness, attract political support to a
conservation issue and obtain funding (Clucas et al., 2008; Barnes et al., 2016a).
The data shows no apparent preference for either sexes (table 1.2.; figure 1.2.). This
may be due to the fact that one third of the study objects were not specified as a specific
sex, and therefore could ‘tip the scales’ in favour of the male sex, if the sex was known.
Also, if the male sex is targeted more frequently by hunters, due to their generally larger
body size and proportionally larger armaments than females, it could lead to an
intensification on trait-selectivity. The pressure would thus be greater for males, instead
of being divided between both sexes (Milner et al., 2006).
Mating success for most ungulate species is correlated with body and horn size and
sexual selection has therefore favoured large armaments (Garel et al., 2007; Mysterud,
2012). If this relationship is distorted, there is a strong support for the ‘intensive-
harvest’ hypothesis, where the harvest of males is gradually shifting the age structure
thus younger and smaller males become the dominant group (Monteith et al., 2013).

31
Fenberg and Roy (2008), found in their literature review that a total of 108 species of
fish, invertebrates and terrestrial vertebrates had been subject to size-selective
harvesting. Ungulates made up the majority of the terrestrial vertebrate group, which is
similar to the findings in this thesis (table 1.2.; figure 1.2.). They emphasise that aquatic
species are primarily those who are being affected by size-selective harvesting. Thus,
larger terrestrial vertebrate species are only a minor group when considering the number
of species affected.
Trait-selective harvesting has up until now not been very well studied with only a
limited amount of papers mentioning the phenomenon. Not all of the studies that were
found in the literature review were directly focused on uncovering the effects of hunting
and had vastly different setups; the length of the study periods varied as well as start
year and end year of the observations, studies were conducted on different age groups
and the populations were subject to different management strategies.
Irrespective of the relative roles of phenotypic alterations and plasticity, the serious
extent of the change happening and the rapidness of this shift, can have overwhelming
consequences. The studies in this review indicate that the pressure on certain traits lead
to changes in the species morphology and life history. Human harvesting has the benefit
of being able to select specific traits and keep the pressure on these traits by maintaining
and adjusting their efforts. Exploited species will experience life history alterations and
ecological dynamics will be inflicted. Interacting predators and prey will have to adjust
and keep pace with possible declines in harvestable biomass and population instability.
This issue is in need of long term studies that span worldwide and focus on multiple
classes. Focus should especially be on species who are known to be hunted, whether or
not this is done sustainably, including determining the recovery rate of species who are
experiencing morphological change due to harvesting.

33
INTRODUCTION
The literature review of 21 studies showed that while evidence of the effect of selective
hunting exists, the data originated from different studies with incomparable setups.
Thus, based on the review it was not possible to assess any general trends for exploited
species. To investigate the impact of human use on species further, the Living Planet
Index, or LPI, was included in this analysis on sustainable use of species.
BACKGROUND
The LPI was developed as a WWF project in 1997 to measure the worlds biodiversity
and its change in state over time and aims to measure population trends in vertebrate
species since 1970 (Loh et al., 2005). Thus, the LPI time series data does not tell us
anything about how morphological traits change over time. However, it can give a good
indication of the population trends over time for different taxonomic groups. This,
comprised with the literature concerning morphological change, can begin to draw a
picture of species that are highly controlled by anthropogenic forces in one way or
another. By piecing morphological change over time, together with change in
population trends over time, provides us with an idea of what is happening to species
worldwide.
As species trends can be influenced by numerous factors, the impact of utilisation has
been the main focus. Other factors such as change in precipitation and temperature, or
human disturbances other than hunting such as habitat destruction, air, chemical, noise
and light pollution as well as invasive species, diseases, nutritional state and so on,
should of course also be recognised as factors that influence population fluctuations.
However, it would be impossible to cover them all in this thesis. Therefore, the results
should be seen as an attempt to explain what can be observed in relation to human use
and how that affects population abundance.
Use, such as hunting, fishing and harvesting is a big part of how humans uphold the
need for food and other items that come from animals (Plummer et al., 2009). The

34
extent of this use is also the reason that many believe we are heading towards a massive
extinction crisis, if we do not rectify our direct and indirect use of natural resources and
other ways we negatively impact our planet (Pimm et al., 2001; Ceballos & Ehrlich,
2002; Hoekstra et al., 2005). This is why it is important to monitor the state of all living
things to be able to focus conservation management strategies towards the ones that are
in the most crucial state.
AIM
The aim of the second part of this thesis was to see how population abundance has
changed over time in relation to utilised and not utilised populations, by (1) analysing
the global trend for all vertebrate populations over time, especially mammals and birds,
(2) analysing population trends for all vertebrate populations in the Afrotropical and
Palearctic realm, and (3) analysing population trends for all vertebrate populations in
protected areas.

35
METHODS
THE LIVING PLANET INDEX
The Living Planet Index is a measure of the state of global biological diversity based on
population trends of vertebrate species from around the world and can be accessed via
an online portal. The database currently holds time series data for over 18,000
populations of more than 3,500 mammals, bird, fish, reptile and amphibian species
which are gathered from sources such as journals, online databases as well as
government reports and dates back to 1970 (Loh et al., 2005). The population time
series data is augmented with a variety of information such as population taxonomy,
location and ecology as well as what form of interaction they might have with humans.
The data set has a category that indicates whether a species has been ‘utilised’ or not by
humans and is categorised as such at population level. The definition of ‘utilised’ can be
found in the nomenclature. The LPI is calculated with a generalised additive modelling
framework to determine the underlying trend for each population time series. Average
rates of change are then calculated and aggregated to either species or population level
(Loh et al., 2005; Collen et al., 2009).
Data was pulled from the LPI time series data provided by the Zoological Society of
London and used in R. Seeing as the category ‘utilised/not utilised’ in the data is tagged
to population level, this category was chosen for the next part of the analysis. This gives
a better estimate of the state of different harvested populations, thus, developing an
indicator for use: The Utilised Species Index (Tierney et al., 2014). The index was
calculated using time series data from 1970 to 2014 for 7191 populations, where 2043
populations were coded as ‘utilised’ and 5148 were coded as ‘not utilised’. Following
Loh et al. (2005) and as revised by Collen et al. (2009), a bootstrap re-sampling
technique was used to generate annual 95% confidence intervals (CI) around each index
value.

36
METHODOLOGY
Seeing as the category for ‘utilised’ is fairly new, not all 18,000 populations in the time
series data have been assigned a value (0=not utilised, 1=utilised, 2=unknown), limiting
some of the combinations for ‘utilised’, ‘class’ and ‘realm’. Unknown entries were
excluded from the analysis. For overview of data points for all combinations of the data
selection see appendix 3-7.
The analyses were conducted by selecting three categories. First, either ‘utilised’ or “not
utilised” was chosen. Then all vertebrate species were included; Actinoopterygii, Aves,
Mammalia, Reptilia, Amphibia, Cephalaspidomorphi, Elasmobranchii, Holocephali,
Chondrichthyes, Myxini and Sarcopterygii, henceforth all vertebrate populations or all
vertebrate species. All regions were then selected to give a global trend (step one, figure
2.1). The same was done for mammal populations and bird populations worldwide. To
further analyse trends happening within the combinations in step one, a diagnostics
package in R was used to disintegrate the results seen for these combinations.
The same procedure as in step one, was followed for the two realms, the Afrotropical
and the Palearctic (step two), where instead of selecting all regions, either the
Afrotropical realm or the Palearctic realm was selected in the category ‘T_realm’
(appendix 2). Next, the diagnostics package was again used to additionally investigate
the two realms (step two, figure 2.2.).
Utilised, Not Utilised
All vertebrate
species, mammals,
birds
Global trend
Utilised, Not Utilised
All vertebrate
species, mammals
and birds
Afrotropical realm
and Palearctic realm
Figure 2.1. Overview of the process of the data selection (step one).
Figure 2.2. Overview of the data selection (step two).

37
Additionally, the LPI data set allows for the selection of protected areas. Thus, an
analysis on protected areas was carried out, based on the procedure in step one. This
analysis was done by selecting for either ‘utilised’ or ‘not utilised’, where all species
had to be combined to ensure enough data points. In the category ‘Protected_status’,
‘yes’ was selected (Protected status + yes = Protected area) (step three). Again, a
diagnostics package was used.
Further, the average height and weight of utilised and not utilised bird species was
found as well as the proportion of ungulates and armaments in utilised and not utilised
mammal species (appendix 8-11).
Another factor to acknowledge is the proportion of different threat statuses among the
populations. The LPI data set has information on the threat status of populations based
on the IUCN red list, which meant that the ‘threatened categories’ such as ‘Critically
endangered’, ‘Endangered’ and ‘Vulnerable’ could be selected to give an estimate of
how big a proportion of populations were under threat (appendix 12-16).
Utilised, Not Utilised All vertebrate species Protected areas
Figure 2.3. Overview of data selection (step three).

38
RESULTS
GLOBAL TRENDS
The global trend for all utilised vertebrate populations shows a decline of c. 25%
between 1970 and 2014 (figure 2.4., left panel; 2014 Utilised Species Index = 0.75, CI =
0.59-0.98). The global trend for not utilised vertebrate populations shows an increase of
c. 90% between 1970 and 2014 (figure 2.4., right panel; 2014 Utilised Species Index =
1.90, CI = 1.64-2.10).
Utilised Not Utilised
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1.8
2.0
1970
1975
1980
1985
1990
1995
2000
2005
2010
1970
1975
1980
1985
1990
1995
2000
2005
2010
years
Index(1970=1)
group
Utilised
Not Utilised
Figure 2.4. Trends (± 95% CI) for all species in the world. Left: utilised
populations. Right: not utilised populations. The lighter colour shading represents
confidence intervals. The y-axis indicates the percentage change from baseline level
at 1.

39
The global trend for utilised mammal populations shows an increase of c. 3% between
1970 and 2014 (figure 2.5., left panel; 2014 Utilised Species Index = 2.03, CI = 0.60-
1.62). The global trend for not utilised populations of mammals shows an increase of
over 100% from 2002 (figure 2.5., right panel; 2002 Utilised Species Index = 2.0, CI =
1.64-N/A).
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1.8
2.0
1970
1975
1980
1985
1990
1995
2000
2005
2010
1970
1975
1980
1985
1990
1995
2000
2005
2010
years
Index(1970=1)
group
Utilised
Not Utilised
Figure 2.5. Trends (± 95% CI) for mammal populations around the world. Left:
utilised populations. Right: not utilised populations. The lighter colour shading
represents confidence intervals. The y-axis indicates the percentage change from
baseline level at 1.

40
The global trend for utilised bird populations shows a decline of c. 45% from 1970 to
2012 (figure 2.6., left panel; 2012 Utilised Species Index = 0.55, CI = 0.24-1.2). The
global trend for not utilised bird populations increases by c. 80% from 1970 to 2012
(figure 2.6., right panel; 2012 Utilised Species Index = 1.80, CI = 1.65-N/A).
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1.8
2.0
1970
1975
1980
1985
1990
1995
2000
2005
2010
1970
1975
1980
1985
1990
1995
2000
2005
2010
years
Index(1970=1)
group
Utilised
Not Utilised
Figure 2.6. Trends (± 95% CI) for bird populations on a global scale. Left: utilised
populations. Right: not utilised populations. The lighter colour shading represents
confidence intervals. The y-axis indicates the percentage change from baseline
level at 1.

41
REALMS
The Afrotropical realm shows a decline of c. 90% for utilised populations from 1970 to
2010 (figure 2.7., left panel; 2010 Utilised Species Index = 0.10, CI = 0.10-0.19). The
trend for not utilised populations in the Afrotropical realm shows an increase of c. 30%
from 1970 to 2010 (figure 2.7., right panel; Utilised Species Index = 1.30, CI = 0.85-
2.20).
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1.8
2.0
1970
1975
1980
1985
1990
1995
2000
2005
2010
1970
1975
1980
1985
1990
1995
2000
2005
2010
years
Index(1970=1)
group
Utilised
Not Utilised
Figure 2.7. Trends (± 95% CI) for populations of all species in Afrotropical realms.
Left: utilised populations. Right: not utilised populations. The lighter colour shading
represents confidence intervals. The y-axis indicates the percentage change from
baseline level at 1.

42
The Palearctic realm shows for utilised populations an increase of over 100% from 2005
(figure 2.8., left panel; 2005 Utilised Species Index = 2.0, CI = 1.40-N/A). The trend for
not utilised populations in the Palearctic realm shows an increase of over 100% from
1993 (figure 2.8., right panel; 1993 Utilised Species Index = 2.0, CI = 1.60-N/A).
Figure 2.8. Trends (± 95% CI) for populations of all species in the Palearctic realm. Left:
utilised populations. Right: not utilised populations. The lighter colour shading represents
confidence intervals. The y-axis indicates the percentage change from baseline level at 1.
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1.8
2.0
1970
1975
1980
1985
1990
1995
2000
2005
2010
1970
1975
1980
1985
1990
1995
2000
2005
2010
years
Index(1970=1)
group
Utilised
Not Utilised

43
PROTECTED AREAS
The Utilised Species Index shows a decline of c. 35% in utilised populations in
protected areas from 1970 to 2014 (figure 2.9., left panel; 2014 Utilised Species Index =
0.65, CI = 0.35-1.24). The Utilised Species Index for the not utilised populations
increases with over 100% from 1970 to 2010 (figure 2.9., right panel; 2005 Utilised
Species Index = 2.0, CI = 1.7-N/A).
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
1.6
1.8
2.0
1970
1975
1980
1985
1990
1995
2000
2005
2010
1970
1975
1980
1985
1990
1995
2000
2005
2010
years
Index(1970=1)
group
Utilised
Not Utilised
Figure 2.9. Trends (± 95% CI) for populations of all species worldwide in protected
areas. Left: utilised populations. Right: not utilised populations. The lighter colour
shading represents confidence intervals. The y-axis indicates the percentage change
from baseline level at 1.

44
DISCUSSION
REALMS
Notably the utilised populations in the Afrotropical realm are declining with over 90%,
almost nearing extirpation. The decline in utilised populations is in itself not surprising
as ‘utilised’ indicates an off-take, which would be expected to be apparent in a trend for
utilised populations. However, the magnitude of the decline is excessive illustrating a
grim outcome for future perspectives as 55.8% of the same group of vertebrates are also
threatened species (appendix 15). In comparison, the LPI by biogeographic realms
(WWF, 2014) showed for the Afrotropical realm a decline of 19%. However, this
finding does not distinguish between utilised and not utilised species. Further, the
number of not utilised populations is four times larger than the number of utilised
populations (appendix 4), thus when both groups are combined, the general trend could
smooth out the steep decline in utilised populations. The Afrotropical realm shows trend
lines in utilised and not utilised populations similar to African mammal and bird
populations (appendix 23-24).
The Palearctic realm showed an increase in utilised and not utilised populations, in stark
contrast to the trends we see for populations in the Afrotropical realm. The overall
increase for the trend could perhaps be due to the successful conservation efforts in the
Arctic driving the trend upwards (Tierney et al., 2014). People are perhaps more likely
to rely on use of species that are more common and therefore more easily available.
Moreover, utilised species might be more effectively regulated and provided with
greater protection than not utilised species to ensure supply for the demand. This is in
line with findings that the increased sport hunting in the Arctic areas has become much
more regulated and sustainable (Tierney et al., 2014). Correspondingly, the utilised
populations from the Utilised Species Index experience a relatively low threat status at
9.8% (appendix 15).
Even though not utilised populations are not under threat from human use, there are
other human disturbances such as forest loss and overall habitat degradation that could

45
be putting pressure on this group (Norris et al., 2010). The pool of not utilised
populations is also larger than the utilised populations and may be a lot more diverse in
their morphology and life history than utilised populations (Sæther & Bakke, 2000;
Hoffmann et al., 2010). The trend lines will show a consistent tendency, if species have
similar life histories, perhaps eliciting same responses to threats, where confidence
intervals might be smaller than expected even though number of data points are limited.
For example, 86% of utilised mammals in Africa were ungulates and 74% of utilised
mammal populations in Africa are known to bear armaments, compared to not utilised
mammal populations in Africa where 78.5% were ungulates and only 54.5% are known
to bear armaments (appendix 10-11). Similarly, there was a difference in height and
weight of utilised and not utilised bird populations in Africa, where not utilised birds
tended to be smaller but heavier than utilised birds who were bigger but lighter
(appendix 8-9). Also, a large proportion of the utilised bird populations were black-
footed penguins (appendix 8). The large proportion of ungulates and armaments in
utilised mammal populations and black-footed penguins in the utilised birds could be
the reason for consistent trend lines with subsequent small confidence intervals,
compared to the not utilised populations.
The challenge faced when making comparisons between realms and other areas, is that
estimates of total population size can be difficult to obtain. Estimates may only be
available for parts of the population and in parts of its range and therefore not fully
representing the species trend. Further, the representation of all vertebrate classes,
populations and realms in the index are not always accurate (Tierney et al., 2014).
Moreover, there may be other factors than utilisation impacting the populations in the
Utilised Species Index, such as in the case of the African elephant (Loxodonta
africana), where the primary threat is listed as ‘habitat loss’ and ‘exploitation’ is listed
as a secondary threat, or in the case of the European roe deer (Capreolus capreolus),
where ‘invasive species’ are listed as the primary threat, and ‘exploitation’ as a
secondary threat. Therefore, it can be difficult to determine to what extent usage is the
main driver (Tierney et al., 2014).

46
PROTECTED AREAS
The Utilised Species Index showed that protected areas saw a 37% decline in utilised
populations and a 100% increase in not utilised populations from 1970 to 2014. In
comparison, the terrestrial LPI of populations inside protected areas showed a decline of
18% between 1970 and 2010 (WWF, 2014). However, the report does not distinguish
between utilised and not utilised populations. Utilised species seem to be faring better
on a global scale than in protected areas, as the global trend for utilised populations saw
a decline of 25%. Though, this comparison should be seen in the light of the underlying
data as protected areas are also part of the global trend, and the composition of
vertebrate classes, regions and so on, are largely dissimilar between the two groups. For
example, the region with the greatest amount of utilised populations in protected areas
is Latin America and the Caribbean, whereas the region with the most utilised
populations on a global scale is North America.
The amount of protected areas in the world has risen tenfold from 1950 to 2009,
however it does not seem to have been of much help to the worlds species raising
concern about the quality of protected areas and the management systems (Kareiva et
al., 2012; Geldmann et al., 2015). Craigie et al. (2010) reported on a 59% decline in
African mammals and stated that there has clearly been a failure in protected areas in
Africa, especially in the western parts, to protect its mammals. It can be argued that
protected areas are non-strategically placed compared to threatened species and that
they should be better connected (Juffe-Bignoli et al., 2016; Santini et al., 2016). Further,
it seems that small to intermediate sized species who are under threat from utilisation
are impacted particularly badly due to management efforts and external funding being
prioritised for larger-bodied, charismatic flagship species (Barnes et al., 2016a). The
socio-economic context of the management and maintenance of protected areas and the
wildlife within those boundaries is critical (Barnes et al., 2016a). Thus, the measures
taken should be more case specific and more focused on threat mitigation (Chape et al.,
2005; Santini et al., 2016).

47
Protected areas can deliver successful outcomes but it seems there is an unwillingness
for countries to declare new protected areas in areas where biodiversity is the most
threatened and at the same time ensure a management form that is consistent and
committed to providing a positive outcome for the biological community (Barnes et al.,
2016b).
GLOBAL TRENDS
The Utilised Species Index trend for all vertebrate populations on a global scale showed
a decrease of almost 25% from 1970 to 2014, in line with expectations, where the
terrestrial LPI shows a decline of 39% between 1970 and 2010 (Hoffmann et al., 2010;
WWF, 2014). The primary threats to LPI populations are exploitation (37%) and habitat
degradation and habitat change (31.4%) according to WWF (2014). The Utilised
Species Index data showed a decline in fish population abundance, where fish made up
77% of utilised populations, in line with the general perception of diminishing fish
stocks (Hutchings & Reynolds, 2004). Since birds were the predominant group with
64% in the not utilised populations, the trend for all vertebrates seems to be highly
driven by the not utilised bird populations.
A study showed a decline of 14% for utilised species at a global level between 1970 and
2007 (Tierney et al., 2014) (appendix 25), compared to the 25% decrease found for all
utilised vertebrate populations in this thesis. Both results show that populations are
being unsustainably exploited and the difference between the two results could perhaps
be explained by the difference in class composition. Tierney et al. (2014) reported that
88% of the time series data used to generate the global index relate to birds, while the
time series data used in this thesis is represented by 77% birds. Further, Tierney et al.
(2014) did not define the usage to population level.
For both birds and mammals, not utilised populations followed the same increasing
trend. However, utilised bird populations decrease with 45% between 1970 and 2010,
whereas mammal populations stayed near the baseline level from 1970. The primary
threat for 87% of utilised bird populations was habitat degradation or habitat loss, where

48
utilisation was indicated as a secondary threat. Findings suggests that bird populations
have declined by c. 22% from 1700s to 1990s and that the cause of this decline is
mostly due to habitat degradation from change in land use by humans (Gaston et al.,
2003). The primary threat for mammal populations was exploitation where the
proportion of threat status in mammals whether utilised or not utilised were very
similar, 21% and 26% respectively. These findings are consistent with Hoffmann et al.
(2010) who showed that 25% of mammals are classified as threatened. This indicates
that mammals have on a global scale not changed significantly in abundance which
might be driven by the 21-26% of threatened species being overridden by species who
are not threatened and overall doing well.
LIMITATIONS AND STRENGTHS TO THE LPI
In order to halt and reverse the loss of global biodiversity and population abundance
there needs to be in place a tool for measuring and assessing the loss. Conservation
efforts should be judicious and monitored in order to help future efforts. The LPI could
be one of the tools needed to accurately be able to estimate if objectives, such as the
SDGs and the Aichi Biodiversity Targets, are being met. However, the limitations to the
index should be recognised. The taxonomic scope of the index is heavily dominated by
birds and mammals with very little data on amphibians, which is the taxonomic group
that is most in decline (Hoffmann et al., 2010). Reptiles and fish are also largely
underrepresented. The geographic coverage of the LPI is weighted so that regions
should be equally addressed, however regions that are better studied make up a larger
proportion of the data and therefore an imbalance between different regions can not be
completely ruled out (Loh et al., 2005). Further, it should be noted that the trends for
populations in most recent years might be exaggerated because of reduced sample sizes.
The LPI consists of thousands of different studies and this might be both the biggest
strength but also the biggest weakness of the index. It is very useful to be able to
aggregate global population time series trends for a large number of species, however
the way information has been collected can turn the underlying data into smoke screens.
Data may be collected for large and wide-ranging populations one year and smaller and

49
less stable populations another year. Species may be long-lived or short-lived and have
different life strategies concerning the trade-off between quantity and quality of
offspring and different ways of responding to threats (Taylor et al., 1990). All this the
LPI does not account for in its weighted system (Loh et al., 2005; Collen et al., 2009).
Nevertheless, the way that the LPI can be fragmented into subsets and the large amount
of information that has been augmented for each entry is one of its great strengths. This
is useful knowledge and can be an effective indicator for future use in conservation
efforts. The LPI is already a useful instrument for monitoring global sustainable targets
however, the index should be continuously updated and augmented with new
information so that it can become an even more reliable tool.

50
GENERAL DISCUSSION
Since the Stockholm conference in 1972 and later the Brundtland report, issues
concerning humans’ use of the environment have been laden with appeals to manage
natural capital sustainably. Probably the biggest challenge faced when addressing
sustainability issues is the uncertainty about what actions should be taken, an issue to be
addressed at all levels. Locally, environmental managers should be able to follow
simple guidelines that do not require development of new technologies or heavy
paperwork and should be tailored to the type of area that it is concerning to avoid
confusion for the management. Nationally, the measures should comply with the
existing policies and way of living otherwise the task may be too hard to fulfil as
political agendas may be conflicting. Globally, it can be hard to obtain an agreement
and even harder to enforce this as ways of measuring and assessing improvements are
still far apart (Dovers, 1997). Another challenge is the way our economy through
market trade and lack of pricing-in the cost of nature is pressuring biodiversity and
ecosystems. This is happening in such a way that it is becoming clear that this
relationship between humans and nature cannot go on, if future generations are to have
the same amount of resources as previous generations (Bishop, 1993).
In 2015, the Millennium Development Goals (MDGs) expired after having been the
focus of global policy debates for 15 years. The MDGs have been a step in the right
direction for global efforts to be made for widespread societal concerns including
environmental degradation. The substantial progress, and some failures, towards
achieving the MDGs, sparked the establishment of the SDGs. The SDGs include both
developing and developed countries and they state much clearer what the goals aim to
do. Especially goal 12, 14 and 15 have incorporated sustainable objectives deeper into
plans for biodiversity and ecosystems (Sachs, 2012; Griggs, 2013). Similarly, the Aichi
Biodiversity Targets were adopted in 2011 at the 10th
Conference for Parties to the
Convention on Biological Diversity and will expire in 2020. The targets, particularly 11
and 12, describes its aims to ensure improvements to biodiversity by sustainable use and
how conservation networks should look like for terrestrial and inland water as well as

51
coastal and marine areas. Further, extinction of known threatened species should be
prevented and species in decline should be improved and sustained. However, both the
SDGs and Aichi Targets lack ways to monitor the progress (Juffe-Bignoli et al., 2016).
The direct use of wild living resources still remains essential for many people’s
livelihoods, while the indirect use of ecosystems is crucial for all humans. Both
practices mostly have been and still are, unsustainably conducted. The separation of
humans and nature seems difficult as 300 million people in traditionally organised
societies occupy almost one-fifth of Earth’s surface where they depend heavily on
fishing, gathering and hunting of terrestrial resources (Hutton & Leader-Williams,
2003). It is still believed that protected areas are a good conservation tool (Geldmann et
al., 2015) and a ‘cornerstone of biodiversity conservation strategies’ (Brink et al.,
2016), where Aichi Target 11 wants to include 10% of Earth’s surface within protected
areas. Although opting for areas to become ‘strictly protected’ is only realistic in a very
small portion of the Earth and therefore a more pragmatic approach would be to
encompass humans into nature management (Hutton & Leader-Williams, 2003).
The seemingly trade-off between nature and people derives from a conservation strategy
that has focused on hotspots. Areas that are deemed hotspots are then established as a
national park or reserve to protect animal and plant life, at the expense of local people
who are often displaced or lose the access to resources that their livelihoods depend on.
Conversely, the ‘ecosystem services’ strategy is becoming more popular. This strategy
focuses on areas that are being degraded while still including the community in the
conservation plans (Kareiva & Marvier, 2007). In order to really have an impact on
improving sustainability the support from the affected local communities in the area is
important to establish. The local communities should be equipped with the
responsibility and ownership of the land-area as well as incentives for land use
strategies that will benefit species (Hutton & Dickson, 2000; Apensberg-Traun, 2008).
Not only will biodiversity and ecosystems benefit from this, but it will also force the
international political arena to recognise that local people are a part of these ecosystems
and therefore also a part of the solution. Further, by empowering them, the livelihoods

52
for present and future generations in the local communities will be greatly improved,
which is a step closer to the SDG on eradicating poverty everywhere (Scheyvens, 1999).
The role of private sectors providing funding for protected areas is an increasing
practice, as protected areas are often having difficulties covering their costs. To
generate revenue, the private sector can organise hunting tours of wild animals. Of
course, the exploitation of species is concerning, yet it also creates an incentive for the
management to maintain populations at a level that still makes hunting profitable,
providing that the tenures are long-term (Brink et al., 2016). Further, the areas used for
hunting of wild animals will subsequently avoid being converted into agricultural land,
the practice will protect the species living there and can benefit local people (Lindsey et
al., 2006; Brink et al., 2016). Lindsey et al. (2006) stress that trophy hunting should
only be an alternative when conditions are unsuitable for ecotourism, and when there
are not enough tourists to generate revenue. Still, there is a need for an effective
regulatory framework as Africa’s protected areas do not adequately conserve
biodiversity. McShane et al. (2011) calls for realistic expectations to the apparent ‘win-
win’ approaches in conservation plans where both wildlife and humans are benefitted.
Trade-offs seem inevitable when so many factors need to be taken into account.
The threats that species are subject to should be characterised to ensure a basis for better
decision making. Thus, assembling morphology, function, physiology, behaviour,
habitat use, reproduction, adaptability and life history for species, should make it
possible to figure out the relationship of biological traits to disturbances such as
increased hunting pressure, habitat degradation and climate change, whilst factoring in
ways to improve or maintain local communities’ livelihoods (Turak et al., 2016).

53
CONCLUDING REMARKS
It is no longer debatable that humanity’s actions are becoming a problem for not only
every living entity around us, but for ourselves as well. Sadly, the latter is probably
what will generate the biggest effort. The SDGs are perhaps the greatest attempt to date
at fighting this unfortunate development and the objectives of the SDGs should be seen
as interacting synergistically with each other as well as the Aichi Targets, taking the
experience with the MDGs into account, where measurability is key.
Although the LPI has faults that can easily be misinterpreted it still stands as an
excellent measuring tool. Many of the problems with over-representation of some
classes and regions can be weighted differently in a way that allows for improvements.
Further, the information that the index holds is being expanded on continually (Loh et
al., 2005). The implementation of the SDGs should thus be supported by the many
attempts in science to quantify the changes that are seen (Griggs et al., 2014). Future
recommendations therefore include the need for policies and science to interact at a
larger scale and draw benefits from each other. To continue to develop techniques for
ways of measuring the state of our planet and its natural capital and to expand on
existing databases, such as the LPI initiative. And finally, to make these databases
available and accessible to all, for example via an online portal as with the LPI,
allowing knowledge to be transparent and to be circulated in order to encourage
accountability.

54
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APPENDICES
Appendix 1: List of search terms for literature review in part 1.

61
Appendix 2: World map of realms and biomes.

62
Appendix 3: Data points for regions and classes for utilised and not utilised
populations.
Appendix 4: Data points for utilised and not utilised populations in realms.
Class Realms Datapoints
(Util/Not Util/
Unknown)
Util/Not Util
All Afrotropical 1008 43/178
All Antarctic 3 0/1
All Australasia 265 7/145
All Indo-Malayan 290 12/64
All Nearctic 3108 62/1487
All Neotropical 529 6/161
All Oceania 82 0/32
All Palearctic 1915 113/575
Utilised/Not Utilised
World, Birds 151/3308
World, Mammals 234/825
World, All species 2043/5148
Africa, Birds 21/126
Africa, Mammals 43/121
Africa, All species 196/322

63
Appendix 5: Data points for utilised and not utilised birds in realms.
Class Realms Datapoints (Util/Not
Util/ Unknown)
Util/ Not Util
Aves Afrotropical 111 2/40
Aves Antarctic 2 0/0
Aves Australasia 192 5/105
Aves Indo-Malayan 117 6/22
Aves Nearctic 2515 20/1282
Aves Neotropical 301 1/50
Aves Oceania 69 0/25
Aves Palearctic 1244 15/350
Appendix 6: Data points for utilised and not utilised mammals in realms.
Class Realms Datapoints
(Util/Not
Util/
Unknown)
Util/Not Util
Mammalia Afrotropical 872 41/116
Mammalia Antarctic 1 0/1
Mammalia Australasia 35 2/6
Mammalia Indo-Malayan 172 6/42
Mammalia Nearctic 491 41/141
Mammalia Neotropical 119 4/67
Mammalia Oceania 5 0/0
Mammalia Palearctic 634 98/194

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