Reproductive success in the Iberian

Lynx (Lynx pardinus): Effects of prey

abundance, individual age and
population sex-ratio

Guillermo Uceda Gómez (5581664)

Minor Research Project
Environmental Biology Master Program
Utrecht University
2017
Abstract
The Iberian lynx (Lynx pardinus, Temminck 1827) is the most endangered felid species in the

world. It is at the verge of extinction, despite of all the effort made up to today. The species

relies almost exclusively on the European rabbit (Oryctolagus cuniculus L.), whose populations

have sharply declined in the Iberian Peninsula in the last 50 years. Prey abundance, individual

and population traits are important factors in determining reproduction success. Thus, this study

aims to explore the combine effect of rabbit abundance, individual female age and population

traits (sex-ratio and density) on the reproduction of the Iberian lynx. The results suggest that

Iberian lynx reproductive success (probability of producing offspring) is positively correlated

with prey abundance and female age. However, litter size was not significantly influenced by

any of the study factors, although the model averaging approach showed that prey abundance

was by far the most important factor (relative importance 0.59) followed by sex-ratio (0.35) and

female age (0.27). The present study emphasizes the importance of optimal prey abundances

for lynx reproduction success. However, further research is needed to clarify whether other

factors may be modulating Iberian lynx litter size.

Key words: Iberian lynx, extinction, reproduction, population and individual traits, and prey
abundance

Note: Front cover picture taken by Dr. Alfonso San Miguel Ayanz
TABLE OF CONTENTS

1. Introduction .............................................................................................................. 3
2. Materials and methods ............................................................................................. 8
2.1. Study species ...................................................................................................... 8
2.1.1. Feeding ecology.......................................................................................... 8
2.1.2. Habitat requirements................................................................................... 9
2.1.3. Reproduction ............................................................................................ 10
2.2. Study area: Doñaña-Aljarafe............................................................................ 11
2.3. Data collection ................................................................................................. 13
3. Results ..................................................................................................................... 18
4. Discussion ............................................................................................................... 24
5. Conclusions. Future perspectives for the Iberian Lynx ....................................... 27
6. Acknowledgements ................................................................................................. 28
7. References ............................................................................................................... 29
8. Appendix ................................................................................................................. 33
 1. Introduction

In the past decades, wild animal populations have been subject of an exacerbated human

pressure that has led them to a strong decline in their numbers and habitat range (Hoffmann et

al., 2010). This process of animal biodiversity loss is manifested in species extinction rates that

surpass background rates by two or three orders of magnitude (Pimm et al., 1995). Almost one-

quarter of the world’s mammal species are globally considered threatened or extinct

(www.iucnredlist.org; Galetti & Dirzo (2013)). If the current rates of species extinction

continue, Earth will witness the sixth massive extinction in two centuries (Barnosky et al.,

2011). The major drivers of this loss are habitat destruction, alien species introduction, climate

change and pollution (Wilcove et al., 1998).

Species with small population sizes, slow life histories (e.g. small litters), small

geographical ranges, large home ranges and that are at high trophic levels are more vulnerable

to enter in extinction vortexes (Purvis et al., 2000; Cardillo et al., 2005; Davies et al., 2016).

Thus, predators, and in particular specialists, such as the Iberian lynx (Lynx pardinus,

Temminck, 1827) are more prone to extinction due to their life history traits (Bruno &

Cardinale, 2008).

Predators are keystone species in maintaining the structure and stability of communities and

their removal can have several cascading, indirect effects on the entire ecosystem (Duffy, 2003).

Duffy (2003) referred to this process as trophic skew, when a strong shift in the number of

species between consumer and prey trophic levels occurs. This has the potential of altering the

structure and dynamic of the entire food web. However, predator extinctions do not occur alone,

instead, they are usually accompanied by the insertion of other predators, along with, the

addition and/or removal of other species at other trophic levels (Bruno & Cardinale, 2008).

Thus, predator extinction can strongly modify the functioning of the entire ecosystem, in terms

of biomass distribution and energy flow among trophic levels (Shurin et al., 2002).

3
 Iberian lynx, a top predator in the Mediterranean Iberian Peninsula, is susceptible to

extinction due to its life history and ecological traits. In fact, it is considered the most threatened

felid species in the world (Nowell & Jackson, 1996) and listed as endangered by the

International Union for the Conservation of Nature (Rodríguez & Calzada, 2015). Due to its

endangered status, the species is subject to a long-term conservation project (Iberlince LIFE

Project; www.iberlince.eu; Fig. 1). The species still thrives in eight subpopulations spread

around the south-west Iberian Peninsula, harbouring 483 individuals (Simon, 2016). Currently

the species geographic range expands 1600 km2 only in Andalusia, the region that shelters most

individuals.

Lynx decline started in the early 20th century when hunting and trapping were the major

sources of mortality, as well as, the associated predator control and the exploitation of the

European rabbit (Oryctolagus cuniculus) (Rodriguez & Delibes, 2004). However, the Iberian

lynx populations experienced a sharper decline during the 70s. During these years, the lack of

prey, the European rabbit (accounts for 80 to 90 % of the lynx diet), constrained the growth of

lynx populations (Simón et al., 2012). The European rabbit was affected by two diseases, the

myxomatosis and the Viral Haemorrhagic Disease (VHD) that made drop rabbit populations

very rapidly. Thus, Lynx individuals have to look for food further away risking their lifes. In

this sense, more recently, road casualties have produced several deaths each year, for instance

15 in 2016 (Simón, 2016). Moreover, Ferreras et al. (2010) stated that the transformation of the

landscape into agricultural land has remarkably contributed to the decline of the species. This

trend has reduced the amount of suitable habitat for the lynx, as well as, the amount of

appropriate areas for reintroduction.

In addition, current populations have an insufficient effective size to sustain long-term

viable populations (Simón et al., 2012). Hence, the extant populations are menaced by the

deleterious consequences of inbreeding and the loss of genetic diversity (Abascal et al., 2016).

4
Biased sex-ratio, decreased litter size and age of territory acquisition, together with an increased

mortality due to natural causes are among these consequences (Palomares et al., 2012).

The Iberian lynx conservation project has launched many actions to recover the lynx

populations, as well as to improve its habitat. In this regard, the project has actively work to

increase the carrying capacity of the lynx areas by means of increasing the amount of available

prey. Furthermore, the project has implemented some measures to reduce road casualties such

as ecological corridors to facilitate the dispersion of the individuals (Simón et al., 2012).

Awareness campaigns have also been launched in and around lynx areas, as well as, in

reintroduction sites (Rodríguez & Calzada, 2015). Thus, the program raises awareness about

the benefits of preserving the lynx (i.e. ecosystem services).

Given its critical situation status, the Iberian lynx conservation program has also planned

translocation and reintroduction schemes (Simón et al., 2012). Thus, a few individuals have

been translocated to increase the genetic diversity of remnant subpopulations. Moreover, every

year 60 to 70 individuals are reintroduced from the ex-situ program since its kick-off in 2005

(Vargas, Beston, & House, 2009).

Despite all the efforts made up to day, the species is still threatened with extinction. Thus,

it is still necessary to keep on conducting fundamental and applied research for generating

instrumental information to improve the management plans and to mitigate the decline of the

species. Little is known about the combined effects of lynx population traits (density and sex-

ratio) and prey population features (abundance) on the reproductive success (production of

offspring) of the Iberian lynx. Thus, it is important to know which factors affect Iberian lynx

reproduction and to determine their relative importance. Palomares et al. (2005) pointed out

that the reproductive success of the breeding females is not affected by the abundance of

resources, however, other authors have stated the opposite (Simón et al., 2012). On the other

hand, prey availability affected the reproductive output of other species of carnivores (e. g. Red

5
fox (Vulpes vulpes) in north-eastern Spain (Villafuerte et al., 1996)). In this sense, litter size

and the recruitment rate (the number of surviving cubs) became smaller when the available prey

was low. Additionally, prey availability may affect the pregnancy rate of females (Fuller &

Sievert, 2001), thus, less females get pregnant when the abundance of prey is low. Furthermore,

Ferreras et al. (1997) stated that reproduction in Iberian lynx females, as expected, was

influenced by individual female age due to the bad effects of reproductive senescence.

Consequently, the aim of this study is to test whether rabbit abundance (main prey of the

Iberian lynx) and lynx population (density and sex-ratio) and individual (age) traits have any

influence on the reproductive output (fertility and litter size) of territorial Iberian lynx females.

For this purpose, data from one of the oldest remnant subpopulations (Doñana-Aljarafe) will be

analysed. I hypothesized that rabbit abundance will be positively correlated with reproduction

success, measured as both fertility rate and litter size. Similarly, I hypothesize that lynx

population traits (density and sex-ratio) will mediate in the overall reproduction success. As

Iberian lynx is considered a monogamous species (Ferreras et al., 1997), I expect greater

reproduction success at balanced sex-ratio (1:1). Since sex-ratio and population density are

usually highly correlated in mammal populations (Kruuk et al., 1999), I will examine the

possible relationship between both populations traits and attempt to disentangle their effect on

the overall reproduction success of Iberian lynx. Finally, female age is expected to significantly

influence both, reproductive success and litter size. Thus, I expect that age will negatively affect

reproductive success and litter size, due to the effect of reproductive senescence.

6
Figure 1. Map of the Iberian lynx metapopulation in Spain and Portugal. Different colours depict
the location of the different subpopulations of Iberian lynx in 2016. Orange, Toledo Mountain
range; Yellow, Eastern Sierra Morena; Dark purple, Guarrizas Valley; Red, Andujar-Cardeña;
Light green, Guadalmellato Valley; Light purple; Matachel Valley, Marine blue, Guadiana
Valley (Portugal); and Dark blue, Doñana-Aljarafe (IberLince LIFE Project, 2016;
www.iberlince.eu).

7
2. Materials and methods

2.1. Study species

2.1.1. Feeding ecology

The Iberian lynx is a super-predator specialized in preying upon European

rabbit. According to several studies the European rabbit accounts for 80% to

90% of its diet (Gil-Sánchez et al., 2006). However, the Iberian lynx sometimes

also preys upon geese, ducks, rats, hares, partridges, and young red, roe and

fallow deer, as well as mouflons (Vargas, Beston, & House, 2009).

The minimum rabbit density necessary to sustain a viable lynx population

varies depending on the time of year (Palomares et al., 2001). Thus, in autumn

the minimum density required is 1 rabbit/ha, whereas at the end of the spring

(late lactation period and early supply of kittens with food) this figure raises up

to four or five rabbits per hectare. Due to the fact that, the lynx feeds almost

exclusively on rabbits, when the abundance of the lagomorph drops, the lynxes

increase their home ranges to compensate this lack of prey (Simón et al., 2012).

Likewise, when rabbit densities plummet below a crucial limit the lynxes are

forced to abandon the area. Moreover, Simón et al. (2012) stated that the lynx is

able to regulate the amount of secondary prey items depending on rabbit

abundance.

Although counterintuitive, the presence of the Iberian lynx favours the

European rabbit (San Miguel, 2015). In this regard, the lynx is considered a

super-predator and thus it puts away other generalist predators such as the red

fox (Vulpes vulpes) (Palomares & Caro, 1999). Thus, these other predators

8
 change their use of the space and abandon lynx territories. Consequently, the

pressure put on the European rabbit populations decreases.

2.1.2. Habitat requirements

The Iberian lynx is highly adapted to Mediterranean scrubland where it finds

refuge and prey (Palomares et al., 2000). This type of ecosystem is present in at

least 75% of the areas occupied by the lynx (Vargas, Beston, & House, 2009).

In these areas, the average scrub cover is 55%. The remaining areas are strips of

land located between the scrubland and open habitats such as grasslands, edges

or dehesas (Mediterranean savannah-like formations) or even olive (Olea

europaea) groves (Vargas, Beston, & House, 2009). The main habitat

requirement is the presence of 1 to 5 rabbits per hectare depending on the season

of the year (Palomares et al., 2001). Another essential factor for the survival of

the species is the presence of suitable habitats in which the lynx finds refuge

and sets its breeding dens (Fernández, Delibes, & Palomares, 2006; Palomares

et al., 2000). This is essential because the species tends to avoid open habitats.

Moreover, the presence of rocky outcrops (e.g. Limestone, marls, or plasters)

favours the presence of the European rabbit (Fernández, Delibes, & Palomares,

2006). Consequently, the species needs vast extensions with high abundance of

rabbits and with enough available refuge. This is due to the fact that the mean

home range of the felid varies between 250 and 2100 ha (Simón et al., 2012).

9
2.1.3. Reproduction

The Iberian lynx presents a monogamous mating system, although when the

carrying capacity of the ecosystem is reached, they tend to polygyny (Simón et

al., 2012). Thus, the current mean male:female ratio is 0.83 and 0.91, for the two

main subpopulations, Doñana-Aljarafe and Ándujar-Cardeña, respectively (for

the period 2010-2016, IberLince LIFE Project, 2016). The species is sexually

mature after its first year of life. However, the females do not start breeding until

the age of three and remain fertile until the age of nine years old (San Miguel,

2015). The females are monooestrous and the oestrous period takes place

between December and February (Simón et al., 2012). The pregnancy lasts for

63 to 65 days (Vargas, Beston, & House, 2009) and they give birth to 2 to 4 cubs

per litter (Simón et al., 2012).

The place chosen by the females to give birth are normally rocky formations

but when these lack they may do it elsewhere, as in Doñana-Aljarafe. Thus,

sometimes Iberian lynx females give birth in tree holes (i.e. Cork oak (Quercus

suber) or Ash tree (Fraxinus angustifolia)) or in between tall and dense bushes

(Fernández & Palomares, 2000). Moreover, after the lactation period, females

supply its kittens with food (Aldama, 1993), increasing their consumption rate

of rabbits. Hence, the female must capture two or three rabbits per day (Simón

et al., 2012).

10
 2.2. Study area: Doñaña-Aljarafe

The study site is located in the south-western part of Spain, Huelva province (37º 01´ N,

6º 26´W). This area encompasses 3500 km2 where 550 km2 belong to Doñana National Park

(Fig. 2). The area is characterized by sandy flat terrains at sea level. The climate is thermo-

Mediterranean dry, sub-humid (Rivas-Martínez, 2007). Thus, it has a marked seasonality

having mild and rainy winters and hot and dry summers. Mean annual rainfall is 500-600 mm.

The area comprises several types of ecosystems, mainly sand dunes, marshlands,

Mediterranean scrubland, and an ecotone that connects all the former ones. Its singularity is due

to the great diversity of biotypes and species that it harbours. The marshland is a highly

productive ecosystem and is of great importance for many African and European bird species.

The vegetation in the area is predominantly covered by Mediterranean scrubland where

Pistacia lentiscus bushes and Halimium halimifolium shrubs prevail (Rivas-Martínez, 2011).

There are also scattered trees of Quercus suber and Olea europaea. Riparian vegetation can

also be found along streams (e. g. Fraxinus angustifolia, and Tamarix africana; Castaño Corral,

Mateos Mateos, & Rivera Silva (2003)). Pinus pinea and Eucalyptus spp. plantations, as well

as, pastureland are also present in the area. Additionally, the area is subject to an intensive

agricultural activity.

This area represents one of the oldest extant subpopulations that still thrives in the

Iberian Peninsula. Currently, this subpopulation comprises 74 individuals and covers and

extension of 667 km2 (IberLince LIFE Project, 2016) (see Table 1 for further information on

population dynamics).

11
Figure 2. Map of the study site. A. Map of the Iberian Peninsula. B. Zoom into the study area of
Doñana and its surroundings. The green area represents the Doñana National Park and its area of
influence.

12
 Table 1. Demographic parameters of the Doñana-Aljarafe subpopulation for the period 2010-
2016. The number of individuals per year, the number of territorial females (reproductive females
and/or resident in a given area), the number of cubs, the area of presence (in km2), the sex-ratio
(males/females), and the density (lynxes/km2) are presented in the table (adapted from IberLince
LIFE Project, 2016).

Year 2010 2011 2012 2013 2014 2015 2016

Individuals 78 86 88 93 80 76 74
Females 17 19 25 25 23 26 24
Cubs 23 20 26 24 17 17 16
Presence 445 591 556 518 594 681 667
Sex-ratio 0.67 0.72 0.88 0.72 1.00 0.90 0.93
Density 0.18 0.15 0.16 0.18 0.14 0.11 0.11

2.3.Data collection

In this study, I examined data from 43 territorial females from the Doñana National Park

area. The data was collected during a 7-year period, from 2010 to 2016, by the technicians of

the Iberlince LIFE Project. Demographic data (litter size, production of offspring (at least one

cub), age, population density and sex-ratio) was obtained through capture and radio tracking,

track surveys, and camera trapping, following the methods described in Ferreras et al. (1997)

and Simón et al. (2012). Rabbit abundance in each of the territories occupied by the females

was estimated following Palomares (2001).

Response variables. The two response variables, Litter size and Reproductive success (females

with at least one cub), used in this study were obtained using two complementary

methodologies:

13
 Camera-trapping. This technique consists on setting up camera-traps on the areas of

known presence of lynxes. This methodology is very successful when studying secretive

species which are difficult to spot by an observer (O´Conell et al., 2011). Each individual can

be identified based on its recognizable fur pattern (Garrote et al., 2010). With this technique,

the Iberlince LIFE Project controls all the territories of known presence, as well as, those of

new formation. Thus, it serves to retrieve individual information on presence/absence,

reproduction success of the females, and litter size. However, it has certain limitations. In this

sense, the camera-trapping technique allows detecting a great number of territorial females but

it only detects the cubs after a certain age. Hence, those cubs that die in early stages of their life

are not considered when calculating the reproductive success and litter size of the females.

The cameras used are equipped with passive infrared sensor (PIR) (see Gil-Sánchez et al.

(2011)for further information about the cameras). The models used are Leafriver® and DCL

Cover®. One camera- trap is set in each trapping-station. Moreover, additional camera-trap

stations are located in lynx´ latrines and corridors that the animals often use. Every year a

trapping campaign is conducted. In Doñana-Aljarafe, the campaign is spread all year round,

containing five to seven sampling bouts. The sampling effort is one camera per km2,

approximately equivalent to four cameras per lynx territory. Thus, each year a maximum of 566

trapping stations are simultaneously functioning in each subpopulation.

Remote sensing and radio collaring. The radio collaring is a complementary technique to

camera-trapping. The radio collars used by the programme are GPS-GSM collars that permit

the location of the individuals both via satellite and via VHF emission. The models used are

Tellus-Televit and Microsensory Systems. The remote sensing programme tries to obtain three

to five radiolocations each week. The VHF emitters are geolocated by triangulation, taking as

valid cutting angles 60º and 120º. During the geolocation, the activity of the individuals is

considered. Hence, its triangulation is not retrieved until the animal is resting.

14
Predictor variables:

1) Rabbit abundance. This predictor variable was obtained through the count of cumulative

signs of the presence of European rabbit (burrows, scratchings and latrines) in fixed

transects. The calculation of the rabbit abundance is based on signs (relative densities) and

can detect population fluctuations. This sampling method estimates the distribution and

abundance of wild rabbits in large areas. The sampling of the transects was carried out in

meshes of 2.5 km X 2.5 km (IberLince LIFE Project, 2011). This mesh size was chosen

because is close to the average vital domain of the Iberian Lynx (Gil-Sánchez et al., 2011).

The surveys are conducted in those areas which have greater potential of rabbit presence.

Thus, four equidistant linear transects of 750 m are surveyed in each mesh. During the

survey, all the rabbit latrines within a width of 2 m from the transect line are counted. The

surveys were carried out from May to June, coinciding with the greatest rabbit densities.

All signs of rabbit presence were recorded in a field chart and georeferenced. Moreover,

these data were included in a Geographical Information System (GIS) and stored in two

different shp. files containing the number of signs recorded and the transects conducted.

Finally, the Kilometric Index of Abundance (KAI) is calculated as:

KAISquare= Totallat/Total km,

where KAISquare is the kilometric index of abundance of each mesh surveyed, Totallat is

the sum of all cumulative sign encountered during the survey, and Total km is the sum of

all transect lengths within that mesh square.

2) Age. The age of the individuals was estimated with camera-trapping. The technique is based

on the unique fur pattern that each individual has. Thus, every reproductive season all the

cubs are identified individually and therefore their date of birth is known.

15
3) Population sex-ratio. The sex-ratio of the study populations was calculated as the

proportion of males to females during the study period. In this calculation, the number of

adult males and females (sexually mature) was included. These data were obtained using

camera trapping.

4) Population density. The density of the metapopulation was calculated as the number of

individuals per km2 during the study period. The number of individuals present in the

subpopulation is obtained by camera-trapping as described above. To define the area of

presence of the lynx in the metapopulation, radio collaring was used.

2.4. Statistical analysis

First, all the variables within the dataset were checked for collinearity. Hence, only those

predictors with no or low correlation were included in the model (-0.30 < r < 0.30). Lynx density

and population sex-ratio were found to be highly correlated (r = -0.72). Thus, a set of two

models was created to decide which predictor to include in the subsequent statistical analysis.

The decision was taken based on the Akaike Information Criterion (AIC) weights. Finally, lynx

density (AIC= 216.2) was excluded from the analysis given the lower AIC of population sex-

ratio (AIC= 214.2).

Generalized Linear Mixed Models (GLMM) were used to test the influence of a series

of predictors on the response variables Reproductive success and Litter size. For both models

fixed effects were: rabbit abundance, age, and population sex-ratio. The only random effect

included in the models was individual nested in year. This random setting of the model allowed

controlling for differences in the reproductive performance of territorial females due to

population and individual traits. In the model for Litter size the distribution of the data was set

to “Poisson” family (count data), whereas for Reproductive success was set to a “Binomial”

distribution (binary data; female with or without offspring).

16
 The dataset was fitted to those statistical models using R 3.4.0 (www.r-project.org). A

model averaging approach was used to summarize all the possible models and to make stronger

interpretation because model-averaged parameters work in a more constant mode under low to

moderate levels of collinearity (Grueber et al., 2011). This approach implies the calculation of

a weighted average for each predictor present in a set of models. The weighting is given by the

Akaike Information Criterion weights. The AIC weights can also be used to calculate the

importance of each predictor, by summing them up in all the models in which the predictor

appears. Firstly, the maximal model, including all predictors was fitted. Subsequently, the

model comparison was performed using the AIC weights. For model comparison, the “dredge”

function within the “MuMIn” package of R was used. To finally obtain the importance of each

predictor (from 0 to 1), the “model.avg” function in the same package was used.

Accordingly, to represent the results from the GLMM of Reproductive success combined

graphs were used (histograms plus logistic curves). To obtain these graphs the “plot.logi.hist”

function from the “popbio” R package was used. In these graphs, the response variable

Reproductive success was plotted against each of the predictor variables (rabbit abundance,

age, and population sex-ratio).

17
 3. Results

Throughout the seven-year study period, the number of reproductive females varied among

years. The highest proportion of reproductive females occurred in 2010 when 48 % of the

females successfully reproduced with at least one cub (11 out of 21 territorial females). On the

other hand, during 2014 only 16 % of the territorial females reproduced (4 out 25 territorial

females; Fig. 3). The mean proportion of reproductive females during the study period was 35

% ± 11 (mean ± SD; all years pooled).

Proportion of reproductive females

0,50
0,45
0,40
0,35
0,30
0,25
0,20
0,15
0,10
0,05
0,00
2010 2011 2012 2013 2014 2015 2016
Year

Figure 3. Bar chart showing the proportion of reproductive females

(with at least one cub) between 2010 and 2016.

The mean litter size during the study period was 2.30 ± 0.91 (mean ± SD; all years pooled),

with range of 1-4 cubs per reproductive female. The litters consisted of one cub (N= 11), two

cubs (N= 23), three cubs (N= 16), and four cubs (N= 6), respectively. The minimum age of

reproduction was 2, whereas the maximum was 11 years old. However, the mean age of

reproduction for the study females was 6.32 ± 2.62 (all years pooled; mean ± SD), although,

the biggest proportion of females that reproduced occurred at the age of 11 years old (see

Appendix for further information on the age of reproduction).

18
 The mean rabbit abundance during the study period was 14.45 ± 12.57 (mean ± SD; all

years pooled) as expressed by the KAI. The maximum rabbit abundance occurred in 2012

(KAI= 21.74 ± 15.25), whereas the minimum occurred in 2016 (KAI= 10.51 ± 7.81). The sex-

ratio of the adult population varied from 0.44 ± 0.10 adult males/females (mean ± SD), in 2010,

to 0.77 ± 0.10, in 2015 (Fig.4). The mean sex-ratio during the study period was 0.67 ± 0.10

(mean ± SD; all years pooled).

Sex-ratio Doñana-Aljarafe
1,20

1,00

0,80

0,60

0,40

0,20

0,00
2010 2011 2012 2013 2014 2015 2016

Adults Population

Figure 4. Bar chart showing the variation in the sex-ratio between 2010 and
2016 for lynxes from Doñana-Aljarafe subpopulation.

The results from the model selection approach for the response variable reproductive

success showed that the best fitting model was the one that included all predictors (w= 0.301;

AIC= 206.7). The subsequent model averaging assigned to all predictors a relative importance,

based on the sum of model weights across all models containing each predictor. Thus, the

relative importance of rabbit abundance was 0.75, 0.77 for age, and 0.52 for population sex-

ratio (see Appendix for further information on the model averaging approach)

19
 The analysis also showed that age had a statistically significant effect on the probability of

reproduction of Iberian lynx females (p-value= 0.0453; see Table 2). Thus, individual female

age (2-11 years old) was positively correlated with reproductive success (Fig. 5).

Figure 5. Fitted regression line showing the probability of reproductive success

(females with at least one cub) against the predictor variable age. The histogram shows
the observed data and the line shows the predicted probability.

Moreover, the analysis also showed that the predictor variable rabbit abundance also had a

significant effect on the probability of reproduction of Iberian lynx females (p-value= 0.0468;

see Table 2). Therefore, with increasing rabbit abundances the probability of reproduction also

increases (Fig. 6). On the other hand, population sex-ratio was not found to be statistically

significant on the probability of reproduction (p-value= 0.1490; see Table 2; Fig. 7).

20
Figure 6. Fitted regression line showing the probability of reproductive success (females
with at least one cub) against the predictor variable rabbit abundance (KAI index). The
histogram shows the observed data and the line shows the predicted probability.

Figure 7. Fitted regression line showing the probability of reproductive success (females
with at least one cub) against the predictor variable population sex-ratio. The histogram
shows the observed data and the line shows the predicted probability.

21
 Table 2. Summary of the GLMM model averaging to analyse the factors affecting
reproductive success in Lynx pardinus territorial females. Fixed effects are shown in a decreasing
order of relative importance. Bold type highlights those variables that had a significant effect on
reproductive success (females with at least one cub).

Fixed R. Importance Avg. Std. Error z-value P-value

effect Estimate
Age 0.77 0.142 7.05 x10-2 2.002 0.045
Rabbit 0.75 0.028 1.38 x10-2 1.988 0.046
abundance
Sex-ratio 0.52 -2.557 1.76 1.443 0.149

The results from the model selection approach for the response variable litter size showed

that the best fitting model was the one containing only the predictor rabbit abundance (w=

0.268; AIC= 175.5). However, a model averaging approach was performed to assign a value of

relative importance to each predictor, based on the sum of model weights across all models

containing each predictor (see Appendix for further information on the model averaging

approach). Thus, the relative importance of rabbit abundance was 0.59, much greater than the

relative importance of female age (0.27) and sex-ratio (0.35). The analysis also showed that

none of the variables had a statistically significant effect on the number of cubs that each of the

females had (see Table 3). Despite of these results, there is a tendency that at higher rabbit

abundances the litter size increases accordingly (p-value= 0.106; marginally significant) (see

Fig.8).

22
Table 3. Summary of the GLMM model averaging to analyse the factors affecting litter size
in Lynx pardinus territorial females. Fixed effects are shown in a decreasing order of relative
importance.

Fixed R. Importance Avg. Std. Error z-value P-value

effect Estimate
Rabbit 0.59 0.011 6.41 x10-3 1.614 0.106
abundance
Sex-ratio 0.35 -0.722 8.07 x10-1 0.873 0.382

Age 0.27 0.006 3.42 x10-2 0.166 0.868

Figure 8. Boxplot showing the relationship between rabbit abundance and litter size.

23
 4. Discussion

The relationship between reproductive output and prey abundance is well documented in

mammals (Fuller & Sievert, 2001). Moreover, these authors stated that the population

recruitment is a function of the proportion of productive females, litter size and offspring

survival, and these demographic parameters should be higher when prey is abundant. This

statement is partially in agreement with the hypotheses presented in this study.

The results from the General Liner Mixed Models for the response variable reproductive

success showed that the probability of reproduction is affected by the abundance of prey (Fig.

6). In a food manipulation experiment with terrestrial vertebrates, Boutin (1990) found similar

results. In his experiments, Boutin (1990) showed that the proportion of reproductive females

increased with increasing prey abundances. Additionally, Fuller & Sievert (2001) in their

review found compiling evidences of prey abundance affecting reproduction in six species of

carnivores (i.e. Coyotes; Canis latrans; Canada lynx; Lynx canadiensis; Wolverine; Gulo gulo;

etc). Furthermore, Brand & Keith (1979) found a 33% decrease in pregnancy rates of female

Canada lynxes, during years of prey scarcity. However, López-Bao et al. (2010) in another food

manipulation experiment with the Iberian Lynx in Doñana National Park, found opposite

results. These authors found that the abundance of prey did not affect the proportion of

reproductive females (with at least one cub). They noticed a slight decreased in this proportion

for resident adult females. Moreover, in another study that involved the Iberian lynx, Palomares

et al. (2005) did not find any correlation between the proportion of breeding females and the

abundance of European Rabbit. Yet, these authors only studied 3 breeding females throughout

a nine-year study period. Thus, the sample size might have influenced the results of the study.

These contrasting results between the present study and the above mentioned, might be

caused by other factors. For example, the low genetic variability of the Iberian lynx in Doñana

(Rodríguez & Delibes, 2002), might be limiting the reproductive performance of the territorial

24
females. Despite the contrasting evidences, its seems reasonable and highly supported for other

vertebrates in general that the amount of food available for the females during the reproductive

season might influence their body condition and, therefore their reproductive success, as

occurred in other lynx species (e.g. Lynx rufus in “Nova Scotia”; Bonenfant et al. (2002)).

Another factor that was found to influence the reproductive success of Iberian lynx females

in Doñana was the age of individual females. Age was positively correlated with the probability

of successful reproduction, thus older females showed a higher probability of reproductive

success, producing at least one cub (Fig. 5). The positive correlation between age and the

probability of reproduction might be the result of a better body condition of older breeding

females. Thus, the hunting skills of older females are better, enabling them to obtain greater

body condition. Reproductive females were between 2 and 11 years old. The age of

reproduction of the studied females was similar to that in other lynx species (Kvam, 1991), and

to that in other Iberian lynx studies (Gaona, Ferreras, & Delibes, 1998). Reproductive

senescence was not observed in the studied subpopulation. These results are in agreement with

a study of captive and wild Eurasian lynxes (Lynx lynx) were senescence was observed at the

age of 12 years old or older (Henriksen et al., 2005). In the present study, the biggest proportion

of females that produced at least one cub occurred between 8 and 10 years old (Fig. 9 in the

Appendix). However, Ferreras et al. (1997) observed that females older than 9 years old could

not reproduce in Doñana National Park and were displaced by younger females. An increased

age of reproduction might be due to an increased carrying capacity of the breeding territories,

with respect to 1997. The increased carrying capacity was achieved by means of reintroduction

of European rabbits into lynx territories (Simón et al., 2012). Thus, senescence in the studied

population might occur at later stages of the Iberian lynx female life history (> 11 years old).

25
 Contrary to the expectations, none of the factors included in the statistical model

significantly influenced litter size. In the present study, the mean litter size was 2.30 ± 0.91.

This figure was similar to that of other lynx species; Henriksen et al. (2005), captive Eurasian

lynx and, Kvam (1991), wild Eurasian lynx. Palomares (2003) did not find any relationship

between cub survival and rabbit abundance during the denning period. However, Villafuerte

(1996) found that red fox litters were smaller after an outbreak of rabbit haemorrhagic disease,

thus litter size was dependent on the amount of prey. Additionally, Herfindal et al. (2005) in

their study of Eurasian lynxes found that prey density affected home-range size and this

indirectly affected female reproductive success. Despite of these contradictory results, the

relative importance of rabbit abundance was found to be much greater than the rest of the

variables (0.59). Thus, it is reasonable to assume that at higher rabbit abundances, females could

have bigger litters (Fig. 8).

On the other hand, there are several other aspects to be considered. First, the data used in

the present study as litter size corresponds to the period after the denning phase of the cubs.

Hence, neonatal mortality, due to scarcity of resources and/or non-natural mortality, is not

considered. The main reason for this inaccuracy in the data is the difficulty to spot the breeding

dens from Iberian lynx females. Moreover, the amount of data available for this study could

have influenced the results. In this sense, rabbit abundance can be considered marginally

significant (p-value= 0.106) and this might be the result of a small sample size. However,

Iberian lynx seems to have little plasticity to respond to changes in prey abundance as observed

in other lynx species (Brand & Keith, 1979).

Also, Solokov, Nidenko & Serventuk (1994) demonstrated that fights between cubs at early

stages of their life, greatly influence litter size of Eurasian lynx breeding females. This

phenomenon has also been reported to occur in the Iberian lynx (Antonevich et al., 2009), but

was not considered in the present study. Yet, this phenomenon has only been observed in

26
captivity thus, its occurrence in the wild is still controversial. Moreover, there might be other

factors that do not directly affect the litter size but the breeding female. Hence, body condition

prior to parturition could be an important factor affecting the fecundity of the breeding females

(Bonenfant et al., 2002). Likewise, it has been observed that the impoverished genetic diversity

of the subpopulation in Doñana, might affect life history parameters such as, litter size

(Rodríguez & Delibes, 2003).

5. Conclusions. Future perspectives for the Iberian Lynx

The present study stresses the existence of a threshold for abundance of prey that allows

Iberian lynx reproduction. On the other hand, more research should be conducted to clarify

which are the factors that modulate litter size in the Iberian lynx. Thus, long-term monitoring

of the breeding dens should be undertaken.

The Doñana subpopulation might be entering an extinction vortex (Palomares et al., 2012).

In this vortex demographic (reduced litter size, reduced reproductive rate) and genetic

(inbreeding and vulnerability to disease) factors might be interacting to extirpate the

population. It is therefore necessary, to keep on conducting basic and applied research to clarify

which factors affect Iberian lynx reproduction, as well as other aspects of its biology and

ecology. Hence, if the Iberian lynx is to survive active and evidence-based conservation must

be undertaken. Nevertheless, the focus should be on: (1) restoring the ecological conditions that

allow Iberian lynx settlement and reproduction (i.e. Adequate prey abundances); (2) tackling

down the threats associated with inbreeding, or higher susceptibility to infectious diseases of

wild populations (i.e. Increasing the ecological connectivity of lynx populations); (3) assessing

the response of free-ranging lynx to the conservation measures.

In this regard, a lot of effort should be put into creating the optimal conditions for the

European rabbit to settle in lynx territories. Habitat management (e.g. artificial building of

breeding warrens, increase pasture and shelter availability for rabbits, etc) should continue,

27
hence, improving the carrying capacity of lynx territories and allowing the dispersion of young

individual into new territories. Likewise, lynx habitat should be managed properly creating the

ideal conditions for the lynx to settle (i.e. increased presence of breeding dens and adequate

rabbit abundances). Moreover, translocations and reintroduction programmes should go on to

avoid the deleterious consequences of inbreeding. This is especially crucial in Doñana where

the heterozygosity of the population has plummeted. Thus, ecological connectivity of the

subpopulations should be improved, building eco-corridors that allow individuals to disperse.

Additionally, is it also of great importance to decrease non-natural mortality, by means of

awareness campaigns in lynx populated areas. Finally, to assess the impact of these measures

the IberLince LIFE Project should set up clear goals and monitor the efficacy of it conservation

and management practices regularly.

6. Acknowledgements

First, I would like to acknowledge the supervision I have received from Dr. Prof. Alfonso

San Miguel and from Dr. Ramón Perea from the Technical University of Madrid. Also, I would

like to acknowledge Miguel Angel Simón, director of the Iberlince LIFE Project, and to Dr.

Luis García, from the Technical University of Madrid for giving me the opportunity to conduct

the present study on such iconic species. In addition, I would like to acknowledge all the advices

and support that I have received from Dr. Aida López Gómez and MSc Marta Peláez. I would

also like to acknowledge the nice working environment in which I have been involved all this

months. This, has served to me to realize that despite all the difficulties, good research is being

conducted in Spain.

28
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8. Appendix

Proportion of reproductive females:

Figure 9. Proportion of reproductive females in each age class, with respect to the
total number of females in that age class in the study population.

Reproductive females: Doñana-Aljarafe

11
10
9
8
7
Age

6
5
4
3
2
0,00 0,10 0,20 0,30 0,40 0,50 0,60 0,70
Proportion

Reproductive females

Results from the model averaging approach for the two sets of models:

A. Reproductive success.

Table 4. Summary table of the GLMM model averaging approach for the response
variable reproductive success. All the possible models are presented and which predictor
variable do they contain. In bold the best fitting model based on its AIC value and Weight.

Model Intercept Age Sex- Rabbit df logLink AIC delta Weight

ratio
1 -0.600 - - - 3 -102.721 211.4 4.74 0.028
2 -1.395 0.134 - - 4 -100.707 209.4 2.71 0.077
3 1.028 - -2.449 - 4 -101.635 211.3 4.57 0.031
4 0.401 0.145 -2.790 - 5 -99.358 208.7 2.02 0.110
5 -1.017 - - 0.027 4 -100.516 209.0 2.33 0.094
6 -1.845 0.138 - 0.028 5 -98.430 206.9 0.16 0.278
7 0.496 - -2.245 0.026 5 -99.649 209.3 2.60 0.082
8 -0.163 0.147 -2.569 0.027 6 -97.350 206.7 0.00 0.301

33
 B. Litter size.

Table 5. Summary table of the GLMM model averaging approach for the response
variable litter size. All the possible models are presented and which predictor variable do they
contain. In bold the best fitting model based on its AIC value and Weight.

Model Intercept Age Sex- Rabbit df logLink AIC delta Weight

ratio
1 0.834 - - - 3 -85.033 176.1 0.54 0.204
2 0.823 0.002 - - 4 -85.032 178.1 2.54 0.075
3 1.213 - -0.584 - 4 -84.765 177.5 2.01 0.098
4 1.191 0.007 -0.617 - 5 -84.744 179.5 3.97 0.037
5 0.648 - - 0.010 4 -83.761 175.5 0.00 0.268
6 0.614 0.005 - 0.010 5 -83.749 177.5 1.98 0.100
7 1.143 - -0.788 0.011 5 -83.291 176.6 1.06 0.158
8 1.100 0.011 -0.835 0.011 6 -83.238 178.5 2.95 0.061

34