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Tesis doctoral
Carolina Soto Navarro
Sevilla, Diciembre 2012
Director de tesis
Dr. Francisco Palomares Fernndez
Profesor de Investigacin
Departamento de Biologa de la Conservacin
Estacin Biolgica de Doana-CSIC (EBD-CSIC)
Sevilla-Espaa
Tutor
Dra. M Jos LeivaMorales
Profesora Titular
Departamento de Biologa Vegetal y Ecologa
Universidad de Sevilla
Sevilla-Espaa
ndice
11
INTRODUCCIN........................................................................................
12
Contexto de la tesis.............................................................................
Marco terico......................................................................................
Teora de la seleccin de hbitat.............................................
13
El concepto de hbitat.............................................................
21
25
28
29
31
Caractersticas ecolgicas de las especies y mecanismos de
coexistencia.............................................................................
32
De cmo simplemente encontrar el hbitat ms idneo suele
no ser suficiente.......................................................................
36
39
41
21
21
41
rea de estudio........................................................................
46
65
93
135
157
185
CONCLUSIONES........................................................................................
209
AGRADECIMIENTOS...............................................................................
213
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Contexto de la tesis
El trmino biosfera fue acuado por el gelogo Eduard Suess en 1875, pero el concepto ecolgico
de biosfera procede de 1920 con Vladimir I. Vernadsky, precediendo a la introduccin en 1935 del trmino
ecosistema por Arthur Tansley. En esta tesis empleo la definicin de biosfera como ecosistema global.
En ecologa, la teora de estados estables alternativos predice que los ecosistemas pueden existir
en mltiples estados (conjunto de condiciones biticas y abiticas nicas). Estos estados alternativos no son
transitorios y por lo tanto se consideran como estables a escalas de tiempo ecolgicamente relevantes. Los
ecosistemas pueden sufrir una transicin de un estado estable a otro, en lo que se conoce como un cambio de
estado (a veces llamado un cambio de fase o cambio de rgimen), cuando son sometidos a perturbaciones. Uno
de los cambios de estado ms rpidos del planeta y el ms reciente, ha sido la transicin desde la ltima era
glacial al presente perodo interglacial (Scheffer et al. 2009, Lenton 2012) que se produjo a lo largo de milenios
(Hoek 2008).
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convertido en una parte integral de su entorno. Esta influencia tiene efecto a diferentes
escalas espaciales y niveles biolgicos, tales como la distribucin geogrfica de las
especies, la organizacin espacial de las poblaciones y el comportamiento individual
a escala fina. Por ejemplo, la fragmentacin de hbitats puede implicar la divisin
de las actuales poblaciones en subpoblaciones o metapoblaciones4 alterando as su
dinmica (por ejemplo, Banks et al. 2005). Este escenario puede restringir a muchas
Existe una amplia diversidad de definiciones para el trmino fragmentacin de hbitats. En esta tesis
lo definir como el proceso por el que un hbitat grande se transforma en pequeos parches del mismo hbitat
(vase la revisin de Fahrig 2003).
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especies a reservas naturales y reas adyacentes en gran parte del mundo (Woodroffe
& Ginsberg 1998). No obstante, las reas protegidas no estn exentas en muchos
casos de las alteraciones relacionadas con las actividades humanas, bien porque
actividades humanas de alteracin del medio han tenido lugar de forma comn o
bien porque sufren las consecuencias de la influencia de las reas que las circundan.
Adems, para muchos de los ambientes y ecosistemas no existe una informacin
precisa ni fiable sobre las complejas interrelaciones que regulan y determinan la
distribucin y abundancia de las especies, ni los efectos que diferentes factores
tienen sobre el individuo, las poblaciones o las comunidades.
junto con los cambios en la sociedad y las consecuencias de los niveles de explotacin
anteriores (Conover 2002), pero el reconocimiento de que la biodiversidad desempea
un papel esencial en el bienestar humano y en el equilibrio de los ecosistemas es
creciente, y ha llevado a la generacin de medidas urgentes para incrementar la
conservacin de especies y hbitats en todo el mundo, lo que tambin se traduce en
la cristalizacin de la era moderna de la ciencia y poltica conservacionistas. Desde
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que se acu el trmino en 1978 y se cre esta disciplina5, los estudios centrados
en la Biologa de la Conservacin han crecido exponencialmente hasta alcanzar la
relevancia actual que tiene esta rea de investigacin (Figura 1).
para las poblaciones e individuos, pero me referir a las especies en esta seccin
por conveniencia de lectura) surge de la interaccin entre eventos deterministas
y estocsticos (Corsi et al. 2000). De hecho, es el resultado de la interaccin
entre eventos biolgicos (por ejemplo, alimentacin, reproduccin o dispersin;
deterministas) y eventos impredecibles (por ejemplo, incendios, tormentas;
estocsticos). Durante estos eventos biolgicos, los animales pueden elegir las
reas que mejor satisfagan sus requerimientos ecolgicos (por ejemplo, reas con
abundantes recursos en pocas de escasez), es decir, pueden elegir hbitats a travs
de sus movimientos. En un contexto ecolgico, los requerimientos de las especies
se identifican por tanto en el marco de la seleccin de hbitat, un concepto que
desarrollar ms adelante.
El trmino Biologa de la Conservacin fue introducido como el ttulo de una conferencia que tuvo
lugar en la Universidad de California, San Diego en La Jolla, en 1978, organizado por los bilogos Bruce
Wilcox y Michael E. Soul. Con posterioridad, en 1987, se cre la primera sociedad cientfica profesional (The
Society for Conservation Biology); The Society is a response by professionals, mostly biological and social
scientists, managers and administrators to the biological diversity crisis that will reach a crescendo in the first
half of the twenty-first century. We assume that we are in time, and that by joining together with each other
and with other well-intentioned persons and groups, the worst biological disaster in the last 65 million years
can be averted. Although we have varying philosophies, we share a faith in ourselves, as a species and as
individuals, that we are equal to the challenge. For these reasons we join together in professional alliance, in
the service of each other, but also in the service of the less articulate members of our evolutionary tree (Soul
1987:4-5).
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Figura 1. Tendencia en el uso del trmino Biologa de la Conservacin en las publicaciones cientficas.
Los resultados proceden de una bsqueda en la ISI Web of Knowledge (http://isiknowledge.com/), con el
trmino conservation biology como topic keyword. La lnea vertical se fij en el ao 1978.
He utilizado el trmino relativamente para subrayar que hoy en da, debido a la alteracin humana
del paisaje, la expresin de hbitat idneo debe ser interpretada con cautela. En mi opinin, se refiere a lo
mejor dentro de lo malo ms que simplemente a una calidad de hbitat buena para la especie segn sus
requerimientos. En esta tesis emplear la idoneidad de hbitat en este sentido.
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empleado en el sentido de un grupo de especies que utilizan recursos similares y por lo tanto, pueden
competir.
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densidades de venados mula o bura (su principal especie presa), una mayor presin
de herbivora y en consecuencia un descenso en el reclutamiento de lamos de
rivera, un aumento en las tasas de erosin de los mrgenes de rivera y una reduccin
resultante de la abundancia de especies tanto acuticas como terrestres (Ripple &
Beschta 2006). Otro ejemplo relevante de su importancia es el proceso conocido
como liberacin de mesodepredadores (mesopradator release)9 (por ejemplo,
Palomares et al. 1998, Gehrt & Prange 2007). Es decir, los carnvoros presentan un
papel claro en el mantenimiento directo o indirecto de la biodiversidad, mediante el
control de mesodepredadores y diversificacin de las presas (Terborgh et al. 1999,
Miller et al. 2001). Estos estudios constituyen ejemplos de investigaciones holsticas
a nivel de ecosistema y estimulan enfoques similares en diferentes biomas, a fin de
comprender plenamente el papel de los mamferos carnvoros en el funcionamiento
de los ecosistemas y su representatividad como especies clave (keystone species)10.
Esto slo se puede lograr con investigaciones a largo plazo basadas en protocolos
estrictos de monitoreo de las especies y su medio ambiente (Yoccoz et al. 2001).
10
Las ideas relativas a la liberacin de mesodepredadores se remontan varias dcadas, cuando los
ecologistas comenzaron a observar que la eliminacin de depredadores originaba explosiones poblacionales
de otras especies inferiores (por ejemplo, Paine 1969, Pacala & Roughgarden 1984). El trmino fue acuado
por Soul et al. (1988) para describir un proceso mediante el cual las poblaciones de mamferos carnvoros
de tamao intermedio se hacan ms prevalentes en ausencia de un carnvoro superior, y las poblaciones de
diversas aves se vean deprimidas como consecuencia de ello. En esta tesis empleo el trmino de una manera
ms amplia segn Brashares et al. (2010) para definir la expansin en densidad o distribucin, o el cambio en
comportamiento de un predador de rango medio como resultado de la disminucin en densidad o distribucin
de un predador superior. Aunque la liberacin de mesodepredadores se emplea normalmente en el contexto de
la teora trfica de cascadas (por ejemplo, Berger et al. 2008, Brashares et al. 2010), se trata esencialmente de
una interaccin intragremial entre depredadores.
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Marco terico
Teora de la seleccin de hbitat
El concepto de hbitat
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E1
Niche
(used)
Environment
(available)
E3
E2
Figura 2. Representacin esquemtica del nicho ecolgico. Las flechas negras representan variables
ambientales (e.g. cobertura de matorral, disponibilidad de presas), y por tanto el espacio ecolgico.
La elipse gris oscuro se corresponde con los valores de esas variables que estn disponibles para
la especie (poblacin o individuos). La elipse gris claro representa el rango de valores usado por la
especie, es decir, su nicho ecolgico.
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a menudo con el concepto de hbitat. Fue desarrollado por primera vez por Grinnell
(1917) para referirse a todas las caractersticas del medio ambiente que le permiten a
una especie sobrevivir y reproducirse (ntese la similitud con la definicin de hbitat
mencionada, por ejemplo, Hall et al. 1997). Posteriormente, Elton (1927) introdujo
el papel funcional de la especie dentro de su comunidad, en su nueva definicin
del concepto. Estos autores estn detrs de las controversias pasadas y actuales.
Tenemos en cuenta el impacto de las especies en su medio ambiente y comunidad o
slo el efecto del ambiente en la especie, es decir, el efecto de factores limitantes11en
la especie? Esto tambin depende del contexto. En 1957, Hutchinson formaliz el
concepto de nicho con un modelo geomtrico. Defini el nicho como el hipervolumen
en el espacio multivariado de variables ambientales (el espacio ecolgico, Figura 3)
donde una especie puede persistir (Figura 2). Esta definicin hace hincapi en la gama
de condiciones ambientales necesarias para la persistencia de la especie, es decir, el
nicho Grineliano12 (que es similar al concepto de hbitat). En este contexto, el nicho
ecolgico representa la posicin de la especie en la gama de condiciones ambientales,
de manera que cada dimensin del nicho se corresponde por tanto con un subconjunto
de este rango potencial o realmente importante para la especie. Hutchinson no obstante
reconoci el papel potencial de la especie en su comunidad mediante la descripcin de
dos tipos de nicho: el nicho fundamental y el nicho realizado.
11
los factores limitantes son cualquier proceso [o factores] que afectan de una manera cuantificable
el crecimiento de una poblacin (Messier 1991), tal y como recursos trficos, refugios o condiciones climticas.
.
12
definido por Grinnell 1917.
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E1
(a)
(b)
E3
E2
Figura 3. (a) Espacio geogrfico y (b) ecolgico. La localizacin (normalmente definida por dos
coordenadas en el espacio; longitud y latitud) de una especie se emplea con frecuencia para analizar
sus propiedades ecolgicas en el espacio ecolgico de variables ambientales (E1 a E3). (Adaptado
de Calenge 2005).
Un biotopo es un rea fsica con condiciones ambientales uniformes dnde viven un conjunto
especfico de plantas y animales.
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ms atpicas (valores extremos del gradiente de una variable), mientras que las
especies no marginales usarn condiciones ambientales medias. La especializacin
es la anchura del nicho, es decir, el grado de tolerancia de la especie al gradiente
ambiental. Cuanto ms grande sea el nicho, mayor tolerancia presentar la especie,
mientras que cunto ms estrecho sea, ms especializada estar la especie en el
uso de ciertos recursos. Estos conceptos son particularmente tiles para describir y
cuantificar la relacin entre una especie y el medio ambiente disponible para ella.
En los ltimos aos, se han desarrollado numerosos anlisis para la estimacin del
nicho ecolgico (Guisan & Zimmermann 2000, Calenge & Basille 2008). Aunque
este concepto ha sido desarrollado y utilizado a nivel de especie o poblacin,
tambin puede ser generalizado a nivel individual.
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especies especialistas, para las que el hbitat en el rango geogrfico puede ser de
crucial importancia. Sin embargo, como la mayora de los procesos ecolgicos, la
seleccin de hbitat a menudo se produce a ms de una escala (Levin 1992).
Los diseos de tipo II, III y IV se utilizan para estudios a nivel individual
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Las escalas de seleccin definidas por Johnson (1980) ayudan a reducir esta
subjetividad, ya que tienen una base biolgica, pero no la eliminan por completo
(Erickson et al. 2001). Por ejemplo, en la escala de establecimiento del rea de
campeo en la distribucin geogrfica de una especie, los lmites del rea de estudio
a menudo abarcan el rea en la que se distribuye la poblacin.
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muy variada (generalistas de hbitat y dieta), mientras que otras tienen tolerancias
ambientales muy especficas y estrechas y slo consumen ciertos recursos
(especialistas de hbitat y dieta). Estas dos categoras de especies tienen diferentes
dinmicas poblacionales (Kolasa & Li 2003). Por ejemplo, la variacin en la
densidad de poblacin es mayor en los especialistas que en los generalistas (Kolasa
& Li 2003). Del mismo modo, los especialistas de hbitat utilizan unidades de
hbitat ms pequeas anidadas dentro de unidades de hbitat ms grandes (Kolasa
& Pickett 1989). Esto tiene otra consecuencia ya que las especies que utilizan
pequeas unidades de hbitat tienden a tener bajas densidades poblacionales como
consecuencia de la disminucin de la eficiencia en la bsqueda de los parches
adecuados y de la mortalidad durante la dispersin (Kolasa & Romanuk 2005).
As, la disponibilidad de hbitats adecuados parece afectar ms a las especies
especialistas que a las ms generalistas, las cuales utilizan una gama ms amplia
de tipos de hbitats para satisfacer sus necesidades (Munday et al. 1997, Bean et
al. 2002). Obviamente existe un gradiente continuo en el nivel de especializacin
de una especie entre el especialismo ms extremo (como el caso del lince ibrico;
Lynx pardinus) y el completo generalismo (como el caso del zorro comn; Vulpes
vulpes).
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de individuos puede ser un indicador pobre de la calidad del hbitat para algunas
especies.
con modelos de seleccin de hbitat puros hago referencia a modelos en los que no se tienen en
cuenta las potenciales interacciones competitivas existentes entre especies pertenecientes al mismo nivel trfico
que la especie bajo estudio, sino solamente las caractersticas anatmicas del paisaje que las rodea.
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17
entindase como recurso cualquier fuente de la cual la especie obtenga un beneficio, es decir tanto
las caractersticas del paisaje como la vegetacin o disponibilidad de presas, como el propio espacio en s
mismo.
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presa especfica) y con ello reduce la oportunidad de usar ese mismo recurso a
otra especie. La competencia por interferencia implica sin embargo interacciones
comportamentales entre las especies como la predacin intragremial (Polis et al.
1989) en el caso ms extremo o bien mecanismos ms sutiles como evitar los
hbitats ms usados por el predador principal y mostrar preferencias por hbitats
menos productivos (Harrison et al. 1989, Thurber et al. 1992, Durant 1998, Fedriani
et al. 1999, Fuller & Keith 1981), ajuste de los patrones de actividad para reducir
los encuentros con el predador principal (Litvaitis 1992, Johnson et al. 1996) o
bien formar grupos para competir de una manera ms exitosa por los recursos
y/o obtener ventajas antipredadoras (Kruuk 1975, Eaton 1979, Lamprecht 1981,
Gittleman 1989).
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por cuatro especies nativas (el lince ibrico Lynx Pardinus, el tejn europeo
Meles meles, el zorro comn Vulpes vulpes y el gato monts Felis silvestris) y dos
introducidas en tiempos histricos (la gineta comn Genetta genetta y el meloncillo
Herpestes ichneumon) (Figura 4) en un rea protegida del suroeste de Espaa; el
Parque Nacional de Doana.
El tejn europeo (7-8 Kg.) es un carnvoro social y territorial con una amplia
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Al igual que en el caso del meloncillo, la gineta (2 Kg.) tambin en una especie
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(Rosalino & Santos-Reis 2002, Galantinho & Mira 2009). Debido a que presentan
ciertos patrones de seleccin de hbitat definidos en funcin de la zona y cierta
especializacin local en la dieta, la gineta se considera entre el tpico generalista y
el tpico especialista de hbitat y dieta (Virgs, Llorente & Corts 1999). El tamao
medio del rea de campeo en Doana de la especie es de 541 hectreas (Palomares
& Delibes 1994).
El gato monts (3-7 Kg.) es un carnvoro del cual, desde un punto de vista
cientfico, se ha sabido bastante poco hasta aos muy recientes. Aunque puede
vivir en una gran variedad de hbitats, tradicionalmente se ha considerado como
una especie tpicamente forestal (Guggisberg 1975, Ragni 1978, Blanco 1998). No
obstante, en climas ms secos como la cuenca Mediterrnea la especie se encuentra
en paisajes constituidos por mosaicos de matorral y pastizales con abundantes cursos
de agua y presas adems de una alta cobertura de arbustos a escala de microhbitat
(Lozano et al. 2003). Se considera como un especialista facultativo de dieta; siendo
el conejo de monte (Oryctolagus cuniculus) la presa ms abundante en su dieta
cuando est presente, pero consumiendo una alta proporcin de roedores cuando los
conejos son escasos o ausentes (Moleon & Gil Snchez 2003, Lozano et al. 2006,
Sarmento 1996).
del planeta (UICN 2008); es endmico de la Pennsula Ibrica (Rodrguez & Delibes
1992, Ferreras et al. 2010) y en la actualidad existe nicamente en dos poblaciones
estables y reproductoras: Doana y Sierra Morena oriental (Ferreras et al. 2010).
El lince ibrico es un especialista trfico, estrictamente dependiente del conejo de
monte (Delibes et al. 2000). La densidad de conejos determina la densidad de linces
(Palomares 2001, Palomares et al. 2001). Adems, asociado entre otras cosas a su
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Figura 4. Especies de estudio en el Parque Nacional de Doana; (a) tejn europeo (Meles meles),
(b) zorro comn (Vulpes vulpes), (c) gato monts (Felis silvestris), (d) lince ibrico (Lynx pardinus),
(e) gineta comn (Genetta genetta) y (d) meloncillo (Herpestes ichneumon).
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rea de estudio
mayor con paisaje de matorral y bosque situado en el entorno de las marismas del
ro Guadalquivir en el suroeste de Espaa, entre las provincias de Huelva y Sevilla
(Figura 5). El inters despertado por la fauna de este enclave conlleva la creacin
de un rea protegida en 1964, as como un centro de investigacin, la Estacin
Biolgica de Doana. En 1969 el rea bajo proteccin aumenta hasta las 35.000
ha con la creacin del Parque Nacional de Doana. En 1982 el rea protegida ve
aumentada de nuevo su superficie con la creacin del Parque Natural de Doana en
el entorno del Parque Nacional (Garca-Novo & Martn-Cabrera 2005).
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47
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48
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Figura 7. Mosaico de ecosistemas del Parque Nacional de Doana. (a) monte blanco, (b) monte
negro, (c) pastizal en La Vera de Doana, (d) laguna temporal inundada en poca invernal, (e)
pastizal en zona inundable desecada durante la poca estival, (f) zonas de repoblacin de pinos
pioneros, (g) eucaliptar, (h) marisma inundada en poca invernal, (i) inicio del tren de dunas
mviles, (j) pinares en corrales entre dunas, (k) dunas mviles (cerro de Los nsares de Doana),
(l) vista de pinares de repoblacin desde el inicio del tren de dunas mviles, (m) vista general de
zona de sabinares y matorral mediterrneo (n) vista general de La Vera de Doana con pastizal y
alcornoques remanentes del bosque primigenio de Doana.
El tren de dunas mviles de Doana, que avanzan hasta 6 metros por ao,
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CHAPTER 1
66
Chapter 1
ABSTRACT
Track census is a widely used method for rapid faunal assessments, which
67
Chapter 1
RESUMEN
cuadrculas de 2 x 2 km2 localizadas dentro del rea de matorral del Parque Nacional
de Doana (situado en el suroeste de Espaa). Nuestros resultados mostraron
diferencias en el nmero de rastros detectados en los censos dependiendo del
observador, as como una interaccin significativa entre el observador y el perodo
del da en el que se realiz el censo. Adems, las variables que incrementaban la
calidad del sustrato (una mayor humedad ambiental y una menor velocidad del
viento, as como un nmero bajo de das transcurridos desde la ltima lluvia),
permitieron una mayor deteccin de rastros de carnvoros. No obstante, dependiendo
del tamao de la especie, otras variables como la distancia del transecto al borde
de la vegetacin tambin afectaron los resultados. Recomendamos restringir los
censos a ciertas condiciones climticas determinadas cuando se planteen realizar
68
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69
Chapter 1
INTRODUCTION
Few methods are suitable for monitoring elusive, low-density species (Mills
The broad application of natural sign surveys such as track counts has firmly
established their use as a tool for wildlife detection. Track surveys do not rely upon
special technology or equipment, can be relatively straightforward and quick to
conduct, and can easily incorporate multispecies and large geographic area objectives.
Moreover, track counts do not require a behavioural response to attractants or other
survey equipment, thus there are potentially fewer species-specific limitations
and biases inherent to track surveys (Long et al. 2008). Nevertheless, field-based
species identification may be ambiguous or unfeasible so additional efforts and
highly skilled and experienced trackers are needed to validate the identification of
species or individuals. This weakness related to species identification combined
with the limited availability of appropriate tracking mediums or conditions, the
70
Chapter 1
obtained by spotlight counts, and population size have been previously reported
(e.g. Newsome et al. 1989; Short et al. 1997; Garel 2010), it is important to highlight
that track censuses can only be taken as indices of presence, relative abundance
or density estimators (Anderson 2001) and that such indices are rated closely to
true animal abundance across habitat types, observers, and other factors (see Gibbs
200). Herein, the number of tracks of certain species encountered on a transect will
depend on biological factors, such as their abundance, food density and distribution,
vegetation structure and intra- or interspecific interference, including humans (e.g.
Odonoghue et al. 1997; Shapira et al. 2008; Bayne et al. 2008; Blaum et al. 2009)
but there are other classes of variables that affect the index (Buckland et al. 1993).
These variables are related to the observer, including the observers training and
experience, eyesight and fatigue level, the environment (i.e. climatic conditions and
local habitats) and aspects of the species itself, such as their body size (Anderson
2001; Mackenzie and Kendall 2002). Among the variables associated with the
environment, such as wind speed, temperature, humidity, cloud cover, time of
sunrise or days from the last rain or snow, many have been previously suggested as
potential influences on the results of track sampling (Norton 1990; Hayward et al.
2002; Long et al. 2008). These non-biological factors constitute an important source
of error, as they affect the probability of detection and therefore the count. If they are
not considered when designing a monitoring program they can increase variance or
uncertainty for the estimates of relative abundance indices (Thompson et al. 1989).
71
Chapter 1
Despite the potential influence that the above-mentioned factors may have
on track counts, specific studies on the subject are scarce (Jennelle et al. 2002;
Karanth et al. 2003, but see Stander 1998; Balme et al. 2009; Zielinski and Schlexer
2009). Nevertheless, there is a growing suggestion to include measures of precision
and estimates of the detection probability when using indices values (usually raw
counts) purporting to measure relative abundance (e.g. Anderson 2001; Rosenstock
et al. 2002; Engeman 2003).
Here, we studied how methodological and climatic factors affected the number
72
Chapter 1
METHODS
Study area
The study was carried out in Doana National Park in southwestern Spain
(379N, 626W). This is a 550 km2 flat sandy area at sea level bordered to the south
and west by the Atlantic Ocean and to the east by the Guadalquivir River mouth.
The climate is Mediterranean subhumid (i.e. characterised by mild wet winters
and hot dry summers), with an average annual rainfall of 500-600 mm. There are
three main environmental units in the park: marshland, dunes and Mediterranean
scrubland (Fig. 1). Track censuses were restricted to the scrubland area, which
is mainly characterised by heterogeneous patches of xerophytic species such as
Halimium sp. and Cistus sp., and hygrophytic ones such as Erica sp., with some
patches of Juniperus phoenicea and Pistacia lentiscus shrubs. Interspersed with the
scrubland there are scattered cork oak trees (Quercus suber) and wild olive trees
(Olea europea), and a few patches of pine Pinus pinea and eucalyptus Eucalyptus
sp. plantations. The Mediterranean scrubland represents approximately half of the
National Park surface area.
Carnivore species in our study area are the red fox (Vulpes vulpes), the
Eurasian badger (Meles meles), the Egyptian mongoose (Herpestes ichneumon), the
common genet (Genetta genetta), the polecat (Mustela putorius) the Iberian lynx
(Lynx pardinus), the European otter (Lutra lutra), wild and domestic cat (Felis sp.),
and domestic dog (Canis familiaris). Polecats and otters were excluded from our
study because of their low abundance.
carnivore community and sampled in our study were small mammals (i.e. Garden
dormouse (Eliomys quercinus), Southern Water Vole (Arvicola sapidus), bush rat
73
Chapter 1
(Rattus rattus), Long-tailed Field Mouse (Apodemus sylvaticus) and other mice
(Mus spp.), but the most common are A. sylvaticus (Kufner and Moreno 1989)),
European rabbits (Oryctolagus cuniculus), red partridges (Alectoris rufa), domestic
cows (Bos Taurus) and horses (Equus caballus) and wild ungulates such as the
fallow deer (Dama dama), the red deer (Cervus elaphus) and the wild boar (Sus
scrofa).
Track sampling
factors affecting wild and domestic carnivore abundance and distribution within
Doana National Park, during the wet seasons of 2007-08 and 2008-09 (from
Figure 1. Map of the study area showing Doana National Park in the southwestern
Spain and the 2x2 km2, where carnivore and prey track censuses were carried out
74
Chapter 1
November 2007 to May 2008 and from October 2008 to April 2009) three and two
observers sampled 59 and 57, respectively, 2 x 2 km squares all with at least 40% of
their surface located within the scrubland area of the park (Fig.1). Marshland area
was not sampled as its clay soils make it unsuitable for track censuses. The squares
with 40% area of open dune were excluded for the present study since vegetation
is clearly different from the rest of the scrubland area, which would add an extra
source of variability to data. The three observers of the first year were a fieldworker
with 15 years of experience and two without previous experience, but that were
trained for two months by the experienced fieldworker. During the second year, the
experienced observer and one of the others carried out the surveys.
(ca. 1.5 km/h) in at least 3 km-long sandy paths (in car tracks or firebreaks). Once
a continuous track, that crossed side to side across the pathway, was detected, we
georeferenced it using a GPS. We noted location as a grid reference, date, and the
methodological variables, censusing day time (we established three block schedules;
early morning (from 8 a.m. to 12 a.m.), afternoon (from 12 a.m. to 3 p.m.) and
evening (from 5 p.m. to sunset), start time and end time for each census, and the
observer who carried out the census. In order to homogenize the number of tracks
detected per grid and maximize the probability of detection for each carnivore
species, we re-sampled the same path (leaving at least 7 days between samplings) a
second time in a few squares until completing 3 km if during the first sampling there
was not enough available path within the square to achieve this distance. Thus, we
had more censuses than total number of 2x2 km squares. We always carried out
surveys at least 3 days after any rainfall.
75
Chapter 1
variations in abundance for some species (e.g. see Kufner 1986; Palomares et al.
2001 for small mammals and European rabbits, respectively). Thus, we carried out
the sampling of prey tracks in April 2009 along transects in every 2x2 km square
sampled for carnivore tracks. These transects for prey species were walked as they
were for carnivore tracks, were 25 m in length and approximately 1.7 m wide (i.e.
the area of a four-wheel-drive car) and were located in the middle of the census path
and separated by at least 300 m. We recorded the location as a grid reference, date,
observer, and the following methodological variables: distance from nearest border
of census transect to the closest vegetation border (not recorded for carnivore track
censuses as they were carried out by zigzag walking), pathway where transects
were established (firebreaks or car tracks) and quality of the substrate for detecting
tracks based on the presence of grass (we established two categories; good (when
grassy groundcover was less than 10% in any part of the 25 m transect) and fair
(when grassy groundcover in any part of the 25 m transect was between 11-30%).
We considered unsuitable for prey counts transects in which grassy groundcover
was more than 30% in some part of the transect.
(calculated as the average of the maximum temperature measured on the census day
and the maximum temperature measured two consecutive days before the census
day), relative humidity, maximum wind speed, and the number of days since last
rain. The data was obtained from a station located inside Doana National Park
(Control RM1 Meteorological Station; Latitude: 37 118, Longitude: 6 33 17
http://icts.ebd.csic.es).
76
Chapter 1
Data analyses
response for each landscape variable before fitting them into the final equations (Austin,
2002). With this aim we fit Generalised Additive Models (GAMs) (Hastie and Tibshirani,
1990) using carnivore Kilometric Abundance Indexes and the number of prey tracks as
response variables and fitting smoothing splines with 3 degrees of freedom to model
every climatic and methodological continuous effect. The smoothed variables were then
turned into suitable parametric terms guided by visual inspection of the partial residual
plots (Crawley, 2005). The postulated models were then fit to the track-census dataset
using general linear models (GLM) with the log link, negative binomial error structure
and linear and non-linear responses to fixed effects in accordance with the GAM results.
We analyzed the effect of methodological variables observer (observer) and censusing
day time (day_time), and climatic variables maximum temperature (max_temp), average
relative humidity (humidity), maximum wind speed on census day (wind_speed), and
days since last rain (last_rain) on the carnivore Kilometric Abundance Index (KAI)).
We also included in the models the interactions between observer and day_time, as the
number of samplings carried out by each observer in each daily time period was different.
Correlations between predictors were always low (r < 0.6) so we fitted full models (i.e.
models including all the methodological and climatic variables). As for some 2x2 km
squares we carried out more than one census, our sampling unit was the census and not
the square. We examined the effect of the above variables on total carnivore abundance
index, small carnivore (from 1-5 kg of body mass) abundance index (including the
common genet, wild and domestic cats and the Egyptian mongoose) and medium-sized
carnivore (>7 kg of body mass) abundance index (including the red fox, the Eurasian
badger, the Iberian lynx and the domestic dog).
77
Chapter 1
variables contributing least to the model (i.e. variables with P > 0.3) before models
were refitted. Only variables with P 0.05 were interpreted as statistically significant.
Overdispersion was not a problem ( was close to 1 (1.14 - 1.21)) for any of the models
(Zuur at al. 2009). We analyzed data separately for each study year as the number of
observers changed and to maximize the variability between conditions, which could
affect the number of tracks detected on sand substrates.
We also performed general linear models (GLM) with negative binomial errors
and log link function to analyse the effect of observer, distance from border of census
transects to the closest vegetation border (dist_veg.), type of path (place) where prey
censuses were carried out (firebreaks or car tracks), quality of the substrate (quality) and
climatic variables last_rain, max_temp, humidity and wind_speed on track counts data
of prey species. We also included in the models the interactions between observer and
quality and observer and place. Prey data were grouped as total prey, small prey (small
mammals), medium-sized prey (rabbits and partridges) and large prey (cows, horses and
ungulates). Correlation between predictors was low (r < 0.5), so we fitted full models.
Overdispersion was not a problem ( = 1.04 - 1.24). The sample unit to adjust GLM was
the 25 m transects.
regression-model selection procedure excluding variables with P > 0.3, and then refitted
the models.
All statistical analyses were performed using the SAS 9.2 statistical software
(SAS Inst. Inc., Cary, NC), GAM and GLM were fitted using the gam and genmod
procedures, respectively.
78
Chapter 1
RESULTS
A total of 471 km were walked and 8,373 carnivore tracks were found during
surveys, with the red fox, the Eurasian badger and the Egyptian mongoose being
the most recorded species (Table 1). For prey, 5,000 tracks were detected on 11,575
m sampled (Table 2). Prey species more often detected were ungulates (i.e. fallow
deer, red deer, wild boar) and rabbits.
For the first year, there were differences in the number of tracks detected
among observers for all small and medium carnivores, and the number of tracks
decreased when wind speed increased for total carnivores, increased when humidity
was higher for small carnivores, and was highest during the afternoon for mediumsized carnivores (Table 3, Fig. 2). A significant interaction was also detected between
the censusing day period and the observer, with the three observers finding more
medium-sized carnivore tracks in the evening than in the morning or afternoon
(Table 3, Fig. 2d).
Table 1. Carnivore kilometric abundance index (tracks / km; KAI) in the scrubland area of
Doana National Park during the wet seasons of 2007-2008 and 2008-2009.
Species
Positive
censuses
2007-2008
Total
number of
tracks
KAI
(meanSD)
Range
Positive
censuses
2008-2009
Total
KAI
number
(meanSD)
of tracks
Range
Lynx
pardinus
19
65
0.4 0.9
0 - 5.4
14
75
0.4 0.9
0 - 5.0
Meles meles
68
599
3.9 5.1
0 - 20.6
53
666
3.7 3.8
0 - 17.3
60
631
4.0 4.4
0 - 21.1
48
544
2.8 3.3
0 - 12.5
76
3,138
17.7 9.1
2.4 - 44.2
61
2,333
11.9 6.5
1.2 - 32.6
21
160
0.8 2.3
0 - 15.9
20
66
0.4 0.7
0 - 2.8
Felis sp.
25
0.2 0.8
0 - 5.7
17
27
0.1 0.3
0 - 1.3
Canis
familiaris
12
23
0.2 0.7
0 - 5.9
11
21
0.1 0.3
0 - 2.0
Herpestes
ichneumon
Vulpes
vulpes
Genetta
genetta
Chapter 1
During the second year, more tracks were detected when humidity increased for total
and medium-sized carnivores, and fewer small carnivore tracks were recorded when
daily maximum temperature and days since last rain increased (Table 3, Fig. 3).
Most of the variables considered affected the total number of prey tracks
detected (Table 4). Thus, the number of tracks for total prey increased when the
daily maximum temperature, humidity and days since last rain increased (Fig.
4c), and decreased when wind speed and distance to vegetation edge was highest.
Furthermore, the number of total tracks was higher when samplings were carried
out in car tracks than in firebreaks (Fig. 4b), one observer detected more tracks
than the other (Fig. 4a), and there was a significant interaction between observer
and place (Fig. 4d). Some interesting exceptions to this general pattern were found
when data were separated by type of prey (Table 4). For small prey, number of
tracks was not affected by wind speed, humidity, maximum daily temperature and
days since last rain, and the interaction between observer and place was not found.
For medium-sized prey the number of tracks found was not affected by wind speed,
daily maximum temperature and days from the last rain, but in this case quality of
the road did affect results, with a higher number of tracks being detected at transects
without grass. Finally, the number of large prey tracks was not affected by daily
maximum temperature, distance to vegetation, type of path, or observer, and there
was no interaction between observer and place.
DISCUSSION
Chapter 1
sound monitoring programmes. Indexes derived from track surveys are partially a
function of animal abundance, but are also a function of a long list of methodological
and climatic variables and characteristics of the species being surveyed. Our results
support the hypothesis that non-biological factors can affect the number of animal
tracks detected in surveys, and must be taken into account when planning to study
animal abundance, distribution or the biological factors determining them.
The aim of this study is not to establish a standard protocol to carry out
No of tracks
233
2,132
260
210
2,165
Mean SD
0.5 1.4
4.6 9.4
0.6 1.3
0.4 1.6
4.7 9.4
Range
0 - 13
0 - 115
0 - 11
0 - 18
0 - 33
81
Chapter 1
America or South Africa where track surveys occurring in dust, mud or sand are more
broadly employed, many authors have also tried to standardize survey design and
Table 3. Results of GLM analysis to test for the effect of several methodological and climatic
variables on abundance indeces of total, small and medium-sized carnivores in Doana
National Park. Standard errors have been omitted to simplify the table. Variables with P > 0.3
excluded from the models are represented as (-). Least squares means (LS-Means) of the categorical
fixed effects observer and day time are shown. Non-est. means that the model could not calculate the
parameter because of little data. (*P < 0.05; **P < 0.01; ***P < 0.001; ns not significant).
Effects
Intercept
wind_speed
humidity
2007-2008
2008-2009
Small
carnivores
Mediumsized
carnivores
Total
carnivores
Small
carnivores
Medium-sized
carnivores
Total
carnivores
3.7238 ***
0.5065
3.2957***
2.4779**
2.0154
1.4906
-0.1329*
-0.1316
0.1071
-0.1751
-0.0711
0.0269**
0.0194**
0.0261**
-0.0376
-0.0937**
-0.0095
-0.0160
-0.048**
-0.0023
early morning
24.5214
15.1443**
16.5645
15.1596
afternoon
28.3601
22.8087**
20.3801
16.6357
evening
observer
35.5493
27.8652**
29.2495
23.1351
20.7509***
3.4544***
16.1252**
3.5478
20.0273
27.8207***
2.4517***
25.1415**
2.4450
16.1825
38.8229***
11.5468***
28.3257**
ns
***
max_temp
last_rain
day_time
observer*day_time
82
Chapter 1
There was some exception to the general rule stated. A positive correlation
was found between number of tracks recorded and days since last rain for large prey
species. Their large size rendered their tracks easily recognizable in poor substrate
conditions.
Table 4. Results of GLMM analysis to test for the effect of several methodological and climatic
variables on abundance indeces of total, small and medium-sized and large prey in Doana
National Park. Standard errors have been omitted to simplify the table. Variables with P > 0.3
excluded from the models are represented as (-). Least squares means (LS-Means) of the categorical
fixed effects place, quality and observer are shown. Non-est. means that the model could not
calculate the parameter because of little data. (*P < 0.05; **P < 0.01; ***P < 0.001).
Effects
Total prey
Small prey
Medium-sized prey
Large prey
Intercept
2.4520***
-2.7311***
0.7317*
2.4978***
wind_speed
-0.0005**
-0.0004
-0.0006**
humidity
0.0001***
0.0001**
0.0001**
max_temp
0.0002**
0.0002
last_rain
0.0439***
0.0622***
distance_veg
-0.0666***
-0.1018*
-0.1234***
2.4818**
2.8317**
-1.0304**
-2.3403**
1.2910*
1.7615*
place
firebreak
car track
quality
without grass
with grass
observer
1
2
2.5409
2.7726
-1.3780
-1.9927
1.8563**
1.1962**
1.5986***
2.3358***
3.1145***
2.1989***
-1.2665**
-2.1042**
2.3213***
0.7312***
observer*place
***
***
Non-est.
Non-est.
Non-est.
Non-est.
observer*quality
83
Chapter 1
of the species sampled. A higher number of prey species tracks were detected when
transects ran near a vegetation border or in car tracks. This result could be caused
by two different reasons. Vegetation must exert a protective effect against wind and
maintain a higher level of moisture on the sand, thereby increasing substrate quality
for detecting tracks. Also, medium and small prey may prefer to remain near a
vegetation edge to decrease predation risk (Hughes and Ward 1993, but see Moreno
et al. 1996).
Figure 2. Effects of wind speed on total carnivore tracks per km detected (a), of observer (least
square means and their standard errors are represented) on total carnivore tracks per km (b), of
relative humidity on small carnivore tracks per km (c), and differences in the number of mediumsized carnivore tracks per km detected by observers in each period of the day given as estimated
least square means and their standard errors (d) during the first study year. F values computed as
MSModel / MSError (i.e. Mean SquareModel/Mean SquareError), and their respective p-values are shown.
84
Chapter 1
among size groups or between years. This could be due to the fact that many of the
variables that affect detectability, and therefore the count, exhibit time trends (i.e.
vary between years) further confounding the value and interpretation of the index.
This is an important issue as it makes track censuses incomparable across different
climatic and methodological conditions.
Figure 3. Effects of maximum temperature on small carnivore tracks per km (a), and of days
since last rain on small carnivore tracks per km detected (b) for the second study year. F
values computed as MSModel / MSError, and their respective p-values are shown.
85
Chapter 1
depending on skill level of observers (Bider 1968; Smallwood and Fitzhugh 1995;
Anderson 2001; Wilson and Delahay 2001; Silveira et al. 2003). For this reason,
suggestions have been recently proposed to decrease differences among observers
or to evaluate observer skills (Sadlier et al. 2004; Evans 2006, 2009; Zielinski and
Schlexer 2009).
Figure 4. Effects of observer (least square means and their standard errors are represented) on
total prey tracks per 25 m transects (a), of censusing place on total prey tracks per km (least square
means and their standard errors are represented) (b), of days since last rain on number of large prey
tracks (c), and the interaction between observer and type of pathway sampled given as estimated
least square means and their standard errors (d). F values computed as MSModel / MSError, and their
respective p-values are shown.
86
Chapter 1
day, days from the last rain or snow, observers involved in the sampling, or distance
to vegetation borders will also help to diminish variability in the number of tracks
recorded. Some of these suggestions have been approached by maintaining constant
substrate quality through the use of artificial substrates and by erasing and resampling
newly left tracks on transects for a given fixed number of days (Gruber at al. 2008;
Watts et al. 2008; Russell et al. 2009; Zielinski and Schlexer 2009).
These results could be applied to a variety of research fields, both for testing
validity of pre-existing data and improving the suitability and performance of future
studies based on track surveys. Although the indices derived from track censuses
are only partially a function of animal abundance (Anderson 2001), if the variables
associated with the observer, the environment, and characteristics of the species
being surveyed are controlled, the reliability of the information extracted from these
methods may be improved.
87
Chapter 1
ACKNOWLEDGMENTS
of Education and Science) and 17/2005 (Spanish Ministry of the Environment; National Parks
Research Programme). Land-Rover Espaa S.A. lent two vehicles for this work. We are very grateful
especially to J.C. Rivilla for their assistance during fieldwork. C. Soto was also supported by a JAEPredoc grant from the CSIC.
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Zielinski WJ, Kucera TE, Barrett RH (1995) American marten, fisher, lynx, and wolverine: survey
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methods for their detection. USDA forest service, Pacific Southwest Research Station
Zielinski WJ, Schlexer FV (2009) Inter-Observer Variation in Identifying Mammals from Their
92
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93
94
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ABSTRACT
Chapter 2
segregation along the ecological niche dimensions have been shown as key
mechanisms for the most specialist species (genets and lynxes) allowing coexistence
whereas for the most generalist species such as mongooses, foxes and badgers,
another mechanisms such as spatial-temporal segregation of activity patterns might
help to explain coexistence among them.
96
Chapter 2
RESUMEN
Chapter 2
98
Chapter 2
INTRODUCTION
1960) states that two ecologically similar species cannot coexist unless they differ
sufficiently in niche separation, that is, in the way they use resources. Thus, some
degree of partitioning has to occur in the realized niche of coexisting species which
can occur at the temporal, trophic and/or habitat selection level. This is particularly
obvious in heterogeneous landscapes where spatial heterogeneity foments
coexistence between similar species (i.e., within the same trophic level), selection
of different habitats being one of the main processes that promotes sympatry (Levin
1974, Rosenzweig 1984). Nevertheless, mechanisms that facilitate coexistence
between similar species in apparently homogeneous landscapes where a general
habitat type dominates have been poorly explored.
apparently homogeneous landscapes (i.e., where the spatial variation in the species
biotic or abiotic environment is low) may help to understand the way species
exploit resources. Hence, besides fine-scale habitat use segregation, coexistence is
therefore possible if species exhibit differences in their life history traits that allow
niche differences in space (Brown and Wilson 1956, Hutchinson 1959, Chesson
2000). In other words, cohabitation may be privileged when species with different
degrees of habitat and/or trophic specialisation are present.
99
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Chapter 2
generalisation (i.e., the red fox). The Eurasian badger and the red fox are habitat and
thropic generalist species (Carvalho and Gomes 2001, Cavallini and Lovari 1991,
Kruuk and Parish 1985, Roper and Lups 1995, Balestrieri et al. 2004, Rosalino et
al. 2005, Amores 1975). The Egyptian mongoose is a habitat specialist but trophic
generalist (Palomares and Delibes 1991, Palomares and Delibes 1993, Zapata et al.
2007,) whereas the Iberian lynx is a highly habitat and trophic specialist species
(Palomares et al. 1991, Delibes et al. 2000, Palomares et al. 2001). The genet
meanwhile is considered between typical generalists and typical specialists (Virgs
et al. 1999).
multiple variables due to their higher degree of habitat and trophic specialisation
whereas badgers and foxes should have fewer requirements and respond to single
explanatory factors such as prey availability or vegetation and landscape structure
because of their omnivorous diets and habitat generalism. For mongooses, as
habitat specialists but trophic generalists vegetation and landscape structure should
explain their pattern of habitat use. Therefore, genets and lynxes should show the
most marginal (i.e., located in extreme values of variable gradients) and narrowest
realized niches whereas foxes and badgers would have the most widespread and
widest niches among those possible. Mongooses meanwhile may show a nonmarginal niche, wider than lynxes and genets but narrower than foxes and badgers.
101
Chapter 2
Figure 1. The study area. Doana National Park and its location in south-western Spain.
102
Chapter 2
METHODS
Study area
This study was located in Doana National Pak (DNP), a fully protected
area in southwestern Spain (550 km2 379N, 626W) (Fig.1). DNP is a flat sandy
area at sea level bordered to the south and west by the Atlantic Ocean and to the
east by the Guadalquivir River mouth. The climate is Mediterranean subhumid (i.e.,
characterized by mild wet winters and hot dry summers), with an average annual
rainfall of approximately 550 mm. Approximately half of the surface of DNP is
covered by Mediterranean scrubland, and the other half by marshland. There is
a dune system in the southern part of the scrubland area. Track censuses were
restricted to the scrubland biotope.
Human access into the park is regulated, but some low impact traditional
uses are maintained under control including cattle-raising, apiculture, and fishing
with traditional methods. The northern and western edges of the protected area are
103
Chapter 2
in close contact with human settlements, crop fields and a highly intensive use paved
road. There are two villages close to the edge of the park limits. Populations in the
main suburban settlement (situated in the western vicinity)vary greatlybetween
winterand summer, as this area ismainlya summer resort occupied by about
250.000 people during the summer seasons. The other village situated in the north
is occupied by about 1,635year-round residents, although on holydays the numbers
of people increase considerably and duringa spring pilgrimage may even reach up
to one million people. There are also private large and medium-sized farms used for
agriculture as well as six visitor centers, hiking and cycling paths, recreation zones
and bird observatories in the nearby area.
Sampling protocol
coordinates. Quadrants were sampled throughout the wet seasons, from November
2007 to May 2008 and from October 2008 to April 2009. In each quadrant, a 3 kmlength survey route was slowly walked searching for carnivore and prey tracks. The
survey routes were located along firebreaks and car roads between 2 and 12 m wide.
Once a continuous track that crossed from one side to the other across the pathway
was detected, we georeferenced it using a global positioning system. We resampled
21 squares over the two sampling periods a second time (leaving at least 7 days
between samplings) until completing 3 km, if during the first sampling there were
insufficient available paths within the square to achieve this distance. Thus, as we
had more censuses than quadrants we averaged track counts to get a single value per
quadrant. We always carried out surveys at least 3 days after any rainfall.
104
Potential target prey species of the carnivores studied were also sampled
Chapter 2
by track censuses. Preys sampled were small mammals (mostly long-tailed field
mouse (Apodemus sylvaticus) according to Kufner and Moreno 1989), European
rabbits (Oryctolagus cuniculus), red partridges (Alectoris rufa), domestic cows
(Bos Taurus) and horses (Equus caballus) and wild ungulates such as the fallow
deer (Dama dama), the red deer (Cervus elaphus) and the wild boar (Sus scrofa).
Wild and domestic ungulates are rarely prey of any of the carnivore species studied
here, but we sampled them since they may provide on occasions an important food
source as carrion for some of them. For the first study year, the prey censuses were
carried out at each quadrant at the same time as the carnivore censuses. For the
second study year, we concentrated prey sampling in a one-month period to avoid
particularly apparent inter-monthly variations in abundance for some species (i.e.,
small mammals and European rabbits) (Kufner 1986, Palomares et al. 2001). Thus,
we carried out the sampling of prey tracks in April (corresponding to the intraannual abundance peak in both species) along transects in every quadrant sampled
for carnivore tracks. As Kilometric Abundance Index (KAI) of prey calculated
for each quadrant was obtained from censuses carried out in different months,
we applied a correction in order to homogenize prey indexes for the two study
years relativizing the first years Kilometric Abundance Indexes to April using
the abundance trend curve of the species that likely exhibit higher inter-monthly
variation in their relative densities (rabbits and small mammals) throughout the year
(Moreno et al. 2007, Moreno and Kufner 1988, Villafuerte et al. 1993, Villafuerte
and Moreno 1997). These transects for prey species were located in the middle of
the same routes walked for carnivore tracks, were 25 m in length and approximately
1.7 m wide (i.e., the area of a four-wheel-drive car) and separated by at least 300 m..
Thus, between 7 and 10 prey censuses were carried out per quadrant.
105
Chapter 2
where we censused prey species. There, we visually estimated the cover of short
shrubs (species such as Halimium sp. and Cistus sp.), tall shrubs (species such as
Erica sp., Juniperus phoenica and Pistacia lentiscus) and trees in a circle of 25 m
diameter around the sampling point. In addition we also measured other variables
related to habitat structure: average tree height and average tall- and short-shrub
height.
Predictive variables
Fourteen variables were selected to study habitat selection patterns for each
The distance to water, distance to La Vera and distance to the anthropic edge
(see Table 1) were calculated from digitized roads, urban settlements, and water
source cover layers in DNP using a Euclidean distance-based approach (Perkin and
Conner 2004, Benson and Chamberlain 2007). Roads included firewalls and car
roads inside DNP. Urban settlements included towns and villages surrounding the
National Park. Water sources included natural and artificial ponds (i.e., dug for the
cattle in zones were the water table is higher) permanently flooded. Traffic index
per quadrant was derived from data on a previous study on the effect of traffic
on biodiversity in DNP (Romn et al. 2010). Ecotones between pastureland and
scrubland were defined using a 1:10 000 fine-scale vegetation map for the years 1996106
Chapter 2
Table 1. Explanatory variables used to model relative abundance of Egyptian mongoose, Eurasian
badger, and red fox and the probability of Iberian lynx and common genet presence.
Variable
Code
Definition
Units
Vegetation
tall shrub
%B
trees
%T
Landscape
distance to water
DW
distance to La Vera
DV
Prey availability
m/ha
rabbits
Ra
small mammals
SM
Total prey
Tot
distance to antropic
edge
DH
Human disturbance
Traffic index
Humidity
Observer
car/m
%
no units
107
Chapter 2
2006 obtained from the Sistema de Informacin Ambiental de Andaluca for the
Doana area. We reclassified vegetation units or polygons based on four vegetation
attributes of physiognomy, species composition and density of each vegetation layer
within the polygon: (1) trees (P. pinea, Quercus suber and Eucaliptus spp.); (2)
tall shrubs (subsequently referred to as bushes) of mature Mediterranean shrubland
(e.g., P. lentiscus, M. communis) and also tall, thicket Erica spp.; (3) short shrubs
(H. halimifolium, Ulex spp., Stauracanthus genistoides) and (4) pastures. The result
was a reclassified vegetation digital map with 316 polygons and four vegetation
attributes per polygon. The projection for all GIS layers and data was UTM 30S,
datum European 1950 (ED50). Hawths tools and Geopreocessing extension in
ArcInfo 9.3 (ESRI, Redlands, California, USA) was used to calculate distancebased variables, to identify ecotones, and to calculate their density (Table 1).
Statistical analysis
view that ecological inference can best be approached by weighing evidence for
multiple working hypotheses simultaneously (Hilborn and Mangel 1997, Burnham
and Anderson 1998, Johnson and Omland 2004). In essence, these methods consist
of identifying a priori the alternative hypotheses for habitat selection and their
mathematical formulation, and then testing their support by fitting the relevant
equations to species distribution data and examining penalized maximum-likelihood
estimates (e.g., Fernndez et al. 2003, Johnson et al. 2004).
predict species habitat use and distribution in DNP, therefore restricting the model
selection process to a few meaningful combinations of predictors of the species. For
108
Chapter 2
Chapter 2
a typical generalist and a typical specialist species (Virgs et al. 1999). Genets need
bushes and hollow trees as sites for nocturnal and diurnal resting and feed mainly on
small mammals such as the long-tailed field mouse (Apodemus sylvaticus) (Delibes
1974, Palomares 1986, Palomares and Delibes 1988, Palomares and Delibes
1991). We also hypothesized that distance to permanent water resources may be an
important factor for all species particularly during the hottest months when many
surface water sources dry out. Additionally, proximity to humans and infrastructure
derived from their activity as well as traffic index inside the protected area may
be detrimental to all the species because they produce higher mortality, degrade
the original Mediterranean ecosystems and involve a higher risk of predation or
competition with non-native carnivores (i.e., domestic dogs).
Highly correlated predictors (r > 0.6) were never included in the same
Models (GLMM) in SAS 9.2 with logit-link and binomial (for lynxes and genets) or
negative binomial (for badgers, mongooses and foxes) error structure (McCullagh
and Nelder 1989). Observer and quadrant were modelled as random effects,
humidity as an additional explanatory variable (Soto et al. 2012), and the distance
110
Chapter 2
covered per quadrant during track censuses as an offset in all the models.
where yj is the number of tracks at quadrant j and wj is the inverse Euclidean distance
between locations i and j. Hence, an autocovariate at location i is defined as a
weighted sum of observation records y at locations j in a neighbourhood determined
by Ni (Miller et al. 2007). The neighbourhood size may be informed by biological
111
Chapter 2
parameters, such as the species dispersal capacity (Knapp et al. 2003) if the cause
of spatial autocorrelation is known (or at least suspected). We hypothesised that
autocorrelation in our data may partially originated from movement of censused
individuals between sampling sites so we set the neighbourhood size to two quadrants
from each quadrant border to capture the average home range for all species. We
incorporated each autocovariate as an additional explanatory variable in the GLMM
models to account for the variation explained by space while maintaining the same
variable selection procedures as for spatially invariant models. Finally, we tested
autocovariate models for autocorrelation in the Pearson residuals, using Morans
I correlograms and semivariograms of the most parsimonious model. We used
variogram procedure in SAS 9.2 to conduct these tests.
Chapter 2
the study area. The niche breadth or species tolerance is measured by an additional
variance term provided by this method. Low values of species tolerance mean that
a species is distributed across habitats with a limited range of conditions (specialist
species), while high values imply that a species is distributed across habitats with
widely varying environmental conditions (generalist species). Residual tolerance is
the variation in species occurrence not accounted for by the main gradient. Outlying
mean index is robust to unimodal, linear, or a mixture of species response curves and
is not biased against species-poor or low-abundance sites on the synthetic gradient.
Its interpretations are also robust to multicollinearity among the explanatory
variables (Doldec et al. 2000). We determined significance of the outlying mean
index analysis at =0.05 based upon a Monte Carlo simulation (Metropolis and
Ulam 1949), in which observed marginalities were statistically compared to 10,000
random permutation values of species marginalities or the null hypothesis that
species are distributed equivalently in relation to the environmental variables. The
OMI analysis was performed with the ADE4 library (Thioulouse et al. 1997) in the
R Software (R Development Core Team 2005).
RESULTS
We surveyed 471 km and 8,373 carnivore tracks were found, with foxes,
badgers and mongooses being the most frequent species (Fig. 2, Table 2). For prey,
5,000 tracks were detected on 11.6 km sampled. The most common prey species
were wild ungulates and rabbits (Table 2). The variables Ra and Tot were correlated
(r = 0.669, P < 0.001).
residuals, habitat use models for badgers, mongooses and foxes showed spatial
113
Chapter 2
Figure 2. Maps of scaled abundances (0-1) (tracks/km) per quadrant of the five carnivore
species for both study years in DNP. (a) Genetta genetta; (b) Lynx pardinus; (c) Herpestes
icheumon; (d) Meles meles and (d) Vulpes vulpes.
114
Chapter 2
autocorrelation while it this was not detected for lynxes and genets. We therefore
fitted spatial negative binomial generalised models (i.e., using an autocovariate)
for badgers, mongooses and foxes and non-spatial logistic generalised models for
lynxes and genets.
The best approximating models (AICc < 2) for mongooses, badgers and
foxes belonged to the set of candidates designed with the hypothesis of vegetation
and landscape structure + prey availability as well as with the hypothesis of prey
availability (Table 3). For mongooses, top models included as predictors of their
relative abundance DV, DW, Tot, Ra, %SB, %B and eBP. Variables DW and DV had
the highest weights and were positively associated with the number of mongoose
tracks (Table 5). Variables Tot and %B were also positive and had relatively high
weights (wj > 0.4) (Table 5). For badgers, models included as predictors Ra, SM,
%SB, %B, %T, DW, V and eBP. All of these predictors were positive except for the
distance to La Vera. Ra was the variable with the highest weight (wj = 0.577) (Table
5) and was positively associated with the number of badger tracks. For the red fox
only one model was supported by data (Table 3). This model included as predictors
%B, %T, DW and SM and were all positively associated with the number of fox
tracks. Only two variables showed high weights; %B (wj = 0.809) and DW (wj =
0.871) (Table 5).
The best approximating models for lynxes and genets belonged to the set
116
61
55
29
12
15
14
69
69
49
43
21
69
69
Vulpes vulpes
Herpestes ichneumon
Meles meles
Genetta genetta
Felis sp.
Canis familiaris
Lutra lutra
TOTAL
Oryctolagus cuniculus
Small mammals*
Alectoris rufa
Domestic ungulates**
Wild ungulates***
TOTAL
65
65
27
44
52
65
65
14
12
21
22
53
50
61
17
2nd
73 (1.7)
2nd
4312
37 (0.9)
29 (0.7)
35 (0.8)
83 (1.9)
665 (15.4)
617 (14.3)
3.25.8
0.20.7
0.20.8
0.10.5
1.22.5
2.63.7
3.94.6
17.29.1
0.30.9
1st
Average
210 (4.2)
260 (5.2)
233 (4.7)
2.5.2
12.521.6
6.7.4
0.3 0.9
2nd
Average
0.0-5.0
2nd
Range
0-0.09
0-0.3
0-0.2
0-1.1
0.0-42.1
0.0-3.8
0.0-5.9
0.0-3.9
0.0-15.9
0.0-19.5
0.0-21.2
3.2.3
4.0.7
3.6.0
32.844.6
2.64.45
0.20.4
0.20.5
0.20.3
0.40.8
3.33.4
3.13.7
0-0.1
0-0.07
0-0.1
0.007-1
0.0-77.4
0.0-2.2
0.0-3.4
0.0-1.4
0.0-3.1
0.0-17.3
0.0-17.2
0.0-5.4
1st
Range
KAI
9132
5000
26.927.9
0-1.1
15.417.0
0-1
168 (1.8)
848 (9.3)
454 (5.0)
6639
49 (0.7)
38 (0.6)
48 (0.7)
280 (4.2)
786 (11.8)
996 (15.0)
74 (1.1)
1st
N tracks (%)
*Eliomys quercinus, Arvicola sapidus, Rattus rattus, Apodemus sylvaticus, Mus spp.
17
69
Lynx pardinus
#1st
Specie
Positive quadrants
Table 2. Number of positive 2x2 km quadrants (for 69 and 65 sampled in each year), total tracks found (N tracks) and percentage regarding total
tracks found (%), and number of tracks per km (KAI) for each carnivore and their potential prey species censused in Doana National Park during
the wet season for every study year.
Chapter 2
Chapter 2
presence of genet tracks while DW was negatively associated. Finally, for lynxes
top models included as predictors %SB, %B, DW, eBP, V and Ra. Variables included
in the best models showed high weights (wj>0.8) (Table 5). %SB, %B, DW, eBP
and Ra were positively associated with the presence of lynx tracks while DV was
negatively associated.
To illustrate niche separation between species, we used the first two axes
of the outlying mean index analysis, which accounted for 99.95% of the total
explained environmental variability (Table 1, supplementary information). The
overall outlying mean index analysis (i.e. sensitivity of carnivores to environmental
variables) was significant (P < 0.0001).
The position of the species along the first two OMI axes is presented in Fig.
3. Genets and lynxes were clearly separated from mongooses, badgers and foxes.
Figure 3. OMI analysis. Ecological position of the five carnivore species in the n-dimensional
hypervolume representing fine-scale environmental variability in DNP. (a) Lynx pardinus,
(b) Genetta genetta, (c) Herpestes ichneumon, (d) Meles meles and (d) Vulpes vulpes.
117
118
5. DW, DV
862.91
8. Tot
9. DH, T
9.94
6.82
0.00
0.02
859.79
7. SM
Human disturbance
858.83
6. Ra
0.03
5.86
3. %B, %T, DW
Prey availability
16
11
12
15
14
17
Herpestes ichneumon
AICc
i
Wi Ranking
1. Intercept only
Null model
Model
889.94
861.13
861.61
858.97
890.03
888.92
886.03
885.97
910.64
AICc
30.97
2.16
2.64
0.00
31.06
29.95
27.06
27.00
51.67
0.00
0.11
0.09
0.33
0.00
0.00
0.00
0.00
0.00
Meles meles
i
Wi
14
15
13
12
11
17
Ranking
4.99
4.71
4.16
0.03
0.04
0.05
1222.05
1221.77
1221.22
16
14
12
11
13
17
Vulpes vulpes
i
Wi Ranking
AICc
Table 3. Summary of spatial negative binomial predictive models for Egyptian mongoose, Eurasian badger and red fox abundance and model
selection estimators; -2 log(L)=-2 log-likelihood estimates; AICc = second order Akaikes Information Criterion; i = (AICc)i - (AICc)min; Akaike
Wi = Akaike weights. Top models (AICc2) are shown in bold.
Chapter 2
855.37
852.97
0.02
6.59
859.56
0.05
0.48
0.14
0.02
0.25
4.61
0.00
2.40
5.95
1.33
Global models
858.92
13
10
894.15
867.54
864.74
863.31
865.11
863.36
860.64
859.94
35.18
8.57
5.77
4.34
6.14
4.39
1.67
0.97
0.00
0.00
0.02
0.04
0.02
0.04
0.14
0.21
16
10
2.86
4.33
6.01
3.86
2.05
0.00
3.86
0.10
0.05
0.02
0.06
0.15
0.42
0.06
1219.92
1221.39
1223.07
1220.92
1219.11
1217.06
1220.92
15
10
Chapter 2
119
Chapter 2
Table 4. Summary of non-spatial logistic predictive models for common genet and Iberian lynx
occurrence, and model selection estimators; -2 log(L)=-2 log-likelihood estimates; AICc = second
order Akaikes Information Criterion; i = (AICc)i - (AICc)min; Akaike Wi = Akaike weights. Top
models (AICc2) are shown in bold.
Genetta genetta
Model
AICc
Wi
Lynx pardinus
Rank
AICc
Wi
Rank
Null model
1. Intercept only
17
151.89 44.15
0.00
15
12
107.74
0.00
0.52
3. %B, %T, DW
163.47
0.02
141.70 33.96
0.00
10
13
121.26 13.52
0.00
5. DW, V
14
125.16 17.42
0.00
6. Ra
16
151.71 43.97
0.00
14
7. SM
15
152.87 45.13
0.00
16
8. Tot
163.08
6.44
0.02
148.24 40.50
0.00
13
166.58
9.94
0.00
11
155.17 47.43
0.00
17
8.05
0.01
109.53
1.79
0.21
157.25
0.61
0.34
142.21 34.47
0.00
11
162.11
5.47
0.03
122.23 14.49
0.00
162.67
6.03
0.02
110.99
3.25
0.10
162.32
5.68
0.03
110.02
2.28
0.17
156.64
0.00
0.47
146.19 38.45
0.00
12
161.11
4.47
0.05
122.93 15.19
0.00
164.91
8.27
0.01
10
122.72 14.98
0.00
Prey availability
Human pressure
9. DH, T
Vegetation and landscape structure
+ prey availability
Global models
14. %SB, %B, DW, eBP, V, Ra,
DH, T
15. %B, %T, DW, SM, DH, T
Genets and lynxes had the highest marginality values and the lowest tolerance
to average habitat conditions of the synthetic gradients (Table 2, supplementary
information). Badgers and foxes had the lowest marginality and the highest
120
Chapter 2
tolerance values whereas mongooses exhibited relatively low marginality and high
tolerance. Although the overall response of mongooses was non significant their
marginality and tolerance levels was between that of genets/lynxes and badgers/
foxes as predicted.
Ordination diagrams on the first two axes of the OMI describe the
environmental gradients that best discriminated the occurrences of the five carnivore
species in DNP (Fig. 1, supplementary information). The first OMI axis was most
influenced by prey availability and shrub cover far from human nuclei and the
proportion of tall shrubs and trees at the opposing end of the gradient. The second
OMI axis was most influenced by distance to the main ecotone between marshland
and scrubland (La Vera) and prey availability and percentage of ecotones between
pastureland and scrubland at the opposite end of the gradient (Table 2, supplementary
information). The presence of genet tracks was negatively associated with axis 1
(Fig. 3), and thus was positively related to patches distant from human nuclei and
with high tree and tall shrub coverage. Lynxes were positively associated with axis
1 and negatively with axis 2 and thus positively related to patches with high prey
availability (mainly rabbits) close to La Vera. Conversely, mongooses, badgers and
foxes were more likely to be found across a wide range of habitat types with varying
environmental conditions.
DISCUSSION
different axes over which species can differ), thus broadening opportunities for
coexistence. However, in quite homogeneous landscapes where a general habitat
type dominates, the distribution of the resources available for species is limited, and
121
122
autocov
HUM
DH
Tot
SM
Ra
DW
eBP
%T
%SB
%B
Variables
0.22
0.07
0.61
0.16
0.01
2.96
0.02
8.08
0.17
0.04
0.08
0.21
0.08
0.24
-0.27
-0.10
0.12
0.17
1.00
1.00
0.64
0.32
0.93
0.96
0.29
0.44
0.48
Herpestes
ichneumon
SE()
wj
9.71
0.02
3.27
0.01
0.01
0.00
0.02
0.01
0.03
0.00
0.19
0.14
0.10
0.26
SE()
0.05
-0.01
0.25
0.42
0.14
0.33
Meles meles
1.00
1.00
0.21
0.25
0.58
0.26
0.46
0.24
0.29
0.45
wj
1.91
0.01
0.00
0.00
0.29
0.35
0.00
0.02
0.00
0.01
0.22
0.25
SE()
wj
1.00
1.00
0.47
0.51
0.87
0.81
Vulpes vulpes
0.02
-0.16
0.01
0.60
-0.12
0.05
0.10
0.02
0.14
0.04
0.18
0.07
0.01
0.06
SE()
wj
1.00
0.56
0.56
0.47
0.98
0.83
0.95
Genetta genetta
0.02
0.02
-2.56
0.48
1.85
1.69
0.32
0.02
0.00
0.70
0.37
0.69
1.2
0.12
SE()
wj
1.00
0.38
1.00
1.00
0.90
0.90
0.90
Lynx pardinus
Table 5. Variable weight (wj), average model coefficient () with average standard error (SE()) for variables included in top models.
Variables are described in Table 1.
Chapter 2
Chapter 2
opportunities for coexistence are likely to be the most restrictive. Hence, subtle
patterns of habitat partitioning as well as coexistence between species with different
degree of specialisation may promote ecological separation and therefore sympatry
under this scenario. A parsimony-based strategy for confronting different model
hypotheses allowed us to assess fine-scale landscape attributes linked to five
carnivore species in a protected area where species coexist in a largely similar
habitat type, the Mediterranean scrubland. These species exhibited spatial storage
or separate niches along a gradient of environmental variability according to their
degrees of specialization.
On the basis on OMI analysis, lynxes and genets are marked specialists and
showed the narrowest and most marginal niches throughout DNPs environmental
conditions (Fig. 3). Additionally, lynxes and genets showed niche segregation
as a result of their differences in ecological preferences (Table 2 supplementary
information), which is supported by the fine-scale habitat use results. Results
of fine-scale habitat use models for lynxes and genets were probably related to
the availability of prime resources for both species such as refuges and prey.
Lynxes and genets showed the most restricted habitat use pattern depending on
vegetation, landscape, prey and human-related variables. Lynxes were associated
with zones of high density of bushes likely representing areas of late-successional
Mediterranean communities, an uncommonm microhabitat in the area due to the
human transformation of the autochthonous vegetation (Garca-Novo and MartnCabrera 2005). The density of ecotones between bushes and pastureland, which in
turn favour abundance of the main prey of lynx, the European rabbit (Fernndez
et al. 2006, Fernndez et al. 2007), was also a robust predictor of the presence of
lynx tracks, as well as the distance to the ecotone band between the Mediterranean
123
Chapter 2
scrubland La Vera, where refuge and grass availability favour a high rabbit density
in Doana (Villafuerte and Moreno 1997, Fernndez and Palomares 2000). As
expected, genets showed association to areas with high small mammal abundance
(a community mostly composed of long-tailed field mouse), important prey items
for the species in Mediterranean areas (Palomares and Delibes 1991), as well as
preference for pine forest areas with undergrowth bushes. These results agree with
previous findings for the radio-tracked population of the Iberian lynx (Palomares et
al. 2000, Palomares 2001, Fernndez et al. 2003) and the common genet (Palomares
and Delibes 1988, 1991, 1994, Palomares et al. 1996) in the protected area. The low
abundance and the spatially structured distribution of both species in DNP (Fig.
2) could therefore be explained by the fact that genets and lynxes may be limited
in their home ranges to areas of simultaneous convergence of high small mammal
or rabbit availability as well as a high percentage of tall shrub cover and overall
understorey or a high percentage of tree cover, respectively.
Foxes and badgers showed a non-specific habitat use pattern and a wide
Mongooses were positively associated with dense bush cover and high
availability of different prey items as a whole (i.e., rabbits, small mammals and
124
Chapter 2
and badgers had broader niches than mongooses (Fig. 3, Table 1 supplementary
information). In fact, mongooses exhibited and intermediate niche position and
niche breadth between that of genets/lynxes and that of badgers/foxes (Table 2,
supplementary information). These results suggest that mongooses exhibited certain
degree of habitat specialization.
The wider spatial niches exhibited by mongooses, foxes and badgers may
fine-scale habitat selection only for the most specialist species (lynxes and genets).
Badgers and foxes meanwhile did not show any clear pattern of specialization at
a fine scale as expected due to their high levels of habitat and trophic generalism.
Although mongooses were more specialised in habitat selection than badgers and
foxes, for areas such as that studied here the three species can coexist with no
apparent differences in habitat use and/or realized niche segregation. Differences in
activity patterns of these three species might help to explain the coexistence among
them. Mongooses are almost exclusively diurnal in the study area (Palomares and
125
Chapter 2
Delibes 1993) while badgers are exclusively nocturnal (Revilla and Palomares
2002) and foxes nocturnal and crepuscular. Therefore, the carnivore community
structure in DNP seems not to be defined only by habitat use partitioning.
Interspecific interactions among predators are another aspect that should
be explored and can also greatly shape the community structure. Mongooses, for
example, were strongly negatively associated with distance to La Vera, which might
be related with the high use that lynxes made of this area (Palomares et al. 1991,
Viota et al. 2012).
ACKNOWLEDGEMENTS
Education and Science) and 17/2005 (Spanish Ministry of the Environment; National Parks Research
Program). Land-Rover Espaa lent us two vehicles for this work. We are very grateful especially to J.C.
Rivilla and S. Desnia for assistance during fieldwork and to M. Gonzlez for her valuable suggestions
on earlier versions of the manuscript. C. Soto was also supported by a JAE-Predoc grant from the CSIC.
126
Chapter 2
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132
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Supplementary information
Table 1. Results of the outlying mean index analysis depicting relationships of occurrences of
Carnivora species to the suite of environmental variables measured across sampling sites in DNP.
Inertia = variance or weighted sum of squared distances to the origin of the environmental axes;
OMI = outlying mean index (marginality) or the deviation of a particular species distribution
from the overall mean habitat conditions (origin of outlying mean index axes), described by the
environmental variables; Tol = tolerance index, which is analogous to niche breadth or spatial
variance of an organisms niche across the measured environmental variablesa function of
all sampling sites with which the species is associated; RTol = residual tolerance. Italicized terms
represent the percentages of variability corresponding to a specific statistic. P = frequency based
on number of random permutations (out of 10,000) that yielded a higher value than the observed
outlying mean index (P 0.05 indicates a significant influence of the environmental variables for a
species).
Species
OMI
Tol
5.73
39.30
17.40
43.30
0.00040
1.61
6.54
26.40
31.90
41.70
0.00010
0.35
2.30
7.45
2.70
18.00
79.30
0.09329
11.79
0.25
1.69
9.83
3.00
13.60
83.40
0.06429
10.89
0.04
4.99
9.51
0.30
12.40
87.30
0.63804
Inertia
OMI
Tol
Lynx pardinus
13.22
5.19
1.35
Genetta genetta
15.66
4.13
Herpestes ichneumon
9.39
Meles meles
Vulpes vulpes
Rtol
Rtol
Table 2. Loadings of environmental variables for the first two axes of the outlying mean index
analysis. Values represent the best linear combination that explains occurrences of Carnivora at
DNP. Variables are described in Table 1.
Variable
Axis 1
Axis 2
%SB
0.2014
0.0251
%B
-0.2361
-0.1876
%T
-0.2138
0.1877
DH
-0.5163
-0.1776
Traf
0.1279
0.3091
eBP
0.0237
-0.5020
DW
0.2807
0.2649
DV
-0.0605
0.5255
Ra
0.2431
-0.3693
SM
0.0365
0.0165
Tot
0.2635
-0.2575
Eigenvalue
0.3766
0.0003
0.9986
0.0009
133
Chapter 2
Figure 1. Contribution of environmental, human and prey variables to the first axes produced by
the OMI analysis. The length of the arrow describes the relative importance of each variable in the
analysis, and the direction of the arrow indicates among-variable correlations.
DV
Traf
DW
T
SM
DH
SB
Axis 1
B
Tot
Ra
eBP
134
CHAPTER 3
135
136
Chapter 3
ABSTRACT
137
Chapter 3
RESUMEN
Chapter 3
INTRODUCTION
of the quality of that habitat for the species is a widely accepted assumption in
habitat use studies (but see VanHorne 1983). Nevertheless, some species may not
exhibit the highest densities in their potentially best optimal habitats according to
the species life-history traits, as their relative abundances and pattern of habitat use
can be affected by interspecific interactions with other species at the same trophic
level (Case and Gilpin 1974, Estes and Palmisano 1974, Menge and Sutherland
1987, Durant 1998).
interspecific interactions (Palomares and Caro 1999). With the exception of some
species that prey in packs (e.g., Rogers and Mech 1981, Paquet and Carbyn 1986),
interference competition is inflicted by a larger carnivore species on a smaller
one killing it (termed as intraguild predation (Polis et al. 1989)) or excluding it
from habitat patches or prey carcasses (Palomares and Caro 1999). These highly
asymmetrical interactions mediated by body size are common (e.g. Fedriani et al.
1999, Kamler et al. 2003, Durant et al. 2010) being the smaller species almost
invariably the loser due to their subordinate position (Palomares and Caro 1999,
Prugh et al. 2009, Roemer et al. 2009). Nevertheless, smaller members of a guild
may also kill cubs, young or subadult individuals of the larger species (Palomares
and Caro 1999). In the absence of predator species, small carnivores should be
ideally distributed based on habitat quality and preferred food availability (Van der
Meer and Ens 1997, Roemer et al. 2009), but in most natural communities smaller
species are potentially subject to top-down effects that mediate their ability to use
preferred habitat and limit their access to high-quality foraging areas (Palomares
139
Chapter 3
and Caro 1999, Ritchie and Johnson 2009). Moreover, in some carnivores,
intraguild predation has a considerable impact on small carnivore mortality rates
and consequently a negative effect on the relative abundance of the species in
the presence of the larger predator (Ralls and White 1995, Sovada et al. 1998).
In such cases, due to these potential predation-driven direct effects or fear-driven
indirect effects, the relative abundances of some predators may not be related to
prey availability or vegetation type. Densities of smaller members of a guild may
represent a trade-off between suitable areas that satisfy their basic requirements and
areas where probability of encounters with larger predators are null or lower (i.e., to
diminish the risk of being killed).
sympatric carnivores on the habitat use pattern of five small and medium-sized
carnivore species in a Mediterranean protected area in southwest Spain, namely
Doana National Park (DNP). On previously fitted fine-scale habitat use models for
a guild of five carnivore species we analysed whether the inclusion of the relative
abundances of any carnivore species improves the explanatory power of habitat
models (Johnson and Omland 2004, Burnham and Anderson 2002). The studied
carnivore guild is composed by the Iberian lynx (Lynx pardinus, 9-15 kg), Eurasian
badger (Meles meles, 7-8 kg), the red fox (Vulpes vulpes, 5-7 kg), the Egyptian
mongoose (Herpestes ichneumon, 3 kg) and the common genet (Genetta genetta,
2 kg). In DNP these species form a community of continuum degree of habitat
and trophic specialization predators with different degree of habitat use overlap.
It has been described aggressive interactions between some of these carnivores in
DNP, which always involved lynxes as the top predator. The Iberian lynx uses to
kill genets, mongooses and foxes (Palomares et al. 1996, Fedriani 1997), but no
140
Chapter 3
aggressive interactions between lynx and badger has been reported. On the other
hand, aggressive interactions between badgers, foxes, mongooses and genets have
been unreported, although according to the theory differences in body size among
some of these species could support such interactions (Palomares and Caro 1999,
Fedriani et al. 2000).
Specifically, we tested the hypothesis that in our carnivore guild the spatial
METHODS
Study area
DNP is a fully protected and flat sandy area at sea level in southwestern
Spain (550 km2 379N, 626W), located on the west bank of the Guadalquivir
River mouth. The climate is Mediterranean sub-humid, characterized by dry, hot
summers and mild, wet winters. Average annual rainfall is 500-600 mm. There are
three main biotopes in the park: scrubland, dunes, and marsh (Valverde 1958). The
dune area is situated at the western border of the protected area limited by the
141
Chapter 3
Atlantic Ocean and the marsh area at the northern and eastern borders limited by the
Guadalquivir River. The Mediterranean scrubland represents approximately half of
the National Park surface area and is mainly characterised by heterogeneous patches
of xerophytic species such as Halimium sp. and Cistus sp., and hydrophytic species
such as Erica sp., with some patches of Juniperus phoenica and Pistacia lentiscus
shrubs. Interspersed among the scrubland are scattered cork oak trees (Quercus
suber) and wild olive trees (Olea europea), and a few patches of pine Pinus pinea
Figure 1. Sketch of the possible expected interrelationships among the different carnivore
species studied here. Single-headed arrow and straight lines represent aggressive interactions that
have been previously reported in the literature for those species in the study area; single-headed
arrow and dashed lines represent potential aggressive interactions based in the body size of the
involved species but that have been previously unreported; and double-headed arrow and dashed
lines represent potential neutral expected interactions between similar-sized species in the DNP
carnivore guild.
142
Chapter 3
In addition to the studied species (i.e., red fox, Eurasian badger, Egyptian
mongoose, common genet and Iberian lynx), there are other carnivores such as
least weasel (Mustela nivalis), European polecat (Mustela putorius), Eurasian otter
(Lutra lutra) and wild cat (Felis silvestris), but they were excluded from the study
because of their scarcity in the area.
Sampling protocol
densities for foxes, badgers, lynxes, mongooses and genets we divided the study
area into 69 2 x 2 km quadrants following UTM coordinates (Figure 2). Quadrants
were sampled throughout the wet seasons of 2007-2008 and 2008-2009. In each
quadrant, a 3 km-length survey route was slowly walked searching for carnivore
tracks. The survey routes were located along firebreaks and car roads between 2 and
12 m wide. Once a continuous track that crossed from one side to the other across
the pathway was detected, we georeferenced it using a global positioning system.
For the two sampled seasons altogether we resampled 21 squares (leaving at least 7
days between samplings) a second time until completing 3 km, because there were
insufficient available paths within the square to achieve this distance. Thus, as we
had more censuses than quadrants we averaged track counts to get a single value per
quadrant. We always carried out surveys at least 3 days after any rainfall.
143
Chapter 3
Data analysis
given species according this outlined in Figure 1, we first fitted fine-scale habitat
use models for each species (Chapter 2). After, we added the relative abundance
(Kilometric Abundance Index; KAI) or presence of the different carnivore species
to the best approximating habitat model describing abundance of each species
(hereafter null models) and observed whether models improved (i.e. a decrease of
the Akaike Information Criterion corrected for small sample size; AICc). According
to the information-theoretic approach, models with AICc < 2 from the model with
the lowest AICc are considered to be virtually the same plausible models (Burnham
and Anderson 2002). Hence, in spite of the existence of certain model selection
uncertainty in multimodel inference, statistical models that decrease the AICc of
a reference model more than 2 units could be considered as more parsimonious
models and to best fit the data. Thus, for the specific aims of our study, when AICc
< -2 and model coefficient () was negative we considered that there were support
to the predictions about the abundance or presence of a given species affected the
abundance or presence of the target species; when AICc < -2 and model coefficient
() was positive we considered that both species could coincide in habitat resources
used, but no negative interaction occurred between them; and finally when -2 >
AICc < 2 we considered that abundance or presence of a given species did not
affect the abundance or presence of the target species. Note that if our hypothesis
is right, the inclusion of the relative abundance of presence of smaller species in
the model of habitat selection of larger species should have either no effect or if
any (i.e., AICc > 2), should be positive. Hence, we included as a control of
hypothesis all possible carnivore species in null models for each target species to
144
Chapter 3
Chapter 3
Variable
Code
Definition
Mean cover of bushes per quadrant calculated by averaging values
obtained at the different sampling points of vegetation censuses
Mean cover of trees per quadrant calculated by averaging values
obtained at the different sampling points of vegetation censuses
Measured in meters using a Euclidean distance-based approach
from the quadrant centre to the nearest permanently flooded natural
or artificial pond (i.e. dug for the cattle at zones were the water table
is higher) in a digitized water sources cover layer of DNP
Measured in meters from the quadrant centre to the ecotone between
the marshland and the Mediterranean scrubland (locally called La
Vera)
Units
tall shrub
%B
trees
%T
distance to
water
DW
distance to
La Vera
DV
small
mammals
SM
rabbits
Ra
distance to
antropic
edge
DH
Mean daily traffic (MDT), adjusted for road length (m) and type of
road (i) in the quadrant (= (MDTi * mi)) derived from Romn et
al. 2010
car/m
Traffic
index
Humidity
Hum
Observer
Obs
146
%
%
m
%
no
units
Chapter 3
RESULTS
< -2) after including the occurrence and/or presence of lynxes (Table 2). The
three species were less abundant in areas where lynxes exhibited a higher relative
abundance (Fig. 3a, c and e). Habitat models of foxes, genets (Fig. 3b) and badgers
also improved after including the relative density of mongooses (Table 2) but the
Figure 2. Map of the DNP showing its location in southwest of the Iberian Peninsula and the 22
km2 quadrants where carnivore track censuses were carried out during the wet seasons of 2007-2008
and 2008-2009. The dashed zone represents the marshland area of the DNP.
147
Chapter 3
three species exhibited higher relative densities in areas where mongooses were
also more abundant.
The inclusion of the presence of genets and the relative abundance of badgers
(Fig. 3d) also improved the habitat model of mongooses that were more abundant
in areas where genets were detected or where the occurrence of badgers was higher
(Table 2). The inclusion of any of the smaller members of the guild significantly
improved the habitat model of the Iberian lynx (Table 2).
DISCUSSION
Our results support the hypothesis that in a carnivore guild, the occurrence
of some species exerts a significant effect on the relative abundance of other guild
members after controlling by species-specific habitat use patterns. As we previously
hypothesized, the relative abundance of foxes, mongooses and genets in DNP was
negatively affected by the presence or occurrence of lynxes, the larger reported
intraguild predator for all of them (Palomares and Caro 1999). In fact, it had been
previously described for Doana as foxes, mongooses and genets avoided temporal
or spatially highly used areas by lynxes (Palomares et al. 1996, 1998, Fedriani et
al. 1999, Viota et al. 2012). Similar results have been obtained in other carnivore
communities involving African wild dogs Lycaon pictus and cheetahs Acinonyx
jubatus with lions Panthera leo and spotted hyenas Crocuta crocuta (Laurenson
1995, Creel and Creel 1996, Durant 2000).
Also as predicted, the relative abundance of smaller species did not affect
the relative abundance of larger ones; in our case lynxes, badgers and foxes.
However, although theory predicts that larger species such as foxes and badgers
might negatively interact with genets and mongooses, results did not confirm
148
Chapter 3
Table 2. Results from negative binomial (foxes, badgers and mongooses) and logistic regression
models (genets and lynxes) investigating the effects of the occurrence and/or presence of predator
species on each carnivore species in the DNP. We report the small sample-sizeadjusted Akaikes
information criteria (AICc), the difference in AICc between each model and the null model (AICc),
the model coefficient () with standard error (SE) as well as its P-value of those models that
decreased the null model 2. Variables are described in Table 1.
Species
Models
SE() P-value
AICc
AICc
0. Intercept only
1. Null model
%B + %T + DW + SM + DH
+ Traf + Hum + autocov
2. Null model + KAIl
3. Null model + KAIb
4. Null model + KAIf
5. Null model + KAIIm
6. Null model + Lynx
175.23
12.55
162.68
0.00
159.75
162.07
161.07
153.01
160.62
-2.94
-0.61
-1.61
-9.67
-2.06
-1.14
.
.
0.26
-1.13
0.72
.
.
0.08
0.59
0.05
.
.
0.00
0.08
0. Intercept only
1. Null model
DW + DV + Tot + Hum +
autocov
2. Null model + KAIl
3. Null model + KAIb
4. Null model + KAIf
5. Null model + KAIg
6. Null model + Lynx
7. Null model + Genet
918.52
38.76
879.76
0.00
877.23
874.89
879.74
879.63
881.56
863.83
-2.53
-4.87
-0.02
-0.13
1.80
-15.93
-0.31
0.06
.
.
.
0.82
0.14
0.02
.
.
.
0.19
0.03
0.01
.
.
.
0.00
0. Intercept only
1277.53
24.66
1252.87
0.00
1253.99
1254.09
1250.46
1254.84
1249.69
1250.91
1.12
1.22
-2.41
1.97
-3.18
-1.96
.
.
0.03
.
-0.27
.
.
.
0.01
.
.
0.03
.
0.02
.
Genetta genetta
Herpestes
ichneumon
Vulpes vulpes
1. Null model
%B + %T + DW + SM + Hum
+ autocov
2. Null model + KAIl
3. Null model + KAIb
4. Null model + KAIm
5. Null model + KAIg
6. Null model + Lynx
7. Null model + Genet
0.12
.
149
Chapter 3
Meles meles
910.64
9.29
901.35
0.00
901.98
903.12
890.65
903.04
900.83
902.95
0.63
1.77
-10.70
1.69
-0.52
1.60
.
.
0.11
.
.
.
.
.
0.03
.
.
.
.
.
0.00
.
.
.
0. Intercept only
153.89
1. Null model
%B + %SB + DW + eBP + DV
111.92
+ Ra + Hum
2. Null model + KAIb
112.27
3. Null model + KAIf
111.00
4. Null model + KAIm
113.92
5. Null model + KAIIg
110.43
6. Null model + Genet
112.21
41.97
0.00
0.35
-0.92
2.00
-1.49
0.29
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
0. Intercept only
1. Null model
%SB + %B + eBP + DW + DV
+ Ra + Hum + autocov
2. Null model + KAIl
3. Null model + KAIf
4. Null model + KAIm
5. Null model + KAIg
6. Null model + Lynx
7. Null model + Genet
Lynx pardinus
Chapter 3
The same explanation might apply to the positive association found between
mongooses and genets; two similar-sized species. Genets and mongooses partially
overlap in their habitat use in DNP (Chapter 2). Furthermore, previous studies have
also shown that both mongooses and genets use similar microhabitats such as dense
bushes or thickets for activity and resting in Doana (Palomares and Delibes 1993),
and that a large fraction of their diets may coincide (Palomares and Delibes 1991).
On the same way, although mongooses exhibit a certain higher degree of habitat
specialization than foxes and badgers, the niche overlap of all three predators is large
probably due to their trophic and/or habitat generalism (Chapter 2). Additionally,
mustelids are not species with a high aggressive nature (Palomares and Caro 1999),
so it is reasonable to think that they can coexist with smaller predators without any
interference.
Our results demonstrate that the inclusion of the relative abundance of other
carnivore species in habitat models for a given species may significantly improve
them. Models based on species relative densities that excluded other predators
occurrence could be a misleading proxy of habitat quality when studying patterns of
habitat use by species, and this may be particularly important for smaller predators
that very often are intraguild prey in natural communities. Smaller species might be
more abundant at sub-optimal or marginal habitats due to predation-driven direct
or fear-driven indirect effects from larger predators. There are some examples of
other carnivore species that have been reported as exhibiting limited patterns of
distribution and abundance due to interference competition with larger predators
(Berger and Gese 2007, Peterson 1995, Thurber et al. 1992, White and Garrott
1997, Burrows 1995, Durant 2000). However, habitat use models for carnivore
species at the apex of their ecological communities that are based on vegetation
151
Chapter 3
Figure 3. Relationships between the probability of genet presence and the numbers of lynx (a) and
mongoose tracks (b) detected; between the numbers of mongooses tracks and that of lynxes (c) and
badgers (d); and between the number of fox tracks and the presence of lynx (least square means and
their standard errors are represented) (e).
152
Chapter 3
ACKNOWLEDGMENTS
This study was financed by the projects CGL2004-00346/BOS (Spanish Ministry of Education and
Science) and 17/2005 (Spanish Ministry of the Environment; National Parks Research Programme),
and sponsored by Land-Rover Espaa S.A. We are especially thankful to J.C. Rivilla and S. Desnia
for their assistance during fieldwork. C. Soto received a JAE predoctoral grant from CSIC (Spanish
National Research Council).
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CHAPTER 4
158
Chapter 4
ABSTRACT
Chapter 4
RESUMEN
Los efectos de los perros en la fauna nativa en las reas protegidas debe
km2 y analizamos los factores ambientales y/o humanos que pudieran determinar
su presencia a travs de modelos de regresin logsticos. No detectamos perros en
zonas alejadas de los lmites antrpicos del rea protegida a pesar del esfuerzo de
muestreo (ms de 470 Km. de caminos censados). Segn nuestros resultados, los
perros presentes en el Parque Nacional de Doana deben ser perros domsticos
que hacen incursiones de manera ocasional en el rea protegida desde la matriz
antrpica circundante. La deteccin de rastros de perros dependi de la presencia
de humanos y consecuentemente de los recursos que proporcionan. Los efectos
directos de los perros en la fauna nativa no deben por tanto mostrar la misma
intensidad en toda la extensin del rea protegida o a lo largo de toda la longitud
160
Chapter 4
161
Chapter 4
INTRODUCTION
the causes of native species decline and endangerment (e.g., King 1985, Soul
1990, Williamson 1999, IUCN 2012). Due to human population growth, urban
sprawl and the rapid urbanization of natural landscapes, humans and with them
their companion animals such as domestic dogs Canis familiaris may be closely in
contact with wildlife in many areas (Ordeana et al. 2010) including places where
nature conservation is a priority, such as protected areas and national parks created
to protect populations of vulnerable or threatened species. These non-native species
introduced by human across the globe (Wandeler et al. 1993) may therefore increase
their ranging activity towards the remaining natural landscapes extending within
those areas the deleterious human-associated effects.
dogs may pose distinct threats including competition, interbreeding, predation and
disease. Dogs harass and kill wildlife exhibiting in many cases a surplus killing
behaviour (e.g., Iverson 1978, Kruuk & Snell 1981, Manor & Saltz 2004, Banks
& Bryan 2007), compete with wildlife (e.g., Boitani 1983, Butler & du Toit 2002,
Butler et al. 2004, Vanak et al. 2009) and spread disease like rabies, parvovirus or
canine distemper (e.g., Sillero-Zubiri et al. 1996, Cleaveland et al. 2000, Fiorello
et al. 2004, Fiorello et al. 2006, Vanak & Gompper 2009). Moreover, dogs such
as mid-sized canids, can also exert a top-down influence on smaller carnivores
through interference competition or intraguild predation (e.g., Glen & Dickman
2005, Mitchell & Banks 2005, Vanak & Gompper 2009). Studying where domestic
dogs came from in protected areas is needed in order to manage them and prevent
their potential impacts on native fauna. Dogs at protected areas may exhibit varying
162
Chapter 4
levels of human reliance, from domestic dogs whose needs are satisfied directly
or indirectly by people (i.e., owned dogs, urban or rural free-roaming dogs and/
or village dogs (Vanak & Gompper 2009)) and spend time sporadically in natural
protected areas, to feral dogs living and reproducing freely in protected areas. The
effects of dogs on wildlife may therefore depend on their nature (i.e., domestic dogs
vs. feral dogs), on where they are found and on the factors controlling their numbers
and space use.
densities in areas close to human residences or places with a high density of houses
(e.g., Odell & Knight 2001, Ordeana et al. 2010) as well as in areas near natural
reserve borders with agriculture, where rural human residences are nearby. In
contrast, they may exhibit lower densities in areas contiguous to large tracts of
native forest, which may be acting as a buffer to the entrance of dogs (Srbek-Araujo
& Chiarello 2008).
patch edges (Torres & Prado 2010). Hence, if we consider protected areas as large
patches, presence and therefore direct negative effects of domestic dogs on native
fauna (i.e., predation and/or competition) may decrease from the anthropogenic
matrix to the protected area interior reaching their maximum at the borders. This
could strengthen the negative anthropogenic edge effect associated with these
artificial border areas (Woodroffe & Ginsberg 1998, Revilla et al. 2001) diminishing
therefore the reserves effectiveness in conserving wildlife.
Feral dogs are meanwhile completely wild and independent of human-derived
materials as food sources (Nesbitt 1975, Green & Gipson 1994). They depend
almost exclusively on wild-caught food (e.g., Marsack & Greg 1990, Glen &
163
Chapter 4
Dickman 2005, Mitchell & Banks 2005, Campos et al. 2007, Glen & Dickman
2008) and might not exhibit any human association. The direct threats of feral dogs
to wildlife may therefore spread throughout entire protected areas.
In this context we studied the patterns of detection of dog tracks and the
Our main research goals were to answer: (1) where are dogs present at
DNP? And (2) what factors predict dog presence? A priori, we hypothesised that
dogs using DNP might come either from a domestic dog population formed by
individuals that incur occasionally from the surrounding matrix, or from a feral
dog population living and reproducing freely. In the first case, we would expect
that dogs are heavily dependent on humans and are more abundant at the edges
of the protected area close to human settlements than far away from these edges.
In the second case, we would expect that dogs avoid human association, are more
evenly distributed throughout the protected area, and their presence or abundance
related to environmental features describing habitat suitability or potential wild
food availability. DNP is optimal for the design of a study of this type since 1) part
of the protected area is surrounded by human settlements and others are not, and 2)
it is large enough to potentially hold a feral dog population in its interior.
164
Chapter 4
METHODS
Study area
The study was carried out at Doana National Park (DNP), a flat sandy area
located in south-western Spain (550 Km2, 379N, 626W) at sea level. We defined
the anthropogenic DNP edge as the northern and western edges in close contact
with human settlements, crop fields and a high-traffic highway, and the natural DNP
edge as the southern edge limiting with the Atlantic Ocean and the eastern edge
limiting with the Guadalquivir River through a 27,000 ha marshland area (Fig. 1).
The climate is Mediterranean sub-humid, with mild, wet winters, and hot,
dry summers, and an average annual rainfall around 550 mm. There are three main
biotopes in the park: scrubland, dunes, and marsh (Valverde 1958). The dune area
is situated at the western border of the protected area limited by the Atlantic Ocean
and the marsh area at the northern and eastern borders limited by the Guadalquivir
165
Chapter 4
Figure 1. Study area defined by the Doana National Park site and the surrounding anthropogenic
area. Dog tracks detected during track counts in 22 km2 cell grids between November 2007 and
April 2009 are shown below in detail.
166
Chapter 4
Among larger mammals wild boar (Sus scrofa), red deer (Cervus elaphus)
and fallow deer (Dama dama) are frequent. Wild carnivores include red fox (Vulpes
vulpes), Eurasian badger (Meles meles), Egyptian mongoose (Herpestes ichneumon),
common genet (Genetta genetta), least weasel (Mustela nivalis), European polecat
(Mustela putorius), Eurasian otter (Lutra lutra), wild cat (Felis silvestris) and
Iberian lynx (Lynx pardinus). 14 small-medium sized mammal species have been
recorded in DNP, as well as 397 bird species, approximately half of which are
breeding in the park.
Field methods
on sandy paths at 69 2x2 km2 grid cells located within the entire scrubland and dune
areas of the protected area during the wet season of 2007-08 and 2008-09.
We sampled for dog tracks in each square by slowly walking (ca. 1.5 km/h)
at least 3 km along available pathways (i.e., sandy roads and firebreaks). Once a
track was detected, we georeferenced it using a GPS. We re-sampled the same path
(leaving at least seven days between samplings) a second time in a few squares until
completing 3 km if during the first sampling there were insufficient available paths
167
Chapter 4
within the square to achieve this distance. We always carried out surveys at least
three days after any rainfall.
was assessed by sampling potential prey availability and general habitat structure.
Feral dogs are habitat generalists and opportunistic foragers depending almost
exclusively on a wide variety of wild-caught food (e.g., Boitani et al. 1995, Marsack
& Greg 1990). Potential prey availability was estimated by counting tracks of small
mammals, European rabbits (Oryctolagus cuniculus), red partridges (Alectoris rufa),
domestic cows (Bos taurus) and horses (Equus caballus) and wild ungulates such
as the fallow deer (Dama dama), the red deer (Cervus elaphus) and the wild boar
(Sus scrofa). Prey such as small mammals, rabbits, partridges and young of wild
ungulates might be hunted by feral dogs, but adults of many species are consumed
as carrion (e.g., Sillero-Zubiri & Macdonald 1997, Butler et al. 2004, Aiyadurai &
Jhala 2006). Prey species were surveyed by walking between 7 and 10 25 m-long
transects of approximately 1.7 m wide (the width of a four-wheel-drive car) and
separated by at least 300 m within each 2x2 km grid. During the first year, prey
transects were carried out throughout the wet season, when tracks from dogs were
surveyed, but during the second year, we concentrated samplings within the month
of April to avoid possible intermonthly variations in abundance for some species
(e.g., see Kufner 1986, Palomares et al. 2001 for small mammals and European
rabbits, respectively.
Mediterranean scrubland), general habitat structure was recorded for the first year.
We estimated visually the percentage of open ground cover, and the percentage
and modal height of three vegetation categories: short shrub (xerophytic species
168
Chapter 4
such as Halimium sp. and Cistus sp), tall shrub (Erica sp., Juniperus phoenica and
Pistacia lentiscus shrubs) and trees. The estimation of these variables related to
habitat structure were carried out in a circle of 15 m radius around the sampling
point every 300 m on transects walked for dog and prey tracks. For each square
sampled, we averaged the value obtained at the vegetation sampling points.
Data analyses
explaining the detection of dog tracks within each 2x2 km grid at DNP using
generalised linear models with a binomial error distribution and a logit link function
(logistic procedure in SAS 9.2 (SAS Inst. Inc., Cary, NC)). Non-biological factors
(i.e. methodological and climatic variables) have been previously reported as
potentially affecting results of track censuses (Soto et al. 2012); therefore we also
incorporated in the model fitting these factors to control for their potential effect.
Each grid cell was associated with a set of habitat variables as vegetation
type (dunes (> 60% of open groundcover on average inside the grid), pine forest (>
60% of pine vegetation on average) and Mediterranean shrub (> 60% of shrub (short
or tall) vegetation on average) and prey abundance (kilometric abundance index
of total prey) and with variables describing their location in DNP by calculating
Euclidean distance from the grid cell centre to every infrastructure.
Chapter 4
respectively) (Obs), relative humidity (%) on census day (Hum), days since last rain
(Rain), year (Year) and the maximum temperature (C) calculated as the average of
the maximum temperature on the census day and the maximum temperature two
consecutive days before the census day (Temp). Climatic data was obtained from a
meteorological station located inside DNP (Latitude: 37 118, Longitude: 6 33
17) http://icts.ebd.csic.es.
Initially, the correlation among variables was explored using Kendalls tau
statistics, in order to eliminate highly correlated variables (tau > 0.4) and among
them; we retained the more ecologically meaningful ones.
We used the Akaike Information Criterion (AIC) corrected for a small sample
size (AICc) and the difference in AICc between each model and the model with the
lowest AICc (AICc) to rank the models according to their capacity to describe
the data parsimoniously (Burnharm & Anderson 2002). The model with the lowest
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Chapter 4
AICc and those with AICc 2 were considered to be supported. AICc values
were used to compute Akaikes weights (i), which is the weight of evidence that a
model is the best approximating model given the model set (Burnham & Anderson
2002) and is defined as
i =exp(-1/2i)/R r=1exp(-1/2r).
determined as the sum of the i across all models where j occurs. Larger j values
indicate a higher relative importance of variable j compared to other variables.
For each hypothesis we used data from both years and we began by fitting all
variables included and then successively removing the terms that decreased the
AIC the most (Crawley 2002).
using the nonparametric estimate of the area under the curve (AUC) of receiveroperating characteristic plots (Hosmer & Lemeshow 2000). AUC indices range
from 0.5 to 1, with ranges from 0.5 to 0.7 indicating poor discrimination, from 0.7
to 0.8 acceptable discrimination, from 0.8 to 0.9 good discrimination, and > 0.9
outstanding discrimination. The AUC measure from the ROC curve is considered
useful for comparing the performance of the detection of dog tracks-absence model
in a threshold-independent fashion (Fielding & Bell 1997).
171
Chapter 4
RESULTS
A total of 471 km was walked and 72 dog tracks were found during surveys
(Fig. 1). We detected dog tracks at 16 and 12 grid cells surveyed in each study year,
respectively. Four anthropogenic variables were strongly correlated. In particular,
distance to the anthropogenic edge of the National Park (D_ANT), distance to
nearest village (D_VIL), distance to nearest paved road (D_ROAD) and distance
to the natural edge of DNP (D_NAT). Analyses were focused on distance to the
anthropogenic edge of DNP and distance to the natural edge of DNP.
biological factors included humidity (Hum) and days since last rain (Rain). Humidity
was positive but non-significant (odds ratio = 1.035, 2 = 2.159, P=0.142) whereas
days since last rain was negatively and significantly correlated with detection of
dog tracks (odds ratio = 0.931, 2 = 4.286, P=0.035). Both predictors were therefore
included as covariables in further analyses.
and prey variables showed that the a priori hypothesis best adjusted to data only
included human-related predictors.
The best model describing the detection of dog tracks at DNP after adjusting
for detection probability variables in the null model included the distance to the
anthropogenic edge of DNP (explaining 27.6 % of the deviance); the next model
included the distance to the anthropogenic edge of DNP and the distance to the
natural edge of DNP (models 1 and 3, Table 1).
Chapter 4
0.06(0.04)Rain
where P is the probability of dog occurrence; values within parentheses are standard
errors.
as the sum of the i across all models where the variable occurred was j=0.999.
The discriminating ability of the top model was AUC = 0.802 (P < 0.0001).
DISCUSSION
Results show that the detection of dog tracks at DNP was associated with
distance from the park boundary with human presence, a synthetic indicator of
Table 1. Selection results from logistic regression models investigating the effects of anthropogenic,
habitat and a combination of all variables on detection of dog tracks at DNP. For the top models, we
report the small sample-size-adjusted Akaikes information criteria (AICc), the difference in AICc
between each model with the lowest AICc (AICc) and the AICc weight (wi).
Model code
Deviance
139.210
AICc
121.107
AICc
0. Null model
wi
12.939
0.000
100.170
108.168
0.000
0.276
97.470
109.469
1.301
0.144
98.240
108.242
0.074
0.266
99.750
109.747
1.579
0.125
112.970
120.971
12.803
0.000
106.720
122.265
14.097
0.000
110.900
122.897
14.729
0.000
97.950
111.953
3.785
0.042
98.850
110.853
2.685
0.072
100.810
110.811
2.643
0.074
Anthropogenic
Habitat
Global
173
Chapter 4
human influence that captures the effect of distance to nearest village and nearest
paved road.
We found a high number of dog records near the reserves borders where
rural and suburban households were closer and we were unable to detect signs of
dogs far away from these anthropogenic DNP edges in spite of our large census
effort. Additionally, dog tracks detectability did not seem to be related to the
environmental variability of DNP such as vegetation type or prey availability.
Chapter 4
hypothesis that dogs using DNP come from a domestic dog population formed by
individuals that arrive sporadically from the surrounding matrix (i.e., owned dogs,
urban/rural free-roaming dogs or village dogs (Vanak & Gompper 2009)) and not
from a feral dog population living and reproducing freely. The lack of association
between detection of dog tracks and wildlife food resources possibly also reveals
the dependence of dogs on human-derived materials, which is typical for the vast
majority of dog populations for which diet has been studied (Atickem 2003, Butler
et al. 2004, Vanak 2008, Vanak & Gompper 2009).
some feral dogs use human garbage for food (Green & Gipson 1994). A feral dog
population established inside DNP could therefore use the edge of the protected
area to access human subsidies. Nevertheless, the primary feature that distinguishes
feral from domestic dogs is the degree of reliance on humans, so if dogs using
DNP come from a feral dog population living and reproducing freely but accessing
human subsidies for food, we would expect dog detectability to be dependent
on habitat suitability and/or wild food availability and marginally dependent on
human-related variables.
In contrast with domestic dogs, feral dogs are highly social living in packs
or groups year round in most cases (Daniels & Bekoff 1989, Green & Gipson 1994).
In our study area, we only detected isolated dogs tracks. Additionally, cameratrapping studies conducted inside DNP during the same period detected only six
dogs, all of which were found near human settlements and identified as domestic
animals based on their external physical appearance (personal observ.).
Additionally, the occurrence of dog tracks restricted to the DNP edge and
the low number of tracks detected far away from these reserves edges supports
175
Chapter 4
the idea that the presence of domestic dogs at Doana may be exacerbating the
anthropogenic edge effect associated with its border areas.
Some previous authors have also reported a higher occurrence of domestic dogs
near the edges of natural reserves compared to their interiors (Butler & du Toit
2004, Srbek-Araujo & Chiarello 2008, Lacerda et al. 2009, Marks & Duncan 2009).
In this sense, domestic dogs could be generally considered as human-derived
edge effect at protected areas. The direct threats of domestic dogs to wildlife may
therefore not spread throughout the entire protected area or along the entire edge
length reaching its maximum near the anthropogenic border areas.
fragmented landscapes because they may affect key population parameters, such
as survival and reproduction (Murcia 1995, Noss & Csuti 1997). The peripheries
of reserves thus function as population sinks (Revilla et al. 2001) and the resulting
edge effect can cause the decline or the extinction of protected carnivore populations
(Woodroffe & Ginsberg 1998). The higher occurrence of dogs in these border areas
due to human presence in the surrounding matrix might exacerbate this effect.
dwellings (Vanak & Gompper 2009, Butler & du Toit 2002) and appear to exhibit
a range mainly limited to reserves borders, their daily activity pattern may involve
free-ranging that can bring them into contact with wildlife, especially when their
movements are not confined to a proscribed outdoor area (Butler et al. 2004, Vanak
2008).
Diseases from dogs that affect wild species (e.g., Woodroffe & Ginsberg
1999, Cleaveland et al. 2000) could be transmitted across the border of reserves
worsen therefore the direct edge effects represented by domestic dogs in protected
176
Chapter 4
areas (i.e., the eventual predation and displacement of some species near the park
border). In consequence, domestic dogs maintaining highly virulent, multi-host
pathogens can induce mortality to wild carnivores owing to interspecific disease
transmission between susceptible wild carnivores in the community even when the
wildlife population of interest may never have contact with them.
DNP houses one of the last metapopulation of the most globally endangered
felid species, the Iberian lynx (Lynx pardinus) (Palomares et al. 2011) so dogs
living or spending time inside the area come into contact with this wild endangered
carnivore posing a serious risk for its conservation (Ferreras et al. 1992, Meli et al.
2009 and 2010, Milln et al. 2009a and 2009b).
Domestic dogs are present in large numbers in urban, suburban and rural
areas, so, due to their high numbers, they can have a substantial impact on wildlife,
even when they do not need to hunt to survive.
Chapter 4
In the specific case of DNP, controlling the dog populations is urgent and key
for the wildlife protection in this National Park. The detection of dog tracks at DNP
was dependent on the presence of people and consequently on the resources they
provide such that the potential direct effects of dogs on wildlife may be stronger on
these anthropogenic boundaries. Management of domestic dogs in protected areas
where feral dogs populations are not established may therefore be focused at borders
and neighbourhoods close to human dwellings. We suggest that control measures
must include the restriction of domestic dogs free-roaming activity through local
public awareness focusing on responsible ownership and biodiversity conservation,
the removal of un-owned dogs through systematic campaigns on reserve boundaries
near human settlements, as well as strengthening of pet policies.
ACKNOWLEDGMENTS
This study was financed by the projects CGL2004-00346/BOS (Spanish Ministry of Education and
Science) and 17/2005 (Spanish Ministry of the Environment; National Parks Research Programme),
and sponsored by Land-Rover Espaa S.A. CS received a JAE predoctoral grant from CSIC
(Spanish National Research Council). We are especially thankful to J.C. Rivilla and S. Desnia for
their assistance during fieldwork, to N. Fernndez for his useful inputs for analysis of the data and
valuable suggestions, and to C. Dickman and P. Ferreras for their comments on earlier versions of
the manuscript.
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ANNEX II
Domestic dogs sighted within Doana National Park and photo-trapped during the study period.
184
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185
186
Chapter 5
ABSTRACT
The wildcat Felis silvestris is a protected species in Europe but the lack
187
Chapter 5
RESUMEN
188
Chapter 5
INTRODUCTION
The European wildcat Felis silvestris is one of few wild felids in Europe,
In order to develop action plans for the conservation of the wildcat and
Chapter 5
wildlife have been protected for more than five decades (that is, a priori low human
pressure and persecution), and where habitat and prey availability should be
potentially favourable to wildcat.
abundance of the species has been very low long ago. Fifty years ago Valverde
(1967) considered the species as low abundant in the area. More recent available
information refers to a litter of three kittens found within the Doana National
Park in 1997 when looking for Iberian lynx litters (N. Fernndez, pers. comm.),
three individuals captured in 1999, 30 pictures of wildcats photo-trapped between
2000 and 2007 and a total of 52 direct sightings of the apparently wildcats between
1989 and 2000 collected by all personal working in the Doana National Park
(Centro International de Estudios y Convenciones Ecolgicas y Medioambientales
(CIECEM)) (see Annex III). Note that for the same period, other carnivores such as
foxes or lynxes were sighted in 1211 and 669 occasions, respectively.
The main objective of this study was to assess the wildcats current status
in the Doana area and to discuss about the factors explaining its abundance.
Given how little is known about wildcat biology and the vulnerability of Iberian
populations, we aimed to provide baseline data to promote further research and
conservation of the wildcat in southern Spain.
METHODS
Study area
the south and west by the Atlantic Ocean and to the east by the Guadalquivir River
mouth. The area is flat and mostly near sea level; soils are predominantly sandy and
190
Chapter 5
of marine origin. The climate is Mediterranean subhumid and has marked seasons:
winters are mild and wet, and summers are hot and dry. Mean annual precipitation is
approximately 550 mm. Its situation between the Atlantic Ocean and Mediterranean
Sea and in southern Europe promotes some of the highest biological diversity on
the continent, particularly of vertebrate animals and vascular plants (FernndezDelgado 1997). There are three main biotopes in the park: scrubland, dunes, and
marsh (Valverde 1958). The dune area is situated at the western border of the
protected area where it is limited by the Atlantic Ocean and the marsh area lies at the
northern and eastern borders limited by the Guadalquivir River. The Mediterranean
scrubland represents approximately half of the National Park surface area and
is mainly characterised by heterogeneous patches of xerophytic species such as
Halimium sp. and Cistus sp., and hydrophytic ones such as Erica sp., with some
patches of Juniperus phoenica and Pistacia lentiscus shrubs. Interspersed among
the scrubland are scattered cork oak trees (Quercus suber) and wild olive trees (Olea
europea), and a few patches of pine Pinus pinea and eucalyptus Eucalyptus sp.
plantations. Vegetation in bare sand dunes is scarce and dune hollows are colonized
by pines Pinus pinea and varied scrubland species.
DNP is fully protected and access to the core area and the dirt-road network
inside DNP is restricted to the park staff and researchers. The northern and western
edges of DNP are in close contact with human settlements, crop fields and a high
use paved road (Fig. 1). These surroundings support intense human activity, with
private large- and medium-sized farms used for agriculture as well as six visitor
centers, hiking and cycling paths, recreation zones and bird observatories in the
nearby area.
191
Chapter 5
Field methods
Chapter 5
Table 1. Variables used to compare means between quadrants with no evidence of cat presence,
quadrants where wildcats were photographed and quadrants where cat tracks were detected during
track censuses by non-parametric tests.
Variable
Vegetation
Code
Definition
Units
Short shrub
Short shrub
height
Tall shrub
%S
S_h
%B
B_h
Trees
%T
Tree height
T_h
Landscape
Distance to water DW
Distance to La
Vera
DV
Ecotones between
pastureland and
eBP
scrubland
m/ha
Prey availability
Rabbits
Ra
Small mammals
SM
Total prey
Tot
Human disturbance
Distance to
antropic edge
DH
Predators occurrence
Kilometre
abundance index
of domestic dogs
Kilometre
abundance index
of lynxes
m/ha
tracks/km
* calculated by averaging values obtained at the different sampling points within quadrants
193
Chapter 5
where cat tracks were detected during track censuses. Camera traps were set
in the borders of car tracks and firebreaks, or in the edges of patches of dense
Mediterranean shrub with pasturelands (habitat potentially favourable for wildcats
(Lozano et al. 2003). In order to maximize the detection of wildcats we used scent
lures of valerian, catnip and canned sardines sprayed on a piece of cotton attached
to a wooden stake at a 3050 cm height, wet fish (sardines), live prey (reported as
the most efficient lure for sampling some felid species (Guilt et al. 2010, Garrote et
al. 2012)) such as rock pigeons (Columba livia) and rabbits (Oryctolagus cuniculus)
in wire cages inaccessible to wildcats, as well as no attractants. Cages of live prey
were approximately 100 x 50 x 50 cm and supplied with ample food and water
at least twice a week. On average, we set up 15 digital camera traps with passive
infrared motion sensors and automatic flash (Cuddeback Digital Scouting Model
Expert) per quadrant. Cameras were placed 20 cm above ground, at a distance
of 2-4 m from the lure with 300-400 m between them. We set camera traps with a
delay time of 1 min between successive photos, and checked at least twice per week
to replace attractant lures and twice a month for battery replacement.
Differentiation between wildcats and domestic cats was based on the general
physical appearance and on the pelage pattern of the individuals. Studies on the
European wildcat have indicated that camera trapping can be used to some extent
to determine the presence and abundance of this species and that individuals are
identifiable based on their morphology (Ragni and Possenti 1996, Monterroso et al.
2005, Anile et al. 2007, Karanth et al. 2004). We considered photos as independent
events when taken more than 4 hours apart for the same individuals or if different
individuals could be identified (OBrien et al. 2003).
194
Chapter 5
Figure 1. Locations of cat tracks detected during track censuses in 2 x 2 km2 quadrants during
2007-2009 (a); locations of the 166 trapping stations set during 2008-2010 (b), and the camera trap
stations that provided pictures of wildcats and domestic cats, respectively, as well as the home range
of the wildcat radio-tracked in DNP (c) are shown.
195
Chapter 5
project that aimed to study the effectiveness of red fox control actions within DNP.
The captured animal was chemically immobilized with a 0.75 ml dose (100mg*ml-1)
of tiletamine- zolazepam (Zoletil, Virbac, Spain), measured, weighted, checked
for any sanitary disorders and sexed. Genetic analysis from blood samples collected
revealed that the individual was pure or without any indication of parental
domestic heritage (Alves, P.C., pers. comm.). After handling, the individual was
maintained in the dark and returned to the capture location for release after complete
recovery of reflexes (13 h). The individual was fitted with a radio-collar (Wildlife
Materials, Inc., Carbondale, Illinois, USA), radio-tracked between December 2009
and March 2010 and located on average twice per week between 9:00 am and 2:00
pm. We used triangulation to determine the position of the individual (White and
Garrott 1990) and the minimum convex polygon method to estimate the home
range size based on all available locations (Mohr 1947, White and Garrott 1990)
196
Chapter 5
with Hawths Analysis Tools (Beyer 2004) in ArcGIS (ESRI, Redlands, CA). We
determine the main vegetation types included within the individual home range
from a 1:10 000 fine-scale vegetation map for the year 1996-2006 obtained from the
Sistema de Informacin Ambiental de Andaluca (http://www.juntadeandalucia.es/
medioambiente/site/rediam/)
RESULTS
censused in each year, respectively (Fig. 1). We set cameras at 166 different points
in 25 quadrants. Camera effort was 5761 trap-days with an average of 24.4 day/
cameras (SD= 8.02, range=1-33). We obtained a total of 2173 photographs in
which we identified mammals (n = 2050) or birds (n = 123). The red fox was the
most common species photographed followed by the Egyptian mongoose and the
common genet (Table 2). Thirty pictures of cats were taken at 12 of the 166 points
where we set camera trap stations. Twenty-eight of these pictures were of wildcats
and two pictures were of domestic cats. These camera traps were baited with
pigeons (n = 8) and wet sardines (n = 6) (Table 2). Wildcats were photographed
between 20:00 and 07:00 hours. Six different wildcats were identified (Fig. 2) from
12 camera trap records (Table 3) and all were photographed only once. None of
the individuals could be sexed, nor could the presence of more than one individual
per trap-site be ascertained. Two-hundred six camera-trap days were required on
average to document the presence of an individual.
approximately 24 km2 (Fig. 1). More than 60% of the individual home range
included areas of Mediterranean short scrubland (i.e., species such as Halimium sp.
197
Chapter 5
and Cistus sp.) and approximately 18% pine woodlands with understorey vegetation
(i.e., short shrubs and tall shrubs such as Erica sp., Juniperus phoenica and Pistacia
lentiscus).
quadrants with only cat tracks and quadrants with no evidence of cat presence for the
Table 2. Total number and percentage of pictures taken during camera-trapping surveys at DNP.
Specie
Felis silvestris
28
1.3
Felis catus
0.1
Lynx pardinus
0.1
Genetta genetta
60
2.8
Herpestes ichneumon
152
7.0
26
1.2
Meles meles
Vulpes vulpes
1284 59.1
Canis familiaris
Others*
0.3
611
28.1
Table 3. UTM coordinates (29S) of the camera-trap positions where wildcat were photographed.
The period of time the camera was active and the number of wildcat pictures taken and baits used
are also provided.
Trap station
198
Pictures
Trap-days
Bait
185585
4106480
22
Pigeon
186451
4106914
22
Pigeon
185780
4100419
10
Pigeon
186990
4099547
29
Wet sardine
184710
4101010
29
Wet sardine
185476
4107561
29
Wet sardine
185967
4108394
29
Pigeon
185895
4108264
29
Wet sardine
186503
4109315
29
Wet sardine
10
183961
4117788
20
Pigeon
11
190451
4105106
22
Pigeon
12
189275
4105888
30
Pigeon
Chapter 5
Figure 2. Individual wildcats distinguished on the basis of their external aspect in the Doana
National Park between 2008 and 2010.
variable distance to the anthropic edge of the protected area (Table 4). Mann-Whitney
U post-hoc tests revealed that quadrants where wildcats were photographed were
closer to the anthropic edge of the protected area than quadrants with no evidence of
cats and than quadrants where tracks were detected (Table 5). No differences were
found for any of the remaining variables analysed.
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DISCUSSION
wildcats throughout the DNP and reveals a priori surprising low abundance of the
species in the protected area. Wildcats like autochthonous Mediterranean scrubland
areas with scrubpastureland mosaics (Lozano et al. 2003), and may rely for
feeding on rabbits (Gil-Snchez et al. 1999, Lozano et al. 2006)) or small mammals
(Sarmento 1996, Molen and Gil-Snchez 2003, Carvalho and Gomes 2004). These
habitats and prey are common in many parts of the DNP, so one would expect that
wildcats were more abundant in the area than what we have found. Furthermore,
protection of the DNP for more than 5 decades has provided a safe place for wildcats.
Therefore, DNP should hold one of the largest wildcat populations in south western
Spain and be one of the most important areas for conservation of the species.
Wildcat scarcity in DNP might be due to several factors. First, during the
last decades wild rabbit population has decreased everywhere and in the DNP due to
diseases such as myxomatosis and Rabbit Hemorrhagic Disease (RHD) (Thompson
and King 1994, Villafuerte et al. 1995) and to changes in scrubland management
(Moreno and Villafuerte 1995). Abundance of the wildcat population might have
diminished for this reason. In fact, the large home range size of the radio-tracked
individual in our study area suggests low food abundance. However, there are some
areas where rabbits are abundant within the park, and wildcats can also consume
other alternative prey species such as small mammals (Lozano et al. 2003, Malo et
al. 2004), so this reason not fully explain the low abundance of wildcats in the area.
Secondly, the isolation of DNP from the nearest natural areas (Sierra Morena and
Cdiz) due to human settlements, widespread field crops or the Guadalquivir River
may also contribute to the low abundance of the species. Wildcats disappeared
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Chapter 5
from many regions across its range and reached minimum levels at the beginning
of the 20th century (McOrist and Kitchener 1994). The recovery of the species
in several places was possible in the 1990s when anthropic pressure on wildcat
populations and their habitat was reduced (e.g., Parent 1975, Easterbee et al. 1991)
but the isolation and fragmented distribution of the species in Doana may have
prevented the recovery in spite of the reduction in potential threats. Additionally,
Table 4. Results of the Kruskal Wallis Test to test for differences in the means of several variables
between (0) quadrants with no evidence of cat presence, (1) quadrants where wildcats were
photographed and (2) quadrants where cat tracks were detected. Significant variables are represented
in bold.
Mean rank
Variable
Chi-square P-value
Vegetation
%SB
2.574
0.276
%S
1.265
0.531
S_h
4.229
0.121
%B
2.590
0.274
B_h
0.796
0.672
%T
0.796
0.672
T_h
0.193
0.908
6.637
0.036
eBP
3.065
0.216
DW
2.257
0.324
DV
5.168
0.075
KAId
2.437
0.296
KAIl
1.731
0.421
0.606
0.739
SM
4.963
0.084
Tot
0.312
0.856
Human disturbance
DH
Landscape
Predators
Prey
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Chapter 5
an increased mortality rate over time due to potential disease transmission from
domestic cats (e.g., McOrist et al. 1991) could be another causal factor explaining
wildcat scarcity in Doana. The domestic cat population in the Doana area might
have increased over the last decades due to the urbanization of the surrounding
neighbourhoods of the protected area. Hence, as in other natural areas (e.g., Ferreira
et al. 2011), feral and/or domestic may spend time sporadically within DNP during
their free-ranging activity potentially transmitting diseases to their wild relatives.
In fact our results confirm the presence of domestic cats within the DNP close to
human settlements (Fig. 1, Table 4). Finally, although our results do not confirm
that the relative abundance of the Iberian lynx can negatively affect the detection
of wildcats, interspecific interactions between wildcats and the Iberian lynx may
have also led to the competitive exclusion of the species as carnivores persisting
at low population densities have been suggested to experience an increase in the
effect ofintraguildpredation (Creel and Creel 1998, Creel 2001, Creel et al. 2001,
Creel and Creel 2002). The European wildcat and the Iberian lynx may potentially
overlap in habitat use of Mediterranean scrubland but intraguild interaction with
the Iberian lynx may have resulted in habitat partitioning in that wildcats will avoid
habitat patches of high lynx densities to the detriment of its own success in prey
acquisition and access to the most suitable habitats. In turn, wildcats may have
been forced to enlarge their home ranges to continue hunting and may even roam
in human-occupied areas increasing their mortality risk. In fact our results revealed
that wildcats were detected more frequently near the anthropic edge of the area
suggesting that individuals could be ranging outside of DNP. Nevertheless, in spite
of the continued decrease of the Iberian lynx population in the Doana area in
recent decades (Palomares et al. 2012) wildcats have not increased its abundance
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Chapter 5
in the area confirming the hypothesis that the low occurrence of the species in DNP
can not be only attributed to Iberian lynx decrease.
In summary, although the DNP is optimal for a large wildcat population the
potential threats explained above may shed light on the low occurrence of the species
in the area. Nevertheless, many threats to the species may remain unidentified in
Doana, so although its low population density makes field studies and direct
observation difficult, more research and detailed information on occurrence as well
as on wildcat habitat requirements based on radio-tracking efforts are necessary to
provide guidelines for management and conservation of the species in the area.
ACKNOWLEDGEMENTS
Education and Science) and 17/2005 (Spanish Ministry of the Environment; National Parks Research
Program). Land-Rover Espaa lent us two vehicles for this work. We are very grateful especially to
J.C. Rivilla and S. Desnica for assistance during fieldwork. C. Soto was also supported by a JAE
Predoc grant from the CSIC.
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ANNEX III
b), wildcat radio-tracked in the study area between December 2009 and March
2010 (c) and wildcat photographed in March 2011 by the personal working in the
Doana National Park (Centro International de Estudios y Convenciones Ecolgicas
y Medioambientales (CIECEM)) (d). Credit pictures: CIECEM (a, b and d) and C.
Soto (c).
208
Conclusiones
209
Conclusiones
Conclusiones
5. Modelos de uso del hbitat a escala fina indicaron que la presencia de lince
estuvo asociada a zonas con una alta cobertura de matorral alto y densidad
de ecotonos entre matorral alto y pastizal, mientras que la presencia de
ginetas se asoci a zonas de pinares con vegetacin densa de sotobosque
as como con una alta abundancia de micromamferos. Los zorros, tejones
y meloncillos mostraron un patrn de seleccin de hbitat menos marcado,
siendo la abundancia de presas totales y de conejo de monte, o la cobertura
de matorral alto algunos de los predictores de su abundancia y distribucin.
Conclusiones
10. Para un control efectivo de perros que usan el interior del Parque Nacional
debera actuarse sobre los ncleos urbanos circundantes, as como intensificar
las medidas de control en los lmites del Parque Nacional.
12. La baja abundancia de gato monts en el Parque Nacional podra explicarse por uno
o ms factores tales como el descenso de las poblaciones de conejo, el aislamiento
de la poblacin, una posible alta incidencia de enfermedades debido al contacto
con gatos domsticos, y la competencia por interferencia con el lince ibrico.
212