KORA Bericht Nr. 13 e
Juni 2002
ISSN 1422-5123
Spatial and Temporal Distribution of Coat Patterns
of Eurasian Lynx (Lynx lynx) in two Re-introduced
Populations in Switzerland
Karin Thüler
KORA
Koordinierte Forschungsprojekte zur Erhaltung und zum Management der Raubtiere in der Schweiz.
Coordinated research projects for the conservation and management of carnivores in Switzerland.
Projets de recherches coordonnés pour la conservation et la gestion des carnivores en Suisse.
KORA, Thunstrasse 31, CH-3074 Muri. Tel +41-31-951 70 40, Fax +41-31-951 90 40, Email: info@kora.ch, http://www.kora.unibe.ch
2
KORA Bericht Nr. 13
KORA Bericht Nr. 13 e:
Spatial and Temporal Distribution of Coat Patterns of Eurasian Lynx (Lynx lynx) in two reintroduced Populations in Switzerland
Autoren
Auteurs
Authors
Karin Thüler
Amselweg 2
3322 Schönbühl
karinthueler@bluemail.ch
Bearbeitung
Adaptation
Editorial
Damiano Torriani (Karten)
Adrian Siegenthaler (Layout)
Bezugsquelle
Source
Source
Kora, Thunstrasse 31, CH-3074 Muri
T +41 31 951 70 40 / F +41 31 951 90 40
info@kora.ch
Titelfoto
Photo de la page de titre
Front cover picture
Different coat pattern types of Eurasian lynx
Anzahl Seiten/ Pages: 35
ISSN 1422-5123
©KORA Juni 2002
Juni 2002
3
Spatial and Temporal Distribution of Coat Patterns of
Eurasian Lynx (Lynx lynx) in two reintroduced
Populations in Switzerland
Diploma Thesis
Faculty of Science of the University of Bern
presented by
Karin Thüler 2001
Supervisor of the work:
Prof. Dr. Marcel Güntert, Natural History Museum Bern
4
KORA Bericht Nr. 13
Acknowledgements
I am grateful to the following persons who supported me during this study: Special thank to:
•
Dr. C. Breitenmoser-Würsten and Dr. U. Breitenmoser.
•
My supervisor Prof. Dr. M. Güntert who made-possible this diploma-thesis and for their comments and contributions.
•
J. Laass for helping with database and pictures.
•
E. Grégorova, Zoo Bojnice, Slovakia, C. Stanisa, Stara cerkev, Slovenia, D. Huber, Veterinary Faculty, University of Zagreb, Croatia and Aloizije Frkovic, Zagreb, Croatia for sending information about native lynx.
•
The Museum of Natural History in Berne, Dr. P. Lüps and M. Troxler for showing and helping me with skins,
L. Schäublin for helping me with take pictures, V. Andres for helping me with literature.
the Natural History Museum Basel, Dr. R. Winkler, Museum Gruérien Bulle, D. Buchs, Bündner Natural Museum Chur, Mr. Bardill, Natural Museum Thurgau, H. Geisser, Natural History Museum Fribourg, M. Beaud,
Museum of Natural History Geneva, A. Keller, Museum of Natural History Lugano, S. Fossati, Natural Museum Lucerne, R. Heim, Museum of Natural History Neuchatel, Mr. Zimmerli, Natural Museum Olten, Dr. P.
Flückiger and R. Leuenberger, Museum of Natural History Sion, J.-C. Praz, Natural Museum Solothurn, M.
Winistörfer, Natural Museum St.Gallen, Dr. J. Barandun, Collection of Natural Science Winterthur, Mr. Fasnacht, Museum of Zoology University of Zurich, Dr. C. Claude.
•
Damiano Torriani for create overview Swiss-maps, colleagues from KORA (Coordinated research projects for
the conservation and management of carnivores in Switzerland, Muri b. Berne, Switzerland), Kim Baumann for
correct my English, my family and friends.
Digitale geographische Daten:
Gewässer und politische Grenzen: © BFS GEOSTAT, © Bundesamt für Landestopographie;
Ortschaft und Wald: Vector 200, © Bundesamt für Landestopographie;
Höhenmodell: DHM25: © Bundesamt für Landestopographie; RIMINI: © BFS GEOSTAT.
Juni 2002
Content
5
Spatial and Temporal Distribution of Coat Patterns of Eurasian Lynx
(Lynx lynx) in two reintroduced Populations in Switzerland
Content
Abstract
6
1. Introduction
6
1.1.
Evolutionary history of coat patterns in felids
6
1.2.
History of lynx in Switzerland and current status
6
2. Material and Methods
7
2.1.
Available materials
7
2.2.
Classification of Coat Patterns
8
2.3.
Evaluation ans statistical analyses
8
3. Results
8
3.1.
Identification of coat pattern types
8
3.2.
Assignment of coat pattern types
9
3.3.
Distribution of the different coat pattern types
9
3.3.1. Spatial distribution of coat patterns in Switzerland
10
3.3.2. Temporal changes in pattern type frequencies
10
4. Discussion
20
4.1.
Description of coat pattern types
20
4.2.
Comparisons with other descriptions of coat patterns in lynx
20
4.3.
Spatial and temporal distribution of coat patterns in Switzerland
21
5. References
24
6. Appendices
25
I
Review on Coat Patterns.
25
II
Form with all necessary information needed per individual and guidelines to fill out the form
31
III
Spots-Counting rules
32
IV
Database of Lynx - Pictures
33
6
Abstract/Introduction
KORA Bericht Nr. 13
Abstract
Lynx had become extinct throughout most of Central
and Western Europe at the end of the 19th century. This
was also the case for Switzerland. However, in 1971
the Eurasian lynx was reintroduced into the Swiss
Alps. Further re-introduction programs followed in the
Swiss Jura Mts, Slovenia and Croatia. All animals released came from the same source population of the
Carpathian Mts in Slovakia, and some of them have
been closely related. As small, isolated populations are
theoretically vulnerable to genetic drift, where alleles
with low frequency are likely to disappear from the
population gene pool, a change in the occurrence of
different coat patterns can be an indication for this. In
the recent population of the Swiss Alps a temporal
change of the occurrence of different coat pattern types
was found. Another possible indication is the loss of
the non-spotted type in the Jura Mts population.
In this study, a classification system for coat patterns in lynx was developed and five different coat patterns were defined: large spots, small spots, without
spots, rosettes and small spots with rudimentary rosettes. The frequency of occurrence of these coat patterns was then compared between the two reintroduced populations in Switzerland and the source
population in the Carpathian Mts of Slovakia and another re-introduced population in Slovenia/Croatia. Additionally coat patterns of historic lynx from Switzerland were analyzed. The dominant coat pattern type in
the historic Swiss population was non-spotted and in
recent populations large-spotted. Rosettes were found
only recently. There existed a correlation between spatial and temporal distribution of the different coat pattern types.
1. Introduction
1.1. Evolutionary history of coat patterns in felids
Carnivores show a wide variety of pelage colors and a
great diversity of markings on their coat – including
spots, stripes, bands and patches (Ortolani & Caro
1996) (Appendix I, 1.). The current theory of felid coat
pattern evolution proposes that the primitive pattern is
one of relatively large spots which have a tendency to
break down, first by forming a lighter center and then
by breaking up into smaller spots spaced into rosettes
and later individually. At each step in this general decay of the basic pattern, striped patterns may develop
(Weigel 1961). Werdelin & Olsson (1997) coded the
coat patterns of felids into six discrete categories: uniform, flecks, rosettes, vertical stripes, small blotches
and blotches. It is generally accepted that specific color
patterns in mammals are genetically determined but the
actual mechanisms that create the patterns are still unknown.
Lynx belong to the spotted cats. Their coloration
can differ widely. Coats of northern lynx are more
greyish and less spotted than in southern Europe where
the color changes into reddish-brown (Breitenmoser
and Breitenmoser-Würsten 1998). The Eurasian lynx
presents a high phenotypical variability within the species. Grégorova (1997) defined a hypothesis about a
new type of pattern: In principle, it is possible to distinguish two basic types of patterns: spotted (small or
large spots) as the most frequent one and without spots
as the rarest one. The third type is defined as an intermediate type (rosettes). This „new” type has hypothetically evolved from hybridization of the previous two
types. In some felid species (e.g. the pampas cat, Felis
colocolo) there exist a spatial pattern of different coat
patterns (Garcia-Perea 1994) (Appendix I, 2.).
1.2. History of lynx in Switzerland and current
status
Until 1900 lynx was exterminated in Western and
Southern Europe. Some small populations survived in
the Balkan and the Carpathian Mts. In central Switzerland lynx had already disappeared during 17th century.
In the Jura Mts the species survived until the beginning
and in the Alps until the end of the 19th century. Ragni
(1993) analyzed coat patterns of the extinct Alpine
population and recognized a non-spotted type
(concolor) as predominant and concluded that the Alpine lynx represented a differentiated taxon. In 1971,
the Swiss government decided to re-introduce lynx into
Switzerland and four years later the first animals have
been released in the Alps (Breitenmoser 1983, Breitenmoser and Baettig 1992, Haller 1992). The animals
originated from the Carpathian Mts in Slovakia (zoo in
Ostrava). In addition to the official releases in Obwalden, Waadt and Neuenburg there were some illegal releases in the Alps and in the Jura Mts (Breitenmoser
1983, Haller 1992). In the founding population only
very few individuals were released. Additionally, according to Koubek and Cerveny (1996), some of them
were probably closely related (mother-son or siblings).
Today, there are about 75–80 adult and subadult lynx
living in the Swiss Alps. The population in the Jura
Mts and in the Alps are still small and isolated and
therefore theoretically vulnerable to genetic drift,
where alleles with low frequency are likely to disappear from the population gene pool (Griffiths et al.
1996). Beltran and Delibes (1993) found preliminary
evidence for this in the Iberian lynx (Lynx pardinus) in
Coto Doñana, Spain, where the population of approximately 40–50 lynx was isolated since the early 1960s.
A project on genetic variability, analyzing inbreeding
problems, shows that originally three coat patterns
were present in the population, but today no animal exhibits the rare small-spotted pattern (Nowell & Jackson
1996). In Switzerland intensive field studies since 1983
Juni 2002
Introduction / Material and Methods/
had provided extensive material on 87 lynx that were
caught over the years. At each capture, pictures of the
animals were taken. Anecdotal observations on differences in coat patterns over space and time were made
during the last few years.
According to these observations, the following hypotheses were formulated: i) Lynx have clearly distinguishable coat patterns. (ii) The occurrence of these
different types of coat patterns is different between the
two re-introduced populations in Switzerland and has
changed over time since the releases in the early 1970s.
(iii) Genetic drift has occurred since the releases due to
a small founder population, and therefore the occurrence of the different types of coat patterns in the reintroduced populations are different from the source
population in the Carpathian Mts of Slovakia. The aim
of this study therefore was to (i) develop a classification system for the different coat patterns observed and
assign all available lynx individuals to a defined coat
pattern type, (ii) compare the occurrence of these coat
patterns between the two re-introduced lynx populations in Switzerland and the historic population and
(iii) compare the occurrence of these coat patterns in
Switzerland with the occurrence in the source population of Slovakia and another re-introduced population
in Slovenia/Croatia.
7
2. Material and Methods
2.1. Available materials
For this study I collected all available information on
342 individuals of the species Lynx lynx from populations in Switzerland, Slovakia, Croatia and Slovenia
(Table 1). I created a form with all necessary information (location [last origin, place where the lynx were
caught/found/dead, coordinates of caught/found], sex,
age, year of birth, relatives) needed per individual and
sent it to collaborators in Slovakia, Croatia and Slovenia (Appendix II). From museums of natural history in
Switzerland I also received a list with information
about the lynx specimens stored. All this information
was collected in a table where I setup a database
(Microsoft Access) with all available lynx that could
provide information on coat patterns from Switzerland
(Appendix IV).
From the two Swiss populations, pictures from captures, photo traps, video recordings and mortalities
were available. From dead lynx stored in museums, I
first had to take pictures of pelts and mounts. A standardized procedure was applied: Using a grey cloth as
background (only with pelts) and next to every object
on every picture placed a ruler. The camera was a
Nikon FE2 with a 55 macro lens (28-85, 70-210) and a
Table 1. All available specimen.
Material
Type of specimen
Switzerland
Number of specimen
Total 228
Monitored lynx in telemetry project Pictures from captures (various field studies in Switzerland)
87
Photo-traps
Pictures from photo traps (population estimation study, predator identification program)
13
Video
Pictures from videos (predator identification program)
Lynx in museum
Lynx hides and stuffed specimen in museums in Switzerland :
Mounts
Pelts
Animals not yet mounted
43
51
2
Other dead lynx
Pictures from dead lynx (Kora's database of dead lynx), pelts,
mounts
23
Slovakia
9
Total 47
Lynx in Zoo
Pictures of animals
5
Lynx in private
Pictures of pelts
2
Dead lynx
Pictures of pelts
40
Croatia
Lynx pelt of hunter
Total 45
Pictures of pelts and mounts
Slovenia
45
Total 22
Lynx in zoo
Pictures of animals
5
Lynx in wild cat project
Pictures of animals
1
Monitored lynx in telemetry project Pictures of captures
4
Dead lynx
Pictures of pelts
12
Total Objects 342
8
Material and Methods / Results
KORA Bericht Nr. 13
Nikon SB-15 flesh and the film a Kodak Professional
PORTRA, ISO 160vc, 135mm. I took pictures from the
following body/coat areas: left and right, shoulder/
upper arm, forearm, hand, femur/hip, lower leg, foot,
flank and dorsum and the complete left and right side.
The pictures were made in daylight. All pictures were
scanned and saved in a jpg-format. From Collaborators
in Slovakia, Slovenia and Croatia I had already
scanned pictures saved in a jpg-format.
2.2. Classification of Coat Patterns
I intuitively deducted 5 different coat pattern types:
type 1 as “large spots”; type 2 as “small spots”; type 3
as “without spots” and two types of rosettes: type 4a as
“clear rosettes” and type 4b as “spots with rudimentary
rosettes” and chose typical representatives of every
type.
To describe these 5 types I counted the number of
spots on 6 selected areas of the body surface, according
to anatomical criteria: left/right, shoulder, upper arm
and forearm (L/R1); left/right flank and back (L/R2),
left/right femur, hip and lower leg (L/R3) (Figure 1). In
this areas the typical coat pattern type was represented.
The area next to the backbone and the ventral-side
(white coat color) depending on the posture of the individual, was not visual on several pictures. So I did not
use these areas for analyses. Spots were counted after
defined rules. Large or small spots: every identifiable
single spot and spots in V-form were counted as one
spot. Spots in rosettes or in rudimentary rosettes: every
identifiable spot arranged in rosettes/small blotches
was counted as one spot. Spots arranged in groups:
every identifiable spot arranged in strips or other arrangements was counted as one spot. Strips where no
interception was identifiable was counted as one spot.
Coats without spots: no counting of spots (Appendix
III). To verify the measurements and countability of
counting the first 10 objects I counted three times and
calculated the mean value. Also coats with undistinguishable or bleached patterns I counted three times.
As reference to the degree of bleaching of pattern I
used the black tail color and its degree of bleaching. To
distinguish between large and small spots I randomly
measured the diameter of 10 spots (randomly distributed over the body) of 12 individuals, 6 typical large
and 6 typical small spots and calculated the mean
value.
2.3. Evaluation and statistical analyses
The differences in number of spots in the six body/coat
area were tested by Kruskal-Wallis-Test. The Principal-Component-Analysis was used to test differences
between the coat pattern types based on the total number of spots and the existence of rosettes. To compare
data of the five populations (historical and recent Swiss
Alps, Jura Mts, Slovakia, Slovenia/Croatia) I used the
χ-square-test. For tests of normality I used the Kolmogorov-Smirnov-test.
Figure 1. The six selected areas of the body/coat (dotted areas): left/right, shoulder, upper arm and forearm (L1/R1);
left/right flank and back (L2/R2) and left/right femur, hip
and lower leg (L3/R3).
All available lynx were assigned after a created
path-diagram based on the description of the five different coat pattern types. To evaluate the spatial distribution I only used individuals with known location
(last origin, place of caught/found/dead, coordinates of
caught/found) and for temporal analyses individuals
with known year of birth.
3. Results
3.1. Identification of coat pattern types
From 148 individuals, I counted the number of spots in
six different body/coat areas. This method proved to be
useful and applicable. The mean value was calculated
of the same area between the left and right side
([L1+R1]/2, [L2+R2]/2, [L3+R3]/2) and the total number of spots of every individual.
The number of spots per body/coat area of type 1
and 3 were significantly different to all other types
(Mann-Whitney-U-test, P < 0.1, bilateral) and also between each other. Type 2, 4a and 4b were significantly
different to type 1 and 3 but tested against each other
there was no significant difference between the number
of spots per body/coat area (Mann-Whitney-U-test, P >
0.1, bilateral) (Table 2, Figure 2). To distinguish between the five different types another criterion was
needed: the Principal-Component-Analysis assorted the
individuals in groups which showed the intuitive deducted coat pattern types, based on the total number of
spots per individual and the existence of rosettes. The
first principal component explained 82.76 % of variance and the second 6.58 % (Figure 3). The untypical
allocations showed juveniles which have less spots according their smaller bodies. One untypical allocated
type 3 showed an individual which had no spots on
shoulders, flanks and hips but more than the average on
the upper arms, femurs and legs.
Juni 2002
Results
9
Table 2. Statistical analysis of the total number of spots per coat pattern type (n = 148).
Median
25% Quartile
75% Quartile
min
max
Type 1
316.50
294.00
349.75
130.00
430.00
Type 2
528.00
477.00
660.00
309.00
812.00
Type 3
183.50
136.50
219.00
84.00
492.00
Type 4a
630.00
586.00
714.00
335.00
812.00
Type 4b
581.00
520.00
648.00
168.00
804.00
160.00
100.00
50.00
Type 4b L/R3
Type 4b L/R2
Type 4a L/R3
Type 4b L/R1
Type 4a L/R2
Type 3 L/R3
Type 4a L/R1
Type 3 L/R2
Type 3 L/R1
Type 2 L/R3
Type 2 L/R2
Type 2 L/R1
Type 1 L/R3
Type 1 L/R2
0.00
Type 1 L/R1
number of spots per body area
Coat pattern type
coat pattern type and body area
Figure 2. Number of spots per body/coat area of the five
coat pattern types (L1/R1: left/right shoulder, upper arm and
forearm; L2/R2: left/right flank and back; L3/R3: left/right
femur, hip and lower leg, Figure 1).
With all this information the five coat pattern types
were consequently definitively defined (Table 3): type
1 with large spots (Figure 4), type 2 with small spots
(Figure 5), type 3 without spots on shoulders, flanks
and hips but spots on upper arms, femurs and legs
(Figure 6). In type 4 (rosettes) I distinguished two
types: type 4a with clear rosettes or small blotches
(single spots were no more visual and the rosettes
showed circle-forms) (Figure 7) and 4b with small
spots and rudimentary rosettes (single spots were visual within rosettes) (Figure 8). The definition contained
size, form, number, color and distribution of spots in
the following body/coat areas: shoulder / upper arm,
forearm, hand, femur / hip, lower leg, foot, flank and
dorsum.
3.2. Assignment of coat pattern types
To assign all available individuals to one of the five
coat pattern types it was important to look at all body/
coat areas. Each coat pattern type showed variation and
only the analysis of the complete body allowed the assignment to one type. According to the analysis of
numbers of spots and the definition of types I created a
path-diagram (Figure 9). First question: “are spots vis-
ual on flanks or not?” (type 3 can be separated from the
other four types); second question: “are rosettes identifiable or not?” (type 1 and 2 can be separated from
type 4a and 4b). The question decision is between type
1 and 2, concerns the size and number of spots and between type 4a and 4b about the clearness of rosettes.
From a total of 342 individuals, 4 individuals (1.2%)
could not be assigned to one of the five types. Between
females (n = 70) and males (n = 53) (unknown n = 28)
(df = 4, χ2 = 2.6, P > 0.5) and young (juveniles &
subadults, n = 55) and adult (n = 82) (unknown n= 14)
animals (df = 4, χ2 = 1.89, P > 0.5) there was no significant difference. Consequently all individuals were
pooled, including those with unidentifiable age and
sex.
3.3. Distribution of the different coat pattern types
Data from six different populations were available
(historical Swiss population, n = 21; recent population
of the Swiss Alps, n = 128; Jura Mts population, n =
44; Slovakian population, n = 47; Slovenian population, n = 22; Croatian population, n = 45; total individuals: 306, from 32 of total 338 assigned individuals
no location data were available or they originated from
zoo populations, which I did not use). Lynx from Slovenia and Croatia belong to the same re-introduced
population. As the occurrence of different coat patterns
did not differ between the two subsamples ( df = , χ2 =
5.52, P > 0.05), these animals were subsequently
pooled. Rosettes have shown up only in recent populations. However, coat pattern type 4a and 4b have only
reached a considerable proportion in the re-introduced
population in the Alps (46% vs. 18-28%). For further
analyses type 4a and 4b were also pooled to “type 4”.
The animals re-introduced into Switzerland and Slovenia/Croatia originated from the same source population in the Carpathian Mts of Slovakia. The comparison between the source population in Slovakia and the
re-introduced population in the Jura Mts (df = 3, χ2 =
5.01, P > 0.05) and the population in Slovenia/Croatia
(df = 3, χ2 = 2.96, P > 0.05) showed a similar occurrence of the four different coat pattern types (Figure
10). In all three populations type 1 was dominant (over
60%). Type 4 was represented between 18-28% and
Type 2 and 3 about 10%. A different development was
found in the population of the Swiss Alps (df = 3, χ2 =
8.58, P < 0.05). Predominant was type 4 (46%). Type 1
10
Results
was represented with 35% and type 2 and 3 with together with 18.75%.
The comparison between the re-introduced population of Slovenia/Croatia and the population of the
Swiss Alps showed a significant difference between
the different coat pattern types (df = 3, χ2 = 20.29, P <
0.001). The occurrence of type 4 was percental higher
in the population of the Swiss Alps than in the Slovenian/Croatian population. No difference was found between the re-introduced Jura Mts population and the reintroduced population of Slovenian/Croatian (df = 3, χ2
= 6.00, P > 0.1). The occurrences of the 4 coat pattern
types showed same ratios.
3.3.1. Spatial distribution of coat patterns in Switzerland
Analyses of the spatial distribution in Switzerland resulted in a significant difference in the frequency of the
four coat pattern types between the Jura Mts population
and population of the Swiss Alps (df = 3, χ2 = 24.35, P
< 0.001). In the Jura Mts type 1 was dominant (77%)
and type 4 were found in only 9 of 44 (20%) individuals. In the Swiss Alps the frequency showed the opposite: type 1 was found in 35% and type 4 in 59 of 128
(46%) individuals (Figure 10). In the population of the
Swiss Alps all of the five types (1, 2, 3, 4a, and 4b)
were found. In the Jura Mts the coat pattern type without spots (type 3) is no longer present and the one with
small spots (type 2) is very rare (2%).
The range of the re-introduced population of the
Swiss Alps is highly topographically structured. To
analyse the spatial distribution of the different coat pattern types, the range was divided into four regions
(Figure 11). The occurrence of different types in the
region of Berne, Waadt, Fribourg (NWA) and Wallis
(SA) showed no difference (df = 3, χ2 = 1.59, P > 0.5).
KORA Bericht Nr. 13
Between the Central Alps (CA) and NWA/SA the frequency of coat patterns was just not significant (df = 3,
χ2 = 6.57, P > 0.05). A high significantly different frequency of coat pattern types was found between the
population of the Swiss Alps and the Jura Mts population (df = 3, χ2 = 27.2, P < 0.001).
3.3.2. Temporal changes in pattern type frequencies
Before 1908, in the historic population 15 of 21 (71%)
individuals showed type 3 (without spots) and only 4
(19%) type 1 (large spots). Type 4 did not exist. Reintroduced lynx (after 1972) were of completely different pattern types than the historic individuals (Swiss
Alps, df = 3, χ2 = 55, P < 0.001; Jura Mts, df = 3, χ2 =
45.6, P < 0.001). In the recent population of the Swiss
Alps 45 of 128 (35%) individuals showed type 1 and
only 10 (8%) type 3, moreover type 4 was found in 59
of 128 (46%) individuals. The frequency of type 2 was
low in all populations (historical Alps: 2 of 21[9.5%],
recent Alps: 14 of 128 [11%], Jura Mts: 1 of 44 [2%]).
During the fist decade after the re-introduction the
variety of types increased, and after 20 years all five
types were present in the Swiss populations. To analyse this change more in detail frequencies were analyzed in 10 year intervals (1970-1979, 1980-1989 and
1990-1999). In the late 70s type 1 had become rare and
type 3 had completely disappeared. During the 90s the
frequency of type 4 increased distinctly. These temporal changes were only found in the population of the
Swiss Alps. In the Jura Mts the database was too small.
The temporal change correlated with the spatial distribution. The frequency of coat patterns in the 1970s
showed the distribution in Central Switzerland. The
distribution in the North-west and South Alps reflected
the 1990s.
Figure 3. Principal-Component-Analyse. Every
symbol shows a single individual which is assorted after the total number of spots and the existence of rosettes. The first principal component
explains 82.76 % of variance, the second 6.58 %.
Total spots:
Ø: 184
Type 3:
Without spots
Total spots:
Ø: 528
Type 2:
Small spots
Total spots:
Ø: 317
Type 1:
Large spots
irregular
small spots
Ø: < 1,5 cm
round, rarely
strip-formation
distribution
size
form
round
small spots
Ø: < 1,5 cm
regular
black
round
form
black
regular
color
distribution
number
no spots but
small spots
some small
Ø: < 1 cm
spots on the
elbow,
spots Ø: < 1 cm
size
regular
black
Ø: 33 spots (L/R 1)
round
regular
regular
distribution
black
black
color
Ø: 87 spots (L/R 1)
black
color
number
round
large spots,
Ø: 1,6-3 cm
Ø: 58 spots (L/R 1)
round, V-form
(open to the
head) or other
different spotformations
form
number
large spots,
Ø: 1,6-3 cm
Shoulder / up- Forearm
per arm
size
Coat pattern type Spots
Table 3. Description of Coat Pattern Types
Regular
black
Ø: 25 spots
round
small spots,
Ø: < 0,5 cm
regular
black
Ø: 25 spots
round
small spots,
Ø: < 0,5 cm
regular
black
Ø: 25 spots
round
small spots,
Ø: < 0,5 cm
Hand
round
small spots
Ø: < 1,5 cm
regular
black
regular
black
round
regular
black
Ø: 32 spots (L/R 3)
round
no spots but
small spots,
some small
Ø: < 0,5 cm
spots on the
knee,
spots Ø: < 1 cm
regular
black
Ø: 100 spots (L/R 3)
regular
black
round
round
large spots,
Ø: 1,6-3 cm
Lower leg
Ø: 55 spots (L/R 3)
small spots
Ø: < 1,5 cm
regular
black
round
large spots,
Ø: 1,6-3 cm
Femur / hip
-
-
-
no spots
-
-
-
-
no spots
-
-
-
-
no spots
Foot
large spots,
Ø: 1,6-3 cm
Dorsum
black or brownish
regular
black
Ø: 19 spots (L/R 2)
round
no spots but some
small spots on the
ventral side,
spots Ø: < 1 cm
regular
round, rarely stripformation
small spots
Ø: < 1,5 cm
regular
black
-
-
-
no spots
regular
black
Ø: 82 spots (L/R 2)
round, rarely stripformation
small spots
Ø: < 1,5 cm
irregular
black
Ø: 48 spots (L/R 2)
round or spots organ- round or spots organized into strips of 5 or ized into strips of 7 or
fewer spots
fewer spots
large spots,
Ø: 1,6-3 cm
Flank
Juni 2002
Results
11
Total spots:
Ø: 581
Type 4b:
Small spots with
rudimentary rosettes
Total spots:
Ø: 630
Type 4a:
Rosettes
round or irregu- round
lar spot- and
rudimentary
rosettes
(small spots
organized into
patterns of 3-7
spots)
form
black
irregular
distribution
regular
black
Ø: 95 spots (L/R 1)
small spots
Ø: < 1 cm
regular
black
Ø: 25 spots
round
small spots,
Ø: < 0,5 cm
regular
black
Ø: 25 spots
round
small spots
Ø: 1-2 cm
Lower leg
small spots
Ø: < 1 cm
irregular
black or
brownish
irregular
black
irregular
black
Ø: 102 spots (L/R 3)
round or irregu- round
lar spot- and
rudimentary
rosettes
(small spots
organized into
patterns of 3-7
spots)
small spots
Ø: < 1 cm
irregular
spots: black
rosettes: inside
brownish
Ø: 116 spots (L/R 3)
rosettes: small round or
spots organized washed out
into patterns of
3-7 spots,
irregularly spotformations
small spots
Ø: 1-2 cm
Femur / hip
-
-
-
-
no spots
-
-
-
-
no spots
Foot
spots: black
rosettes: inside
brownish
irregular
black
round or irregular
spot- and rudimentary
rosettes
(small spots organized into patterns of
3-7 spots)
small spots
Ø: < 1 cm
irregular
spots and strips: black
rosettes: inside
brownish
irregular
black
Ø: 93 spots (L/R 2)
round or irregular
spot- and rudimentary
rosettes (small spots
organized into patterns of 3-7 spots)
small spots
Ø: 1-2 cm
irregular
rosettes,
round spots or strips
irregular small or larger spots or strips,
spots: Ø: 1-2 cm
strips: w: 0,5-1 cm,
l: 3-7 cm
Dorsum
Ø: 103 spots (L/R 2)
rosettes: small spots
organized into patterns of 3-7 spots,
irregularly spot- formations
small spots
Ø: 1-2 cm
Flank
Results
color
number
small spots
Ø: 1-2 cm
size
regular
irregular
distribution
black
spots: black
rosettes: inside
brownish
Ø: 100 spots (L/R 1)
color
number
rosettes: small round or longspots organized ish
into patterns of
3-7 spots,
irregularly spotformations
form
Hand
Irregular small small spots,
and larger spots Ø: < 0,5 cm
small spots
Ø: 1-2 cm
Shoulder / up- Forearm
per arm
size
Coat pattern type Spots
Table 3. (cont.) Description of Coat Pattern Types
12
KORA Bericht Nr. 13
Juni 2002
Results
13
Type 1: large spots
Shoulder / upper arm
Flank
Back
Figure 4. Description of type 1: large spots.
Forearm / hand
Femur / hip
Lower leg / foot
14
KORA Bericht Nr. 13
Results
Type 2: small spots
Shoulder / upper arm
Flank
Forearm / hand
Femur / hip
Lower leg / foot
Back
Figure 5. Description of type 2: small spots.
Juni 2002
Results
15
Type 3: without spots
Shoulder / upper arm
Femur / hip
Flank
Forearm / hand
Back
Figure 6. Description of type 3: without spots.
Lower leg / foot
16
KORA Bericht Nr. 13
Results
Type 4a: rosettes
Shoulder / upper arm
Flank
Forearm / hand
Femur / hip
Lower leg / foot
Back
9. Thereofwas
significant
Figure 7. Fig.
Description
typeno4a:
rosettes. correlation between the distances from the den sites to the water bodies and
the precipitation (water shortage).
Juni 2002
Results
17
Type 4b: small spots with rudimentary rosettes
Shoulder / upper arm
Flank
Forearm / hand
Femur / hip
Lower leg / foot
Back
9. Thereofwas
significant
correlation
between the
distances from the den sites to the water bodies and
Figure 8. Fig.
Description
typeno4b:
small spots
with rudimentary
rosettes.
the precipitation (water shortage).
18
KORA Bericht Nr. 13
Results
Spots on flanks
no
yes
Type 3
Spots in Form of
Rosettes
no
yes
Clear rosettes
(small blotches)
Spots < Ø 1.6 cm
Number of spots > Ø 422
no
yes
Type 1
no
Type 2
yes
Type 4a
Type 4b
Number of Individuals
Figure 9. Path-diagram for assigning all available lynx to one of the five coat pattern types. Type 1: large spots, type 2: small
spots, type 3: without spots, type 4a: rosettes and type 4b: small spots with rudimentary rosettes.
? ? Type 1: large spots
? ? Type 2: small spots
? ? Type 3: without spots
? ? Type 4a: clear rosettes
? ? Type 4b: small spots with
52
48
44
40
36
32
28
24
20
16
12
8
4
0
28
4
6
Slovakia
rudimentary rosettes
7
2
(n=47)
45
29
14
10
Alps CH
(n=128)
30
52
48
44
40
36
32
28
24
20
16
12
8
4
0
Number of Individuals
52
48
44
40
36
32
28
24
20
16
12
8
4
0
Number of Individuals
Number of Individuals
Source-Population
34
4
1
0
Jura CH
(n=44)
5
52
48
44
40
36
32
28
24
20
16
12
8
4
0
44
10
7
4
2
Slovenia / Croatia (n=67)
Figure 10. Frequency of the various coat pattern types in Swiss and European populations (recent population of the Swiss
Alps; Jura Mts population; Slovakian population; Slovenian/Croatian population; total individuals: n = 307).
Juni 2002
Results
19
Figure 11. Distribution of coat pattern types in recent lynx populations in Switzerland (CA: Central Alps; NWA: North west
Alps with Berne, Fribourg and Waadt; SA: South Alps, Wallis; n = 185).
? ? Type 1: large spots
? ? Type 2: small spots
? ? Type 3: without spots
? ? Type 4a: clear rosettes ? ? Type 4b: small spots with rudimentary rosettes
100%
Numbers of Individuals in %
1
80%
7
3
1
2
1
1
4
1
1
60%
1
1
1
3
1
2
8
2
3
2
4
3
2
4
1
2
2
4
6
6
20%
6
8
1
4
40%
8
5
2
3
7
1
1
8
4
1
12
5
10
7
1
1
Time Periode
Figure 12. Changes in the frequency of the different coat pattern type from 1802 – 1999 in Switzerland (n = 175).
1998-1999
(n=18)
1996-1997
(n=30)
1994-1995
(n=28)
1992-1993
(n=15)
1990-1991
(n=12)
1988-1989
(n=6)
1986-1987
(n=9)
1984-1985
(n=4)
1982-1983
(n=10)
1980-1981
(n=7)
1978-1979
(n=3)
1976-1977
(n=6)
1974-1975
(n=3)
1972-1973
(n=3)
1970-1971
(n=0)
1851-1900
(n=11)
1802-1850
(n=10)
0%
20
Discussion
KORA Bericht Nr. 13
4. Discussion
4.1. Description of coat pattern types
Werdelin & Olsson (1997) coded felid coat patterns
into six discrete categories: flecks (small spots not organized into patterns), rosettes (small spots arranged
into patterns of six or fewer spots), small blotches
(small irregularly shaped areas of dark on a usually
lighter background), blotches (large areas of variable
color framed by dark and set on a lighter background),
vertical stripes (dark, dorsoventrally or anterodorsallyposteroventrally directed stripes on a lighter background) and uniform (no distinguishable pattern)
(Figure 13). My definitions of the five coat pattern
types corresponds to Werdelin & Olsson’s following
types: type 1 and 2, “large and small spots” correspond to “flecks”; type 3, “without spots” to “uniform”; type 4a, “clear rosettes (small blotches) to
“small blotches” and type 4b “small spots with rudimentary rosettes” correspond to “rosettes” (Figure 14).
Werdelin and Olsson (1997) looked at the coat patterns of living cat species in conjunction with the evolutionary relationship within the cat family to estimate
how often each type of pattern had given rise to each of
the others and used clad. meth. of phyl. inf. to reconstruct historical events in a phylogenetic framework.
Because small flecks gave rise to large spots, rosettes,
stripes and blotches much more frequently than any of
the other possible transitions, they suggest that the
common ancestor of modern cats, rather than having
large spots which broke apart later, was patterned with
small flecks. As cats evolved, the flecks coalesced into
larger blotches, rosettes or stripes. Lynx coat pattern
may have developed after this hypothesis. On the other
hand, Weigel (1961) proposed a felid coat pattern evolution from a primitive form with large spots to smaller
spots spacing into rosettes, and then striped patterns
may have developped (Figure 15).
4.2. Comparisons with other descriptions of coat
patterns in lynx
Several authors observed and described different coat
patterns of Eurasian lynx during the last century (Table
4). Most of them observed three different types of patterns: large spots, small spots and without spots but the
rosette-spotted types has not been described in details.
The descriptions differ widely. Only the large spotted
type was always exactly recognized and described. In
most of the descriptions variations of the same type
were defined as other types. Grégorova (1997) defined
a hypothesis about a new type of pattern: intermediate
(rosettes) type. This type has hypothetically evolved
from hybridisation of a spotted and a non-spotted type.
Different frequencies of coat patterns in the Eurasian lynx in different areas of its range were analyzed
by several authors. The observed types were large-,
small- or non-spotted. The frequencies differed immensely but geographical trends have been found
(Table 5). In the Carpathian Mts the large-spotted type
showed a high frequency. In Western and Northeastern Siberia and in Central Asia type 3, without
spots, was mostly observed. Type 2 showed generally a
low frequency.
Figure 13. The six different coat patterns in felids. Left column, top to bottom, flecks, rosettes, vertical
strips; right column, top to bottom, small blotches, blotches, uniform. After Werdelin & Olsson (1997).
Juni 2002
Discussion
21
Figure 14. The four different types of spots in lynx coat patterns: spots correspond to flecks, (included small
and large spots), without spots correspond to uniform, clear rosettes (small blotches) correspond to small
blotches and rosettes or rudimentary rosettes correspond to rosettes. Left pictures top to bottom after Werdelin
& Olsson (1997).
4.3. Spatial and temporal distribution of coat patterns in Switzerland
In 1971 the Eurasian lynx was re-introduced into the
Swiss Alps and initially, had spread quite fast over the
western Alps and the Jura Mts but since the mid 1980s,
the population expansion came to a halt, even though
there were still large areas of suitable habitat not yet
occupied in eastern Switzerland (Breitenmoser 1983,
Haller 1992). Though the re-introduced individuals in
the Jura Mts and population of the Swiss Alps originated from the same source population, the Carpathian
Mts of Slovakia, the development of the frequency in
different coat pattern types differed widely. Before the
extermination the range of the Eurasian lynx extended
throughout Europe (Nowell & Jackson 1996) therefore
the historic Swiss population belonged to the same
Carpathian population as the Slovakian population.
The change of the occurrence of different types developed from type 3, without spots, in the historical population to type 1, large spots, in the early re-introduced
population, then to type 2, small spots, and finally to
type 4a and 4b, rosettes and rudimentary rosettes, in
late re-introduced or recent populations. Ragni (1993)
analyzed coat patterns of the extinct Alpine population.
22
Discussion
He recognized a non-spotted type (concolor) as being
predominant.
In the source population all five types were present,
also in the re-introduced populations in Slovenia/
Croatia and the Swiss Alps but the frequency is different. In the Jura Mts, the bottleneck created by the reintroduction lead to the loss of one coat pattern type:
type 3. The population of the Iberian lynx (Lynx pardinus) with approximately 40-50 individuals was isolated
since the early 1960s. Originally thee coat patterns
were present but today no animal exhibits the rare
small-spotted pattern (Beltran and Delibes, 1993). In
the Swiss Alps population, the frequency of the coat
pattern types had significantly changed compared with
the source and the historical population. A probably
aggravated exchange is possible between the Central
and NW-Alps but between Jura Mts and Alps no exchange is possible because of the topography. Therefore the two populations are still small and isolated.
KORA Bericht Nr. 13
Effects of genetically drift is found in both population
in Switzerland: in the population of the Swiss Alps, a
postponement of frequencies of coat pattern types was
found. In the Jura Mts the genetically drift resulted in a
loss of one coat pattern type.
Small, isolated sub-populations are vulnerable to
genetic drift. Alleles with low frequency are likely disappeared from the population gene pool (Griffiths et al.
1996).
Color patterns in felids are genetically determined
but the actual mechanisms that create the patterns are
still unknown. Definitive conclusions about the reason
for the change of the occurrence of the different coat
pattern types can be made after genetically analyses, so
the results will be linked to a study on populations genetics of Swiss populations. Analysis of known pedigrees from field studies in Switzerland and in zoos
could contribute to the knowledge of the mechanisms
behind the transmission of coat patterns.
Figure 15. The current hypothesis of coat pattern evolution in Felidae. After
Weigel (1961).
Juni 2002
Discussion
23
Table 4. Descriptions of coat patterns of Eurasian Lynx.
Ognev (1935)
Central Russian lynx: a) some with more dark patterns of bands and spots; b) some have an almost
uniform color without spottiness on trunk.
Stollman (1963)
Carpathian Lynx: a) characteristic spots (the spots form long strips on the dorsum and on the flanks
the spots are almost round); b) less spotted; c) without spots (uniform: the spots appear only indicated, on the legs, shoulder and belly the spots appear clearer).
Vasiliu & Decei (1964)
Rumanian Carpathian Mts: a) spots (well developed all over, specially on the dorsum and flanks); b)
small spots; c) without spots (on the dorsum and only pale spots on the flanks).
Matjuschkin (1978)
Coat pattern types: a) large and clear spots; b) small but clear spots and small stripes on the back; c)
without spots on the back but with spots on the legs; d) and without spots all over.
Miric (1978)
Balkan lynx: a) large spots (intensive colored spots, rarely placed); b) small spots (pale, small and
densely placed spots); c) weak spots (lack of spots on the back and flanks).
Ragni et al. (1993)
Eurasian lynx: spotted (permanent black spots), striped (the pattern is arranged in horizontal stripes
and bars of permanent-evanescent) and concolour (without markings in the somatic regions or with a
pattern so scattered and obsolescent that it cannot be defined).
Grégorova (1997)
Coat pattern types: a) spotted (small or large spots), b) without spots, c) intermediate type (rosettes,
hypothetically evolved from hybridization of the previous two types).
Table 5. . Frequency of the different coat patterns of Eurasian lynx in different areas of its range. Description of coat pattern
types a–d: see Table 4. x = frequency unknown.
Region
Author
large spots
a
small spots
b
The Balkans
Miric (1978)
Carpathian Mts
Stollmann (1963) (CSSR)
90.00 %
x
Kunc (1971)
67.70 %
22.60 %
Matjuschkin (1978)
Kaukasus
x
x
Western Siberia
c
d
31.83 %
33.00 %
Iberian lynx
not spotted
17.00 %
x
9.70 %
x
-
x
14.00 %
x
-
-
60.00 %
20.00 %
3.00 %
Sajanen/Baikal
-
27.00 %
Siberia
36.00 %
38.00 %
Jakutien/Amur
-
-
x
x
Altai
-
-
x
x
x
65.00 %
-
-
27.00 %
73.00 %
Formosow (1929)
-
-
x
x
Bannikow (1954)
-
-
x
x
Tienschan-Saur/Altau
10.00 %
Central Asia
Northern Mongolia
North-eastern Siberia
Kistschinski (1967)
22.20 %
77.80 %
24
References
KORA Bericht Nr. 13
5. References
Bannikow, A. G. 1954. Die Säugetiere der MVR. Trudy
Mong. kom. Ak. Nauk SSSR Moskau 53.
Beltran, J. F. and M. Delibes, 1993. Physical characteristics
of Iberian Lynxes (Lynx pardinus) from Doñana, southwestern Spain. J. Mamm. 74 (4): 852–862.
Breitenmoser, U. 1983. Zur Wiedereinbürgerung und Ausbreitung des Luchses (Lynx lynx L.) in der Schweiz.
Schweiz. Z. Forstwes. 134: 207–222.
Breitenmoser, U. and M. Baettig, 1992. Wiederansiedlung
und Ausbreitung des Luchses Lynx lynx im Schweizer
Jura. Revue suisse Zool. 99: 13–176.
Breitenmoser, U. and Ch. Breitenmoser-Würsten, 1998. Der
Luchs. Biologie einheimischer Wildtiere 1/10a, Zürich.
Formosow, A. N. 1929. Die Säugetiere der nördlichen Mongolei nach dem Material der Expedition vom Jahre 1926.
Predvar. Otcet zool. Eksped. V Sev. Mongoliju. Leningrad.
Garcia-Perea, R. 1994. The Pampas Cat Group (genus Lynchailurus Severtzov, 1858) (Carnivora: Felidae), a systematic and biogeographic review. American Museum of
Natural History Nr 3096: 1–35 (New York).
Garcia-Perea, R. 1994. Pampas Cats: how many species?.
Cat News 20: 21–24.
Grégorova, E. 1997. Lynx, Zoological Garden Bojnice.
Griffiths, A. J. F. et al. 1996. An Introduction to Genetic
Analysis. Sixth Edition. W. H. Freeman and Company,
New York.
Haller, H. 1992. Zur Ökologie des Luchses (Lynx lynx) im
Verlauf seiner Wiederansiedlung in den Walliser Alpen.
Mammalia depicta 15: 1–62.
Kistschinski, A. A. 1967. Zur Verbreitung und intraspezifischen Variation des Wolfs, Vielfrasses und Luchses auf
dem Kolyma-Plateau und im Kolyma-Gebirge. In: Ekologija mlekopitajuschich i ptic. Pages 10–16. Moskau.
Koubek, P. and J. Cerveny, 1996. Population development
and recent distribution of the lynx (Lynx lynx) in the
Czech republic. ACTA Scjentiarum natura 3: 2–15, Lium
academiae scienti arum bohemicae Brno.
Kunc, L. 1971. Individualni variabilita zbarveni rysa ostrovida (Lynx lynx) Karpatske oblasti. Lynx n. s. 12: 60–65.
Matjuschkin, E. N. 1978. Der Luchs. A. Ziemsen Verlag,
Wittenberg Lutherstadt, Germany, 160 pp.
Miric, D. J. 1978. Die Luchspopulationen der Balkanhalbinsel. Serbian Academy of Sciences and Arts, Vol.
DXXXIX, (55): 150 pp.
Nowell K. and P. Jackson 1996. Wild Cats. International
Union for Conservation of Nature and Natural Resources,
110 pp.
Ognev, S. L. 1935. Mammals of USSR and adjacent countries. Vol. 3: 165–186, Carnivora, Moscow.
Ortolani, A. and T. Caro, 1996. The Adaptive Significance
of Color Patterns in Carnivores: Phylogenetic Tests of
Classic Hypotheses. In: Gittleman J. L. (ed.) Carnivore
Behavior, Ecology, and Evolution, Vol. 2: 132–188.
Ithaca, Cornell University Press.
Ragni, B., M. Possenti and S. Mayr, 1993. The Lynx in the
Italian Alps. Cat News 19: 21–25.
Stollmann, A. 1963. Beitrag zur Kenntnis des Luchses, Lynx
lynx in den tschechoslowakischen Karpaten. Folia Zoologica, Brünn (Brno), 12: 301–316.
Vasiliu, G. D. and P. Decei, 1964. Über den Luchs (Lynx
lynx) der rumänischen Karpaten. Säugetierkundliche Mit-
teilungen, München, 12: 155–183. (Bukarest).
Weigel, I. 1961. Das Fellmuster der wildlebenden Katzenarten und der Hauskatze in vergleichender und stammesgeschichtlicher Hinsicht. Säugetierkundliche Mitteilungen,
München, 120 pp.
Werdelin, L. and L. Olsson, 1997. How the Leopard got its
spots: a phylogenetic view of the evolution of felid coat
patterns. Biological Journal of the Linnean Society 62:
383–400.
Juni 2002
Appendix I
25
Review coat patterns (supplementation of the Introduction)
1. General statements on coat patterns in mammals
Mammals exhibit a remarkable variety of coat patterns
(Murray 1988). Especially carnivores show a wide variety of basic pelage colors and a great diversity of
markings on their coat – including spots, stripes, bands
and patches (Ortolani & Caro 1996). Therefore they
are an exciting taxonomic group against which evolutionary theories of coloration can be examined.
At the turn of the century, naturalists began to
speculate about the survival value of the pelage and
skin color (e.g. Roosevelt 1911). Work on animal coloration since 1940 has concentrated primarily on the
development of pelage patterning in mammals (e.g.
Murray 1981), the mechanisms by which animals
match their background (e.g. Endler 1978) and the
theories underlying the evolution of coloration patterns
(e.g. Endler 1988). It is generally accepted that specific
color patterns in mammals are genetically determined
but the actual mechanisms that create the patterns are
still unknown. Weigel (1961) believes that all coat patterns evolved from a dark-spotted type. The spots subsequently broke up and fused in a variety of ways to
give the modern array of coat markings (Figure I.1).
2. Present status of the research
2.1. Individual identification and population estimation
In studies on the behavior of individual animals, spot
patterns have been used as the key feature to distinguish individuals from one another in the field (Caro &
Durant 1991). It is known that individual cheetahs vary
in both coat color and pattern even within small populations and tail markings of cheetahs born in the same
litter resemble each other more closely than those born
in different litters.
These natural markings of coat patterns were also
used for population estimations and studies of density.
Karanth and Nichols (1998) estimated densities of wild
tiger populations using photographic capture-markrecapture models in four ecologically distinct study
sites. Based on the fact that tigers are individually identifiable from their stripe patterns (Schaller 1967),
Karanth (1995) has demonstrated the potential for estimating their population size using photographic „captures”, within the theoretical framework of formal capture-mark-recapture theory. Tigers were identified
from photographs by comparing shapes of specific individual stripes and positions of several such stripes
relative to each other on the animal body. Laass (1999)
used the sight-resight method by photocaptures for a
quantitative monitoring of a lynx population in the
northwestern Swiss Alps.
2.2. Developmental biology and adaptive value
Some striped animals are very difficult to detect in
their natural habitat, whilst others are clearly conspicuous. The pattern composed of regularly repeated stripes
were conspicuous and the irregular stripes were often
cryptic (Godfrey et al. 1987). The coat coloration plays
an important role for the camouflage, for example a
lynx with a reddish coat and black spots disappears in a
beech forest (Breitenmoser and Breitenmoser-Würsten
1990). Coloration can yield concealment through three
different means: i) general color and pattern resemblance or background invoke the similarity between an
animal’s color and that of the natural background in
which it lives; ii) disruptive coloration, sharply contrasting colors and irregular markings break up the
form of the animal, making regions of its body appear
mutually discontinuous and iii) countershading, lightening of the ventral surface and darkening of the dorsal
surface of the animal is believed to counteract the effects of shade and light. Striped patterns for example
may camouflage in two ways: First, the stripes may
Figure I.1. Lynx systematics after Weigel (1961), Eurasian and Canada lynx.
26
Appendix I
simply mimic the pattern of the surrounding scenery
and secondly, stripes may run across the natural contours of the animal `s body, and being visually strong
features, disrupt the characteristic outline of the animal
(Godfrey et al. 1987).
Concealing coloration can also be divided into protective resemblance and aggressive resemblance, respectively reflecting the need to be concealed from
predators and the need to approach prey undetected.
Intuitively, aggressive background resemblance might
be expected to be more prevalent in carnivores, but
many species in this order are themselves subject to
predation (Ortolani & Caro 1996). Although genes
control the processes involved in coat pattern formation, the actual mechanisms that create the patterns are
still unknown. It would be attractive from the viewpoint of both evolutionary and developmental biology
if a single mechanism would be found to produce the
enormous assortment of coat patterns found in nature
(Murray 1988).
Murray’s (1988) mathematical model describes how
these patterns may be generated in the course of embryonic development. An important feature of the
model is that the patterns it generates bear a striking
resemblance to the patterns found on a wide variety of
animals such as the leopard, cheetah, jaguar, zebra and
giraffe. The model predicts that the patterns can take
only certain forms, which in turn implies the existence
of developmental constrains and begins to suggest how
coat patterns may have evolved. Physically, spots correspond to regions of differently colored hair. Hair
color is determined by specialized pigment cells called
melanocytes, which are found in the basal, or innermost, layer of the epidermis. The melanocytes generate
a pigment called melanin that then passes into the hair.
In mammals there are essentially only two kinds of
melanin: eumelanin, from the Greek words eu (good)
and melas (black), which results in black or brown
hairs, and phaeomelanin, from phaeos (dusty), which
makes hairs yellow or reddish orange.
Turing (1952) postulated a chemical mechanism for
generating coat patterns. He suggested that biological
form follows a prepattern in the concentration of
chemicals he called morphogens, which can react with
one other and diffuse through cells. Spatial patterns of
morphogen concentrations can arise from an initial uniform distribution in an assemblage of cells. In reactiondiffusion models one starts with two morphogens that
can react with each other and diffuse at varying rates.
If the morphogens are now allowed to diffuse at equal
rates, any spatial variation from that steady state still be
smoothed out. If the diffusion rates are not equal, diffusion can be destabilizing: the reaction rates at any
given point may not be able to adjust quickly enough to
reach equilibrium. If the conditions are right, a small
spatial disturbance can become unstable and a pattern
begins to grow. The type of pattern that results depends
on the various parameters of the model and can be ob-
KORA Bericht Nr. 13
tained from mathematical analysis. Closely related species can exhibit similar color markings and behavioral
ecology as a result of shared ancestry. In spite of this
fact, by simply testing the correlation between coloration variables and behavioral-ecological variables, adaptations could not be distinguished from homologies
(Ortolani & Caro 1996).
2.3. Systematics and biogeography
Felids have engendered considerable systematic controversy. The cause of the problem is: small samples,
lack of information, ambiguous original descriptions,
and authors ignoring earlier publications. Different generic classifications resulting from the study of a variety of characters (e.g., morphological, morphometrical,
behavioral, biochemical, and cytogenetic) have resulted in the recognition of 4 to 9 genera. The tendency
during recent years to group many felid species into
large cosmopolitan genera (e.g., Ellermann and Morrison-Scott, 1966) has simplified the nomenclature, but
these assemblages are not supported by original data
(Garcia-Perea 1994). In the study with the pampas cat,
Felis colocolo, Garcia-Perea (1994) found some unusual patterns of variation within the population identified as colocolo while conducting a phylogenetic study
on the living species of felids. An examination of all
available museum specimens revealed that this assemblage consisted of three closely related species. Their
diagnostic characters and geographic distribution has
been the subject of that study. 86 specimens of Pampas
cats have been examined, consisting of 72 skins and 51
skulls, from eight large North American, South American, and European collections. The evaluation of morphological variation included descriptions of pelage
variations, which emphasized the characteristics and
distribution of the markings. Spotting patterns are important in felid systematics. Closely related cats sometimes show similar patterns, which has been misinterpreted (e.g., Lynx lynx has been confused with Lynx
pardinus in the Carpathian and Caucasus Mts.). Color
is also important, but is unreliable in old skins because
of fading. Garcia-Perea (1994) recognized five units
for the morphological analysis. Each represent an apparently continuous population.
Felids show great variation of coat patterns. Similarity between species has often been used to indicate
phylogenetic relationships, and variation within species
as taxonomic criteria for subspecific differentiation.
The study of the distribution and characteristics of pelage markings revealed a significant amount of variation, which partially explains the large number of taxa
described for this group. Garcia-Perea distinguished
eight parts of color and pattern variation for the parts of
the head, body, and tail (face, ears, spinal crest, throat,
chest and abdomen, tail, legs and feet). The description
of basic spotting types showed three general patterns
(types 1, 2, and 3). The different coat pattern types are
distributed geographically.
Juni 2002
Appendix I
With felids, it is often difficult to find morphological characters that are completely diagnostic, especially
in closely related species, because atypical character
stated commonly appear at low frequency. For this reason, distinctive morphological gaps in single character
states may not prove useful for detecting genetic discontinuities between species. The level of geographic
variation of the pampas cat observed within the “pajeros” group and the moderate variation shown within
the less extensive “braccatus” and “colocolo” populations suggest that they have been genetically isolated
for a long period. The extent of variation within each is
comparable to that found between recognized subspecies in other felids.
2.4. Heritage of coat patterns
The color patterns in mammals are genetically determined but very little is known about the heritage of
coat patterns. Robinson (1976) suggested that the
change of the spotted pattern to blotches in the cheetahs is comparable to that of the so-called „striped
tabby“ to the „blotched tabby“ in the domestic cat Felis
catus. This implies that the king coat color pattern results from a mutation inherited as a single autosomal
recessive allele. The expression of the aberrant coat
color results from the action of an autosomal recessive
allele (van Aarde & van Dyk 1986).
The variations of the marking-color system of the
Felids are genetically controlled and, in this case, it is
possible to recognize a genetic homology with the multiple allelism to the „Tabby” locus (Robinson 1977). In
particular: The spotted coat and the striped coat can be
considered a gradient of phenotypic expression of the „
spotted-striped” or „mackerel tabby” (Ts) allele, while
the allele controlling the concolor coat is referable to
the „Abyssinian tabby” (Ta). Both alleles belong to the
homologous genetic series well known in Felis and, to
a lesser extent, in other genera of the Felidae family:
Panthera, Acinonyx, Leptailurus (Robinson 1978). A
Felidae-wide study is in progress to determine the
genes coding for coat patterns at the Laboratory of Genomic Diversity, National Cancer Institute, Fredrickton, VA (Director S. J. O'Brien; E. Eizirik, pers.
comm.). Eizirik developed a study of genetic basis of
melanism in leopards and other cats as part of a
broader investigation on the evolution of coat color
genes in the Felidae.
2.5. Coat patterns in Eurasian lynx
Within the lynx family there are four recent species:
the Eurasian lynx, the Pardel lynx, the Canadian lynx
(Lynx canadensis) and the Red lynx (Lynx rufus). In
Switzerland the native lynx is the Eurasian. His present
range extends throughout Europe and Siberia. The
habitat are primarily forested areas which have good
ungulate populations. In central Asia, lynx occur in
more open, thinly wooded areas (Matjuschkin 1978).
Lynx are probably found throughout the northern
27
slopes of the Himalayas, and have been reported both
from thick scrub woodland and from barren, rocky areas above the treeline. On the better-forested southern
Himalayan slopes, the only record is a sighting in alpine tundra (4.500 m) from the Dhaulagiri region of
Nepal. Lynx occur locally over the entire Tibetan plateau, and are found throughout the rocky hills and
mountains of the central Asia desert regions.
The Eurasian lynx has one of the widest ranges of
all cat species, with approximately 75 % of the range
within the borders of Russia. Lynx have been recorded
as far north as 72° N, near the edge of the continental
landmass (Nowell & Jackson 1996). Due to the extension of the present range the habitats of lynx – primarily forested areas inhabited by large ungulate populations – may differ considerably and consequently it is
not astonishing that they show different coat patterns.
In central Asia, lynx occur in more open, thinly
wooded areas (Matjuschkin 1978). Lynx are probably
found throughout the northern slopes of the Himalayas,
and have been reported both from thick scrub woodland and from barren, rocky areas above the treeline.
On the better-forested southern Himalayan slopes, the
only record is a sighting in alpine tundra (4.500 m)
from the Dhaulagiri region of Nepal. Lynx occur locally over the entire Tibetan plateau, and are found
throughout the rocky hills and mountains of the central
Asia desert regions.
The habitats of lynx are very different and consequently it is not astonishing that they show different
coat patterns. The following overview presents the different description of coat patterns of Eurasian lynx of
several authors. Most of them observed three types of
patterns (large spots, small spots and without spots).
The leopard-like pattern has not yet been described in
detail. But in some pictures of several studies, the leopard-like pattern can be recognized (Figure I.2, I.3, I.4).
28
Appendix I
Ognev
(1935):
KORA Bericht Nr. 13
Color variegated: Individuals encountered with various degrees of development of pattern consisting of
longitudinal bands on back and spots on trunk. Uniformly colored or almost uniformly colored lynx encountered together with individuals with fur marked by distinct design. Lynx of the same geographical
regions have numerous color variations. Spots on flanks are very marked, but lose intense black shade.
Spots again more marked on limbs. Blackish hair added here to spots of cinnamon-brown hair. Paws of
massive legs lack spots. Three longitudinal bands formed of closely adjacent elongated spots which are
quite distinct, particularly in middle and posterior region of back, visible on rear of neck. Numerous deviations exist from this most common type of winter fur colors of the Central Russian lynx, some with
more dark patterns of bands and spots, while some have color almost uniform above without spottiness
on trunk.
The coloration of an animal depends on the wideness and number of the hair-bands, on the intensity of
their coloration and on the position of this bands. The longitudinally striped- pattern is the primitive pattern of the vertebrates, because most of the juveniles show this pattern. Spots break down into smaller
spots and rosettes while at the same time leading to various striped patterns as sidelines. Coat pattern
types: spots almost disappeared, pale spots and large spots.
Weigel
(1961):
Stollman (1963):
Variations of coloration of the Carpathian Lynx (three types): a) characteristic spots (the spots form long
strips on the dorsum and on the flanks the spots are almost round), b) less spotted and c) without spots
(uniform: the spots appear only indicated, on the legs, shoulder and belly the spots appear clearer). It’s
not known if the differences in the coat coloration and coat pattern correlated with the age of the individual or if it’s an adaptation at the environment or if it’s just an individual variation.
Vasiliu & Decei
(1964):
Coat pattern types (three types in the Rumanian Carpathian): a) spots (well developed all over, specially
on the dorsum and flanks), b) small spots and c) without spots (on the dorsum and only pale spots on the
flanks). The spots show different forms. Three coat patterns were distinguished: wolf-lynx (on the dorsum with weak spots), fox-lynx (uniform with weak spots only on the flanks) and cat-lynx (large spots
with almost strips on the dorsum).
Matjuschkin
(1978):
Matjuschkin described in his monography four coat pattern types: a) large and clear spots, b) small but
clear spots and small stripes on the back, c) without spots on the back but with spots on the legs, d) and
without spots all over (Figure I.5)
Miric
(1978):
Coat pattern types of Balkan lynx: a) large spots (intensive colored spots, rarely placed), b) small spots
(pale, small and densely placed spots), c) weak spots (lack of spots on the back and flanks).
Ragni et al.
(1993):
Within the known variation of the coat marking-color system of the Eurasian lynx, the authors recognized three types: spotted (permanent black spots), striped (the pattern is arranged in horizontal stripes
and bars of permanent-evanescent) and concolor (without markings in the somatic regions or with a pattern so scattered and obsolescent that it cannot be defined). These patterns are related to the somatic regions: occipis-cervicalis, scapularis, dorsalis, lateralis, omeralis and femuralis (Figure I.6).
Figure I.3. Intermediate coloration. Grégorova (1997).
Figure I.2. Coloration of the back and
flanks. Vasiliu & Decei (1964).
Figure I.4. Coloration of the back and
flanks. Ragni (1991).
Juni 2002
Appendix I
Figure I.5. Variability of coat patterns of Eurasian lynx: A) large and clear spots, B) small but clear spots and small
stripes on the back, C) without spots on the back but with spots in the legs, D) without spots all over. After Matjuschkin (1978).
Figure I.6. Variability of coat patterns of Eurasian lynx: A) large and clear spots, B) small but clear spots and small
stripes on the back, C) without spots on the back but with spots in the legs, D) without spots all over. After Matjuschkin
(1978).
29
30
Appendix I
References
Breitenmoser, U. and Ch. Breitenmoser-Würsten, 1990.
Status, conservation needs and reintroduction of the lynx
Lynx lynx in Europe. Nature and environment Series, No.
45. Strasbourg: Council of Europe.
Caro, T. M. and S. M. Durant, 1991. Use of quantitative
analyses of pelage characteristics to reveal family resemblances in genetically monomorphic cheetahs. Journal of
Heredity 82: 8–14.
Eizirik, E. Pers. Comm. Laboratory of Genomic Diversity,
National Cancer Institute, Frederickton, USA.
Endler, J. A. 1978. A predator’s view of animal colour patterns. Evol. Biol. 11: 319–364.
Endler, J. A. 1988. Frequency-dependent predation, crypsis
and aposematic colouration. Proc. Trans. Roy. Soc. London. (ser. B) 319: 505–523.
Garcia-Perea, R. 1994. The Pampas Cat Group (genus Lynchailurus Severtzov, 1858) (Carnivora: Felidae), a systematic and biogeographic review. American Museum of
Natural History Nr 3096: 1–35 (New York).
Garcia-Perea, R. 1994. Pampas Cats: how many species?.
Cat News 20: 21–24.
odfrey, D., J. N. Lythgoe and D. A. Rumball, 1987. Zebra
stripes and tiger stripes: the spatial frequency distribution
of the pattern compared to that of the background is significant in display and crypsis. Biological Journal of the
Linnean Society 32: 427–433.
Karanth, K. U. 1995. Estimating tiger populations from camera-trap data using capture-recapture models. Biological
conservation 71: 333–338.
Karanth, U. and J. D. Nichols, 1998. Estimation of Tiger
densities in India using photographic captures and Recaptures. Ecology 79 (8): 2852–2862.
Laass, J. 1999. Evaluation von Photofallen für ein quantitatives Monitoring einer Luchspopulation in den Alpen. Diplomarbeit, BOKU Wien, 75 pp.
Matjuschkin, E. N. 1978. Der Luchs. A. Ziemsen Verlag,
Wittenberg Lutherstadt, Germany, 160 pp.
Miric, D. J. 1978. Die Luchspopulationen der Balkanhalbinsel. Serbian Academy of Sciences and Arts, Vol.
DXXXIX, (55): 150 pp.
Murray, J.D. 1981b. A pre-pattern formation mechanism for
animal coat markings. Journal of Theoret. Biol. 88: 161–
199.
Murray, J.D. 1988. How the Leopard gets its spots. Scientific. American 3/1988:62–69.
Nowell K. and P. Jackson 1996. Wild Cats. International
Union for Conservation of Nature and Natural Resources,
110 pp.
Ognev, S. L. 1935. Mammals of USSR and adjacent countries. Vol. 3: 165–186, Carnivora, Moscow.
Ortolani, A. and T. Caro, 1996. The Adaptive Significance
of Colour Patterns in Carnivores: Phylogenetic Tests of
Classic Hypotheses. In: Gittleman J. L. (ed.) Carnivore
Behavior, Ecology, and Evolution, Vol. 2: 132–188.
Ithaca, Cornell University Press.
Ragni, B., M. Possenti and S. Mayr, 1993. The Lynx in the
Italian Alps. Cat News 19: 21–25.
Robinson, R. 1977. Genetics for Cat Breeders. Pergamon
Press. London.
Robinson, R. 1978. Homologous coat colour variation in
Felids. Carnivore 1: 68–71.
Schaller, G. B. 1967. The deer and the tiger. University of
KORA Bericht Nr. 13
Chicago Press. Chicago. Illinois. USA.
Stollmann, A. 1963. Beitrag zur Kenntnis des Luchses, Lynx
lynx in den tschechoslowakischen Karpaten. Folia Zoologica, Brünn (Brno), 12: 301–316.
Turing, A. M. 1952. Phil. Trans. Roy. Soc. London. B. 237,
37.
Vasiliu, G. D. and P. Decei, 1964. Über den Luchs (Lynx
lynx) der rumänischen Karpaten. Säugetierkundliche Mitteilungen, München, 12: 155–183. (Bukarest).
Weigel, I. 1961. Das Fellmuster der wildlebenden Katzenarten und der Hauskatze in vergleichender und stammesgeschichtlicher Hinsicht. Säugetierkundliche Mitteilungen,
München, 120 pp.
Werdelin, L. and L. Olsson, 1997. How the Leopard got its
spots: a phylogenetic view of the evolution of felid coat
patterns. Biological Journal of the Linnean Society 62:
383–400.
Juni 2002
Appendix II
31
Form with all necessary information needed per individual and guidelines to fill out the form
Guidelines to fill out the form
ID Number:
Contact:
Date:
Country:
Population:
Region:
3-letter country code plus running number (e.g. SLK001)
who has filled out the form
when was the form filled out
country where the individual originated from
e.g. Carpathian Mountains or Zoo XY if zoo born
e.g. Mala Fatra
1. Origin of the Object
ID number:
Contact:
Date:
Country:
Population:
Region:
2. Information about the Animal
a.) Type of Object
‫ ٱ‬Monitored lynx in telemetry project
‫ ٱ‬dead lynx of Switzerland registered in a KORA database
‫ ٱ‬Lynx in museum
‫ ٱ‬Lynx in zoo
‫ ٱ‬Lynx pelt of hunters
‫ ٱ‬Random observation
‫ ٱ‬others________________________________________
b.) Location of lynx/pelt
Monitored Lynx
Name of lynx:
‫ ٱ‬alive
‫ ٱ‬dead
‫ ٱ‬unknown
Dead lynx in database
Name of lynx:
Db-Nr:
Lynx in museum
Museum:
Object number:
Origin (last location):
Date of death:
Location of picture:
Address of the museum:
Street, nr.:
Postcode, City:
Phone:
e-mail:
Lynx in zoo
Zoo:
Origin:
Name of lynx:
‫ ٱ‬wild caught
‫ ٱ‬zoo born
‫ ٱ‬if zoo born generation no:
Address of zoo:
Street, nr.:
Postcode, City:
Phone.:
e-mail:
Lynx pelt with hunters
Date when shot:
Random observation:
Data of observation:
c.) Details of the animal
sex:
‫ ٱ‬male
age:
at death:
relatives known:
‫ ٱ‬female
‫ ٱ‬unknown
on picture:
age group:
‫ ٱ‬adult
‫ ٱ‬yes
‫ ٱ‬no
‫ ٱ‬subadult
Date picture taken:
‫ ٱ‬juvenile
‫ ٱ‬unknown
‫ ٱ‬Mother
ID Number:
Name:
‫ ٱ‬Father
ID Number:
Name:
‫ ٱ‬Brother/Sister
ID Number:
Name:
‫ ٱ‬Brother/Sister
ID Number:
Name:
32
KORA Bericht Nr. 13
Appendix III
Spots-Counting rules
Single spots
Large and small spots:
Every identifiable single spot was
counted as one spot.
Spots in V-form were counted as
one spot.
Spots in groups
Large and small spots:
Every identifiable spot arranged in
stripes
other arrangements
was counted as one spot.
Strips were counted as one spot if
no interruptions were identifiable.
Spots in rosettes or small blotches
Rosettes and rudimentary rosettes:
Every identifiable spot arranged in
rosettes or small blotches
rudimentary of rosettes
Without spots
was counted as one spot.
no counting of spots
Juni 2002
Appendix IV
33
Database of Lynx - Pictures
Pictures of lynx taken by KORA members are maintained in a MSAccess database, used to recognize lynx from a
photo-trap picture and to examine coat patterns. All the available lynx could provide information on coat patterns
from Switzerland. The database is composed of all available pictures from captured lynx, recordings lynx found
dead, photo-trap pictures, randomly taken pictures, pictures of mounts and pictures of other animals.
The pictures are saved as a JPEC-files in two sizes: in a previewed form of max. 30 KB (in Fox_k6, in E:/PicDB/
Preview) and in a full-picture version on a CD.
Insert of new data-records
Screenshot of the MSAccess-database Real2: overview and „Path“ and „Switchboard“.
“Path”:
“Switchboard”:
contains the path of the “Previews”.
The following option can be chosen:
- Entry of Pictures
- Entry of new Keywords
- Search in PicDB
- Exit Database
If “Entry of Pictures” is chosen the enter-mask appears. This consists six pages.
34
Appendix IV
KORA Bericht Nr. 13
Screenshot of the enter-mask, Page 1
Page 1:
Enter file-name of the “Preview”-picture (lynxnameDate_number.jpg (e.g.: balu011299_1.jpg): 1)
name of the lynx (four letters), unknown lynx are named
with an “U” and three numbers; 2)
recording date (in ddmmyy); 3) underline; 4) serial picture-number of the same lynx.
Page 2:
This page only appears when “lynx” has been chosen in the index.
The pictures are sorted after the examination-regions: “Alpen80”, “Jura” and “Alpen90”.
In the “Coat”-list, one of the four (intuitively deducted) coat pattern types can be chosen. The criteria of the allocation are in progress.
Page 3:
This page only appears if it is other animals than lynx.
Page 4:
Enter 1) recording date (ddmmyy); 2) coordinates and name of the recording places 3) author; 4)
current location and material-type (slide, negative, photo) of the picture.
Page 5:
Enter the data-name of the full-picture version (same like “Preview”-file).
“CDAlpen90Fang” appears as a default.
Page 6:
Enter the initials of the reader and the date.
Bisher erschienene KORA Berichte
KORA Bericht Nr. 1
Landry, J.M., 1997. La bête du Val Ferret.
KORA Bericht Nr. 2
Landry, J.M., 1998. L'utilisation du chien de protection dans les Alpes suisses: une première analyse.
KORA Bericht Nr. 3
Workshop on Human Dimension in Large Carnivore Conservation. Contributions to the
Workshop 26.11.97 at Landshut, Switzerland, with Prof. Dr. Alistair J. Bath. 1998.
KORA Bericht Nr. 4
Zimmermann, F., 1998. Dispersion et survie des Lynx (Lynx lynx) subadultes d'une population réintroduite dans la chaîne du Jura.
KORA Bericht Nr. 2 d
Landry, J.M., 1999. Der Einsatz von Herdenschutzhunden in den Schweizer Alpen: erste
Erfahrungen.
KORA Bericht Nr. 2 e
Landry, J.M., 1999. The use of guard dogs in the Swiss Alps: A first analysis.
KORA Bericht Nr. 5 d
Angst, C., Olsson, P., Breitenmoser, U., 2000. Übergriffe von Luchsen auf Kleinvieh und
Gehegetiere in der Schweiz. Teil I: Entwicklung und Verteilung der Schäden.
KORA Bericht Nr. 6
Laass, J., 2001. Zustand der Luchspopulation im westlichen Berner Oberland im Winter
2000. Fotofallen-Einsatz Nov./Dez. 2000.
KORA Bericht Nr. 7 e
Breitenmoser-Würsten, Ch., Breitenmoser, U., (Eds), 2001. The Balkan Lynx Population - History, Recent Knowledge on its Status and Conservation Needs.
KORA Bericht Nr. 8
Ryser-Degiorgis Marie-Pierre, 2001. Todesursachen und Krankheiten beim Luchs – eine
Übersicht.
KORA Bericht Nr. 9
Breitenmoser-Würsten Christine, Zimmermann Fridolin, Ryser Andreas, Capt Simon,
Lass Jens, Breitenmoser Urs, 2001. Untersuchungen zur Luchspopulation in den Nordwestalpen der Schweiz 1997–2000.
KORA Bericht Nr. 11 d Breitenmoser Urs, Capt Simon, Breitenmoser-Würsten Christine, Angst Christof, Zimmermann Fridolin, Molinari-Jobin Anja, 2002. Der Luchs im Jura – Eine Übersicht zum
aktuellen Kenntnisstand.
KORA Bericht Nr. 11 f Breitenmoser Urs, Capt Simon, Breitenmoser-Würsten Christine, Angst Christof, Zimmermann Fridolin, Molinari-Jobin Anja, 2002. Le Lynx dans le Jura – Aperçu de l‘état
actuel des connaissances.
KORA Bericht Nr. 12 e Boutros Dominique, 2002. Characterisation and Assessment of Suitability of Eurasian
Lynx (Lynx lynx) Den Sites.
KORA Bericht Nr. 13 e Thüler Karin, 2002. Spatial and Temporal Distribution of Coat Patterns of Eurasian Lynx
(Lynx lynx) in two reintroduced Populations in Switzerland.
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