(PDF) Cranial morphometry of the Eurasian lynx (Lynx lynx L.) from Croatia

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VETERINARSKI ARHIV 80 (3), 393-410, 2010

Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from

Croatia

Tomislav Gomerčić

1

*, Magda Sindičić

2

, Martina Đuras Gomerčić

3

, Goran

Gužvica

4

, Alojzije Frković

5

, Dubravka Pavlović

6

, Josip Kusak

1

, Ana Galov

7

,

and Đuro Huber

1

Department of Biology, Faculty of Veterinary Medicine, University of Zagreb, Zagreb, Croatia

2

Department of Game Biology, Pathology and Breeding, Faculty of Veterinary Medicine, University of Zagreb,

Zagreb, Croatia

3

Department of Anatomy, Histology and Embryology, Faculty of Veterinary Medicine, University of Zagreb,

Zagreb, Croatia

4

Department of Ecogeography, Oikon Ltd, Institute for Applied Ecology, Zagreb, Croatia

5

Kvarnerska 43, Zagreb, Croatia

6

Medoka doo, Zagreb, Croatia

7

Department of Animal Physiology, Faculty of Science, University of Zagreb, Croatia

GOMERČIĆ, T., M. SINDIČIĆ, M. ĐURAS GOMERČIĆ, G. GUŽVICA,

A. FRKOVIĆ, D. PAVLOVIĆ, J. KUSAK, A. GALOV, Đ. HUBER

: Cranial

morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia. Vet. arhiv 80, 393-

410, 2010.

ABSTRACT

The last specimens of the autochthonous Eurasian lynx (

Lynx lynx

) in Croatia were exterminated in

Primorsko-Goranska County in around 1903. Lynx dispersed back to Croatia after six animals were reintroduced

to Slovenia from Slovakia in 1973. Today, lynx are under legislative protection in Croatia and its population

is deﬁ ned as highly endangered. Cranial morphometry was used to analyse a sample of 58 skulls obtained

from animals of the reintroduced population in order to identify geographical variation, sexual dimorphism and

sex-linked cranial characters. Eighty-ﬁ ve cranial measurements were recorded from each skull. Males were

signiﬁ cantly larger than the females in 39 of the 85 cranial measurements, whereas only one measurement, the

postorbital constriction, was greater in females. Our research indicates that lynx in Croatia are more similar to

lynx from Czech than from Norway. No evidence was found for differences in cranial measurements between

the Croatian reintroduced and the Balkan autochthonous lynx population, so there is no reason to deﬁ ne the later

population as a separate subspecies,

Lynx lynx martinoi

ssp.

Key words

: Eurasian lynx,

Lynx lynx

, skull, cranial measurements

*Corresponding author:

Tomislav Gomerčić, PhD, Department of Biology, Faculty of Veterinary Medicine, University of Zagreb, Heinzelova 55, 10000

Zagreb, Croatia, Phone: +385 1 2390 158; Fax: +385 1 2390 158; E-mail: tomislav.gomercic@vef.hr

394 Vet. arhiv 80 (3), 393-410, 2010

Introduction

Once spread all over Europe, Eurasian lynx (

Lynx lynx

L.

)

disappeared from most of

its habitat during the 18

th

and 19

th

centuries, with populations remaining only in remote

areas of Finland, Scandinavia, Siberia, Poland, and the Carpathian and Balkan peninsula

mountains (Kosovo, West Macedonia, Albania) (

BIENIEK et al., 1998; SOLDO, 2001;

HRISTOVSKI, 2001;

PAUNOVIĆ et al., 2001; BEGO, 2001; ZLATANOVA et al., 2001; SPASSOV

et al., 2001;

PANAYOTOPOULOU, 2001; BREITENMOSER-WÜRSTEN and BREITENMOSER,

2001

). Habitat loss, depletion of lynx prey and intensive hunting (

BREITENMOSER et

al., 1998

) were also reasons why the indigenous Eurasian lynx population disappeared

from the Dinaric Mountains. The last specimens of indigenous Eurasian lynx in Croatia

were exterminated in the Primorsko-Goranska County in around 1903 (

FRKOVIĆ, 2001

).

During the 20

th

century lynx were not present in Croatia for over 70 years. However,

Eurasian lynx dispersed to Croatia after three females and three males were reintroduced

to Slovenia from Slovakia in 1973

(

ČOP, 1988

). The size of the recent population in Croatia

is estimated to be up to 60 animals (

MAJIĆ-SKRBINŠEK, 2004

) and lynx in Croatia have

been a strictly protected species since 1995 (

ANON

YM

., 1995

).

Lynx cranial measurements, as one of the relevant ecological traits, have been

studied by numerous authors (

SAUNDERS, 1964; KVAM, 1983; ANDERSEN and WIIG,

1984; GARCIA-PEREA et al.,

1985; WIIG and ANDERSEN, 1986; WIIG and ANDERSEN, 1988;

MIRIĆ and PAUNOVIĆ, 1992;

BELTRAN and DELIBES, 1993; ČERVENÝ and KOUBEK, 2000;

PERTOLDI et al., 2006

), but little quantitative data on the physical characteristics of the

indigenous (extinct) (

MIRIĆ, 1972; MIRIĆ, 1978

) and reintroduced (present) (

GOMERČIĆ

et al., 2009

) Eurasian lynx population from Croatia are published. On the contrary, there

are many scientiﬁ cally unproved theories that animals from the indigenous population

were smaller than the animals from the reintroduced population (

SOVILJ, 2008

).

The goal of this paper was to study the morphometrical characteristics of skulls

belonging to the Croatian reintroduced lynx population in order to identify geographical

variations, sexual dimorphism and sex-linked cranial characters. Especial emphasis was

placed on a comparison of the now present, reintroduced Croatian lynx population with

cranial data of the autochthonous lynx population present in Croatia until the 1900´s.

Materials and methods

Skulls of 58 Eurasian lynx were examined morphometrically, 16 males, 35 females,

and seven specimens of unknown sex. Fifty-four skulls were trophy skulls obtained for

examination from private collections in the Primorsko-Goranska County, Croatia while

four skulls were deposited at the Department of Biology, Faculty of Veterinary Medicine

University of Zagreb, Croatia. The cause of death was known for 47 of the examined

specimens: 43 skulls belonged to animals shot in a hunt, two were caught in traps, and

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

395

Vet. arhiv 80 (3), 393-410, 2010

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

two were killed in trafﬁ c accidents. The location and time of the origin was known for 46

skulls. They were collected in the period from 1980 to 2004 in the Primorsko-Goranska

County, Croatia. Data on the body mass was known for 47 specimens.

Cranial measurements were taken using digital calipers and read to one tenth of a

millimeter. Eighty-ﬁ ve cranial measurements were obtained from each skull (Fig. 1).

Thirty-seven measurements were similar for the left and for the right side of the skull. In

the later analyses the mean value of the summarized left and right side value was used. In

total 43 measurements were related to the cranial bones and 42 to the teeth. The recorded

cranial measurements comprised those used by

VON DEN DRIESCH (1976)

,

WERDELIN

(1981)

,

WIIG and ANDERSEN (1986)

,

GARCÍA-PEREA et al. (1985)

. For comparison with

other lynx species and populations 19 cranial measurements were appropriate.

Fig. 1. Measurements of lynx skull, mandible, and teeth; the same ID number is used for the

beginning and end point of the measurement

396 Vet. arhiv 80 (3), 393-410, 2010

Fig. 1. Measurements of lynx skulls, mandibles, and teeth

1 - Proﬁ le length: from

Prosthion (rostral end of the interincisive suture) to Akrokranion (the most caudal

median point)

2 - (left and right) Condylobasal length: from Prosthion to the most caudal point of the occipital condyle

3 - Median palatal length: from Prosthion to Staphylion (tip of the caudal end of the median palatine suture)

4 - (left and right) Length from the most rostral point of the orbit to Akrokranion

5 - (left and right) Maxillar dental length: from Prosthion to the caudal margin of the fourth upper premolar

(P

4

) alveolus

6 - (left and right) Length from the rostral margin of the upper caninus (C

1

) alveolus to the caudal margin of the

upper third premolar (P

3

) alveolus

7 - Zygomatic width: greatest width between the zygomatic arches

8 - Postorbital constriction: least width caudal to the zygomatic processes of the frontal bones

9 - Interorbital constriction: least supraorbital width

10 - Mastoid width: greatest width between the mastoid processes of the temporal bones

11 - Condylar width: width between the most lateral points of the occipital condyles

12 - Rostral width: between the lateral margins of the C

1

alveoli

13 - Maxillar dental width: between the lateral margins of P

4

alveoli

14 - Skull width between the tips of the zygomatic processes of the frontal bones

15 - Skull width between the tips of the frontal processes of the zygomatic bones

16 - (left and right) Length of the zygomatic arch: from the caudal margin of the infraorbital foramen to the most

caudal point of the zygomatic process of the temporal bone

17 - (left and right) Length from Nasion (junction on the median plane of the right and left nasofrontal sutures)

to the tip of the frontal process of the zygomatic bone

18 - (left and right) Mandibular length: form the most rostral point of the mandible to the caudal end of the

angular process

19 - (left and right) Greatest mandibular length: from the most rostral point of the mandible to the most caudal

point of the condyloid process

20 - (left and right) Mandibular dental length: from the rostral margin of the lower ﬁ rst incisor (I

1

) alveolus to

the caudal margin of the lower ﬁ rst molar (M

1

) alveolus

21 - (left and right) Height of the ramus of the mandible: from the Coronion (dorsal border of the coronoid

process) to the ventral border of the angular process

22 - (left and right) Height from the Coronion to the middle point of the condyloid process

23 - (left and right) Height from the middle point of the condyloid process to the ventral border of the angular

process

24 - (left and right) Height of the body of the mandible at the caudal margin of the M

1

alveolus

25 - (left and right) Height of the body of the mandible at the rostral margin of the lower third premolar (P

3

)

alveolus

26 - (left and right) Length from the rostral margin of P

3

alveolus to caudal margin of the P

4

alveolus

27 - (left and right) Length from the rostral margin of P

3

alveolus to the caudal margin of M

1

alveolus

28a - (left and right) Lower caninus (C

1

)

width: between the lateral and medial surface of C

1

at the level of its

insertion into the alveolus

28b - (left and right) C

1

length: between the rostral and caudal surface of C

1

at the level of its insertion into the

alveolus

28c - (left and right) C

1

enamel height: from the tip of C

1

to the enamel margin

28d - (left and right) C

1

height: from the tip of C

1

to the margin of its alveolus

29a - (left and right) P

3

width: between the lateral and medial surface of P

3

at the level of its insertion into the

alveolus

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

397

Vet. arhiv 80 (3), 393-410, 2010

29b - (left and right) P

3

length: between the rostral and caudal surface of P

3

at the level of its insertion into the

alveolus

30a - (left and right) Lower fourth premolar (P

4

)

width: between the lateral and medial surface of P

4

at the level

of its insertion into the alveolus

30b - (left and right) P

4

length: between the rostral and caudal surface of P

4

at the level of its insertion into the

alveolus

31a - (left and right) M

1

width: between the lateral and medial surface of M

1

at the level of its insertion into the

alveolus

31b - (left and right) M

1

rostral length: from the rostral margin of M

1

to the middle notch

at the level of its

insertion into the alveolus

31c - (left and right) M

1

caudal

length: from the middle notch to the caudal margin of M

1

at the level of its

insertion into the alveolus

32a - (left and right) C

1

width: between the lateral and medial surface of C

1

at the level of its insertion into the

alveolus

32b - (left and right) C

1

length: between the rostral and caudal surface of C

1

at the level of the insertion into the

alveolus

32c - (left and right) C

1

enamel height: from the tip of C

1

to enamel margin

32d - (left and right) C

1

height: from the tip of C

1

to the border of its alveolus

33a - (left and right) P

3

width: between the lateral and medial surface of P

3

at the level of its insertion into the

alveolus

33b - (left and right) P

3

length: between the rostral and caudal surface of P

3

at the level of its insertion into the

alveolus

34a - (left and right) P

4

width without protoconus: between the lateral and medial surface of the P

4

at the level

of its insertion into the alveolus

34b - (left and right) P

4

length: between the rostral and caudal surface of P

4

at the level of its insertion into the

alveolus

34c - (left and right) P

4

width with protoconus: between the lateral and medial surface of P

4

at the level of its

insertion into the alveolus

34d - (left and right) P

4

protoconus length: between the rostral and caudal margins of the protoco

nus

In order to include only adult skulls in the statistical analysis of the cranial

measurements, the age of the examined specimens was determined. Age was determined

by enumeration of root cementum annuli (

KVAM, 1984

). The third upper incisor (I3) was

used for age determination. It was determined that a total of 11 skulls belonged to animals

younger than 3 years, so further analyses included 47 adult specimens.

The cranial measurements were analyzed separately for females and males to detect

sexual dimorphism and sex-linked cranial characters.

Results

The examined lynx skull sample included 16 males (31.4%), 35 females (68.6%), and

seven specimens of unknown sex. The age was successfully determined in 55 animals

(Fig. 2). Forty-four skulls belonged to animals aged three or more years. These skulls

were deﬁ ned as adult. Mean age for adults, animals older than three years, was 8.6 ± 3.4

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

398

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

T. Gomerč

i

ć et a

l

.: Cran

i

a

l

morp

h

ometry of t

h

e Euras

i

an

l

ynx

(Lynx lynx

Lynx

l

ynx

L.) from Croatia

L.

)

from Croat

i

a

Vet. arhiv 80 (3), 393-410, 2010

years and 6.6 ± 2.8 years for males and females, respectively. The oldest female aged nine

years. Six males were older than nine, with the oldest aged 15 years.

In three skulls the age could not be determined but they had hardly visible skull

bone sutures and were deﬁ ned as adult, too. Further analyses of body mass and cranial

measurements (Table 1) included 47 adult specimens, 15 males, 25 females, and seven

of unknown sex.

The body mass of the adult specimens ranged from 15 to 28 kg (mean: 21.9 kg), and

from 12 to 25 kg (mean: 18.4 kg) in males and females, respectively. Adult male lynx had

signiﬁ cantly greater body mass (P<0.01). Their body mass value was on average 16.2%

higher than in females.

Fig. 2. Age distribution of Eurasian lynx (

Lynx lynx

L.) from Croatia

Fig. 3. Zygomatic width/postorbital constriction ratio indicates a sex-linked cranial feature in

Eurasian lynx (

Lynx lynx

L.) from Croatia

Lynx age (year)

Number of individuals

Zygomatic width (mm)

Postorbital constriction (mm)

399

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

T. Gomerč

i

ć et a

l

.: Cran

i

a

l

morp

h

ometry of t

h

e Euras

i

an

l

ynx

(Lynx lynx

Lynx

l

ynx

L.) from Croatia

L.

)

from Croat

i

a

Vet. arhiv 80 (3), 393-410, 2010

Table 1. Age, body mass, and cranial measurements of adult Eurasian lynx (

Lynx lynx

L.) from Croatia (mean, standard deviation,

range). Results of

t

-test for sexually dimorphic characters (p) and sexual difference (%).

Male (n = 15) Female (n = 25)

Male, female and unknown

sex (n = 47) p

Sexual

difference

(%)

Age (year) 8.6 ± 3.4 (4-15) 5.8 ± 1.7 (3-9) 6.6 ± 2.8 (3-15) 0.0025** 32.7

Body mass

(kg) 21.9 ± 3.9 (15-28) 18.4 ± 3.2 (12-25) 19.8 ± 3.7 (12-28) 0.0043** 16.2

Measurement (mm) #

1. 153.2 ± 7.4 (135.6-162.8) 145.5 ± 5.9 (133-154.5) 149.6 ± 7.8 (133-168.3) 0.0008** 5.0

2. 142.8 ± 3.7 (136.3-148.9) 133.5 ± 5.3 (121-141.5) 137.9 ± 6.7 (121-152.9) 0.0000** 6.5

3. 59.1 ± 1.5 (57-62.6) 55.8 ± 1.9 (52.5-58.9) 57.5 ± 2.5 (52.5-63.5) 0.0000** 5.6

4. 122.7 ± 2.9 (116.8-127.5) 115 ± 4.7 (103.4-121.7) 118.6 ± 5.6 (103.4-130) 0.0000** 6.3

5. 61.4 ± 1.5 (58.7-64) 58.3 ± 1.6 (54.8-60.9) 59.8 ± 2.2 (54.8-65) 0.0000** 4.9

6. 30.4 ± 1.1 (28.8-32.5) 28.4 ± 1.1 (25.8-30.4) 29.4 ± 1.5 (25.8-32.5) 0.0000** 6.6

7. 111.3 ± 3.2 (106.5-116.7) 103 ± 4.5 (90.3-108.6) 106.5 ± 5.7 (90.3-117.1) 0.0000** 7.5

8. 39.3 ± 2 (33.6-41.8) 40.5 ± 1.3 (37.9-42.3) 40.1 ± 1.6 (33.6-42.3) 0.0233* -3.1

9. 33.2 ± 1.9 (30.2-36.6) 30.6 ± 2.1 (25.3-33.2) 31.9 ± 2.4 (25.3-36.9) 0.0003** 8.0

10. 67.7 ± 1.9 (65-70.7) 63.9 ± 2.3 (58.5-67.4) 65.6 ± 2.9 (58.5-71.7) 0.0000** 5.6

11. 32.2 ± 1 (30.2-33.6) 31.4 ± 1 (29.2-33) 31.8 ± 1.1 (29.2-33.6) 0.0227* 2.4

12. 40.6 ± 1.5 (37.8-43) 37.5 ± 1.5 (33.6-39.7) 38.8 ± 2.1 (33.6-43) 0.0000** 7.5

13. 62.8 ± 1.4 (60-64.8) 59.9 ± 1.3 (57.1-61.5) 61.2 ± 1.9 (57.1-64.8) 0.0000** 4.6

14. 74.3 ± 2.6 (69.2-77.2) 70 ± 4.2 (58.2-74.4) 71.7 ± 4.2 (58.2-79.1) 0.0039** 5.8

15. 87.1 ± 2.8 (83.2-92.1) 83 ± 3.2 (72.9-87.7) 85 ± 3.6 (72.9-92.1) 0.0003** 4.7

16. 77 ± 2.2 (72.6-79.5) 72.3 ± 3.4 (63-75.8) 74.6 ± 3.9 (63-83.3) 0.0000** 6.0

17. 52.9 ± 1.3 (51.4-55.5) 50.2 ± 2.2 (45.3-53.4) 51.5 ± 2.3 (45.3-55.8) 0.0001** 5.0

18. 106.8 ± 3.5 (101-114.5) 99 ± 4.7 (87.5-104.8) 102.5 ± 5.8 (87.5-114.6) 0.0000** 7.3

19. 106.7 ± 2.8 (102-110.5) 99.4 ± 4.4 (89-105.3) 102.6 ± 5.2 (89-113.1) 0.0000** 6.8

20. 61.6 ± 1.6 (59.3-64.4) 58.3 ± 1.6 (54.9-60.8) 59.7 ± 2.2 (54.9-64.4) 0.0000** 5.4

21. 46.6 ± 2 (43.6-49.6) 42 ± 2.2 (36.7-44.6) 44.1 ± 3.2 (36.7-51.7) 0.0000** 9.8

400

22. 29.7 ± 1.5 (27.1-32) 27 ± 1.7 (22-29.1) 28.2 ± 2.1 (22-32.4) 0.0000** 8.9

23. 17.4 ± 1.1 (15.6-19.7) 15.4 ± 1 (13-17.1) 16.3 ± 1.4 (13-19.7) 0.0000** 11.2

24. 22.1 ± 1.1 (19.7-23.9) 20.3 ± 1.1 (18.1-22.3) 21.2 ± 1.4 (18.1-24.2) 0.0000** 8.2

25. 20.3 ± 0.8 (19-21.5) 19.2 ± 0.7 (17.8-20.2) 19.7 ± 1 (17.8-22.2) 0.0000** 5.7

26. 30.3 ± 1 (28.5-31.8) 29.3 ± 0.6 (28.3-30.5) 29.7 ± 0.9 (28.3-31.8) 0.0001** 3.5

27. 37.7 ± 1 (35.9-39.1) 36.6 ± 1 (34.7-38.8) 37.1 ± 1.1 (34.7-39.1) 0.0007** 3.0

28a. 6.9 ± 0.4 (6-7.6) 6.4 ± 0.2 (5.8-6.8) 6.7 ± 0.4 (5.8-7.6) 0.0000** 7.8

28b. 9 ± 0.5 (8-9.5) 8.4 ± 0.4 (7.6-9.4) 8.7 ± 0.5 (7.6-9.5) 0.0000** 7.5

28c. 17.5 ± 1 (15.4-19.3) 16.9 ± 0.8 (14.5-18.6) 17.1 ± 1.2 (11.9-19.3) 0.0004** 3.7

28d. 20.9 ± 2.7 (11.8-23.6) 19.9 ± 3.3 (5.6-25.9) 20.3 ± 3 (5.6-25.9) 0.3012 5.0

29a. 5.4 ± 0.3 (4.9-6) 5.3 ± 0.3 (4.8-6.1) 5.4 ± 0.3 (4.8-6.1) 0.7021 0.7

29b. 10.3 ± 0.4 (9.7-11) 9.9 ± 0.4 (8.7-10.5) 10.1 ± 0.5 (8.7-11.7) 0.0016** 4.0

30a. 5.9 ± 0.3 (5.5-6.4) 5.8 ± 0.4 (4.9-6.5) 5.9 ± 0.3 (4.9-6.5) 0.2608 2.0

30b. 12.4 ± 0.4 (11.8-13.4) 12.1 ± 0.4 (11.4-12.8) 12.2 ± 0.4 (11.4-13.4) 0.0083** 2.8

31a. 6.6 ± 0.2 (6.2-7) 6.5 ± 0.2 (5.9-6.9) 6.5 ± 0.2 (5.9-7) 0.1338 1.8

31b. 6.8 ± 0.4 (6-7.5) 6.5 ± 0.2 (6.1-7.1) 6.7 ± 0.3 (6-7.5) 0.0083** 4.0

31c. 10 ± 0.3 (9.3-10.3) 9.9 ± 0.3 (9.4-10.6) 9.9 ± 0.3 (9.3-10.6) 0.1156 1.6

32a. 7.8 ± 0.4 (7.2-8.6) 7.2 ± 0.3 (6.6-8) 7.5 ± 0.4 (6.6-8.6) 0.0000** 7.0

32b. 9.7 ± 0.5 (8.8-10.3) 9 ± 0.5 (8.5-10.7) 9.3 ± 0.6 (8.5-10.7) 0.0001** 7.0

32c. 18.4 ± 1.6 (14.5-20.2) 17.4 ± 2.8 (4.9-20.3) 18 ± 2.4 (4.9-20.9) 0.2481 5.2

32d. 23.5 ± 5.1 (6.5-27.7) 22.8 ± 2.9 (11.4-28.3) 23.3 ± 3.6 (6.5-28.3) 0.5399 3.3

33a. 6.4 ± 0.4 (5.6-6.9) 6.3 ± 0.3 (6.1-6.9) 6.4 ± 0.3 (5.6-7) 0.2868 1.6

33b. 11.9 ± 0.5 (10.9-13) 11.6 ± 0.3 (11-12.5) 11.7 ± 0.4 (10.9-13) 0.0085** 3.0

34a. 6.9 ± 0.2 (6.5-7.4) 6.7 ± 0.3 (6.3-7.5) 6.8 ± 0.3 (6.3-7.5) 0.0160* 3.0

34b. 18.4 ± 0.8 (16.6-19.3) 17.9 ± 0.4 (17.4-18.8) 18.1 ± 0.6 (16.6-19.5) 0.0000** 2.6

34c. 8.5 ± 0.4 (8-9.4) 8.2 ± 0.3 (7.7-8.7) 8.3 ± 0.4 (7.7-9.4) 0.0017** 4.2

34d. 3.2 ± 0.3 (2.8-3.9) 3.1 ± 0.2 (2.6-3.5) 3.1 ± 0.28 (2.6-3.9) 0.1306 4.3

*- P<0.05; **- P<0.01;

#

- the list and description of measurements are shown in Materials and methods

Table 1. (Continued)

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

Vet. arhiv 80 (3), 393-410, 2010

401

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

Vet. arhiv 80 (3), 393-410, 2010

Table 2. Cranial measurements and body mass of the genus Lynx

Lynx (Lynx lynx)

from Croatia (this

study)a

Lynx (Lynx

canadensis) from

Newfoundland

(SAUNDERS.

1964)b

Iberian lynx

(Lynx pardinus)

(GARCÍA-PEREA

et al., 1985)

Lynx (Lynx lynx)

from Norway.

(WIIG and

ANDERSEN,

1986)c

Lynx (Lynx lynx)

from Czech.

(ČERVENÝ and

KOUBEK, 2000)

Lynx (Lynx lynx

from the Balkan

(MIRIĆ, 1978,

1981)

Body

mass

(kg)

male 21.9 ± 3.9 n = 15

(15.0-28.0)

23.5 n = 94

(14.0-38.0)

21.8 ± 3.6 n = 25

(14.0-28.0)

female 18.4 ± 3.2 n = 25

(12.0-25.0)

18.9 n = 91

(11.0-26.0)

16.8 ± 2.7 n = 17

(13.0-22.0)

Measurement (mm)d

1.

male 153.2 ± 7.4 n = 15

(135.6-162.8) 129 ± 4.1 n = 23** 133.7 ± 2.5 n = 12

(128.7-137.8)** 153.1 ± 5.1 n = 75 153.3 ± 6.7 n = 43

(137.0-165.1) 151.8 n = 1

female 145.5 ± 5.9 n = 25

(133.0-154.5) 122 ± 3.0 n = 13** 121.8 ± 4.2 n = 8

(113.6-126.2)** 143.0 ± 4.4 n = 56* 145.4 ± 6.3 n = 32

(130.2-157.1)

141.3 ± 1.2 n = 3

(140.0-142.3)

2.

male 142.8 ± 3.7 n = 15

136.3-148.9

121.7 ± 2.6 n = 10

(116.8-125.4)**

139.0 ± 4.4 n =

75**

139.3 ± 9.1 n = 43

(122.0-159.1) 136.5 n = 1

female 133.5 ± 5.3 n = 25

(121.0-141.5)

110.0 ± 4.0 n = 7

(102.6-114.9)**

130.5 ± 3.8 n =

56**

131.4 ± 4.2 n = 32

(121.5-141.5)

127.8 ± 1.4 n = 3

(126.5-129.3)

3.

male 59.1 ± 1.5 n = 15

(57.0-62.6)

52.0 ± 1.6 n = 10

(49.9-55.4)** 58.9 ± 2.1 n = 75 57.1 ± 2.8 n = 43

(51.2-66.4)*

female 55.8 ± 1.9 n = 25

(52.5-58.9)

46.8 ± 2.1 n = 9

(43.8-49.7)** 55.3 ± 1.9 n = 56 54.8 ± 1.9 n = 32

(49.8-58.5)

5.

male 61.4 ± 1.5 n = 15

(58.7-64.0) 60.8 ± 2.0 n = 75 54.7 ± 2.0 n = 43

(50.3-58.3)**

female 58.3 ± 1.6 n = 25

(54.8-60.9) 57.4 ± 1.7 n = 56* 52.9 ± 1.7 n = 32

(48.7-56.0)**

402

7.

male 111.3 ± 3.2 n = 15

(106.5-116.7) 94 ± 3.2 n = 24** 95.6 ± 2.3 n = 10

(92.9-100.8)**

101.3 ± 3.7 n =

75**

107.7 ± 5.6 n = 43

(97.0-118.2)* 106.1 n = 1

female 103.0 ± 4.5 n = 25

(90.3-108.6) 89 ± 1.9 n = 13** 86.5 ± 4.3 n = 10

(80.9-93.5)** 95.4 ± 3.1 n = 56** 102.2 ± 3.4 n = 32

(92.3-108.6)

99.6 ± 2.5 n = 3

(97.8-102.5)

8.

male 39.3 ± 2.0 n = 15

(33.6-41.8)

39.8 ± 1.4 n = 12

(38.1-42.7) 37.8 ± 1.6 n = 75** 40.4 ± 2.7 n = 43

(32.8-46.8)

female 40.5 ± 1.3 n = 25

(37.9-42.3)

40.4 ± 1.1 n = 10

(39.2-42.6) 38.6 ± 1.9 n = 56** 39.2 ± 2.1 n = 32

(34.3-43.4)**

9.

male 33.2 ± 1.9 n = 15

(30.2-36.6) 33.1 ± 1.6 n = 75 34.7 ± 3.3 n = 43

(28.4-44.0) 31.5 n = 1

female 30.6 ± 2.1 n = 25

(25.3-33.2) 31.0 ± 1.4 n = 56** 33.1 ± 2.7 n = 32

(28.5-39.8)**

32.6 ± 1.4 n = 3

(31.2-34.0)

10.

male 67.7 ± 1.9 n = 15

(65.0-70.7)

58.2 ± 1.3 n = 11

(55.4-60.0)** 65.4 ± 2.0 n = 75** 65.9 ± 3.7 n = 43

(56.4-72.2)

female 63.9 ± 2.3 n = 25

(58.5-67.4)

53.5 ± 2.2 n = 8

(50.5-56.3)** 61.2 ± 1.7 n = 56** 62.4 ± 2.8 n = 32

(56.6-68.2)*

11.

male 32.2 ± 1.0 n = 15

(30.2-33.6) 33.9 ± 1.0 n = 75**

female 31.4 ± 1.0 n = 25

(29.2-33.0) 32.7 ± 1.0 n = 56**

12.

male 40.6 ± 1.5 n = 15

(37.8-43.0)

33.7 ± 0.9 n = 13

(32.0-35.1)** 39.2 ± 1.4 n = 75**

female 37.5 ± 1.5 n = 25

(33.6-39.7)

30.7 ± 1.6 n = 10

(28.1-32.4)** 36.7 ± 1.3 n = 56*

13.

male 62.8 ± 1.4 n = 15

(60.0-64.8) 61.6 ± 1.8 n = 75*

female 59.9 ± 1.3 n = 25

(57.1-61.5) 58.9 ± 1.7 n = 56*

Table 2. (Continued)

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

Vet. arhiv 80 (3), 393-410, 2010

403

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

Vet. arhiv 80 (3), 393-410, 2010

14.

male 74.3 ± 2.6 n = 15

(69.2-77.2)

63.8 ± 2.4 n = 8

(58.2-66.2)**

69.5 ± 4.4 n = 43

(57.9-78.2)**

female 70.0 ± 4.2 n = 25

(58.2-74.4)

57.7 ± 3.8 n = 7

(53.5-62.6)**

67.1 ± 2.7 n = 32

(61.6-73.4)**

18.

male 106.8 ± 3.5 n = 15

(101.0-114.5)

89.0 ± 2.1 n = 12

(86.4-94.2)**

102.3 ± 3.5 n =

74** 106.9 n = 1

female 99.0 ± 4.7 n = 25

(87.5-104.8)

79.7 ± 3.2 n = 11

(74.8-84.4)** 95.6 ± 3.0 n = 56** 99.9 ± 1.5 n = 3

(98.5-101.4)

19.

male 106.7 ± 2.8 n = 15

(102.0-110.5)

89.6 ± 1.9 n = 12

(87.5-94.5)**

103.1 ± 3.5 n = 43

(95.0-110.5)**

female 99.4 ± 4.4 n = 25

(89.0-105.3)

81.2 ± 3.1 n = 11

(76.2-85.3)**

99.5 ± 2.7 n = 32

(91.8-103.7)

20.

male 61.6 ± 1.6 n = 15

(59.3-64.4)

48.9 ± 1.0 n = 12

(47.5-50.3)** 56.7 ± 1.7 n = 74**

female 58.3 ± 1.6 n = 25

(54.9-60.8)

44.3 ± 1.6 n = 10

(41.9-46.5)** 53.6 ± 1.6 n = 56**

21.

male 46.6 ± 2.0 n = 15

(43.6-49.6)

37.1 ± 1.1 n = 12

(36.0-39.4)** 46.8 ± 2.2 n = 74 45.3 ± 2.7 n = 43

(39.0-51.2)

female 42.0 ± 2.2 n = 25

(36.7-44.6)

32.4 ± 2.1 n = 11

(29.5-35.0)** 42.4 ± 1.9 n = 56 42.0 ± 2.2 n = 32

(38.4-49.2)

26.

male 30.3 ± 1.0 n = 15

(28.5-31.8)

24.6 ± 0.8 n = 14

(22.7-25.7)**

female 29.3 ± 0.6 n = 25

(28.3-30.5)

22.8 ± 1.0 n = 11

(20.6-23.7)**

34b.

male 18.4 ± 0.8 n = 15

(16.6-19.3)

15.4 ± 0.5 n = 11

(14.4-16.2)** 18.6 ± 0.7 n = 74 18.1 n = 2

(17.5-18.7)

female 17.9 ± 0.4 n = 25

(17.4-18.8)

14.3 ± 0.6 n = 11

(13.5-15.1)** 17.7 ± 0.8 n = 56 18.1 ± 1.3 n = 3

(17.2-19.5)

Table 2. (Continued)

404

34c.

male 8.5 ± 0.4 n = 15

(8.0-9.4)

7.2 ± 0.5 n = 11

(6.3-7.9)**

female 8.2 ± 0.3 n = 25

(7.7-8.7)

6.6 ± 0.4 n = 11

(5.9-7.3)**

* - signiﬁ cantly differs from the Croatian population (P<0.05)

** - signiﬁ cantly differs from the Croatian population (P<0.01)

a - animals at age of 3 years and older

b - adults with fused epiphysis of the humerus

c - animals of age between 17 and 60 months, the measurements corrected to the value of 33.9 months old lynx

d- the list and description of measurements is in Materials and methods

Table 2. (Continued)

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

Vet. arhiv 80 (3), 393-410, 2010

405

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

Vet. arhiv 80 (3), 393-410, 2010

Fig. 4. Maxillar dental width/postorbital constriction ratio indicates another sex-linked cranial

feature in Eurasian lynx (

Lynx lynx

L.) from Croatia

Fig. 5. Prominent external sagittal crest (crista sagittalis externa) in males (a: LS10, 8 years; b:

LS52, 15 years) compared to a female (c: LS51, 8 years)

Postorbital constriction (mm)

Maxilar dental width

406

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

Vet. arhiv 80 (3), 393-410, 2010

Males were signiﬁ cantly larger than the females in 39 (45.88%) of 85 cranial

measurements. Only one cranial measurement, namely the postorbital constriction (8), was

larger in females than in males. The value of the postorbital constriction was on average

3.1% higher in females than in males (Table 1). Values of male cranial measurements

were on average 5% higher than in females, ranging from 0.7% to 11.2%. The main

cranial characters such as the proﬁ le length (1) and the zygomatic width (7), were both

signiﬁ cantly greater in males (P<0.01). Sex-linked cranial features are indicated in the

zygomatic width/postorbital constriction ratio and maxillar dental width/postorbital

constriction ratio. Both ratios are smaller in the males (Fig. 3 and Fig. 4). None of the

nine teeth measurements (28d, 29a, 30a, 31a, 31c, 32c, 32d, 33a, 34d) showed any sexual

dimorphism (P>0.05). The mid-dorsal external sagittal crest (crista sagittalis externa) was

well developed in males, and in most male skulls it was very prominent, extending up to

the caudal border of the frontal bones. The external sagittal crest was not prominent in

any of the female skulls and did not extend as rostral as in males (Fig. 5).

Skull size showed a low correlation to the age and body mass. The proﬁ le length

showed no correlation with the age (R

2

= 0.0775) and a very low correlation with the

body mass (R

2

for both sexes = 0.1053, i.e. 0.148 and 0.116 for males and females,

respectively).

Discussion

Our statistical analysis included 85 cranial measurements of 47 Eurasian lynx skulls

from Croatia, belonging to animals older than 3 years. Namely,

GARCIA-PEREA (1991)

stated that lynx cranial measurements are age related and

ANDERSEN and WIIG (1984)

concluded that lynx skull growth stops at the age of three. The third upper incisor (I

3

)

has been used for age determination, which is different than the canine used in the work

of

CROWE (1972) and KVAM (1984)

. The I

3

has been used for age determination to avoid

signiﬁ cant damage of the trophy skulls. Cranial morphometry was used to identify sexual

dimorphism and sex-linked cranial characters, which have already been described in other

lynx populations (

GARCIA-PEREA et al., 1985; BELTRAN and DELIBES, 1993

).

The Croatian population of Eurasian lynx expressed sexual dimorphism in skull size

but also in body mass. Our results correspond with data of the Norwegian lynx (

WIIG

and ANDERSEN, 1986

), where 15 cranial measurements were signiﬁ cantly larger in males

than females, while only the postorbital constriction was signiﬁ cantly larger in females.

Even though

WERDELIN (1981)

did not establish any statistically signiﬁ cant differences

between male and female lynx skulls, because his study also included juvenile skulls,

both

WIIG and ANDERSEN (1986)

and

ČERVENÝ and KOUBEK (2000)

conﬁ rmed sexual

dimorphism in lynx skull size.

407

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

Vet. arhiv 80 (3), 393-410, 2010

Skulls of male lynx from Croatian can be distinguished from the female skulls by the

more prominent sagittal external crest and by two ratios: the zygomatic width/postorbital

constriction ratio and the maxillar dental width/postorbital constriction ratio, which are

both smaller in males.

Eurasian lynx from Croatia are signiﬁ cantly larger than the Iberian lynx (

Lynx

pardina

) (

GARCÍA-PEREA et al., 1985

) in 14 out of 15 cranial measurements (Table 2).

Cranial measurements of the Canadian lynx (

SAUNDERS, 1964

) are smaller than those

of the Croatian Eurasian lynx (this study) but the body mass does not show a signiﬁ cant

difference between these two lynx species (Table 2). Compared to other Eurasian lynx

populations the Croatian population shows particular differences. The lynx from Croatia

is more similar to lynx from the Czech Republic (

ČERVENÝ and KOUBEK, 2000

) than

those from Norway (

WIIG and ANDERSEN, 1986

). The difference between Norwegian and

Croatian lynx was established in 12 female and 9 male cranial measurements. Norwegian

lynx have only two measurements greater in female skulls and one measurement greater

in male skulls, while all other measurements were signiﬁ cantly smaller in skulls from

Norwegian lynx. There are most similarities in the cranial measurements of the Croatian

and Czech lynx populations. Out of 11 measurements compared only ﬁ ve are signiﬁ cantly

different in males and females, respectively. Only one cranial measurement is larger in the

Czech lynx (Table 2).

In his papers

MIRIĆ (1978, 1981)

deﬁ ned a Eurasian lynx subspecies indigenous to the

Balkan peninsula,

Lynx lynx martinoi

ssp. He based his statement on cranial measurements

of three female and one male lynx from this region. The values of the female skulls

measured by

MIRIĆ (1974)

did not show any statistically signiﬁ cant difference when

compared with female skulls from our study.

MIRIĆ (1978)

measured six measurements

in only one male skull and all of them were within the interval of two standard deviations

of the reintroduced population, meaning that the dimensions of this skull are within the

range of 95% of the dimensions of the skulls from the reintroduced lynx. Out of those six

measurements in total four (1, 9, 18, 34b) are within the range of one standard deviation

of the Croatian population of Eurasian lynx, meaning within 68% of the reintroduced

population.

MIRIĆ (1978)

also stated that the body mass of the new subspecies is not

larger than 25 kg. This value is not signiﬁ cantly different from the Croatian (21.9 ± 3.9

kg, range 15.0 - 28.0 kg) or Czech population (21.8 ± 3.6 kg, range 14.0 - 28.0 kg).

Also when deﬁ ning this new subspecies

MIRIĆ (1978)

stated that 30% of the animals

from Balkan Peninsula have less visible or have no spots at all on their fur, compared to

populations from Slovakia and the Caucasus mountains where only 10% of lynx have this

kind of fur coloration. It is important to emphasize that only few years earlier the same

author published an article saying that certain cranial measurements have proved to be a

very reliable indicator for lynx taxonomy, while fur coloration and body dimensions are

408 Vet. arhiv 80 (3), 393-410, 2010

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

T. Gomerč

i

ć et a

l

.: Cran

i

a

l

morp

h

ometry of t

h

e Euras

i

an

l

ynx

(Lynx lynx

Lynx

l

ynx

L.) from Croatia

L.

)

from Croat

i

a

so variable they cannot be used to distinguish lynx species (

MIRIĆ, 1974

). In the same

paper, based on cranial measurements of two male (one of them damaged and with a large

number of measurements missing) and three female lynx skulls,

MIRIĆ (1974)

concluded

that the Balkan lynx is completely the same as the European lynx, subspecies L.

l. lynx

L.

MIRIĆ

published the same idea in

1972

: “Comparing the craniometrical characteristics

of lynx from Yugoslavia we have seen that they do not differ from the craniometrical

characteristics of the typical subspecies

L. lynx lynx.

”. Even though the number of studied

skulls has not increased, but only the number of studied furs for coloration and spots on

pelts,

MIRIĆ (1978)

deﬁ ned a new Eurasian lynx subspecies

Lynx lynx martinoi

ssp. Our

research established that there is no signiﬁ cant difference between cranial measurements

of the Balkan (

MIRIĆ, 1978

), Croatian (this research) and Czech lynx populations

(

ČERVENÝ and KOUBEK, 2000

). Taking this into consideration and also the small number

of skulls

MIRIĆ (1978)

studied, it is not justiﬁ ed to deﬁ ne the Balkan lynx population as

a separate subspecies.

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osobitosti euroazijskoga risa (

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L

.

) iz

Hrvatske. Vet. arhiv 80, 393-410,

2010.

SAŽETAK

Posljednji primjerci autohtonog euroazijskog risa (

Lynx lynx

) u Hrvatskoj istrijebljeni su oko 1903. godine

na području Gorskog kotara. Ris se u Hrvatsku vratio nakon reintrodukcije šest životinja iz Slovačke u Sloveniju

1973. godine. Ris je danas u Hrvatskoj ugrožena, zakonom zaštićena vrsta. Kranijalna morfometrija rabljena

je u analizi 58 lubanja risova iz reintroducirane populacije, s ciljem određivanja geografskih razlika, spolnog

dimorfizma i spolno vezanih značajki lubanje. Ukupno je 85 mjera izmjereno na svakoj lubanji. Mužjaci su bili

značajno veći u 39 od 85 mjera lubanje, dok je samo jedna mjera, najmanja širina lubanje aboralno od jagodičnih

izdanaka čeonih kostiju, veća u ženki nego u mužjaka. Naše istraživanje pokazuje da je ris iz Hrvatske sličniji

risu iz Češke nego onima iz Norveške. Nisu pronađeni dokazi o postojanju razlika u mjerama lubanje između

reintroducirane populacije risa u Hrvatskoj i autohtone balkanske populacije, te stoga ne postoje temelji za

definiranje balkanskog risa kao odvojene podvrste,

Lynx lynx martinoi

ssp.

Ključne riječi:

kraniometrija, ris,

Lynx lynx

, Hrvatska

Received: 7 April 2009

Accepted: 22 December 2009

T. Gomerčić et al.: Cranial morphometry of the Eurasian lynx (

Lynx lynx

L.) from Croatia

Vet. arhiv 80 (3), 393-410, 2010

Cranial morphometry of the Eurasian lynx (Lynx lynx L.) from Croatia

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