Multi-indicator method for assessing the risk status Case study on Norwegian cattle breeds

Transcription

1 Multi-indicator method for assessing the risk status Case study on Norwegian cattle breeds Jessica Tetteroo

2 Multi-indicator method for assessing the risk status Case study on Norwegian cattle breeds Jessica Tetteroo Internship Animal Breeding and Genetics (ABG-7424) NordGen, Ås Norway November 217 Supervisors NordGen: Anne Kettunen and Peer Berg Supervisor WUR: Bart Ducro i

3 Preface This internship report is written as part of my graduation project for the Master Animal Sciences at Wageningen University and Research in Wageningen. The internship concerns a study on the risk status of six native Norwegian cattle breeds using a multi-indicator method. The internship has been performed from Augustus 217 till November 217 at NordGen section Farm Animals located in Ås, Norway. During my internship I have been guided by my supervisors Anne Kettunen and Peer Berg. I want to express my appreciation to Anne and Peer for their guidance and support during the process. Besides, I would like to thank NordGen for the opportunity to do my internship at NordGen. I enjoyed being part of the NordGen team and the daily activities. I also wish to thank the Norwegian Genetic Resource Centre, and in particular Nina Sæther, for providing me the dataset and the talks about the cattle breeds and the conservation work in Norway. Furthermore, I would like to thank Linn and Cathrine for their support and advise. Jessica Tetteroo Ås, November 217 ii

4 Abstract The aim of this study was to assess the risk status of six native Norwegian cattle breeds using the multiindicator method as proposed in the paper of Verrier et al. (215). The multi-indicator method is based on six indicators: number of breeding females, change in number of breeding females, percentage of crossbreeding, effective population size, breeder organisation and technical support, and socioeconomic context. The pedigree data was obtained from Kuregister, maintained by NIBIO and the Norwegian Genetic Resource Centre. The number of individuals in the datasets varied from 2,869 in Døla cattle to 2,469 in Sided Trønder and Nordland cattle. The programs EVA and ENDOG were used for the analysis. The indicators were scored using the same grids as described in Verrier et al. (215). The risk status was calculated as the simple mean of the six indicators. Based on this the cattle breeds could be considered as not endangered. However, according to the criteria used by the Norwegian Genetic Resource Centre the cattle breeds are endangered. The breeds scored similar for several indicators. Therefore, if the objective was to prioritise between the cattle breeds some indicators would not be relevant. To identify differences that might affect the risk status of the breeds, the indicators and grids should be adapted to the local conditions and the Norwegian conservation landscape. This can be done by redefining the indicators and grids to make them suitable for the Norwegian situation. Another option would be to put more weight on the indicators that are considered most relevant than to those of less importance. Finally, additional indicators reflecting the Norwegian risk landscape could be determined. iii

5 Table of Contents Preface... ii Abstract... iii 1 Introduction Norwegian cattle breeds Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle Conservation Norwegian livestock Breed conservation Problem definition Materials and Methods Population analysis Population size Pedigree Completeness Index Inbreeding Effective population size Trend effective population size Determination of risk status Number of breeding females Change in number of breeding females Percentage of crossbreeding Effective population size Breeder organisation and technical support Socio-economic context Results and Discussion Population analysis Population size Pedigree completeness and inbreeding Pedigree Completeness Index Inbreeding Implications pedigree completeness and inbreeding Effective population size Trend effective population size Determination of risk status Number of breeding females Change in number of breeding females iv

6 3.2.3 Percentage of crossbreeding Effective population size Breeder organisation and technical support Socio-economic context Risk status cattle breeds Usability multi-indicator method Implications and Recommendations References Appendix I Map of Norway v

7 1 Introduction The following paragraph is mainly based on Vangen and Sæther (n.d.). Until approximately 185 the cattle in Norway were very heterogeneous in performance and had a large variety of exterior characteristics. Breeding was solely focussed on optimizing calving time, in particular, calving during spring was favoured. Cows which calved during spring had less feed requirements during winter when the food was scarce, and their high lactation coincided with the unlimited access to pastures. Uniformity in exterior was not of interest yet. In the 185s more focus was put on the improvement of feeding and breeding of Norwegian livestock, inclusive Norwegian cattle in order to enhance milk and meat production. At this point, the breeding strategy shifted from optimizing calving time to exterior characteristics. The prevailing breeding strategy aimed at phenotypic homogenization of the cattle, as animals with a similar appearance were expected to have similar production. To further improve the Norwegian cattle, breeds from abroad were imported. Especially the Ayrshire cattle from Scotland were frequently used for crossbreeding. The Ayrshire cattle were known for their good milk properties. Furthermore, the climate and topography in Scotland were similar to those in Norway. To improve the cattle breeds living in the Norwegian mountains, the local Telemark cattle were considered suitable for crossbreeding with other local mountain breeds. The crossbreeding with Ayrshire and Telemark cattle continued until 188 and was mainly done to improve the milk production with meat as a side product. However, from 188 to 192 the breeding strategy changed. Several leading breeders claimed that local cattle breeds should form the basis for the development of the Norwegian breeds. Therefore, crossbreeding with Ayrshire cattle were no longer allowed in Norway. For the Telemark cattle breeding were mainly restricted to the Telemark region. In the beginning of the 19s there were over twenty Norwegian cattle breeds. Due to inbreeding, small population sizes, and poor production improvement, many of the cattle breeds were merged. In 1935 the studbook of the Norwegian red cattle (Norsk rødt fe, NRF) was established (GENO, 217). During the 196s all Norwegian cattle breeds were merged into the NRF. This resulted in virtually one breeding population of cattle left in Norway. After 196, organized breeding activities had been absent for twenty years for cattle breeds other than NRF. From the old cattle breeds only Telemark cattle and Sided Trønder and Nordland cattle had their own breeding organisations. The remains of the old cattle breeds of Western Fjord cattle, Western red polled cattle, Eastern polled cattle, and Døla cattle have been saved from extinction through conservation work from the late 198s. Currently, the potential of the old cattle breeds in tourism, as caretakers of the cultural landscape, and in niche production of milk and meat is well recognized. 1.1 Norwegian cattle breeds Nowadays, 21 cattle breeds have been reported in Norway (DAD-IS, 217). Seven of these cattle breeds are recognized as native breeds (Table 1.1). In Norway a breed is considered as native when it meets the following criteria (Sæther & Rehnberg, 217): - The breed has to be imported to or established in Norway before The breed should not have any crossbreeding of any imported breeding material, or the import has to follow the Norwegian breeding goals. - The breed has to have or has to had economic or cultural historical importance. This study is based on the six native cattle breeds which are also considered to be endangered. Namely: the Sided Trønder and Nordland cattle, the Western Fjord cattle, the Telemark cattle, the Western red polled cattle, the Eastern red polled cattle, and the Døla cattle. In Table 1.1 the original Norwegian names of the native cattle breeds is given, plus the abbreviation if known. In the following paragraphs a brief description of the history and characteristics of the six breeds is given. 1

8 Table 1.1 Names of native cattle breeds in English and Norwegian, and the associated abbreviation when known. Breed name Original Norwegian name Abbreviation (if known) Sided Trønder and Nordland cattle Sidet trønder- og nordlandsfe STN Western Fjord cattle Vestlandsk fjordfe Telemark cattle Telemarkfe Western red polled cattle Vestlandsk raudkolle Eastern red polled cattle Østlandsk rødkolle Døla cattle Dølafe Norwegian red cattle a Norsk rødt fe NRF a Breed not included in this study Sided Trønder and Nordland cattle The Sided Trønder and Nordland cattle (STN) have his origin in the regions of Trøndelag and northern Norway (see Appendix I for a map of Norway). The breed is a merger of the Nordland cattle and the Sided Trønder cattle breeds, which were until the 192s considered as two separate breeds (Sidet trønderfe og nordlandsfe (STN), 27). Through strong natural selection the breed has developed over the years to a small, fertile and healthy breed (Avlslaget for STN, 28). In 216 the population size of the STN cattle was 1,655 breeding females (Sæther & Rehnberg, 217). The STN is predominantly kept for milk production, with meat production as a side product. In Table 1.2 the milk production values of the STN are given. At adult age the STN has a body weight of 4 kg (Avlslaget for STN, 28). The STN predominantly has a black-and-white coat, while brown-and-white also occurs. The colour distribution varies from entirely white with a few black spots to almost black (Figure 1.1). Usually the coat around the ears, eyes, and noise is black coloured. The STN population is entirely polled. Figure 1.1 Sided Trønder and Nordland cattle (Sæther & Rehnberg, n.d.) Western Fjord cattle The Western Fjord cattle originate from western Norway, in the area ranging from Møre in the north to Hordaland in the south (Appendix I). In the late 18s every fjord had his own local cattle population. However, the many small breeding populations faced problems with inbreeding. Therefore, in the 192s a common studbook was established for the Western Fjord cattle (Laget for Vestlandsk fjordfe, n.d.). In 216 the population size of the Western Fjord cattle was 717 breeding females (Sæther & Rehnberg, 217). The Western Fjord cattle are a dual-purpose breed, kept both for milk (Table 1.2) and meat production. At adult age the Western Fjord cattle have a body weight of 4 kg (Vestlandsk fjordfe, 27). One characteristic of the Western Fjord cattle is a large variation in coat colour and pattern. Colours like black, brown, white, and grey can appear (Figure 1.2). One third of the Western Fjord cattle population has horns. 2

9 Figure 1.2 Western Fjord cattle (Sæther & Rehnberg, n.d.) Telemark cattle The Telemark cattle originate from the Telemark region in southern Norway (Appendix I). The breed, established in 1856, is the oldest Norwegian cattle breed (Telemarkfe, 27). The Telemark cattle have long been used to crossbreed with other local cattle breeds living in the mountains. Therefore, the Telemark cattle have been well established throughout eastern and southern Norway. In 216 the population size of the Telemark cattle was 38 breeding females (Sæther & Rehnberg, 217). The Telemark cattle are a dual-purpose breed, kept both for milk (Table 1.2) and meat production. At adult age the Telemark cattle have a body weight of 5 kg (Telemarkfe, 27). The only permissible coat colour is red-sided or brindled (Figure 1.3). The back should be white from head to tail. The chest, abdomen, hind legs, and tail should also be white. The head should be brown dotted. The entire Telemark cattle population has horns. Figure 1.3 Telemark cattle (Sæther & Rehnberg, n.d.) Western red polled cattle The Western red polled cattle originate from Sørlandet and south-west Norway (Appendix I). In the 18s the crossing of the Lyngdalsfeet cattle with the Rogaland cattle formed the basis for the Western red polled cattle (Vestlandsk raudkolle, n.d.). Through active breeding the Western red polled cattle spread in the 2 th century across western Norway. In 216 the population size of the Western red polled cattle was 153 breeding females (Sæther & Rehnberg, 217). The Western red polled cattle are a dual- 3

10 purpose breed, kept both for milk (Table 1.2) and meat production. At adult age the Western red polled cattle have an average weight of 45 kg (Sæther & Rehnberg, 217). The only permissible coat colour is red, but light brown, red brown or dark brown colour variations are also allowed (Figure 1.4). A number of white markings is allowed. As the name already suggests, the entire Western red polled cattle population is polled. Figure 1.4 Western red polled cattle (Sæther & Rehnberg, n.d.) Eastern red polled cattle The Eastern red polled cattle originate from Østfold and Akershus in south-eastern Norway (Appendix I). The Eastern red polled cattle were the dominant breed of eastern Norway until it got competition of NRF in the 195s. In 1961 the breeding organisation of the red polled cattle merged with the studbook of NRF. Only few farmers in eastern Norway kept several Eastern red polled cattle, which lead to a decline of the breed to approximately ten animals in the late 198s (Østlandsk rødkolle, 27). Crossbreeding with cattle of the Swedish Red was used to restore the breed. In 216 the population size of the Eastern red polled cattle was 356 breeding females (Sæther & Rehnberg, 217). The Eastern red polled cattle are a dual-purpose breed, kept both for milk (Table 1.2) and meat production. At adult age the Eastern red polled cattle have an average weight of 5 kg (Østlandsk rødkolle, 27). The most common coat colour is red, but some white markings are allowed (Figure 1.5). As the name already suggests, the entire Eastern red polled cattle population is polled. Figure 1.5 Eastern red polled cattle (Sæther & Rehnberg, n.d.). 4

11 1.1.6 Døla cattle The Døla cattle originate from Gudbrandsdalen, Østdalen, and Hedmark (Appendix I). In those areas the cattle had access to rich mountain pastures. In the early 19s breeding for a more uniform breed started (Dølafelaget, n.d.-b). In 216 the population size of the Døla cattle was 223 breeding females (Sæther & Rehnberg, 217). The Døla cattle are kept as a dual-purpose breed. In Table 1.2 the milk production values of the Døla cattle are given. The Døla cattle are of the six cattle breeds closest to a beef cattle (Sæther, personal communication, 7 September, 217). At adult age the Døla cattle have an average weight of 5 kg (Dølafe, 27; Dølafelaget, n.d.-a). The variation in coat colour is large with colours like black, brown, red, white, and brindled (Figure 1.6). The coat can be single coloured, spotted, saddled, and seated. One fourth of the Døla cattle population is polled. Figure 1.6 Døla cattle (Sæther & Rehnberg, n.d.). Table 1.2 The average milk production, fat content, and protein content in 216 for the six cattle breeds (Tine, 216, as cited in Sæther & Rehnberg, 217). Average milk production (kg/year) Fat content (%) Protein content (%) Sided Trønder and Nordland cattle 4, Western Fjord cattle 3, Telemark cattle 4, Western red polled cattle 4, Eastern red polled cattle 4, Døla cattle 3, Conservation Norwegian livestock Since 1986 the Animal Genetic Resource committee started with the conservation of Norwegian livestock. In 21, the committees for Plant and Forest were established. In 26, the three committees were merged into the Norwegian Genetic Resource Centre, located in Ås. The goal of the Norwegian Genetic Resource Centre is to prevent the Norwegian livestock breeds from becoming endangered. Therefore, the Norwegian Genetic Resource Centre follows the criteria of the FAO for the risk status of a breed (see paragraph 1.3 for an explanation of the criteria). The Norwegian Genetic Resource Centre has a close cooperation with Norwegian farmers, NordGen, Norwegian Ministry, and the FAO. The activities differ from advising farmers at the choice of the bull to maintaining sustainable breeding. The Norwegian Genetic Resource Centre is responsible for the conservation strategy, while the collecting and storing of cattle semen is done by the breeding organisation of the NRF (Geno). Geno started with collecting and storing semen of the Western Fjord cattle, Western red polled cattle, Eastern red polled cattle, and Døla cattle at The STN and Telemark cattle breeding organisations are independent, 5

12 and responsible for their own collection and storage of semen. For the remaining four cattle breeds the semen of three bulls is collected per year (Sæther, personal communication, 18 August, 217). 1.3 Breed conservation The conservation of breeds is important for maintaining the genetic diversity within and between breeds. Breed conservation includes all activities undertaken by humans to maintain the genetic diversity of a breed. According to the Convention on Biological Diversity of the United Nations (1992) each country is responsible for the establishment of their own conservation programs. Conservation can be done through in situ and ex situ conservation. With in situ conservation all measures aim to maintain live breeding populations in their natural surroundings. This can be done either through the establishment of an active breeding program or measures can be undertaken to ensure the contribution of livestock to the food and agricultural production. Furthermore, with in situ conservation the ecological, cultural, and socio-economic values of a breed can be assessed (Gandini et al., 24). With ex situ conservation the breed is stored outside the environment where the breed developed. This can be done through removing part of the population from the current environment and placing it in a new environment. Other possibility is cryo-conservation in which gametes or embryos are stored in a gene bank. In a gene bank the genetic material of a breed can be stored for the long-term. The genetic material can be used as a back-up in case the breed faces problems with the genetic diversity. Since it is impossible to conserve all livestock breeds, an assessment of the degree of endangerment should be made to enable the prioritisation of the conservation efforts. Breed endangerment is the most important factor for breed prioritisation. Basically it describes the likelihood of a breed to become extinct within a given time period (Gandini et al., 24). However, not all breeds with a high degree of breed endangerment can be conserved. Since the available resources for conservation are often limited. Therefore, a classification system can be used to determine the breed endangerment. One well-known classification system is the one developed by the Food and Agriculture Organization of the United Nations (FAO). In the FAO system the breeds are classified in one of the eight categories: extinct, critical, critical maintained, endangered, endangered maintained, vulnerable, not at risk, and unknown. In the categories a distinction is made between low reproductive capacity species and high reproductive capacity species. Species with a low reproductive capacity, e.g. cattle and horses, take more time and more generations to increase the population size. Therefore, the low reproductivity group is at more risk than the high reproductivity group, e.g. pigs and avian species. Since this study is focussed on cattle breeds, the FAO categories are given specified for the low reproductivity species (FAO, 213): - Extinct: Occurs when there are no breeding males and breeding females nor embryos remaining of the population. Therefore, it is no longer possible to restore the breed. - Critical: The total number of breeding females is less than 3 or the total number of breeding males is less than or equal to five. - Critical maintained: This is the same as for Critical, however, as an additional requirement the breeds are actively managed in this category. - Endangered: The total number of breeding females is between 3 and 3 or the total number of breeding males is less than or equal to twenty and greater than five. - Endangered maintained: This is the same as for Endangered, however, as an additional requirement the breeds are actively managed in this category. - Vulnerable: The total number of breeding females is between 3 and 6 and the total number of breeding males is between twenty and Not at risk: The total number of breeding females is larger than 6 and the total number of breeding males is larger than Unknown: Information on the total number of breeding females and males is not known. For the FAO classification system information of the overall population size, number of breeding females and breeding males, percentage of crossbreeding, and the trend in population size is used for the categorisation (Scherf, 2). Another classification system is the one used by Ruane (2). Ruane (2) performed a case study on the prioritisation of 17 Norwegian breeds. He used seven criteria to determine the risk status of a breed: species, degree of endangerment, traits of current economic value, special landscape value, traits of current scientific value, cultural and historical value, and genetic uniqueness (Ruane, 2). Ruane (2) concluded that the most important criteria for the breed prioritisation is again the breed endangerment. 6

13 1.4 Problem definition The six cattle breeds used in this study form a small proportion of the Norwegian cattle stock. As mentioned briefly before, the conservation of breeds is important for maintaining the genetic diversity. The cattle breeds may carry unique genes which will become valuable in the future to meet changes in market demand or changes in the environment. The aim of this study is to determine the risk status of the six Norwegian cattle breeds. Instead of using the classification system of the FAO (213) or the breed prioritisation framework of Ruane (2), the multi-indictor method of Verrier et al. (215) will be used. The classification system of the FAO and the breed prioritisation framework focus mainly on the demographic factors of a breed. However, the status of a breed is also determined by genetic, sociological, and economic factors (Audiot, 1995, as cited in Verrier et al., 215). With the multi-indicator method of Verrier et al. (215) all these factors are taken into account. In this study, first a population analysis is performed to get an overview of the cattle breeds regarding several general population parameters. Secondly, the risk status of the cattle breeds is assessed using the multi-indicator method. The following research questions will be answered: - What is the population structure of the six cattle breeds? - How valuable is the multi-indicator method in assessing the risk status of the six cattle breeds? - Are the indicators of the multi-indicator method suitable for the Norwegian conservation framework? 7

14 2 Materials and Methods Pedigree data for the six Norwegian cattle breeds were obtained from Kuregister, maintained by NIBIO and the Norwegian Genetic Resource Centre. The data contained information on the unique identification number of the individual, sex, date of birth, date of death, breed, percentage of crossbreeding, and unique identification number of the sire and dam, if known. The number of individuals varied from 2,869 in Døla cattle to 2,469 in STN (Table 2.1). The descriptive statistics of the different cattle breeds are presented in Table 2.1 and Table 2.2. The datasets were first checked for inconsistencies. Offspring born before the parents, individuals with wrong date of births, and wrong sex were corrected. In some cases, one of the parents had to be set as unknown. For each cattle breed several animals appeared as sire or dam but were not recorded as individual in the studbook. Pedigree data was supplemented with those individuals. Year of birth for added individuals was created using information on the birth date of the first progeny and the average age of already recorded dams and sires when their first progeny was born. To create the year of birth of the added sires and dams this average age of sire/dam was subtracted from the year the first progeny was born. In Table 2.2 an overview is given of the number of added individuals and the average age of the sires and dams. Table 2.1 General parameters of the six cattle breed datasets. Number of cattle in dataset a Number of unknown parents Distribution year of births Sided Trønder and Nordland cattle 2,469 1,873 (9.15 %) Western Fjord cattle 1, (5. %) Telemark cattle 8, (5.81 %) Western red polled cattle 4, (8.62 %) Eastern red polled cattle 2, (7.97 %) Døla cattle 2, (6.69 %) a Newly added individuals included. Table 2.2 Characteristics of the added dams and sires. Number of added sires Average age sires (years) Number of added dams Average age dams (years) Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle Population analysis The population analysis was performed with EVA version 3. (Berg, Nielsen, & Sørensen, 26) and ENDOG version 4.8 (Gutiérrez & Goyache, 25). EVA was used to calculate the individual increases of inbreeding and rates of inbreeding. ENDOG was used for calculating the effective population size. In the following section a description of the parameters used for the population analysis is given Population size This parameter indicates the number of offspring born and registered per year. A distinction was made between inbred and non-inbred individuals. 8

15 2.1.2 Pedigree Completeness Index The Pedigree Completeness Index (PCI) is the extent to which the increase in inbreeding depends through time on the proportion of ancestors known in previous generations (MacCluer et al., 1983). The PCI is calculated first separately for the paternal and maternal line of the pedigree: I d.sire or I d.dam = 1 d a i I d.sire and I d.sire indicate the index value respectively for the paternal and maternal line of the pedigree. d indicates the pedigree depth and a i the proportion of ancestors known in generation i. As a consequence of using information of all d generations, each parent is given twice as much weight as their grandparent. The final PCI is defined as: d i = 1 I d = 4 I d.sirei d.dam I d.sire + I d.dam For the PCI the harmonic mean of the paternal and maternal lines is used. Therefore, both the paternal and maternal ancestries should be known to detect the inbreeding. The PCI ranges from to 1. If either one of the parents is unknown, the PCI equals. A PCI of 1 indicates that all ancestors of an individual for the given number of generations are known. In this study the PCI is calculated for five and ten generations. For an accurate estimation of the rate of inbreeding the pedigree information of at least five generations should be completely known (Baumung & Sölkner, 23; Windig & Oldenbroek, 212) Inbreeding Inbreeding is a consequence of mating individuals which are related through a common ancestor (Falconer & Mackay, 1996; Lacy, 1995). The rate of inbreeding ( F) indicates the increase in inbreeding for one generation and is defined as (Falconer & Mackay, 1996): F = F x F x 1 1 F x 1 Where F x is the average inbreeding in the x th generation and F x 1 the average inbreeding in generation x 1. The rate of inbreeding was plotted based on the average inbreeding coefficients per year of birth. The expected rate of inbreeding was also plotted to assess whether mating design deviates from random mating (indicated by alpha in the figures). Inbreeding coefficients lower than expected by random mating indicate intentional avoidance of inbreeding Effective population size The effective population size (N e ) gives an indication of the number of breeding animals leading to the same rate of inbreeding as bred in the idealized population. Characteristics of an idealized population are an infinite population with a constant number of breeding animals, parents have an equal change of contributing offspring, mating occurs randomly, and generations do not overlap (Caballero, 1994). The general formula for calculating N e is (Falconer & Mackay, 1996): N e = 1 2 F The effective population size was computed with ENDOG version 4.8. The program has six different approaches for the calculation of N e. In the following section a description of the six methods for calculating N e is given. For all six cattle breeds the N e was estimated using the six approaches. Based on the results a decision was made which method will eventually be used as indicator of the effective population size in the multi-indicator method. 9

16 Method 1 Effective population size based on equivalent complete generations In this method N e is defined as (Gutiérrez & Goyache, 25): N e = 1 2b Where b indicates the regression coefficient of the individual inbreeding coefficient over the equivalent complete generations. For this method the entire population is used. Method 2 Effective population size obtained from regression on equivalent complete generations In this method a reference population is used. The reference population consists of all individuals with a genetic coefficient higher than two. The genetic coefficient indicates the generation the individual belongs to, based on pedigree information. An individual with unknown parents is considered as a founder and has a genetic coefficient of one. Nonetheless, ENDOG and EVA have a different definition of the base population. While ENDOG gives individuals of the base population a value of zero, EVA gives those individuals a value of one. As a consequence, when ENDOG used a reference population of individuals with a genetic coefficient higher than two, in fact individuals with a genetic coefficient higher than three were used for the values obtained with EVA. N e is computed with the general formula of Falconer and Mackay (1996). However, F is defined as (Gutiérrez & Goyache, 25): F = b 1 (F t b) With F t being the average inbreeding of the reference population and b the regression coefficient of the individual inbreeding coefficient over the equivalent complete generations. Method 3 Effective population size obtained from regression on birth date In this method N e is calculated from the regression coefficient of the inbreeding coefficients over the year of birth. For this method the entire population is taken into account. N e is calculated using the general formula of Falconer and Mackay (1996). However, F is defined as (Gutiérrez et al., 23): F = l b 1 (F t (l b)) Where l indicates the generation interval, b the regression coefficient of the individual inbreeding coefficient over the birth years, and F t the mean inbreeding in the reference population. The generation interval l is calculated as the weighted average: l = ( N x l x ) N total Where N x is the number of individuals in year x, l x the generation interval in year x, and N total the total number of individuals in the dataset. Method 4 Effective population size obtained from logarithmic regression on equivalent complete generations In this method N e is calculated from the logarithmic regression over the equivalent generations in a reference population. The reference population is the same as used for method 2. N e is calculated with the general formula (Falconer & Mackay, 1996). However, F is defined as (Pérez-Enciso, 1995): This can be rewritten as: F t = (1 F) t β = ln (1 F) F = 1 e β Where F t is the average inbreeding in the reference population and β the logarithmic regression of log(inbreeding) on equivalent complete generations. 1

17 Method 5 Effective population size obtained from logarithmic regression on birth date In this method N e is calculated from the logarithmic regression on the birth date. In this method the entire population is used. The steps of the logarithmic regression of (1 F t ) are the same as described for method 4. After estimating, N e is divided by the generation interval (Pérez-Enciso, 1995). The generation interval is again calculated as the weighted average (see method 3). Method 6 Effective population size obtained from individual increase in inbreeding In this method N e is calculated from the individual increase in inbreeding ( F i ), using the same reference population as in method 2. F i is defined as (Gutiérrez et al., 29; 28): F i = 1 t 1 1 F i Where F i indicates the individual coefficient of inbreeding and t the genetic coefficient. The realized effective size (N ), e which is an estimate of N e, can be calculated using the formula (Cervantes et al., 28; Gutiérrez et al., 28): N e = 1 2 F Where F is the average of the F i values of the reference population. This way of calculating N e takes the breeding history of each individual into account instead of the mating policy of the entire reference population (Gutiérrez et al., 28). Since this method involves the individual inbreeding coefficient for each individual, it is possible to calculate the standard error of N e as (Cervantes et al., 28): σ N e = 2N e2 σ F 1 Ne Trend effective population size This parameter indicates the trend in the effective population sizes of the cattle breeds over a certain time period. The effective population size is calculated for five five-year periods, starting with and ending with The trend has only been calculated for the effective population size obtained from individual increase in inbreeding (Gutiérrez et al., 29). The reasoning behind this decision is further explained in paragraph Determination of risk status As mentioned before to determine the risk status of the six cattle breeds, the multi-indicator method of Verrier et al. (215) has been used. The multi-indicator method consists of six indicators: number of breeding females, recent evolution of the number of breeding females, percentage of crossbreeding, effective population size, breeder organisation and technical support, and socio-economic context. To convert the observed numbers into a score, the same grids as described in the paper of Verrier et al. (215) have been used. The scores range from (no risk) to 5 (maximum risk). When the breed had a score of 3 or higher the breed was considered as endangered. In the following paragraphs the calculation of each indicator is explained Number of breeding females For this indicator the observed number of breeding females is calculated for the year 216. Individuals with an unknown dam were not taken into account in the calculations. Adding all observations together gave the observed number of breeding females. In Table 2.3 the grid for converting the observed number of breeding females into a score is given. When the number of breeding females is lower than 7,5 (score 3) the breed is considered endangered (Regulation 445/22 Article 13, as cited in Alderson, 23). 11

18 Table 2.3 Grid for converting the observed number of breeding females into a score. The numbers indicate the bounds for a score (Verrier et al., 215). Cattle 15 1, 7,5 25, 75, Change in number of breeding females For this indicator the change in the observed number of breeding females over the last five years is calculated. The number of breeding females in 211 was calculated the same way as described for the number of breeding females in 216 (paragraph 2.2.1). The recent evolution in the number of breeding females (T 5 ) was defined as: T 5 = Nf t Nf t 5 Nf t 5 Where Nf t is the current number of breeding females in 216 and Nf t 5 the number of breeding females in 211. In Table 2.4 the grid for converting the change in the number of breeding females is given. When the population size of a breed was constant or increasing a score of zero was allocated, since the breed is considered to be at no risk. Table 2.4 Grid for converting the change in the number of breeding females over the last five years into a score. The percentages indicate the bounds for a score (Verrier et al., 215). Cattle -12 % -9 % -6 % -3 % % Percentage of crossbreeding For this indicator the observed percentage of crossbreeding is calculated. Individual was considered as crossbred when the percentage of pure breed was lower than 87.5 %. With this baseline the average proportion of crossbred individuals was calculated from 211 to 216. For the observed percentage of crossbreeding the average from 211 to 216 was taken. Individuals with unknown percentage of crossbreeding were excluded from the calculation. In Table 2.5 the grid for converting the observed percentage of crossbreeding into a score is given. Table 2.5 Grid for converting the observed percentage of crossbreeding into a score. The percentages indicate the bounds for a score (Verrier et al., 215). Cattle % 12.5 % 25 % 37.5 % 5 % Effective population size The effective population size was calculated using the method of Gutiérrez et al. (29) (individual increase in inbreeding; paragraph 2.1.4). The reasoning behind this decision is further explained in paragraph In Table 2.6 the grid for converting the estimated effective population size is given. Table 2.6 Grid for converting the effective population size into a score. The numbers indicate the bounds of a score (Verrier et al., 215). Cattle Breeder organisation and technical support This indicator is divided in five sub indicators (Table 2.7). A score of was given to a sub indicator when the corresponding breed management/ technical support was present. When the corresponding breed 12

19 management/ technical support was absent a score of 1 was given. For several sub indicators a score of.5 was given for intermediate situations. The final score for the indicator is the sum of the five sub indicators. Table 2.7 Sub indicators used to score the breeder organisation and technical support (Verrier et al., 215). Sub indicator Score Breeders organisation present Yes = ; no = 1 In situ management Yes = ; no = 1 Stock in cryobank Yes, with more than ten donor animals = ; yes with less than ten donor animals or no for a species for which cryobanking is technically impossible =.5; no for a species for which cryobanking is technically possible = 1 Technical support present Yes, with local experts and national support = ; yes, with either local experts or national support =.5; no = 1 Cohesion and collective dynamics of breeders Yes = ; intermediate =.5; no = Socio-economic context This indicator is also divided in five sub indicators (Table 2.8). A score of,.5, or 1 was given when the social and economic development was respectively enhanced, intermediate, or adversely affected. The final score for this indicator is the sum of the five sub indicators. Table 2.8 Sub indicators used to score the social and economic development (Verrier et al., 215). Sub indicator Score Young livestock farmers start off raising the breed Yes = ; intermediate =.5; no = 1 Availability of the breed for sale Yes = ; no = 1 Markets for products and services Yes, profitable and diversified = ; yes, average =.5; no = 1 Labels used to distinguish products Yes = ; no = 1 Financial support given to territories Yes = ; no = 1 13

20 3 Results and Discussion 3.1 Population analysis In this section the results for the population analysis are given. Parallel to the results the discussion is given Population size In Figure 3.1 the number of calves born per year is given. The STN has the largest population size, followed by the Western Fjord cattle. The population size of the STN increased exponentially until 28, thereafter the population size stabilised. For the Western Fjord cattle the population size has been increasing linearly over the years. The Telemark cattle had an increase in the population size until 28, thereafter the population size decreased. For the Western red polled cattle the population size steadily increased until 27, thereafter the population size decreased. For both the Eastern red polled cattle and the Døla cattle the population size has been increasing over the years, with a strong increase the last few years. In Figure 3.1 also a distinction is made between non-inbred and inbred individuals. In Table 3.1 the average percentage of non-inbred and inbred individuals, calculated over the last five years, is given. The percentage of inbred individuals was very low for the STN compared to the other cattle breeds. For the other cattle breeds the percentage of inbred individuals ranged from 62 % to 82 %. Table 3.1 Average percentage of non-inbred and inbred individuals calculated from 212 to 216. Non-inbred individuals (%) Inbred individuals (%) Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle Pedigree completeness and inbreeding The results for the Pedigree Completeness Index and inbreeding will be given together. This because the two parameters are closely linked Pedigree Completeness Index In Figure 3.2 the Pedigree Completeness Index (PCI) is given. For all cattle breeds the same trends were observed. The first few years the PCI fluctuated strongly, indicating large variation in the proportion of ancestors known. In addition, for all cattle breeds the PCI for five generations was higher than for ten generations. The PCI for five generations is per definition higher, since with five generations the proportion of ancestors known is most likely higher than with ten generations. For the STN the PCI was very low until 21, thereafter the PCI increased exponentially. For the Western Fjord cattle and the Telemark cattle the PCI increased until respectively 24 and 22. Thereafter, the PCI decreased and stabilized. For the Western red polled cattle, the Eastern red polled cattle, and the Døla cattle the PCI has been increasing over the years. For the three breeds several peaks where observed over the years. To put the different PCI values into more perspective, Figure 3.3 shows the PCI for five generations plotted for the six cattle breeds. The variation when PCI reached the value of.5, indicating 5 % of the ancestors is known, is large between the cattle breeds. For the Telemark cattle 5 % of the ancestors is known since 1991, followed by the Western Fjord cattle in For the Western red polled cattle, the Eastern red polled cattle, and the Døla cattle this is the case since 212. The PCI of the STN was the lowest of all cattle breeds. In 216 only 16 % of the ancestors were known. In conclusion, the pedigree quality is lowest for the STN. On the other hand, the Telemark cattle has the highest PCI. Although the Telemark cattle has most information of the pedigree known compared to the other cattle breeds, the pedigree quality is still not sufficient enough. 14

21 Number of individuals Number of individuals Number of individuals Number of individuals Number of individuals Number of individuals Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle a b c Year of birth Year of birth Year of birth Western red polled cattle Eastern red polled cattle 4 25 d e f Døla cattle Year of birth Year of birth Year of birth Figure 3.1 Population size for the Sided Trønder and Nordland cattle (a), Western Fjord cattle (b), Telemark cattle (c), Western red polled cattle (d), Eastern red polled cattle (e), and Døla cattle (f). Non-inbred individuals Inbred individuals. 15

22 PCI coefficient PCI coefficient PCI coefficient PCI coefficient PCI coefficient PCI coefficient Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle a b c Year of birth Year of birth Year of birth Western red polled cattle Eastern red polled cattle d.8.6 e.7.5 f Døla cattle Year of birth Year of birth Year of birth Figure 3.2 Pedigree Completeness Index (PCI) for the Sided Trønder and Nordland cattle (a), Western Fjord cattle (b), Telemark cattle (c), Western red polled cattle (d), Eastern red polled cattle (e), and Døla cattle (f). PCI for five generations PCI for ten generations. 16

23 PCI coefficient PCI for five generations Year of birth Sided Trønder and Nordland cattle Telemark cattle Eastern red polled cattle Western Fjord cattle Western red polled cattle Døla cattle Figure 3.3 Pedigree Completeness Index (PCI) for five generations for the six cattle breeds Inbreeding In Figure 3.4 the average inbreeding coefficients per year of birth is given. For the STN mating between related individuals often took place. This is indicated by the average inbreeding which was higher than the expected inbreeding. Furthermore, alpha had a positive value. Both indicate that inbred mating was favoured. Especially the high peaks in the first few years indicate that mating between related individuals frequently took place. However, one should keep in mind that the STN had overall the lowest rate of inbreeding compared to the other cattle breeds. As mentioned before, the STN had little information available of the pedigree. Therefore, estimation of the rate of inbreeding is based on incomplete pedigree information. For the Western Fjord cattle mating between related individuals was avoided from 1985 to 1999 (with the exception of 1989). After 1999 inbred mating was favoured, since the average inbreeding was higher than the expected inbreeding. A decrease in inbreeding is a consequence of low PCI. For the same reason, an increase in inbreeding is a consequence of high PCI (Figure 3.2b). If the rate of inbreeding and PCI were plotted in one graph, both curves will follow each other. For the Telemark cattle inbreeding between related individuals was not avoided. The average inbreeding was predominantly higher than the expected inbreeding. The rate of inbreeding followed the progress of the PCI (Figure 3.2c). For the Western red polled cattle mating between related individuals was favoured. The average inbreeding was predominantly higher than the expected inbreeding. Especially during the period from 1992 to 2 inbred mating frequently took place. For the Eastern red polled cattle many different periods can be distinguished with either a preference or avoidance for mating between related individuals. The overall value of alpha approximates zero. The latter indicates that the followed breeding strategy resulted on average in the same level of inbreeding as random mating between individuals would result in. For the Døla cattle also many different periods can be distinguished with either a preference or avoidance for mating between related individuals. However, the overall value of alpha approximates zero, indicating that the followed breeding strategy resulted on average in the same level of inbreeding as random mating between individuals would result in. 17

24 Inbreeding coefficient Inbreeding coefficient Inbreeding coefficient Inbreeding coefficient Inbreeding coefficient Inbreeding coefficient Sided Trønder and Nordland cattle Western Fjord cattle.5.1 a b c Year of birth -.2 Year of birth -.6 Telemark cattle Year of birth Western red polled cattle Eastern red polled cattle d.8.5 e.6.4 f Year of birth -.4 Year of birth -.2 Døla cattle Year of birth Figure 3.4 The average inbreeding, expected inbreeding, and the deviation (alpha) for the Sided Trønder and Nordland cattle (a), Western Fjord cattle (b), Telemark cattle (c), Western red polled cattle (d), Eastern red polled cattle (e), and Døla cattle (f). Average F Expected F Alpha. 18

25 Implications pedigree completeness and inbreeding The average inbreeding coefficient for progeny born in 216 varied between.46 till.685 in the six cattle breeds. The cattle breeds (except the STN) had a large percentage of inbred individuals, indicating that mating between related individuals happened frequently in the cattle breeds. This happens e.g. when inbreeding is used to anchor specific traits in the offspring in order to improve the breed. Another reason could be the frequent use of some bulls within a herd. When the cows and bulls are bred within a certain region and rotation of bulls between different regions is limited, the rate of inbreeding will increase. As a consequence of inbreeding, on the long term the performance of animals can decrease, indicating inbreeding depression (Falconer & Mackay, 1996). Inbreeding negatively affects the lifetime production of milk, fat, and protein (Smith et al., 1998). Besides, inbreeding decreases the fertility (Wall et al., 25) and survival of cattle (Thompson et al., 2). Furthermore, inbreeding will lead to economic losses for the dairy producer (Smith et al., 1998). These results underline the need to continue with the conservation as is currently done by the Norwegian Genetic Resource Centre. For the cattle breeds the pedigree completeness increased over the years. However, for the cattle breeds still part of the pedigree is missing. As mentioned in the Materials and Methods, at least five generations of ancestors should be completely known for an accurate estimation of the rate of inbreeding. For none of the breeds this was the case. An incomplete pedigree leads to an overestimation of the effective population size (Boichard et al., 1997) and underestimation of the level of inbreeding (Boichard et al., 1997; Lutaaya et al., 1999). The incomplete pedigree indicates problems with the registration of cattle in the studbook. When individuals are born their parents were not registered in the studbook, or the parents are registered as dam or sire but further information of the parents was left out. Problems with the registration of cattle were observed in all breeds. The problem was most apparent for the STN. The number of offspring increased exponentially until 28. This exponential growth was not due to the birth of more offspring. A registration problem before 28 is more likely. Besides, the STN had the lowest value for the PCI, indicating little information of the pedigree is known. The incomplete pedigree was reflected in the levels of inbreeding. The rate of inbreeding was considerably lower than the values of the other cattle breeds. The small number of inbred individuals also gave a distorted picture, and was not in line with the results of the other breeds. More likely, the number of inbred individuals is higher but since information of ancestors is lacking, many individuals are considered as unrelated while in fact they have common ancestors. The incomplete pedigree was also reflected in the large standard error for the effective population size of the STN (paragraph 3.1.3). Indicating the effective population size has a certain degree of uncertainty. As comparison, the Telemark cattle has the most complete pedigree of all breeds and the lowest standard error for the effective population size. 19

26 3.1.3 Effective population size In Table 3.2 the results for the effective population size, estimated with the six methods, are given. The table shows a large variation between the estimates obtained with the different methods. The results will only be explained in detail for one breed, the Telemark cattle. The Telemark cattle is chosen since this breed had the most complete pedigree, which made the results more accurate compared to other cattle breeds. With method 1 the effective population size of individuals was estimated (Table 3.2; Figure 3.5). For method 1 the entire population is used. For this reason, individuals with unknown parents (the founders of the population) were also taken into account. The founders have a large influence on the calculation of the effective population size. The founders are assumed to be unrelated to the population, since their pedigree is unknown. Nevertheless, the founders are probably somehow related to the population. As a consequence, the effective population size of individuals does not reflect the actual population size. The effective population size of 19.7 individuals for method 2 was lower than the effective population size estimated for method 1 (Table 3.2). The difference can be explained by the definition of the rate of inbreeding for both methods. As explained in paragraph for method 2 a reference population is used. Only individuals with a genetic coefficient of three or higher were taken into account (Figure 3.6). This implies that founders and other individuals with not enough pedigree information known, were excluded. Therefore, the regression coefficient was higher (.241; Figure 3.6) and the effective population size was lower (19.7; Table 3.2). With method 3 the effective population size of individuals was estimated (Table 3.2; Figure 3.7). For method 3 the entire population is used. As can be seen in Figure 3.7 there are many observations on the x-axis. Those observations are mainly founders for which information on the rate of inbreeding is also missing. Due to the many observations on the x-axis the regression coefficient had a value of.7 (Figure 3.7). The effective population size depends on the regression coefficient. When the slope is small and approaches zero, as is the case for method 3, the effective population size is strongly overestimated. As mentioned before, founders are assumed to be unrelated to the pedigree, but it is questionable whether these individuals are truly unrelated. The founders are assumed unrelated to each other, but the founders are naturally related to later generations. With method 4 the effective population size of individuals was estimated (Table 3.2; Figure 3.8). For method 4 the reference population with only individuals with a genetic coefficient of three or higher was used (Figure 3.8). The effective population size for method 4 was comparable to the value calculated for method 2. The main difference between the methods is that for method 4 a logarithmic regression was performed over the inbreeding coefficients. With method 5 the effective population size of individuals was estimated (Table 3.2; Figure 3.9). For method 5 the entire population is used. The inclusion of the founders leads to many observations on the x-axis. Those observations had a large influence on the regression coefficient of -.8 (Figure 3.9). As mentioned before, due to the small regression coefficient the effective population size is strongly overestimated. For method 6 the effective population size of individuals with a standard error of 5.68 individuals was estimated (Table 3.2). For method 6 the reference population was used. Furthermore, the effective population size was based on the individual inbreeding coefficients of all individuals in the reference population. For the determination of the risk status only one method is used as indicator of the effective population size. Based on the results of Table 3.2 and the aforementioned clarification of the variation between the methods, method 6 is chosen as indicator. Method 1, 3, and, 5 were not applicable since those methods take the entire population into account. As was shown before, for all cattle breeds a proportion of the pedigree was incomplete (Figure 3.2). When pedigree information is incomplete the effective population size will be overestimated (Boichard et al., 1997). Method 2, 4, and 6 used a reference population were only individuals with enough pedigree information were taken into account. Eventually method 6 is chosen since this method directly accounts for differences in pedigree knowledge and completeness at the individual level but also, indirectly, for the effects of mating policy, drift, overlap of generations, selection, migration and different contributions from a different number of ancestors, as a consequence of their reflection in the pedigree of each individual in the analyzed population (Gutiérrez et al., 28; page 361). 2

27 Table 3.2 The effective population size for the six cattle breeds using the six methods (paragraph 2.1.4). Method 1 a Method 2 b Method 3 c Method 4 d Method 5 e Method 6 f Sided Trønder and Nordland cattle (51.24) Western Fjord cattle (9.8) Telemark cattle (5.68) Western red polled cattle (12.57) Eastern red polled cattle (8.1) Døla cattle (11.39) a Method 1: Effective population size based on equivalent complete generations (Gutiérrez & Goyache, 25). b Method 2: Effective population size obtained from regression on equivalent complete generations (Gutiérrez & Goyache, 25). c Method 3: Effective population size obtained from regression on birth date (Gutiérrez et al., 23). d Method 4: Effective population size obtained from logarithmic regression on equivalent complete generations (Pérez-Enciso, 1995). e Method 5: Effective population size obtained from logarithmic regression on birth date (Pérez-Enciso, 1995). f Method 6: Effective population size obtained from individual increase in inbreeding (Gutiérrez et al., 29). Standard error in parentheses. 21

28 Inbreeding coefficient Inbreeding coefficient.5 Method y =.186x Genetic coefficient Figure 3.5 Result for method 1 Effective population size based on equivalent complete generations for the Telemark cattle..5 Method y =.241x Genetic coefficient Figure 3.6 Results for method 2 Effective population size obtained from regression on equivalent complete generations for the Telemark cattle. 22

29 Inbreeding coefficient Inbreeding coefficient Method y =.7x Year of birth Figure 3.7 Result for method 3 Effective population size obtained from regression on birth date for the Telemark cattle..1 Method y = -.261x Genetic coefficient Figure 3.8 Result for method 4 Effective population size obtained from logarithmic regression on equivalent complete generations for the Telemark cattle. 23

30 Inbreeding coeffcient Method y = -.8x Year of birth Figure 3.9 Result for method 5 Effective population size obtained from logarithmic regression on birth date for the Telemark cattle Trend effective population size In Figure 3.1 the trend in the effective population size for the six cattle breeds is given. In Table 3.3 the effective population size and standard error for the different time periods are given. The values for the trend in the effective population size are calculated with method 6 (paragraph 2.1.4). For the STN the effective population increased exponentially the last years. From to the effective population size increased almost threefold and from to the effective population size almost doubled. A small side note needs to be made for the STN. The effective population size had a large standard error. For each time period the standard error was almost half the value of the effective population size. As was shown before, the STN has little pedigree information known. This can be traced back in the large standard error. As a consequence, the effective population size of the STN also has a certain degree of uncertainty. For the Western Fjord cattle the effective population size increased steadily over the years. For the Telemark cattle the effective population size increased steadily until Thereafter, the effective population size stabilised. For the Western red polled cattle the effective population size increased until Thereafter, the effective population size decreased. For the Eastern red polled cattle the effective population size fluctuated. From to the effective population size decreased. Thereafter, the effective population size increased until , which was followed again by a decrease. For the Døla cattle the effective population size decreased from to Thereafter, the effective population size increased steadily. Table 3.3 Trend in the effective population size for the six cattle breeds. Sided Trønder and Nordland cattle (11.85) (17.91) (9.3) (37.55) (53.54) Western Fjord cattle 38.1 (9.16) 38.2 (6.63) (7.14) (9.28) (8.71) Telemark cattle 3.13 (4.4) (4.62) (5.31) (5.91) (5.63) Western red polled cattle (9.4) (11.11) (1.12) (11.39) 6.93 (1.28) Eastern red polled cattle (12.8) (7.62) 35.5 (7.37) (7.3) (5.83) Døla cattle (15.29) (11.42) (8.47) (9.67) 6.79 (9.84) 24

31 Number of individuals Number of individuals Number of individuals Number of individuals Number of individuals Number of individuals Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle 1 6 a b c '92-'96 '97-'1 '2-'6 '7-'11 '12-'16 '92-'96 '97-'1 '2-'6 '7-'11 '12-'16 '92-'96 '97-'1 '2-'6 '7-'11 '12-'16 Time period Time period Time period Western red polled cattle Eastern red polled cattle 6 8 d e f Døla cattle '92-'96 '97-'1 '2-'6 '7-'11 '12-'16 Time period '92-'96 '97-'1 '2-'6 '7-'11 '12-'16 Time period '92-'96 '97-'1 '2-'6 '7-'11 '12-'16 Time period Figure 3.1 Trend in effective population size for the Sided Trønder and Nordland cattle (a), Western Fjord cattle (b), Telemark cattle (c), Western red polled cattle (d), Eastern red polled cattle (e), and Døla cattle (f). The black vertical bars indicate the standard error for the time period. 25

32 3.2 Determination of risk status In this section the results for the multi-indicator method are given. Parallel to the results the discussion is given Number of breeding females In Table 3.4 the observed number of breeding females and the associated scores are given for the six cattle breeds. As mentioned in the Materials and Methods, breeds with a score of 3 or higher are considered as endangered. Based on the number of breeding females all breeds can be considered as endangered. The Western red polled cattle even had the score of maximum risk. Table 3.4 Results for indicator 1 Number of breeding females. Number of breeding females 216 Score Sided Trønder and Nordland cattle 1,334 3 Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle Change in number of breeding females In Table 3.5 the observed number of breeding females in 211 and 216, with the trend and the associated score, are given. For all cattle breeds (except the Western red polled cattle) the trend was positive. Therefore, these breeds were considered to be at no risk. This is in contradiction with the results for the first indicator, number of breeding females, where all breeds were classified as endangered. In Figure 3.11 the number of breeding females is given from 211 to 216. The figure shows that although the overall trend was positive for almost all breeds, the number of breeding females is fluctuating over the years. The number of breeding females increased slowly. In between years the Telemark cattle, Western red polled cattle, and Eastern red polled cattle even had a drop in the number of breeding females. Table 3.5 Results for indicator 2 Change in number of breeding females. Number of breeding females 211 Number of breeding females 216 Trend (%) Sided Trønder and Nordland cattle 988 1, Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle Score 26

33 Trend in number of breeding females Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle Figure 3.11 Trend in the number of breeding females from 211 to 216 for the six cattle breeds Percentage of crossbreeding In Table 3.6 the observed percentage of crossbreeding and the associated score are given. Based on the percentage of crossbreeding only the Western red polled cattle was considered to be as endangered. Nevertheless, for all breeds there was a large variation in the observed percentages of crossbreeding (Figure 3.12). For the STN the percentage of crossbreeding slightly decreased over the years. For the Western Fjord cattle the percentage of crossbreeding increased over the years. For the Telemark cattle the percentage of crossbreeding fluctuated until 214, but increased the last two years. For the Western red polled cattle the percentage of crossbreeding decreased over the years. For the Eastern red polled cattle the percentage of crossbreeding decreased over the years. For the Døla cattle the percentage of crossbreeding fluctuated over the years, but overall there was a decreasing trend. Table 3.6 Results for indicator 3 Percentage of crossbreeding. Crossbreeding (%) Score Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle

34 4% Percentage of crossbreeding 35% 3% 25% 2% 15% 1% 5% % Year of birth Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle Figure 3.12 Percentage of crossbreeding from 211 to 216 for the six cattle breeds Effective population size In Table 3.7 the effective population size and the associated score are given. Based on the effective population size all breeds can be considered as endangered. Both the Telemark cattle and the Eastern red polled cattle had the score of maximum risk. Table 3.7 Results for indicator 4 Effective population size. Effective population size Score Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle Breeder organisation and technical support In Table 3.8 the five sub indicators with the associated scores are given. The first four sub indicators were present for all cattle breeds. All breeds have a breeder organisation and are provided with the same technical support. The only exception was the sub indicator Cohesion and collective dynamics of breeders. Both STN and Telemark cattle scored for this indicator.5. This because some parts of the breeding strategy are not supporting the conservation strategy Socio-economic context In Table 3.9 the five indicators with the associated scores are given. All cattle breeds have labels to distinguish products. However, the labels are not officially recognised. Therefore, the breeds get a score of 1 for this sub indicator. The remaining sub indicators are present for all cattle breeds, leading to the same final score for indicator 6. 28

35 Table 3.8 Results for indicator 5 Breeder organisation and technical support. Sided Trønder Western Fjord Sub indicator and Nordland cattle cattle Telemark cattle Western red polled cattle Eastern red polled cattle Breeders organisation present In situ management Stock in cryobank Technical support present Cohesion and collective dynamics of breeders.5.5 Final score.5.5 The information to score the sub indicators was obtained from an interview with N. Sæther on 18 August 217 and 24 October 217. Døla cattle Table 3.9 Results for indicator 6 Socio-economic context. Sided Trønder Sub indicator and Nordland cattle Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Young livestock farmers start off raising the breed Availability of the breed for sale Markets for products and services Labels used to distinguish products Financial support given to territories Final score The information to score the sub indicators was obtained from an interview with N. Sæther on 18 August 217 and 24 October 217. Døla cattle 29

36 3.2.7 Risk status cattle breeds In Figure 3.13 the end results of the six indicators are given for the cattle breeds. The radar charts show a large similarity among the cattle breeds. Mainly since the breeds had the same score for the change in number of breeding females (except the Western red polled cattle), breeder organisation and technical support (except the STN and Telemark cattle), and the socio-economic context. To determine the risk status of the cattle breeds the average of the six indicators was taken (Table 3.1). The overall score for the STN was equal to 1.58, indicating the breed is not at risk. The main concerns for the STN are the number of breeding females and the effective population size. The Western Fjord cattle and the Døla cattle had the same overall score of 1.83, indicating the breeds are not at risk. The main concerns for the Western Fjord cattle and the Døla cattle are the number of breeding females and the effective population size. The Telemark cattle had an overall score of 2.8, indicating the breed is not at risk. The Telemark cattle received the maximum score for the effective population size. Another concern is the number of breeding females. The Western red polled cattle had an overall score of 2.33, the highest of all breeds. The surface of the radar chart of the Western red polled cattle also was the largest compared to the other breeds. The overall score indicates the Western red polled cattle is not at risk. However, for three of the six indicators the breed had a score of 3 or higher. Solely based on these indicators the Western red polled cattle can be considered as endangered. The Western red polled cattle received the maximum score for the number of breeding females. Other concerns are the percentage of crossbreeding and the effective population size. The Eastern red polled cattle had an overall score of 2., indicating the breeds is not at risk. The Eastern red polled cattle received the maximum score for the effective population size. Another concern is the number of breeding females. In Figure 3.14 the radar charts of the cattle breeds are given, taken into account only four indicators. As mentioned before, the radar charts of the six indicators have a large similarity. In particular the indicators breeder organisation and technical support and socio-economic context are not relevant since almost all cattle breeds had the same score. Therefore, in Figure 3.14 no weight is put on the indicators breeder organisation and technical support and socio-economic context. Although the risk status is more clear in Figure 3.14, the Western red polled cattle is the only breed considered as endangered with an overall score of Table 3.1 Overall score for the cattle breeds based on six and four indicators. Six indicators a Four indicators a Sided Trønder and Nordland cattle Western Fjord cattle Telemark cattle Western red polled cattle Eastern red polled cattle Døla cattle a Breed is considered as endangered when the overall score is 3 or higher. 3

37 a Sided Trønder and Nordland cattle Context Actual Nf Trend Nf b Context Western Fjord cattle Actual Nf Trend Nf Organisation Crossbreeding Organisation Crossbreeding Actual Ne Actual Ne Telemark cattle Western red polled cattle c Context Actual Nf Trend Nf d Context Actual Nf Trend Nf Organisation Crossbreeding Organisation Crossbreeding Actual Ne Actual Ne Eastern red polled cattle Døla cattle e Context Actual Nf Trend Nf f Context Actual Nf Trend Nf Organisation Crossbreeding Organisation Crossbreeding Actual Ne Actual Ne Figure 3.13 Radar charts for the Sided Trønder and Nordland cattle (a), Western Fjord cattle (b), Telemark cattle (c), Western red polled cattle (d), Eastern red polled cattle (e), and Døla cattle (f). Actual Nf = number of breeding females, Trend Nf = change in number of breeding females, Crossbreeding = percentage of crossbreeding, Actual Ne = effective population size, Organisation = breeder organisation and technical support, and Context = socioeconomic context. 31

38 Sided Trønder and Nordland cattle Western Fjord cattle a Actual Ne Actual Nf Trend Nf b Actual Ne Actual Nf Trend Nf Crossbreeding Crossbreeding Telemark cattle Western red polled cattle c Actual Ne Actual Nf Trend Nf d Actual Ne Actual Nf Trend Nf Crossbreeding Crossbreeding Eastern red polled cattle Døla cattle e Actual Ne Actual Nf Trend Nf f Actual Ne Actual Nf Trend Nf Crossbreeding Crossbreeding Figure 3.14 Radar charts for the Sided Trønder and Nordland cattle (a), Western Fjord cattle (b), Telemark cattle (c), Western red polled cattle (d), Eastern red polled cattle (e), and Døla cattle (f). Actual Nf = number of breeding females, Trend Nf = change in number of breeding females, Crossbreeding = percentage of crossbreeding, Actual Ne = effective population size. 32

39 3.2.8 Usability multi-indicator method The multi-indicator method of Verrier et al. (215) was used to assess the risk status of the six cattle breeds. The multi-indicator method is more comprehensive than other classification methods as the one of FAO (213) and Ruane (2). In the six indicators different aspects of demography, genetics, and the social and economic environment are considered. To assess the risk status for the cattle breeds the average of the six indicators was taken. Based on the overall score of the multi-indicator method, the STN can be considered as the healthiest breed and the Western red polled cattle as the most in danger. However, one needs to be careful with the interpretation of the overall score. As mentioned before, the pedigree completeness of the STN is very low, which most likely also has an influence on the overall score of the status of the STN. The real situation of the STN can be more alarming than the overall score suggests. All cattle breeds had an overall score lower than 3, indicating the breeds can be considered as not endangered. The risk status of the breeds is in discrepancy with the risk status determined by the Norwegian Genetic Resource Centre. According to the used criteria of the Norwegian Genetic Resource Centre the cattle breeds are all endangered. The Norwegian Genetic Resource Centre uses the criteria defined by the FAO (213) specified for the low reproductivity group to assess the risk status of the cattle breeds. Based on the number of breeding females, the Telemark cattle, Western red polled cattle, Eastern red polled cattle, and the Døla cattle are considered as critical maintained. Namely, the four breeds have less than 3 breeding females. The breeds are considered as critical maintained instead of critical since the breeds are actively managed by conservation programs. The STN and Western Fjord cattle are considered as endangered maintained. For both breeds the number of breeding females are between 3 and 3 animals. The breeds are considered as endangered maintained instead of endangered since the breeds are also actively managed by conservation programs. The multi-indicator method calculates the risk status as the simple mean of the six indicators. However, it is questionable whether this is the correct method. When a breed receives the maximum score for one indicator, the other indicators indicate no risk, the average will still indicate the breed is not endangered and ignores the negative result for one indicator. This may be an explanation for the difference in the risk status of the cattle breeds as determined in this study and the Norwegian Genetic Resource Centre. The problem could be overcome by putting weight on the indicators one considered to be more important. The diversity in the categories of the multi-indicator method may also be the weak spot. The categories are not related to each other. Therefore, a breed can score no risk on the breed genetics while the social and economic environment are at maximum risk. The FAO method is mainly based on factors related to breed demography. These indicators are more related. If a breed scores low on one indicator, the breed is more likely to also score low on another indicators. 33

40 4 Implications and Recommendations In this study the same definition of the indicators and grids was used as described in Verrier et al. (215). However, the indicators and grids are based on the circumstances in France. The Norwegian conservation landscape may differ in certain aspects from the situation in France. Therefore, the proposed indicators and grids are not adequate for the Norwegian conservation framework. This was reflected in the scores of the cattle breeds for several indicators. Most breeds scored similar for the indicators change in number of breeding females, breeder organisation and technical support, and socio-economic context. The current indicators do not support the prioritisation between the cattle breeds. If one wants to use the multi-indicator method to assess which breed is more endangered and in need for conservation, several of the current indicators are not relevant. Perhaps if the breeds were in an earlier stage of conservation and the establishment of the breeder organisations was not as developed as nowadays, a difference would be visible. To identify differences that might affect the risk status of the breeds, the indicators and grids should be adapted to the local conditions and the Norwegian conservation landscape. This is in accordance with the conclusion of Verrier et al. (215). The indicators and grids could be redefined to make them more suitable for the Norwegian situation. The definition of an indicator can make a large difference. For the number of breeding females the definition used in this study was different from the one used by the Norwegian Genetic Resource Centre. In this study a time frame of one year was used for counting the number of females used as dam. While the Norwegian Genetic Resource Centre used a time frame of three years. As a result the number of breeding females used by the Norwegian Genetic Resource Centre (see paragraph Introduction) were higher than the values calculated in this study. Instead of redefining current indicators and grids an option is to put different weight on the current indicators. As was shown in the Results, when no weight was put on the non-relevant indicators the risk status of the cattle breeds became clearer. Another option is to determine additional indicators which reflect the Norwegian risk landscape more. One indicator could be the number of dairy cows versus suckler cows. In recent years the proportion of dairy cows and suckler cows has changed (results not shown). The acceptance of suckler cows varies among the breeding organisations. Therefore, the proportion dairy cows versus suckler cows may be useful to distinguish between the cattle breeds. Another suggestion for an additional indicator is the production system in relation with the number of organic farms. There seems to be a relation between the percentage of cows used on a farm which belong to one of the six native breeds and the organic character of the farm (results not shown). A large part of the organic farms keeps only purebred native cattle breeds. Another possible indicator could be the production system in relation to the use of marginal land. There seems to be a relation between farms with only purebred native cattle breeds and the use of marginal land (results not shown). The new indicators could be used to recognise early warning signs of factors that might change the status of the cattle breeds in the near future. 34

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44 Appendix I Map of Norway Figure I Map of Norway (University of Texas Libraries, 1971). 38