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In some families of large mammals (Cervidae, Bovidae, Canidae), we examined relationships between the various mating systems adopted and biochemical-genetic variation measured in terms of the mean proportion of polymorphic loci (P), mean heterozygosity (H) and derived coefficients, such as the ratio Pt:P and FIS. Our hypothesis was that genetic variability decreases as the degree of polygyny of the mating system adopted increases. Most of the data were in accordance with this prediction, but also some ambiguous results could be observed. Methodological and practical difficulties connected with our synoptical approach, such as the lack of comparability among most electrophoretic studies and the general scarcity of quantitative behavioural data are critically discussed.
To investigate phylogenetic relationships among 9 genera of the Caprinae (Capra, Ammotragus, Hemitragus, Pseudois, Ovis, Rupicapra, Oreamnos, Nemorhaedus, Capri- cornis) behaviours involved in courtship and mating, aggression, threat, dominance, submission, and marking of adult males were subjected to phylogenetic analysis. Based on all 32 characters and 96 character states investigated, phylogenetic patterns generally were in good agreement with biochemical-genetic data avaliable. Discordance between phylogenetic trees constructed from behavioural and from biochemical-genetic traits as to the position of Ammotragus turned out to be associated with ethological functional categories. Behaviours involved in courtship and mating were identified as the most reliable ones for phylogenetic studies. Courtship displays function as isolation mechanisms among closely related taxa. This is of paramount importance in those forms where secondary sexual characters such as horns are poorly differentiated while in highly evolved taxa size and shape of horns may also trigger readiness for mating in estrous females.
Antler traits (length of the main beam, circumference of the main beam, coronet circumference, and the number of antler points) in roe deer Capreolus capreolus (Linnaeus, 1758) from a population in Casentino (Northern Tuscany) were examined for associations with genotypes at loci coding for enzymes. Significant associations were found only in yearlings. Individuals homozygous for the allele Mpi120 had significantly larger antlers than carriers of other genotypes at Mpi. Individuals homozygous for Pep-2100 had significantly smaller antlers than carriers of other genotypes at Pep-2. In adults the results were essentially the same, but only as a statistically insignificant trend. The data were interpreted in terms of one or more genetic components having a major influence on antler development only in yearlings. This corresponds with behavioural data suggesting that in adults antler size is not related to reproductive success, but in yearlings it is correlated with sexual maturity and the rate of being expelled from the home range of their mothers by territory holders. The situation in the roe deer was compared with previous results on the genetics of antler development in the red deer, where males are social rather than solitary, yearlings do not participate in reproduction, and some antler characteristics are related to reproductive success of adult stags.
In Asian elephant Elephas maximus Linnaeus, 1758 tuskless bulls or maknas are generally rare. Only in Sri Lanka 93% of subadult and adult bulls have been reported to be maknas. Using historical records and computer simulations we demonstrate that this situation is man-made. The following mechanisms were identified to be associated with a loss of tuskers: (1) When using elephants, man has always preferred tuskers. (2) Selective hunting and capturing frequently led to a decrease of tuskers in wildliving populations. (3) The impact of selective hunting and capturing was highest in isolated populations, such as Sri Lanka. (4) Selective removal of tuskers for protecting a maximum wild-living male population resulted in an increase of maknas. The rate of increase in the frequencies of maknas in particular populations with known history could be best explained by a dominant mode of inheritance of tusks in combination with a slight advantage of tuskers in reproduction. For the mainland populations it can be predicted that even in those where tuskers are already largely lacking the allele responsible for the expression of tusks should often be sufficiently abundant to allow the recovery of tusk bearing males.
Restriction fragment length polymorphisms (RFLPs) of mitochondrial (mt) DNA were used for investigating genetic differentiation in chamois (genus Rupicapra). Digestion of the mtDNAs of 58 individuals from 6 populations with a battery of 16 six-base cutting (restriction endonucleases yielded a total of 67 restriction sites. Based on the presence and absence of these restriction sites a total of 8 haplotypes could be defined. Six of them served for assessing genetic diversity within and among 4 local populations of R. rupicapra rupicapra. Estimates of nucleotide divergence among those haplotypes ranged from 0.05% to 0.25%. One chamois from the High Tatra (subspecies R. r. tatrica) was examined and showed the standard haplotype found in R. r. rupicapra. MtDNA in chamois from Catalunya, belonging to R. pyrenaica pyrenaica, was polymorphic for two haplotypes not found in any population of R. rupicapra. Mean nucleotide divergence among haplotypes found in R. rupicapra and R. pyrenaica was 0.56% (SD = 0.16%). Based on this value, an estimated divergence time of about 280 000 years suggests that the mtDNA lineages of R. rupicapra and R. pyrenaica separated prior to the Riss glacial in the later Pleistocene.
Based on previously published electrophoretic data on genetic variability in 31 roe deer Capreolus capreolus Linnaeus, 1758 populations, the proportion of loci polymorphic (P), average heterozygosity (H), and the inbreeding coefficient (FIS) were examined for relationships with the social structure displayed in the various populations. Our hypothesis was that genetic variability is lower and FIS-values are more positive in populations where males maintain a stable pattern of territories during the rutting season (forest dwelling roe deer) than in those characterized by pronounced fluctuations in population structure, both within and among seasons (field or mountain dwelling roe deer). P and H did not show differences among those two groups. FIS was significantly more positive in the 'forest' roe deer than in the more migratory 'type', but only when populations subjected to high culling rates were excluded from the analysis. Highly negative FIS-values in forest populations with high culling rates suggested that considerable perturbations of population structure may be caused by hunting. In conclusion, the 'forest' roe deer and the 'field' roe deer do not represent two distinct ecotypes with a particular genetic integrity, but rather reflect the considerable behavioural plasticity of the species.
A total of 469 brown hares Lepus europaeus Pallas, 1778 from 20 sampling sites in Austria were examined for genetic diversity within and among populations by means of horizontal starch gel electrophoresis. Fourteen out of 54 presumptive structural loci were polymorphic, one of which was excluded from further population genetic analyses due to the occurrence of a null-allele. The mean proportion of polymorphic loci (P) was 15.3% (SD 2.2%), and mean expected average heterozygosity (He) was 4.6% (SD 0.5%). Both relative (Fst = 5.4%) and absolute (mean Nei's 1978 D = 0.0016, SD 0.0016) genetic differentiation among populations were low, suggesting a generally high level of migration. Cluster analysis revealed some separation of brown hare populations in western and northern Austria from those in the east and in the south. In 131 individuals, mtDNA was digested with a battery of 16 restriction endonucleases. Besides the standard type I which occurred exclusively in most of the populations, five additional haplotypes, each of them deviating from type I by one base pair substitution, were detected. Together with rare alleles at allozyme loci, the distribution of variant haplotypes corroborated the spatial pattern obtained by allozyme distances and suggested considerable immigration of brown hares from the adjacent countries in the east and south. Twenty non-metric skull traits were scored in 443 individuals. Character variants were dichotomized (0/1) and the respective frequencies were used to calculate C. A. B. Smith's 'mean measure of divergence' (MMD) among five population groups. Morphological differentiation was in accordance with the major population genetic pattern as revealed by molecular techniques. MtDNA variation (nucleon diversity, nucleotide diversity) and morphological variation (mean of SD in single characters) within populations were not significantly associated with one another, and did not show a relationship with indices of genetic variation obtained by allozyme analysis. These findings suggest that variability in only one of these characters cannot be considered representative for overall gene pool diversity within populations.
Biochemical-genetic variation was studied in springbok Antidorcas m. marsupialis (Zimmermann, 1780) from a large (N > 2000) "wild" population (n = 24) and a small (30 > N > 20) isolated farm population (n = 10) using electrophoretic allozyme analysis. Springbok showed polymorphisms at eight out of 46 loci. The springbok from the large population had a higher proportion of polymorphic loci (P = 15.6%) than those from the small population (P = 8.9%). Average heterozygosity (H = 5.1% and H = 4.1%, respectively) was similar for the two populations. This unexpected result is an artefact of the method for calculating H. H:P ratios are lower for the large population than the small one. The distribution of genotypes differed significantly from Hardy-Weinberg equilibrium for two loci. These were found to have a preponderance of homozygotes. This could not be explained by population fragmentation. The levels of polymorphism and heterozygosity are high compared to results from other African bovids.
A total of 598 Austrian and 1117 Polish skulls of brown haros Lepus europaeus Pallas, 1778 was examined for dental anomalies, occurrence of Wormian bones and formation of a fenestra in the fossa mandibulars assis temporalis. In addition, the degree of ossification of skull sutures and frequencies oi traceable interparietal sutures were analysed in Austrian hares. Frequencies of missing M3 were 1.95% for Austrian and 2.1% for Polish hares from 1986 - 1990. No significant differences as to age or sex were found for incidences of missing M'1; the left side of the tooth row was significantly more often affected by missing M'\ which indicates directional asym­metry. Rotation of the I2 (mainly of low magnitude) was noticed in 27.9% of subadult and adult Austrian hares. Occurrences of all other dental anomalies such as abnormal shape of M', missing I2, missing M3, supernumerary I2, additional upper molar and various cases of malformation or irregular position of teeth were very low, respectively. Ossification of skull sutures was presently rcduced as compared to the material in­vestigated by Cabofi-Raczyftska (1964) and regional differences in the degree of suturai obliteration were found. Temporal fenestration was encountered in 24.6% of subadult and adulL Austrian hares and in 11.6% in Polish hares sampled during 1986 - 1990 (p < 0.001). Frequency of temporal fenestration increased with age: 2.9% in fetuses/ neonates, 16.5% in subadult and 28.1% in adult hares from Austria (p < 0.01, d.f. = 2); 8.7% in subadult and 23.8% in adult Polish specimens. Average frequency of Wormian bones in Austrian hares was 4.8% and 4.4% in the Polish material. Within subadults 3.7% of males and 16.3% of females (p < 0.01) exhibited a traceable outline of the os interparietale.
Red lechwe Kobus leche leche Gray, 1850 (n = 3), black lechwe K. I. smithermani Lydekker, 1900 (n = 10) and Kafue lechwe K. I. kafuensis Haltenorth, 1963 (n = 19) from Zambia were examined for genetic variability and differentiation at 30 pre­sumptive structural loci using horizontal starch gel electrophoresis. Values of polymor­phism (P = 10.0-16.7%) and average heterozygosity (H = 6.3-7.9%) were within the range commonly found in ungulates. Genetic variability was lowest in the red lechwe, which may be due to a genetic bottleneck the Zambia population experienced some 50 years ago. Relative (fst = 21%) and absolute (Nei's 1978, d = 0.020-0.023) genetic differentiation were in accordance with the subspecies status proposed for red lechwe, black lechwe, and Kafue lechwe on the basis of morphological characters.
In order to provide a scale of genetic distances in the Lagomorpha, biochemical- systematic relationships among Lepus europaeus, Lepus timidus, Oryctolagus cuniculus (Leporidae) and Ochotona rufescens (Ochotonidae) were examined by hori­zontal starch gel electrophoresis of 38 isozyme systems. Nei's (1978) genetic distances were calculated over 58 presumptive structural loci and used for the construction of numerical dendrograms. The stability of clusters was examined by the jackknife method and by comparison to a Hennigian cladogram. All these procedures revealed a constant picture of lagomorph relationships, which is in accordance with the conclusions drawn from other evidence. Divergence times were estimated using two fundamentally different approaches. They were in good agreement with paleontological data (0.49myr between the Lepus species, 3.65myr between Lepus and Oryctolagus, 37.5myr between Leporidae and Ochotonidae), but only when calculated in different ways at low and at high taxonomic levels. The results suggest a temporal acceleration of the rate of allozyme evolution in the Leporidae due to rapid adaptive radiation of biochemically highly polymorphic taxa.
A total of 193 brown hares, collected from 7 sampling sites in Poland during 1986 - 1990 were examined for genetic variability and differentiation at 39 presumptive isozyme loci by means of horizontal starch gel electrophoresis. Values of polymorphism (mean P = 0.180, SD 0.039) and average heterozygosity (mean H = 0.047, SD 0.006) were similar to those detected in previous studies on the population genetics of the brown hare. Relative (Cst = 0.041) and absolute (mean D/Nei, 1978/ = 0.0012, SD 0.0013) genetic differentiation among populations were very low, which fits well to the high number of migrant individuals per generation (Nm = 12.7), estimated using the private allele method of Slalkin (1985). Average heterozygosity was examined for associations with geographical distribution, the year of culling, population density, age, sex, body weight and health status, whereby a better survival of heterozygous femals could be detected. According Lo our results, the present decline of the brown hare is noL due to genetic depiction. However, once population sizes drop below a critical threshold, a pronounced inbreeding depression can be expected.
For approximately 100 years blesbok - endemic to South Africa - have been extinct in the wild and confined to fenced game reserves or farms. Biochemical-genetic variation was studied in blesbok from five isolated populations using electrophoretic allozyme analysis. Body weights and liver mineral concentrations were also determined. Material was collected from three localities in the Orange Free State province: a large reserve (PRE, ca 10 000 ha, N = 500-600, n = 23); a smaller reserve (KOP, ca 3 000 ha, N = 150-200, n = 14) with animals derived from the same source; and a farm (MID, ca 4 000 ha, N = up to 700, n = 19). The other two localities were a farm in the northern Cape Province (BEN, ca 10 000 ha, N = 200, n = 18) and another in the southern Cape Province (BRA, ca 150 ha, N = 50-80, n = 27), both with populations derived from small founder stocks. Three loci were polymorphic: Pgm-1, Acy-1, and Gpi-1 but Acy-1 was the only one polymorphic in all five populations. Pgm-1 was polymorphic in two populations derived from the same source and Gpi-1 in the other from the Orange Free State. Calculated over 45 presumptive structural loci the mean proportion of polymorphic loci (P) was 3.5% (SD = 1.2%), and mean expected average heterozygosity (He) was 0.9% (SD = 0.25%). The populations separated out by genetic distance in two distinct groups, those from the Cape Province and those from the Orange Free State. There were considerable differences in mean body weight between some sites. No correlation could be detected with level of heterozygosity. Body weight appeared rather to be related to liver mineral levels. In particular the ratio between copper and molybdenium appears important with those animals high in copper and low in molybdenium having a higher body weight.
European red deer are known to show a conspicuous phylogeographic pattern with three distinct mtDNA lineages (western, eastern and North-African/Sardinian). The western lineage, believed to be indicative of a southwestern glacial refuge in Iberia and southern France, nowadays covers large areas of the continent including the British Isles, Scandinavia and parts of central Europe, while the eastern lineage is primarily found in southeast-central Europe, the Carpathians and the Balkans. However, large parts of central Europe and the whole northeast of the continent were not covered by previous analyses. To close this gap, we produced mtDNA control region sequences from more than 500 red deer from Denmark, Germany, Poland, Lithuania, Belarus, Ukraine and western Russia and combined our data with sequences available from earlier studies to an overall sample size of almost 1,100. Our results show that the western lineage extends far into the European east and is prominent in all eastern countries except for the Polish Carpathians, Ukraine and Russia where only eastern haplotypes occurred. While the latter may actually reflect the natural northward expansion of the eastern lineage after the last ice age, the present distribution of the western lineage in eastern Europe may in large parts be artificial and a result of translocations and reintroduction of red deer into areas where the species became extinct in historical times.
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