Genetic influences on reproduction of female red deer (Cervus elaphus) (1) seasonal luteal cyclicity

This study compared the onset and duration of the breeding season of female red deer (Cervus elaphus scoticus) and its hybrids with either wapiti (Cervus elaphus nelsoni) or Père David's (PD) deer (Elaphurus davidianus). In Trial 1 (1995), adult red deer (n=9), F1 hybrid wapiti x red deer (n=6) and maternal backcross hybrid PD deer x red deer (i.e., 14 PD; n=9) were maintained together in the presence of a vasectomised red deer stag for 12 months. They were blood-sampled daily or three times weekly so that concentration profiles of plasma progesterone could be used to identify the initiation, duration and cessation of luteal events. There was clear evidence of luteal cyclicity between April and September, with the transition into breeding associated with an apparent silent ovulation and short-lived corpus luteum (i.e., 6-12 days) in every hind. A significant genotype effect occurred in the mean time to first oestrus (P<0.05), with wapiti hybrids and 14 PD hybrids being 9 and 5 days earlier than red deer. Between six and nine oestrous cycles were exhibited by each hind, with no difference in mean cycle length (19.5-19.6 days) between genotypes (P0.10). The overall length of the breeding season was significantly longer for wapiti hybrids (143 days) than for either red deer (130 days) and 14 PD hybrids (132 days, P<0.05).In Trial 2 (1998), adult red deer (n=5), 14 PD hybrids (n=5) and F(1) PD x red deer hybrid (n=5) hinds were maintained together from mid-February (late anoestrus) to early May, in the presence of a fertile red deer stag from 1 April. Thrice-weekly blood sampling yielded plasma progesterone profiles indicative of the onset of the breeding season. Again, there was a significant genotype effect on the mean time to first oestrus (P<0. 05), with F(1) PD hybrids and 14 PD hybrids being 13 and 5 days earlier than red deer. However, conception dates were influenced by the timing of stag joining, and were not significantly different between genotypes. The results indicate genetic effects on reproductive seasonality. However, seasonality observed for PD x red deer hybrids more closely approximated that of red deer than PD deer.     

Genetic structure of the Danish red deer (Cervus elaphus)

The red deer (Cervus elaphus) population in Denmark became almost extinct in recent historical times due to over‐hunting. The species has subsequently recovered within remote areas, but non‐Danish individuals have been introduced at several localities. To assess genetic structure, past demographic history, and the possibility of a still existing original stock, we analysed 349 specimens from 11 geographically separate areas and from three enclosed areas, genotyping 11 microsatellite loci. Moreover, an 826‐bp fragment of the control region of the mitochondrial DNA was sequenced for 116 recent specimens and seven museum specimens. There was a significant difference in mean expected heterozygosity (H_E) between the three enclosed areas and the 11 unenclosed areas. Significant departures from Hardy–Weinberg equilibrium were observed in the three enclosed areas and in nine of the unenclosed areas. The overall degree of genetic differentiation among all 14 areas was significant (F_ST = 0.09, P < 0.01), primarily because the mean pairwise F_ST for the three enclosed areas was significantly higher than that for the 11 unenclosed areas. A Bayesian clustering procedure detected three genetically distinct populations and indicated reduced gene flow between the enclosed and unenclosed areas. The individuals in the unenclosed areas show genotypic mixture, presumably as a result of gene flow among them. Markov Chain Monte Carlo simulations, based on the genealogical history of the microsatellite alleles, suggest a drastic decline in the effective population size of the enclosed areas some 188–474 years ago. Mitochondrial DNA analysis of the recent specimens showed seven haplotypes. Individuals from the enclosed Jægersborg Dyrehave contain haplotypes that occur all over Denmark and also are found in Western Europe. A close relationship between Scandinavian and Western European red deer is most likely. Only individuals from the unenclosed Lindenborg Estate and the enclosed Tofte Skov did not group with any other Danish individuals. As six of seven museum specimens had haplotypes also found in modern Danish samples, the current population of red deer in Denmark is genetically close to the original Danish red deer. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 688–701.     

Factors affecting reproduction in red deer (Cervus elaphus) hinds on Rhum

In a population of free-ranging red deer hinds on the Isle of Rhum (Inner Hebrides) we investigated relationships between four aspects of reproductive performance (fertility, calf birth weight, birth sex ratio and calving date) and four variables likely to affect the mother's condition: age, reproductive status, home range area and year of calving. Fertility was significantly related to mother's age, reproductive status and home range area as well as to year of calf's birth. Stag calves were heavier than hind calves and birth weight was significantly related to mother's age, home range area and year of (calf's) birth but not to mother's reproductive status. Birth sex ratio did not differ from parity, and was not significantly associated with any of the four variables examined. Birth date was significantly related to the mother's reproductive status, home range area and year of (calf's) birth but not to mother's age or the sex of the calf.     

Reproductive seasonality in female Iberian red deer (Cervus elaphus hispanicus)

This study characterized the seasonal pattern of luteal cyclicity in Iberian red deer (n=16), by measuring plasma progesterone concentrations in hinds (female red deer) twice per week from calving (May and June) 1996 until May 1997. In eight of these hinds we also examined plasma prolactin profiles to assess seasonal responses to photoperiod. Plasma progesterone concentration in the 16 hinds studied indicated that the reproductive pattern is seasonal, and lasts for 5.73 +/- 0.27 months. After calving, progesterone levels remained basal (no luteal activity) for several months, except in a hind that lost her calf just after calving, and thus did not have to suckle it. This hind showed two consecutive estrus cycles in the month following calving, which suggests that suckling has an inhibiting effect on the resumption of ovarian activity. These results also showed that as long as the hinds do not become pregnant, they show between 5 and 10 estrus cycles per reproductive season (8.06 +/- 0.35), ranging between 105 and 249 days from onset of the first cycle to end of the last one. Uninterrupted cycling lasted for 3.5-6.4 months (mean, 4.6 +/- 0.24). Cyclic luteal activity was found from October to February in all hinds, with a smaller, but notable proportion in September (56.25%) and March (68.8%), whereas it was negligible in the remaining period. Our results show a reproductive season similar to or longer than that recorded by other authors. Prolactin plasma concentrations showed a yearly trend following that of photoperiod, with peak concentrations from April to July, a decrease in August, minimal concentrations from September to February and a sharp increase in March.     

Physiological effects of hunting red deer (Cervus elaphus).

When red deer (Cervus elaphus) were hunted by humans with hounds the average distance travelled was at least 19 km. This study of 64 hunted red deer provides the first empirical evidence on their state at the time of death. Blood and muscle samples obtained from hunted deer after death were compared with samples from 50 non-hunted red deer that had been cleanly shot with rifles. The effects on deer of long hunts were (i) depletion of carbohydrate resources for powering muscles, (ii) disruption of muscle tissue, and (iii) elevated secretion of beta-endorphin. High concentrations of cortisol, typically associated with extreme physiological and psychological stress, were found. Damage to red blood cells occurred early in the hunts; possible mechanisms are discussed. Taken together, the evidence suggests that red deer are not well-adapted by their evolutionary or individual history to cope with the level of activity imposed on them when hunted with hounds.     

Genetic roots of the red deer (Cervus elaphus) population in Eastern Switzerland

Overhunting of red deer (Cervus elaphus) in eastern Switzerland led to its extinction in the second half of the 17th century. Natural recolonization must have taken place later, because red deer were seen again in the canton of the Grisons (eastern Switzerland) in the 1870s. According to historical data, three different populations could have served as the source population. To determine the genetic origin of the eastern Swiss red deer population, we collected samples from five different subpopulations in the canton of the Grisons as well as from four adjacent populations in Germany, Liechtenstein, Austria, and Italy. We analyzed the samples by genotyping 18 microsatellite loci. F(ST) values, assignment tests, correspondence analysis, and fuzzy clustering clearly pointed to Liechtenstein as the most probable source population for the red deer in eastern Switzerland. In addition, our analyses revealed high gene diversity in all examined populations. Gene flow and the high genetic admixture are discussed.     

Genetic diversity in European red deer (Cervus elaphus L.): anthropogenic influences on natural populations

Allozyme, microsatellite and mtDNA (RFLP and sequence) data of European red deer populations were examined as to their capability of indicating anthropogenic influences such as the keeping of animals in enclosures, selective hunting for trophies translocation of specimens to improve trophy quality and habitat fragmentation. Deer in enclosures revealed considerable deviations of allele frequencies from isolation-by-distance expectations but no remarkable loss of genetic diversity. Particular allozyme genotypes were associated with antler morphology, and selective hunting was shown to alter allele frequencies in the expected direction. Habitat fragmentation is reflected by various kinds of genetic markers but due to the lack of information on population histories no unequivocal evidence on particular human activities could be obtained.     

Malignant melanoma in a captive red deer (Cervus elaphus elaphus)

We report a case of malignant melanoma in a captive red deer (Cervus elaphus elaphus). The primary lesion was on the plantar aspect of the right pastern and infiltrated the surrounding skin, but not underlying tendons or bone. Diffuse metastasis to the regional lymph nodes had occurred. Histologically, lesions were characterized by large numbers of variably pigmented epithelioid cells with a high degree of nuclear atypia and many mitotic figures. This is the first reported case of this neoplasm in red deer.     

Phylogeography of red deer (Cervus elaphus) in Europe

Aim To investigate the phylogeographical patterns of red deer (Cervus elaphus) in Europe, and to disentangle the influence of ancient (e.g. Pleistocene ice ages) from more recent processes (e.g. human translocations). Location Europe. Methods In this study we provide by far the most extensive analysis of genetic structure in European red deer, based on analyses of variation at two mitochondrial markers (cyt b and D‐loop) in a large number of individuals from 39 locations. Relationships of mitochondrial DNA haplotypes were determined using minimum spanning networks and phylogenetic analyses. Population structure was examined by analyses of molecular variance. Historical processes shaping the present patterns were inferred from nested clade analysis and nucleotide diversity statistics. Results Within Europe, we detected three deeply divergent mitochondrial DNA lineages. The three lineages displayed a phylogeographical pattern dividing individuals into western European, eastern European and Mediterranean (Sardinia, Spain and Africa) groups, suggesting contraction into three separate refugia during the last glaciation. Few haplotypes were shared among these three groups, a finding also confirmed by F_ST values. Calculations of divergence times suggest that the groups probably split during the Pleistocene. Main conclusions The observed pattern is interpreted to result from isolation in different refugia during the last glaciation. The western and eastern European lineages could be linked to an Iberian and Balkan refugium, respectively. The third lineage might originate from a Sardinian or African refugium. We link local phylogeographical patterns observed in Europe to the post‐glacial recolonization process, shaped by the geographical localization of refugia and barriers to gene flow. Regardless of the importance of red deer as a game species and the tradition of translocating red deer in Europe, we detected few individuals that did not match the trichotomous pattern, suggesting that translocations have occurred mainly at smaller spatial scales.     

Mitochondrial DNA phylogeography of red deer (Cervus elaphus)

In order to understand the origin, phylogeny, and phylogeography of the species Cervus elaphus, we examined the DNA sequence variation of the mitochondrial cytochrome b gene of 51 populations of deer from the entire distribution area of Cervinae with an emphasis on Europe and Asia. Several methods, including maximum parsimony, maximum likelihood, and nested clade analysis, revealed that red deer originated from the area between Kyrgyzstan and Northern India. We found two distinct groups of red deer: a western group consisting of four subgroups and an eastern group consisting of three subgroups. Our mtDNA data do not support the traditional classification of red deer as only one species nor its division into numerous subspecies. The discrepancies between the geographical pattern of differentiation based on mtDNA cytochrome b and the existing specific and subspecific taxonomy based on morphology are discussed.     

Lactation under induced five-month delay reproduction in Iberian red deer (Cervus elaphus hispanicus)

In Iberian deer, calf birth date affects both milk production and composition, as well as the weight change of hind during lactation and calf growth. This study examines a five-month induced delay in calving (autumn group) in a group of 19 Iberian hinds compared to a group of 13 standard calving hinds (spring group). Fertility in the autumn group was half that of the spring one. From the former, a further two calves out of six died before or soon after birth. Successfully lactating hinds showed a greater concentration in protein, fat and lactose (P < 0.001), but did not differ from spring group in total milk yield, calf gain or weight change of hinds during lactation. However, after correcting for the greater calf or hind body weight at calving of the autumn group, this group showed lower values for these variables than the spring group. Results show that at least in Iberian deer, counter-season reproduction is possible, and that the effects on lactation variables are similar to those of a natural delay in calving.     

Spatial organization of female groups in red deer (Cervus elaphus L.)

The spatial organization of female groups in red deer was studied in the periods before and after the rut, in a study area located in the central Sierra Morena (Córdoba, Spain). The study was carried out in terms of age-sex classes, and was based on direct observation and video recording.Movement order proved to be linear, and highlighted the role of the mature hind as group leader and potential group protector. Two tendencies were found in the calf: to place itself behind the adult female, and to take up a central position in the group. With regard to relationships of spatial proximity, the mature hind and the calf tended to keep separate from the rest of the group both in the period before and after the rut. The yearling male became progressively independent of the matriarchal group, tending in the second period of study to associate with individuals from its own age-sex class.     

Activity patterns of female red deer (Cervus elaphus L.) in the Alps

Summary During 1976 to 1978 in the Bavarian Alps (Southern West Germany) the activity patterns of nine female red deer have been studied using radio-telemetry. The daily sum total of activity shows a marked annual cycle, ranging from about 7 h in winter to some 15 h in summer. It becomes obvious that the duration of activity during solar day, night and twilight is not only due to the length of these three periods but also varies independently in the course of the year. The daily pattern of activity reveals a bimodal 24-h rhythm throughout the year. Its two peaks coincide with dawn and dusk. Their temporal position shifts according to the seasonally changing LD-ratio. Also the relative height of the two peaks varies in the course of the year; they decrease in the summer months in favour of more activity during the light time. These daily and annual rhythms rely on corresponding changes in the pattern of activity bursts. In some cases the sequences of activity bursts reflect also influences of weather conditions or of differently structured habitats on the animals' behaviour. The possible influences of some endogenous and exogenous factors on the activity patterns are discussed.     

Fine-scale genetic structure of red deer (Cervus elaphus) in a French temperate forest

Despite the classic population genetic view of a population as a network of sub-populations consisting of randomly mating individuals, the mating system and dispersal patterns of social animals affect the distribution of genetic variation on a local scale. The spatially open, forest-dwelling red deer (Cervus elaphus) population at the Petite Pierre National Reserve in north-eastern France is culled annually, with the management aim of maximising the number of adult males in the population, and is a typical example of an exploited red deer population from continental Europe. Through a change in management policy, the number of adult males in the population has increased over time, leading to a reduction in variance of male reproductive success (Bonenfant et al., 2002). In this study, we investigate the fine-scale genetic structure of the population using 14 microsatellite loci and attempt to find evidence for a change in this genetic structure over time. DNA was extracted from bone powder obtained by drilling into antlers and mandibular condyles. DNA was successfully extracted from up to 30-year-old samples, but it was necessary to genotype samples in duplicate to obtain reliable genetic profiles. Our results point towards a pattern of fine-scale spatial structure amongst female red deer in the study area, but not amongst males, as would be expected for a typical mammalian system with male-biased dispersal and female philopatry. In addition, our results hint at a decrease in spatial genetic structure amongst females over time, which might be related to a change in management policy, but small sample size limited the robustness of this conclusion.     

Onchocercosis in red deer (Cervus elaphus) from Spain.

Onchocercosis, caused by Onchocerca flexuosa, was observed in red deer (Cervus elaphus) from Spain for the first time. Adult specimens of P. flexuosa were found in nodules in subcutaneous tissues in 42 of 125 (33%) red deer between October 1994 and September 1995; intensity of infection +/- SD was 3.93 +/- 5.26 nodules per infected host. A clear seasonal pattern in the distribution of nodules was observed, with higher values of prevalence and intensity in fall and winter in contrast to spring and summer. Significant differences were found among age groups in prevalence, but not in the mean intensity. No differences in infection were apparent between male and female adult red deer.     

The influence of birth date on the development of seasonal cycles in red deer hinds (Cervus elaphus)

The relative importance of age and photoperiod in the timing of seasonal cycles was studied in red deer during the first 18 months of life. Seasonal cycles of appetite, live-weight change, coat growth, oestrus and metabolic rate were examined in two groups of seven hind calves born in early May (group E) or nine weeks later in July (group L).
There was no difference between groups in the timing or amplitude of the seasonal voluntary food intake (VFI) cycle, or in the timing of seasonal changes in rate of live-weight gain, although mean live-weight gain was significantly greater for group L than group E during the first winter of life.
Four animals in group L retained their calf coats throughout the first winter. In group L, the first summer coat fibres appeared approximately four weeks earlier, and mean fibre length for the subsequent winter coat was longer in group L than in group E. There was no difference between groups in the timing or amplitude of the seasonal cycle of plasma prolactin concentration, or in the incidence and mean time of onset of puberty.
There was no difference between groups in mean energy expenditure per unit of live weight at any time of the year. Mean daily energy expenditure was significantly higher in summer than in winter, but values measured when the animals were resting and corrected for differences in VFI were similar throughout the year.
It is concluded that the seasonal cycles of appetite, live-weight change and oestrus in young red deer hinds are not modified by age, although the normal cycle of coat replacement can be disturbed in animals born towards the end of the calving season.