Seasonal migration of male red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) in southern Sweden and consequences for management

Red deer Cervus elaphus is a highly appreciated and intensively managed game species throughout Europe. A common management objective is a sustainable harvest of large trophies. In southern Sweden, management has mainly aimed at preserving the nominate subspecies C. elaphus elaphus. Seasonal migration of red deer males may, however, complicate both harvest management as well as conservation efforts. I used individually identified male red deer in southern Sweden to observe distance travelled from rutting areas to areas used by males in summer and winter. Adult males were identified by antler shape and photo-documented during rut. Photos from the rut were compared to trophies of deer harvested or found dead, to found cast antlers and to stags photographed during summer. From 1969 to 2007, a distance between rutting ground and summer/winter quarters was established for 96 identified stags. An average distance of 14 km and a maximum distance of 47 km were recorded between rut and summer/winter observations. The seasonal migration of males increases the risk of overexploitation of males with harvest in both rutting areas and wintering areas. Harvest management and conservation efforts may fail if males seasonally migrate outside the management unit. The results suggest that seasonal migration must be considered in harvest management and conservation and that there is a need for a regulation of male harvest. Furthermore, the study stresses that the success in deer management of single hunting units, may be largely dependent on the harvest policies in the near surroundings as well as in areas tenths of kilometres away, suggesting that a successful management must rely on co-operation and co-ordination on a landscape scale.     

Liver total lipids and fatty acid composition of shot red and fallow deer males in various reproduction periods

In this study total lipid (TL) content and the fatty acid composition changes in the liver of red deer stags (n=35) and fallow deer bucks (n=19) were examined at various reproductive stages. Samples were taken from mature red deer stags in the rut (September), in the post-rut (October), at the end of the rut (November) and after sexual activity had ceased (January). Sampling from mature fallow bucks was performed in the pre-rut (early October), in the rut (November) and after the reproduction period (January). The results obtained indicated significant (P<0.001) fatty degeneration of the liver in the males at the rutting season. At that time the TL concentration (x+/-S.D. in wet weight) was 156+/-40 g/kg in the red deer stags and 405+/-46 g/kg in the fallow bucks. Subsequent to the rutting season this value decreased to 47+/-15 g/kg by November in the red deer stags and to 51+/-3 g/kg by January in the fallow bucks. Significant changes were also observed in the fatty acid composition of the liver lipids, determined by gas chromatography. Liver samples collected during the rutting season from both red deer stags and fallow bucks showed higher (P<0.001) total proportions of monounsaturated fatty acids and lower (P<0.001) weight percentages of polyunsaturated fatty acids than those taken in January. In comparison with January the liver samples taken in September from red deer stags showed significantly higher (P<0.001) proportions of fatty acids C14:0, C15:0 and C:16:1, and significantly lower (P<0. 001) proportions of C18:0, C20:4 (n-6) and C22:5 (n-3). The proportions of C14:0, C15:1, C17:1 and C18:3 (n-3) in the liver samples taken in November from fallow bucks were higher (P<0.001), while the proportions of C18:0 and C20:4 (n-6) were lower (P<0.001) than those measured prior to rutting (in October) or after the rut (in January).     

Do Red Deer Stags (Cervus elaphus) Use Roar Fundamental Frequency (F0) to Assess Rivals?

It is well established that in humans, male voices are disproportionately lower pitched than female voices, and recent studies suggest that this dimorphism in fundamental frequency (F0) results from both intrasexual (male competition) and intersexual (female mate choice) selection for lower pitched voices in men. However, comparative investigations indicate that sexual dimorphism in F0 is not universal in terrestrial mammals. In the highly polygynous and sexually dimorphic Scottish red deer Cervus elaphus scoticus, more successful males give sexually-selected calls (roars) with higher minimum F0s, suggesting that high, rather than low F0s advertise quality in this subspecies. While playback experiments demonstrated that oestrous females prefer higher pitched roars, the potential role of roar F0 in male competition remains untested. Here we examined the response of rutting red deer stags to playbacks of re-synthesized male roars with different median F0s. Our results show that stags’ responses (latencies and durations of attention, vocal and approach responses) were not affected by the F0 of the roar. This suggests that intrasexual selection is unlikely to strongly influence the evolution of roar F0 in Scottish red deer stags, and illustrates how the F0 of terrestrial mammal vocal sexual signals may be subject to different selection pressures across species. Further investigations on species characterized by different F0 profiles are needed to provide a comparative background for evolutionary interpretations of sex differences in mammalian vocalizations.     

Selectivity of wolf predation on red deer in the Bieszczady Mountains, Poland

A pattern of wolfCanis lupus Linnaeus, 1758 predation on red deerCervus elaphus Linnaeus, 1758 was studied in Bieszczady Mountains in 1991–2002. In total 324 remains of red deer > 4 months old, killed by wolves throughout the year, were found. The sex, age and bone marrow fat content of wolf kills were compared with the same characteristics within the free living red deer population. The overall contribution of calves killed by wolves (24%) in October-May was higher than in the population (17%), and decreased from autumn to spring. Adult males were more vulnerable to wolf predation than adult females: stags constituted 62% and hinds 38% of adult red deer killed by wolves, whereas in the population, the percentages were 37 and 63%, respectively. Stags killed by wolves were younger ([`(x)] = 4.1\bar x = 4.1 years old) than hinds ([`(x)] = 8.9\bar x = 8.9 years old). Wolves killed more > 8 years old hinds and < 5 years old stags than available in the population. In wolf kills, the average fat content in femur marrow was higher among hinds (84.9%) than stags (69.3%) and calves (66.1%). Only 8% of hinds had < 70% femur marrow fat content, whereas 40% of calves and 38% of stags had marrow fat values below that level. Marrow fat content showed seasonal variation and was the lowest in March among all sex-age classes. The monthly share of stags in all kills, and hinds in hind-calf part of the sample was negatively correlated with their monthly average bone marrow fat content, and monthly share of calves was positively correlated with monthly average bone marrow fat content of adults. The segregation of social units (hind-calf and stag groups), except during the rutting season, and the low fat reserves of males from midwinter until spring contribute to the high overall incidence of calves and adult males and the relatively low incidence of adult females among wolf kills.     

Effects of free-ranging,semi-captive and captive management on the acoustics of male rutting calls in Siberian wapiti <Emphasis Type="Italic">Cervus elaphus sibiricus</Emphasis>

In male European red deer Cervus elaphus, rutting calls that are responsible for male reproductive success are higher in fundamental frequency in captivity than in the wild. This study compares the acoustics of stag rutting calls among wild-living, semi-captive and captive stags within an Asian subspecies of C. elaphus, the Siberian wapiti Cervus elaphus sibiricus. Male Siberian wapiti rutting calls (bugles) were collected using automated recording systems in three populations (wild-living, semi-captive and captive), all originated from the Altai/Khakasian region of Central Siberia (Russia). Selected 435 bugles (145 per population) were analysed spectrographically for 14 variables of the bugle high (>1 kHz) fundamental frequency (g0) and scored for shape of g0 contour: trapeze, descending or saddle. Among bugles, 74.3% had the trapeze contour, 23.7% had the descending contour and 2.1% had the saddle contour. The additional low (<0.2 kHz) fundamental frequency (f0) was found in 76.1% of bugles, whereas deterministic chaos was found in 16.8% of bugles. Bugles of captive stags were shortest and highest in frequency. The captive management selectively affected only bugles with the trapeze contour, whereas bugles with descending contour remained unaffected by variations of deer holding regime. Stag rutting bugles are subspecies-specific and may therefore serve as acoustic indicator of subspecies for the Siberian wapiti among other Asian and American subspecies of wapiti.     

Role of Neutral Metabolites in Microbial Conversion of 3β-Acetoxy-19-Hydroxycholest-5-Ene into Estrone

Biotransformation of 3β-acetoxy-19-hydroxycholest-5-ene (19-HCA, 6 g) by Moraxella sp. was studied. Estrone (712 mg) was the major metabolite formed. Minor metabolites identified were 5α-androst-1-en-19-ol-3,17-dione (33 mg), androst-4-en-19-ol-3,17-dione (58 mg), androst-4-en-9α,19-diol-3,17-dione (12 mg), and androstan-19-ol-3,17-dione (1 mg). Acidic metabolites were not formed. Time course experiments on the fermentation of 19-HCA indicated that androst-4-en-19-ol-3,17-dione was the major metabolite formed during the early stages of incubation. However, with continuing fermentation its level dropped, with a concomitant increase in estrone. Fermentation of 19-HCA in the presence of specific inhibitors or performing the fermentation for a shorter period (48 h) did not result in the formation of acidic metabolites. Resting-cell experiments carried out with 19-HCA (200 mg) in the presence of α,α′-bipyridyl led to the isolation of three additional metabolites, viz., cholestan-19-ol-3-one (2 mg), cholest-4-en-19-ol-3-one (10 mg), and cholest-5-en-3β,19-diol (12 mg). Similar results were also obtained when n-propanol was used instead of α,α′-bipyridyl. Resting cells grown on 19-HCA readily converted both 5α-androst-1-en-19-ol-3,17-dione and androst-4-en-19-ol-3,17-dione into estrone. Partially purified 1,2-dehydrogenase from steroid-induced Moraxella cells transformed androst-4-en-19-ol-3,17-dione into estrone and formaldehyde in the presence of phenazine methosulfate, an artificial electron acceptor. These results suggest that the degradation of the hydrocarbon side chain of 19-HCA does not proceed via C₂₂ phenolic acid intermediates and complete removal of the C₁₇ side chain takes place prior to the aromatization of the A ring in estrone. The mode of degradation of the sterol side chain appears to be through the fission of the C₁₇-C₂₀ bond. On the basis of these observations, a new pathway for the formation of estrone from 19-HCA in Moraxella sp. has been proposed.     

The induction of male sexual behavior in red deer (Cervus elaphus) by the administration of testosterone to hinds and estradiol-17β to stags

One-gram implants of testosterone were placed subcutaneously in two adult intact and one castrated red deer hinds, and 100-mg implants of estradiol-17β were given to three castrated and two intact red deer stags. Their sexual behavior was then observed in the wild and in captivity for up to 3 years. Testosterone initially produced prolonged and intense estrous behavior in the hinds; this gradually gave way to behavior normally shown only by rutting stags, such as flehmen, herding threats, and roaring. Testosterone-implanted hinds rose in the dominance hierarchy as measured by competition for food. Pregnancy, or the administration of progestagens, suppressed most of this testosterone-induced behavior. Testosterone also induced development of male secondary sexual characteristics such as formation of small antler pedicles (but not antlers), hypertrophy of the clitoris and neck musculature, and development of a neck mane and male rutting odor. Estradiol-17β mimicked the behavioral effects of testosterone when given to castrated stags, stimulating all components of rutting behavior. In intact stags, the only effects were to abolish antler casting and stimulate roaring; normal rutting behavior continued unchanged. In contrast to the effects of testosterone, estradiol-17β did not influence the social status of either intact or castrated stags. The distinctions between the behavioral effects of androgens and estrogens are ill defined and are determined more by the prior sexual differentiation of the brain and the duration of steroid treatment, than by the nature of the steroid.     

Male red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) dispersal during the breeding season

Breeding dispersal can be of significant ecological and evolutionary importance. Yet, it is seldom considered in mammals. I present data on male red deer (Cervus elaphus) movements between sub-populations in southern Sweden during the rut. I investigated whether these movements could be breeding dispersal driven by mate competition. During the ruts of 1998–2009, I recorded 91 movements of males. The longest movement distance was 18.5 km. Dispersal was not restricted to yearlings or sub-adults, but also observed among adult stags. Of 91 movements observed, 7 were made by yearlings, 46 by sub-adults and 38 by adults. There was a significant move among yearlings and sub-adults towards areas with a higher ratio of females/adult males and towards areas with more females. The movements between rutting areas thereby seemed driven by sexual competition.     

Behavioural factors affecting male reproductive success in red deer (Cervus elaphus)

Variation in reproductive success among adult red deer stags was related to harem size, and the duration and timing of breeding access. All three factors were affected by dominance (fighting success) during the rut. The number of hinds using a stag's rutting area and the duration of individual rutting activity also affected breeding access. Dominance rank in bachelor herds, which may affect access to hinds conceiving after the rut, was correlated with rank in the preceding but not the following rut.     

Role of reduced glutathione in the delta(5)-3-kitosteroid isomerase reaction of liver.

Partially purified rat liver Δ⁵-3-ketosteroid isomerase (EC 5.3.3.1) is profoundly and specifically activated by reduced glutathione (GSH). This stimulating effect shows normal saturating kinetics, and both K_m and V_max are pH-dependent. The binding of GSH is independent of the concentration of Δ⁵-androstene-3,17-dione, whereas the K_m for Δ⁵-androstene-3,17-dione is markedly reduced by saturating levels of GSH. The same catalytic site appears to isomerize both Δ⁵-androstene-3,17-dione and Δ⁵-pregnene-3,20-dione. Several steroidal inhibitors compete with Δ⁵-androstene-3,17-dione, whereas $\text{S}$-methyl-glutathione competes with GSH. This activation of Δ⁵-3-ketosteroid isomerase is also observed in the livers of other species (calf, guinea pig, human), and represents a hitherto unrecognized function of reduced glutathione.     

Identification of genes involved in inversion of stereochemistry of a C-12 hydroxyl group in the catabolism of cholic acid by Comamonas testosteroni TA441

Comamonas testosteroni TA441 degrades steroids such as testosterone via aromatization of the A ring, followed by meta-cleavage of the ring. In the DNA region upstream of the meta-cleavage enzyme gene tesB, two genes required during cholic acid degradation for the inversion of an α-oriented hydroxyl group on C-12 were identified. A dehydrogenase, SteA, converts 7α,12α-dihydroxyandrosta-1,4-diene-3,17-dione to 7α-hydroxyandrosta-1,4-diene-3,12,17-trione, and a hydrogenase, SteB, converts the latter to 7α,12β-dihydroxyandrosta-1,4-diene-3,17-dione. Both enzymes are members of the short-chain dehydrogenase/reductase superfamily. The transformation of 7α,12α-dihydroxyandrosta-1,4-diene-3,17-dione to 7α,12β-dihydroxyandrosta-1,4-diene-3,17-dione is carried out far more effectively when both SteA and SteB are involved together. These two enzymes are encoded by two adjacent genes and are presumed to be expressed together. Inversion of the hydroxyl group at C-12 is indispensable for the subsequent effective B-ring cleavage of the androstane compound. In addition to the compounds already mentioned, 12α-hydroxyandrosta-1,4,6-triene-3,17-dione and 12β-hydroxyandrosta-1,4,6-triene-3,17-dione were identified as minor intermediate compounds in cholic acid degradation by C. testosteroni TA441.     

The scent of the reticulated giraffe (Giraffa camelopardalis reticulata)

The giraffe (Giraffa camelopardalis) emits a scent that can be detected by humans over considerable distances. Dichloromethane extracts of hair samples from adult male and female reticulated giraffes (G. c. reticulata) were analysed by gas chromatography–mass spectrometry. Two highly odoriferous compounds, indole and 3-methylindole, identified in these extracts appear to be primarily responsible for the giraffe’s strong scent. Other major compounds identified were octane, benzaldehyde, heptanal, octanal, nonanal, p-cresol, tetradecanoic acid, hexadecanoic acid, and 3,5-androstadien-17-one; the last compound has not previously been identified from a natural source. These compounds may deter microorganisms or ectoparasitic arthropods. Most of these compounds are known to possess bacteriostatic or fungistatic properties against mammalian skin pathogens or other microorganisms. The levels of p-cresol in giraffe hair are sufficient to repel some ticks.     

Crossing the border? Structure of the red deer (Cervus elaphus) population from the Bavarian–Bohemian forest ecosystem

Anthropogenic impact such as overhunting and habitat fragmentation has reduced the total red deer population (Cervus elaphus) across Europe. In Germany remaining subpopulations are even confined to designated areas with limited or no gene flow among them. Red deer populations inhabiting the Bavarian–Bohemian forest ecosystem had been divided by a fortified State border between Germany and former Czechoslovakia. To assess red deer genetic diversity more than two decades after the removal of the fortifications, we analysed a population from the Bavarian Forest National Park, Germany, and one from the National Park ?umava, Czech Republic, using 11 microsatellite loci and a 910 bp long section of the mitochondrial control region (mtDNA). Bayesian analyses of microsatellite allele frequencies favoured the presence of a single population in the Bavarian-Bohemian forest ecosystem over other population genetic structures. This admixture was supported by a lack of population pairwise differentiation between German and Czech red deer microsatellite genotypes in the analysis of molecular variance (AMOVA, F_ST = 0.009, p = 0.383). Contrastingly, AMOVA revealed a highly significant matrilinear differentiation of mtDNA between the two samples (Φ_ST = 0.285, p = 0.002), whereby German red deer belonged predominantly to haplogroup A (western Europe) and Czech red deer predominantly to haplogroup C (eastern Europe). In combination, these findings indicated a high degree of philopatry by does and extensive gene flow across the former border mediated by stags. They also identified the Bavarian–Bohemian forest ecosystem as part of a suture zone between western and eastern European red deer matrilines.     

The way in which testosterone controls the social and sexual behavior of the red deer stag (Cervus elaphus)

For 10 months of the year red deer stags live together in social groups. These groups then break up, and the animals exhibit a complex pattern of sexual behavior during a short rutting season in September and October. We have utilized this clear separation between social and sexual behavior to try and establish experimentally in free-ranging wild animals the way in which testosterone influences the two types of activity.     

Chemical scent constituents in feces of wild Iberian wolves (Canis lupus signatus)

In spite of the importance of feces in scent marking and intraspecific chemical communication of many mammals, only a few studies have examined the chemical constituents of feces that could have a signaling function. We described here the chemicals found in feces recently deposited in the wild by adult and pup Iberian wolves (Canis lupus signatus). By means of analyses using gas chromatography–mass spectrometry (GC–MS), we identified 77 compounds in feces of adult wolves, mainly heterocyclic aromatic organic compounds, such as indole or phenol, but also steroids, such as cholesterol, carboxylic acids and their esters between n-C₃ and n-C₁₈, aldehydes, alcohols and significant quantities of squalene and α-tocopherol, which would increase the chemical stability of feces on humid substrates. Some of the odoriferous compounds are very likely originated in the anal-sac secretions, and later incorporated onto feces during defecation. However, the abundance of the main odoriferous compounds in feces could be explained by the result of the bacterial action on food components. Feces of pups only had 26 compounds, most of these were also found in adults’ feces, but four compounds were only found in pups’ feces, and there were also age-related differences respect to the proportions of shared chemicals. These differences could be partly due to the absence of anal-sac glands in pups, which would not need to scent mark substrates.     

Spatial and temporal components of the mating systems of kudu bulls and red deer stags

Observations were made on greater kudu (Tragelaphus strepsiceros) in the Kruger National Park in South Africa. Data on movement patterns, courtship, dominance interactions and proportion of time spent in various activities by males during the breeding season are presented. Comparative data on the apportionment of time by red deer (Cervus elaphus) setages were obtained on the Isle of Rhum, Scotland. While spatial components of male reproductive behaviour were similar in the two species, time-investments differed markedly. Kudu bulls showed little change in time spent feeding and moving during the breeding season, did not herd females and tolerated the presence of younger males. In contrast, red deer stags almost ceased feeding and greatly increased their time spent moving and in sexual or dominance-related activities. Thus the spatial and temporal components of male mating strategies can vary independently and require separate evolutionary explanaions. Comparative data for other ungulates are sparse, but suggest that time-investments are influenced by the duration of the breeding season and the breeding sex ratio. However, kudu bulls incur high mortality costs despite their low time-investments in reproductive activities.     

Induction of early rutting in male red deer (Cervus elaphus) by melatonin and its dependence on LHRH

Eight red deer stags, 2 control, 3 control-immunized (i.e. a low titre of LHRH antibodies after active immunization 2 years earlier) and 3 superior cervical ganglionectomized, were given a s.c. implant of melatonin in May at the nadir of the sexual cycle; 5 other stags remained untreated. All the melatonin-treated animals shed the velvet-like skin from the antlers in June or July at least 1 month before the untreated controls, and had an early increase in blood plasma testosterone concentrations. The treated stags were also precocious in the development of rutting behaviour, although this inductive effect was blocked dramatically in the control-immunized stags after a booster immunization against LHRH; these animals failed to show any further reproductive development and cast their antlers. It is concluded that continuous exposure to melatonin in early summer will induce premature seasonal testicular development, an effect dependent on the secretion of LHRH, and similar to that produced by exposure to short daylengths.     

Biotransformation of dehydroepiandrosterone with Macrophomina phaseolina and β-glucuronidase inhibitory activity of transformed products

The biotransformation of dehydroepiandrosterone (1) with Macrophomina phaseolina was investigated. A total of eight metabolites were obtained which were characterized as androstane-3,17-dione (2), androst-4-ene-3,17-dione (3), androst-4-ene-17β-ol-3-one (4), androst-4,6-diene-17β-ol-3-one (5), androst-5-ene-3β,17β-diol (6), androst-4-ene-3β-ol-6,17-dione (7), androst-4-ene-3β,7β,17β?triol (8), and androst-5-ene-3β,7α,17β-triol (9). All the transformed products were screened for enzyme inhibition, among which four were found to inhibit the β-glucuronidase enzyme, while none inhibited the α-chymotrypsin enzyme.     

Smells like debauchery: The chemical composition of semen-like,sweat-like and faintly foetid floral odours in Xysmalobium (Apocynaceae: Asclepiadoideae)

Floral volatiles play an important role in plant communication with both pollinators and antagonists, but remain poorly explored for many plant groups. Asclepiads (Apocynaceae: Asclepiadoideae subtribe Asclepiadinae) represent a diverse group in South African grasslands, but the scents of most species remain unexplored and few genera are sufficiently sampled to allow comparisons between congeners. I used dynamic headspace extraction methods and coupled gas chromatography-mass spectrometry (GC–MS) to examine the scent chemistry of three unusually scented asclepiads in the genus Xysmalobium and then combined these data with previously published data to explore inter- and intraspecific variation in the genus. A total of 74 compounds (33–44 per species) from various compound classes were detected in the species examined here. The sweet but faintly foetid scent of Xysmalobium asperum was dominated by epoxy oxoisophorone in combination with various other terpenoids and aromatics, and small amounts of p-cresol. The sweat-like scent of Xysmalobium tysonianum was dominated by a few aromatics in combination with isovaleric acid and several aliphatic compounds normally associated with microbial degradation or fermentation. The semen-like scent of Xysmalobium parviflorum flowers examined here contained large relative amounts of 1-pyrroline, and comparison with previously published data for dung-scented flowers from a different population revealed clear divergence in the relative amounts of this compound and p-cresol. I also detected 25 compounds that were not shared between the two X. parviflorum populations. Comparison of scent data for eight Xysmalobium species revealed very distinct chemical profiles with limited overlap between species. These results are discussed in relation to the possible roles of these volatiles as pollinator attractants and the evolution of floral scents within the genus.     

3 -Hydroxy- 5 -C 19 -and C 21 -steroid oxidoreductase activity in rat liver

The presence of small amounts of 3β-hydroxy-Δ⁵-C₁₉- and C₂₁-steroid oxidoreductase in the microsomal fraction of rat liver is shown. NAD was the preferred cofactor. K_m for the oxidation of dehydroepiandrosterone (3β-hydroxy-5-androsten-17-one) into androstenedione (4-androstene-3,17-dione) was 3 × 10⁻⁶ M. In similarity to the adrenal and gonadal 3β-hydroxy-Δ⁵-C₁₉-steroid oxidoreductase, but in contrast to the hepatic 3β-hydroxy-Δ⁵-C₂₇-steroid oxidoreductase involved in the biosynthesis of bile acids, the hepatic 3β-hydroxy-Δ⁵-C₁₉-steroid oxidoreductase was inhibited by the 3β-hydroxy-Δ⁵-steroid oxidoreductase inhibitor, 2α-cyano-4,4,17-trimethyl-17β-hydroxy-5-androsten-3-one, and the activity was greatly reduced with microsomes from immature rats.