Male Fallow Deer Research Articles

Is the parallel walk between competing male fallow deer, Dama dama, a lateral display of individual quality?

During competitive encounters protagonists are expected to use signals of individual quality particularly if there is a risk of injury or death. Lateral presentation of body profile, by which information regarding phenotypic characteristics associated with individual quality are displayed, may represent such a strategy. During aggressive interactions, male fallow deer frequently engage in parallel walking which is assumed to represent a mutual display of quality, as mediated by exposure of the maximal profile of the body or antlers. We examined the context and role of the parallel walk during competitive encounters to investigate whether there was evidence that dyads of competing males were assessing differences in phenotypic characteristics. There was no evidence to support the hypotheses that the parallel walk is a lateral display of body size or weaponry or that its use is associated with a reduced level of escalated or risky behaviours during fighting. Total time spent fighting was not shorter when a parallel walk was present than when there was no parallel walk. The parallel walk was highly associated with fighting and it was more likely to be initiated by the subsequent loser. Furthermore, parallel walking frequently followed bouts of fighting and as such may represent a strategy that permits an animal the opportunity to decide whether to continue fighting. Parallel walking was also associated with a failure to resolve contests in favour of one animal indicating that it may be a means of withdrawing from further fighting without incurring a loss in dominance status. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.

Meat quality in male fallow deer ( Dama dama): effects of age and supplementary feeding

Thirty-two male fallow deer, half 18- and half 30-month-old, were slaughtered after a 4-month feeding trial on pasture alone or with daily supplement of 500 g (dry matter) concentrate/head (eight each 18- and 30-month-old). Neither diet nor age produced significant effects on M. longissimus thoracis et lumborum (LM) and on M. semimembranosus (SM) pH and colour. Meat texture properties (Warner–Bratzler–WB Shear Force and compression), water loss and collagen content were unaffected by diet, while the older deer had higher LM WB peak force values (4.15 vs 4.88 kg) and lower collagen solubility (34.61 vs 22.58%). LM and M. semitendinosus (ST) of the supplemented deer had a higher content of fat (pasture vs concentrate feeding: 0.56 vs 0.72% in LM; 0.55 vs 0.78% in ST) and provided lower PUFA, particularly n-3 PUFA, and higher n-6/n-3 ratio (3.30 vs 4.76 in LM; 3.39 vs 4.63 in ST). Thirty-month-olds’ LM and ST were fatter than 18-month-olds’, and provided lower PUFA, both in the n-6 and n-3 fraction.

Some Haematological Values of Farmed Fallow Deer (>Dama dama)

Haematological examinations have been carried out on 62 healthy farmed male and female fallow deer after they had been shot during a routine cull. Comparisons were made between the sexes, between young (up to 2 years) and older (>2 years) animals, and between age groups according to sex. The following parameters (means and SDs) were examined on all samples: erythrocyte count, haemoglobin concentration, packed cell volume, mean cell volume, mean cell haemoglobin, mean cell haemoglobin concentration, total leucocyte count, and numbers of neutrophils, eosinophils, basophils, lymphocytes, band neutrophils and monocytes. Only minor differences in red cell and white cell parameters between different groups of animals were found. However, significant differences in mean cell volume, mean cell haemoglobin concentration, eosinophils and band neutrophils were discovered in different age and sex groups. The values provide useful information for diagnostic purposes in fallow deer.

Age-specific survival and reproductive probabilities: evidence for senescence in male fallow deer (Dama dama).

Survival and reproduction are key features in the evolution of life-history strategies. In this study, we use capture-mark-resighting and multi-state models to examine survival senescence and reproductive senescence in six successive cohorts of fallow bucks that were studied for 16 years. We found that the overall age-specific survival probabilities of males were highly variable and the best-fitting model revealed that fallow bucks have four life-history stages: yearling, pre-reproductive, prime-age and senescent. Pre-reproductive males (2 and 3 years old) had the highest survival. Survival declined sharply after the age of 9 years, indicating that senescence had begun. When we considered reproducing and non-reproducing males separately, there was no evidence of senescence in the former, and steadily decreasing survival after the onset of social maturity in the latter. Reproduction probability also declined in older males, and thus we provide very strong evidence of senescence. Reproducers had a greater chance of reproducing again in the following year than non-reproducers. Furthermore, there were differences in the survival probabilities, with reproducers consistently surviving better than non-reproducers. In our study population, reproducers allocate more to the effort to reproduce than non-reproducers. Therefore our results indicate the generally higher phenotypic quality of reproducing males. These results, along with earlier studies on the same population, could indicate positive relationships between fitness correlates.

Does Lateral Presentation of the Palmate Antlers During Fights by Fallow Deer (Dama damaL.) Signify Dominance or Submission?

A central aim of the study of animal communication is to identify the mode and content of information transferred between individuals. The lateral presentation of the antler palm between male fallow deer has been described as either a signal of individual quality or an attempt to avoid fighting. In the first case two phenotypic features have been proposed by which transmission of individual quality may be facilitated. These are antler size and antler symmetry. The alternative hypothesis proposes that the lateral presentation of antlers occurs as a consequence of averting a threatening posture and may signify a reluctance to fight. We examined whether mature fallow deer use lateral palm presentation as a display during fights to indicate antler size and symmetry. We found no relationship between presentation rate of the antler and antler size and symmetry. Furthermore, males did not preferentially present their larger antler to their opponent. We also investigated whether the rate at which males presented antlers laterally during a fight was related to their ability to win the fight. Our results show that the male who performed more presentations during a fight was more likely to lose it. There were behavioural differences in the way in which a bout of presentation ended; subsequent losers tended to turn their body away from their opponent and subsequent winners tended to lower their antlers to an opponent which we interpret as an invitation to continue fighting. We conclude that the lateral palm presentation serves to de‐escalate fighting between mature fallow deer. It is not a mechanism by which to communicate individual quality but rather an indication that a male is less committed to continuing investment in the current contest.

Carcass quality in male fallow deer ( Dama dama): effects of age and supplementary feeding

Thirty-two male fallow deer, half 18- and half 30-month-old, were slaughtered after a 4-month feeding trial on pasture alone or with daily supplement of 500 g (dry matter) concentrate/head (eight each 18- and 30-month-old). The concentrate was composed of 40% maize, 25% sugar beet pulp, 20% alfalfa, 13% soy flakes, 2% minerals and vitamins. Supplemented deer were heavier (49.9 vs 45.0 kg) and their carcasses produced higher dressing proportions (61.5 vs 57.2%) and bigger amounts of first quality cuts. Supplemented feeding improved muscle development and fat deposition. Thirty-month-olds were heavier than 18-month-olds (53.3 vs 41.6 kg) and their carcasses produced higher dressing proportions (60.9 vs 57.7%) and bigger amounts of first quality cuts; older deer carcasses showed better conformation and bigger amounts of fat deposition.

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).

Fluctuating asymmetry in antlers of fallow deer (Dama dama): the relative roles of environmental stress and sexual selection

Antler lengths were recorded of a total of 250 male fallow deer (Dama dama (L)). Animals sampled were from marked populations where the majority of individuals were of known age. Asymmetry in antler length was normally distributed with a mean not significantly different from zero, confirming that differences in length between the antler pair constitute a true fluctuating asymmetry (FA). We found no clear relationship between the degree of asymmetry in antler length of an individual male and either population density or actual body mass. We did, however, detect a significant relationship between asymmetry and deviation from maximum cohort bodyweight. If deviation from maximum weight within a cohort may be considered some index of competitive success or increasing environmental stress, this may suggest that asymmetry in antler length relates in some way to developmental stress suffered by the individual concerned. The degree of asymmetry recorded in antler length also showed a significant decline with animal age, with antlers of animals of 2 years or older showing significantly greater symmetry. This is consistent with a hypothesis that despite continued competition for resources, there is a changing balance of selection pressure as animals reach maturity, with increased pressure from sexual selection requiring males to produce significantly more symmetrical ornaments.

Fluctuating asymmetry in antlers of fallow deer (Dama dama): the relative roles of environmental stress and sexual selection

Antler lengths were recorded of a total of 250 male fallow deer (Dama dama (L)). Animals sampled were from marked populations where the majority of individuals were of known age. Asymmetry in antler length was normally distributed with a mean not significantly different from zero, confirming that differences in length between the antler pair constitute a true fluctuating asymmetry (FA). We found no clear relationship between the degree of asymmetry in antler length of an individual male and either population density or actual body mass. We did, however, detect a significant relationship between asymmetry and deviation from maximum cohort bodyweight. If deviation from maximum weight within a cohort may be considered some index of competitive success or increasing environmental stress, this may suggest that asymmetry in antler length relates in some way to developmental stress suffered by the individual concerned. The degree of asymmetry recorded in antler length also showed a significant decline with animal age, with antlers of animals of 2 years or older showing significantly greater symmetry. This is consistent with a hypothesis that despite continued competition for resources, there is a changing balance of selection pressure as animals reach maturity, with increased pressure from sexual selection requiring males to produce significantly more symmetrical ornaments

What, if anything, does visual asymmetry in fallow deer antlers reveal?

Fluctuating asymmetry (FA), small directionally random deviations from perfect bilateral symmetry, is thought to reflect individual quality. Furthermore, it has been suggested that FA in secondary sexual characters can be used to assess mate or opponent quality during inter- or intrasexual competition. Studies on fallow deer, Dama dama, have suggested that FA in antlers reflects individual dominance, or the existence of a directional asymmetry (DA) with right antlers being consistently more developed than left antlers. To test these conflicting hypotheses, we analysed relationships between age, dominance and asymmetry in the number of antler points on mature fallow deer males during four rutting seasons in a single population. Age and dominance were only weakly correlated. The number of antler points displayed a pattern of DA (more points on the right than on the left side) that increased with age. Although dominance tended to increase with the total number of antler points, there was no relationship between the level of DA and dominance. These results failed to support the hypothesis that antler asymmetry visually reveals individual quality in fallow deer.

The fighting technique of male fallow deer (Dama dama): an analysis of agonistic interactions during the rut

A population of 560 individually recognizable European fallow deer (Dama dama) was studied in Phoenix Park, Dublin, Ireland, during the rut in order to examine buck fighting technique in relation to age and rank. The percent usage and association among behavioural elements used were analysed in 162 fights and 82 sparrings. During fight sequences, high-ranking bucks tend to use a higher percentage of low-risk behaviours (all behaviours which do not involve antler contact), especially when they interact with lower-ranking deer. Fights between combatants of the same rank class show a higher use of high-risk behaviours (characterized by antler contact), and cluster analysis of a preceding–following matrix revealed that high-risk behaviours are closely associated with each other in these interactions. During sparrings, the low intensity of the interaction allows the deer to test their ability and strength using a higher percentage of behaviours which are generally considered high-risk, without risking injury. The data suggest that the percent usage and the sequences of behavioural elements performed in agonistic interactions are affected by deer rank, probably in relation to the different risks and benefits that different hierarchical positions can offer.

The fighting technique of male fallow deer (Dama dama): an analysis of agonistic interactions during the rut

AbstractA population of 560 individually recognizable European fallow deer (Dama dama) was studied in Phoenix Park, Dublin, Ireland, during the rut in order to examine buck fighting technique in relation to age and rank. The percent usage and association among behavioural elements used were analysed in 162 fights and 82 sparrings. During fight sequences, high‐ranking bucks tend to use a higher percentage of low‐risk behaviours (all behaviours which do not involve antler contact), especially when they interact with lower‐ranking deer. Fights between combatants of the same rank class show a higher use of high‐risk behaviours (characterized by antler contact), and cluster analysis of a preceding–following matrix revealed that high‐risk behaviours are closely associated with each other in these interactions. During sparrings, the low intensity of the interaction allows the deer to test their ability and strength using a higher percentage of behaviours which are generally considered high‐risk, without risking injury. The data suggest that the percent usage and the sequences of behavioural elements performed in agonistic interactions are affected by deer rank, probably in relation to the different risks and benefits that different hierarchical positions can offer.

Social Interactions of Yearling Male Fallow Deer during Rut

Despite a low probability of achieving copulations, yearling male (1.5‐year‐old) fallow deer Dama dama display behavioural changes during rut, interacting with both females and older males. Juvenile interactions in polygynous ungulates may affect further reproductive success through both motor training for future interactions and/or competition for current resources that affect growth. To test this hypothesis, we intensively studied the behaviour of 12 yearling males in two captive populations during a rutting season. Interactions with females were interpreted as motor training aimed at learning to recognize oestrous females, because yearling males rarely interacted with their mother and because interactions essentially consisted of olfactory investigations focused on oestrous females. Because agonistic interactions with older males never involved physical contact and were mainly initiated by the most dominant buck in each population, we concluded that these interactions resulted from the presence of the yearlings close to oestrous females defended by rutting males. Heavier yearlings tended to be dominant over lighter ones and tended to interact more frequently with other animals, suggesting that early growth may influence the ontogeny of social behaviours.

The Epipalaeolithic Faunal Sequence in Israel: A View from Neve David

We carried out a detailed taphonomic, palaeoeconomic and palaeoecological analysis of the faunal remains from Neve David, a major Geometric-Kebaran open air site at the foot of the slope of Mount Carmel. The bone assemblage comprised 15 mammal species, two reptile species and seven genera of molluscs of the Mediterranean Sea. Bone fractures, cut marks and burned bones reflect human activity. The major prey species, as in other Epipalaeolithic sites from Israel, are gazelle and fallow deer (at 60 and 30%, respectively). It is possible that male fallow deer were selectively hunted. Neve David does not differ appreciably from the general Epipalaeolithic faunal sequence. Our attempt to compare the faunal remains of Neve David to other fossil assemblages from the Epipalaeolithic sequence revealed no significant difference between pre-Natufian and Natufian fossil assemblages in the economic characteristics studied, putting in question the basis for various hypotheses regarding the development of early domestication.

Fighting Tactics of Fallow Bucks (Dama dama, Cervidae): Reducing the Risks of Serious Conflict

AbstractCompetition between male fallow deer (Dama dama), during the breeding season was studied to determine if conflict strategies were consistent with the reduction of risk. Agonistic interactions between males were analysed in relation to age, dominance rank and availability of mating opportunities. The breeding season was divided into two main periods: the pre‐rut began when all males had cleaned the velvet from their antlers and ended on the last day before matings were first observed, while the rut refers to the period between the first and last matings. Overall, socially mature males (≥4yr old) were involved in more interactions than immature (≤3yr old) males. Males established dominance rank largely by non‐contact agonistic interactions during the pre‐rut and there was substantial carry‐over of rank to the rut, when it was correlated with mating success. The mating success of males was skewed; mature males achieved 99.4% of the matings and immature males accounted for 0.6%. A mature male was 13 times more likely to fight than an immature male; the mature males that fought most often did so between 0.4 and 0.5 times per hour. During the rut, the number of fights was positively correlated across days with the number of matings. The majority of agonistic interactions (79%) comprising dyads of immature males, involved antler contact. In contrast, mature males engaged antlers in only 42% of their interactions. Fights between mature males lasted more than twice as long as those between immature males and were more likely to occur between opponents with similar dominance ranks. However, towards the end of the rut formerly mismatched opponents were more likely to fight. Thus males operated conditional competitive strategies to decide when to interact and fight. The persistence of rank order from the pre‐rut period to the rut and the tendency for mature males to resolve disputes without antler contact, served to reduce the frequency of fights and therefore the risk of serious injury.

Habitat structure, female distribution, and fallow deer Dama dama mating stands

Male fallow deer Dama dama exhibit inter and intrapopulation variation in mating behaviour, including both territorial and non‐territorial mating tactics. During 1993 I investigated the relationship between fallow deer mating territory location and female distribution in relation to habitat characteristics at the Ottenby nature reserve in southern Sweden where males defend isolated mating stands during the autumn rut. Mating stands were positively associated with dense canopy cover and the presence of hazel Corylus avellana trees, but negatively associated with the number of alder Alnus glutinosa trees. Preference for such sites may be connected with the types of display males engage in during the rut as both auditory and olfactory means of communication can be affected by habitat structure. In relation to one another, the stands were randomly dispersed throughout the wooded region and along the perimeter of fields, with no indication of clumping or overdispersion. Female distribution was not associated with any of the habitat characteristics, or with stand location. The idea that males might be defending areas of favoured resources was ruled out as a possible explanation for stand location, although the idea that males might be defending game trails frequented by females remains as a potential determinant of stand locations.

Male-biased investment in fallow deer: an experimental study

To find the causes of faster growth in males, we studied the growth and behaviour of six male and five female fallow deer,Dama dama, fawns, hand-reared on the same amount and quality of milk, and compared them with naturally reared fawns. The bottle-reared fawns grew more slowly. In particular, the difference in weight gain between bottle- and mother-reared fawns was significantly larger for males during the time when they consumed only milk. Male fawns also sucked harder and were more motivated to obtain milk. These results indicate that male fallow deer fawns receive more milk from their mothers than female fawns under natural conditions and, hence, maternal investment seems male biased in this species. Males grew slightly, but significantly, faster until 10 weeks old, which revealed that sex-differential growth during the period of maternal investment is possible in the absence of differential maternal investment. The results could not reveal whether this small difference in growth is brought about by physiological or behavioural differences between the sexes.

Effects of retinoic acid on pedicle and first antler growth in a fallow buck ( Dama dama L.)

Unilateral injection of 10 mg all-trans retinoic acid (in 1 ml of castor oil) into the early pedicle anlage of a male fallow deer resulted in accelerated growth of the cranial appendage, and altered pedicle and first antler shape on the treatment side, whereas pedicle and antler growth on the control side, injected with 1 ml of vehicle solution only, was normal. It is concluded that retinoic acid is able to alter pedicle and first antler morphogenesis, presumably by affecting the positional information of the periosteal cells covering the incipient pedicle.

Habitat Manipulation Modifies Lek Use in Fallow Deer

AbstractFallow deer leks and single territories often persist in specific locations over many years. In our study area in Tuscany, one site (Lek 1) was used as a lek by fallow deer for at least 30 years. Females passed through the lek year‐round while travelling between sheltering and foraging sites. In 1987–89, forestry operations affected areas near the lek that were used by females as travel routes, but the lek site itself was not affected. After the logging operations, very few females traversed the lek site. Over the next 2 years the lek decreased in size (both in terms of number of territories defended and number of territorial males) until it became a single territory, and after two more years it was abandoned. Another lek (Lek 2) whose access routes were unaffected by forest cuts, continued to exist. Territorial males abandoned Lek 1 and moved to Lek 2 when female visits to Lek 1 became rare. Our results suggest that female movement patterns can be affected by habitat manipulations, and that females will not return to traditional mating sites that become external to their home ranges. Male fallow deer may form leks along female movement routes, where they experience a high female encounter rate as predicted by the hotspot model.

The relationship between plasma testosterone concentrations and the seasonal variation in voluntary feed intake in fallow bucks (Dama dama)

In two experiments conducted at Badgery's Creek, NSW (Latitude 34°S) the roles of testosterone and the thyroid hormones triiodothyronine (T3) and thyroxine (T4) as possible initiators of the change in voluntary feed intake (VFI) associated with the seasonal reproductive behaviour (rut) in male fallow deer were investigated.In Expt 1, the association between changes in these hormones with the onset of the rut was assessed in deer in which the timing of this event was manipulated by changing the photoperiod, or by melatonin administration. Groups were maintained under either natural photoperiod (n=6; control), a constant long daylength (16 h light[ratio ]8 h dark; n=9; LD group) or a constant long daylength and implanted with melatonin capsules (n=5; LD+M group) for 7 months from December until the following July. Blood samples were obtained weekly and VFI recorded.Feed intake decreased by 94% from the last week of March for 3 weeks in the control group. Changes of a similar magnitude were measured in the LD and LD+M groups but these changes were advanced by 1 and 9 weeks respectively in these groups. In all groups, circulating testosterone concentrations increased markedly at a time corresponding with the decrease in VFI. The concentrations returned to basal levels with the resumption in VFI. A distinct decrease in plasma concentrations of T3 and T4 in all three groups was associated with the decrease in VFI, however, the relationship with T3 was less apparent.In Expt 2, the role of testosterone in the regulation of the decline in VFI was investigated. Fallow bucks were treated with testosterone enanthate every 4 or 5 days for 28 days up to 6 weeks prior to the expected onset of the rut. Plasma testosterone concentrations, which were increased 13-fold, resulted in a decline in VFI which was comparable to that observed in the subsequent rut. Plasma free fatty acid concentrations were correlated negatively with the decline in VFI.Thus, the seasonal increase in circulating testosterone concentrations plays an important role in initiating the fall in VFI associated with the rut. As the rut was still apparent in animals maintained under an extended photoperiod, it is possible that factors other than decreasing daylength act as the cue for the timing of the rut.