Effects of public vs. private management on deer antler composition, mechanical and structural variables

Previous studies have shown that antler mineral composition reflects diet. Thus, management involving diet can influence both the mineral composition and mechanical properties of antlers. However, it is not known if reducing population density, which increases availability of food, can compensate mineral deficiencies arising from lack of minerals in plants and, ultimately, in the soil. The present study aimed at assessing if private management often involving a balanced food supplementation produced differences in antler properties compared to both public management and reference antlers from deer farms. We also examined if low population density in a National Park could compensate for mineral deficiencies found in antlers of other public reserves or else, antlers still differed in antler characteristics compared to supplemented deer in private game estates. We used 120 antlers from three public reserves, four private game estates and two deer farms as reference to assess antler composition, mechanical properties, size and structure. Public managed antlers had shorter length compared to private and reference ones, thinner cortex (CT), were less dense, had lower second moment of area (I) and work to fracture (W). In addition, they had content in ash, Ca, Mg, Na, S and Zn lower than antlers from private game estates. In contrast, antlers from public reserves had greater values of Young’s modulus of elasticity (E), Fe, Mn and Si. In most cases, antlers from private management and reference farms showed similar values. When comparing antlers from low population density in a National Park with antlers from private management, differences in antler length, CT, I and ash disappeared, but both differed still in density, E, bending strength, W and content in all minerals mentioned. In conclusion, low density can improve some structural variables, but it cannot compensate for mineral deficiencies whereas food supplementation can.     

Population management and bone structural effects in composition and radio-opacity of Iberian red deer (<Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis>) antlers

Antlers are costly bone structures whose size and external characteristics are influenced by nutrition, climatic variability, and other factors. In this study, we examined the effects of a well-managed captive population vs unmanaged free-ranging herds (greatly differing in food availability and energetic and immunological expenditures) on antler structural characteristics of Iberian red deer. We assessed the effect of management and sample position in the antler on cortical bone depth, bone mineral and protein content, and radiographic bone opacity. Cortical bone depth and mineral percent was greater in captive animals on greater food availability and lower energetic and immunological expenditures. After removing the inverse relationship with mineral content in the model, protein percent was also higher in antlers from captive than in those from free-ranging deer. Management system also influenced radio-opacity indirectly as interaction with other factors and cortical bone depth effects. Structural properties also differed between antler points that are very close in the antler and seem to differ only in the role they play. In conclusion, captive management can affect antler structure and composition, possibly as a result of different availability of food and immunological costs.     

Correlates of variation in deer antler stiffness: age, mineral content, intra-antler location, habitat, and phylogeny

To test correlations between the stiffness of deer antler and aspects of cervid ecology, we measured the stiffness (in bending) of antler from white-tailed deer ( Odocoileus virginianus ) and compared our results to previous measurements from antlers of other cervid species. Stiffness of antler specimens did not correlate significantly with mineral content or the location within the antler from which specimens were taken in O. virginianus. However, antler stiffness in white-tailed deer decreased significantly between two- and three-year-old bucks, matching the time in O. virginianus life history when males shift from sparring with mostly larger individuals to sparring with mostly smaller individuals. Stiffer antlers may enable younger, smaller bucks to have a more effective lever through which their smaller muscular forces can be transmitted during sparring with older, larger bucks. The stiffnesses we measured for white-tailed deer antler are similar to values measured previously from other members of the odocoileine lineage, which are lower than those previously measured from the antlers of deer living in tropical habitats. However, confidence limits for maximum likelihood reconstructions of the ancestral stiffness of deer antler span the range of high stiffnesses found among tropical deer; furthermore, parsimony-based reconstructions of ancestral antler stiffness are equivocal. Thus, the high antler stiffnesses of tropical deer may reflect the retention of an ancestral condition, rather than adaptation to year-round antler use.     

Mineral deficiencies in tule elk, Owens Valley, California

Male tule elk (Cervus elaphus nannodes) are susceptible to high rates of antler breakage in Owens Valley, California. We hypothesized that a mineral deficiency in the diet predisposed male elk to antler breakage. We analyzed elk antler, liver, and forage samples to identify mineral imbalances. We compared the mineral content of livers and antlers from elk in Owens Valley to samples taken from tule elk at Grizzly Island Wildlife Area, a population experiencing normal rates (<5%) of antler breakage. Antler and liver samples were collected from 1989 to 1993, and in 2002, and were tested for calcium (Ca), copper (Cu), iron (Fe), magnesium (Mg), manganese (Mn), molybdenum (Mo), phosphorus (P), sulfur (S), and zinc (Zn). Mineral levels from antler and liver samples were compared to reference values established for elk and deer. We also compared the mineral content of elk forage in Owens Valley, collected in 2002-03, to dietary reference values established for cattle. In antlers, Ca, Fe, and Mg levels were higher in Owens Valley elk than in Grizzly Island elk, although all mineral levels were lower than reference values established for deer antlers. In liver samples, Cu levels from elk in Owens Valley were lower than those from Grizzly Island and lower than minimum reference values; liver Ca and Mo levels were higher in elk from Owens Valley than in those from Grizzly Island. Compared to reference values, elk forage in Owens Valley had high levels of Ca and Mo, and low levels of Cu, P, and Zn. Mineral analyses from antlers, livers, and forage suggest that tule elk in the Owens Valley are Cu and/or P deficient. High levels of Mo and Ca may exacerbate Cu and P deficiencies, respectively. Bone fragility is a symptom of both deficiencies, and an imbalance in Cu, P, or a combination of both, may predispose male tule elk in the Owens Valley to antler breakage.     

Does Cu supplementation affect the mechanical and structural properties and mineral content of red deer antler bone tissue?

The main factors affecting the mechanical (and other) properties of bone, including antler, are the proportions of ash (especially Ca and P) and collagen content. However, some trace minerals may also play more important roles than would be expected, given their low levels in bone and antler. One such trace mineral is Cu. Here, we studied the effects of Cu supplementation on the mechanical and structural characteristics, and mineral content of antlers from yearling and adult (4 years of age) red deer fed a balanced diet. Deer (n=35) of different ages (21 yearlings and 14 adults) were studied. A total of 18 stags (11 yearlings and 7 adults) were injected with Cu (0.83 mg Cu/kg BW) every 42 days, whereas the remaining 17 (10 yearlings and 7 adults) were injected with physiological saline solution (control group). The Cu content of serum was analysed at the beginning of the trial and 84 days after the first injection to assess whether the injected Cu was mobilized in blood. Also, the mechanical and structural properties of antlers and the mineral content in their cortical walls were examined at three (yearlings) or four (adults) points along the antler beam. The effect of Cu supplementation was different in yearlings and adults. In yearlings, supplementation increased the Cu content of serum by 28%, but did not affect antler properties. However, in adults, Cu supplementation increased the Cu content of serum by 38% and tended to increase the cortical thickness of antlers (P=0.06). Therefore, we conclude that, even in animals receiving balanced diets, supplementation with Cu could increase antler cortical thickness in adult deer, although not in yearlings. This may improve the trophy value of antlers, as well as having potential implications for bones in elderly humans, should Cu supplementation have similar effects on bones as those observed in antlers.     

Antler stiffness in moose (Alces alces): Correlated evolution of bone function and material properties?

The material properties of bone can vary considerably among skeletal elements from different parts of the body that serve different functions. However, functional demands placed on a specific type of skeletal element also can vary at a variety of scales, such as between different parts of the element, among individuals of a species, and across species. Variation in bone material properties might be correlated with differing functional demands at any of these scales. In this study we performed three-point bending tests on bone specimens extracted from antlers of moose (Alces alces) to test for three types of variation in bone material stiffness (Young's modulus): within the antler structure, between populations of moose, and between moose and other deer species. Because superficial portions of the antler are exposed to greater bending stress and strain than deeper portions, and because the antler beam (the basal shaft that attaches to the skull) is subjected to greater bending moments than more distal parts of the antler, we predicted that superficial bone and bone from the beam would be stiffer than bone from other parts of the antler. Instead, we identified no significant differences in these comparisons. There were also no significant differences in antler stiffness between moose from Michigan and the Yukon, even though the rapid growth required of antlers from northern latitudes like the Yukon has the potential to compromise bone material properties. However, moose have significantly stiffer antlers (11.6 +/- 0.45 GPa, mean +/- SE) than any other deer in the odocoileine lineage. Moreover, phylogenetic reconstructions of the evolution of antler stiffness in deer indicate a strong potential that high antler stiffness is a derived feature of moose. The unusual palmate shape of moose antlers likely subjects their antler beams to higher bending moments than found in other odocoileines, a factor that may have contributed to the evolutionary divergence of moose antler stiffness from that of other members of this clade. Although similarities in the mineral composition of bone across species likely limit the overall range of phylogenetic variation in bone material properties, our results demonstrate that evolutionary diversity in bone material properties can show correspondence with phylogenetic differences in mechanical or ecological demands on skeletal elements.     

Patterns of antler consumption reveal osteophagia as a natural mineral resource in key periods for red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>)

Osteophagia is a common behaviour among ungulates. It has been considered as a mineral source for cervids, mainly related to phosphorus deficiency. In this study, we aimed to study the seasonality in antler consumption of Iberian red deer (Cervus elaphus) and to assess the differences among sex/age classes. In a game reserve located in South-Eastern Spain, antlers were offered to free-ranging animals, and their chewing behaviour was recorded with camera traps. Red deer was the species observed with greatest frequency (90.5%), and only red deer was observed chewing the offered antlers. Males showed greatest peak of consumption at the end of antler growth; females showed the greatest peak at the beginning of the lactation and calves after delivery of the newborn. All groups showed an important second peak of consumption in September. We conclude that osteophagia might have a role as a natural mineral supplement for red deer, but not for wild boar or foxes; and that each sex/age class uses this resource at different times according to differential needs in calcium and phosphorus, supporting the “seasonally increased requirements hypothesis”.     

Effects of selective harvest on antler size in white-tailed deer: A modeling approach

Selective harvesting in wild deer (Odocoileus spp.) populations is a common practice that may influence antler size. However, in free-ranging populations, response due to selection is unknown or difficult to quantify because antlers are influenced by nutrition and population demographics. We used quantitative genetic models to predict how white-tailed deer (O. virginianus) antlers would respond to selection and what variables (i.e., population size, age structure, mating ratio, and heritability) most affected antler size. We validated our quantitative genetics program by comparing model results with a population of deer used for controlled breeding experiments; modeled antler points (AP) and score increased (2.2–4.3 AP and 48.5–97.7 cm, respectively) after 8 years of selection, similar to observed increases in AP (3.2) and score (92.3 cm) from the controlled population. In modeled free-ranging populations, mating ratio, age structure, and heritability were more important in influencing antler size than size of the population. However, response to selection in free-ranging populations was lower (0.1–0.9 AP) than controlled breeding populations even after 20 years of selection. These results show that selective harvesting of free-ranging white-tailed deer may be inefficient to change population-level genetic characteristics related to antler size. Response of antlers in free-ranging deer will be less than controlled populations, and possibly modeled free-ranging simulations, because individual reproductive success of males is lower, breeding is done by a large group of males, and reproductive and survival rates are lower. These factors, and others, reduce the amount of improvement that can be made to antlers due to selection. Therefore, selective harvesting in free-ranging populations should be justified for managing population demographics and dynamics, but not for changing the genetic characteristics of populations. © 2011 The Wildlife Society.     

Testosterone and estradiol concentrations in serum, velvet skin, and growing antler bone of male white-tailed deer

The growth and mineralization of antlers correlate with the seasonal variation of serum androgens. Whereas seasonal levels of testosterone (T) in plasma are well established, steroid concentrations have not yet been determined in the tissues of growing antlers. Therefore, RIA was used to determine T and 17beta estradiol (E2) in serum, and three areas (tip, middle, and base) of the antler bone and the antler skin, called velvet. Blood and antler tissues of white-tailed deer (Odocoileus virginianus) were collected from May to August. The difference between levels of T and E2 among the sites was calculated using the square root transformation followed by a mixed model analysis with individual deer and an interaction of individual and year (individual(*)year) as a random factor. Concentrations of T in serum (799+/-82 pg/ml) were higher than T values in the velvet (589+/-58 pg/ml, P<0.01) and in the antler bone (538+/-58 pg/ml, P<0.001). Estradiol concentrations differed among antler tissues and serum (P<0.001) and between years (P<0.01). Estradiol concentrations in serum (25+/-25 pg/ml) were consistently lower than those in antler bone (208+/-11 pg/ml, P<0.001) and velvet (150+/-12 pg/ml, P<0.001). The E2:T ratio in serum was 1:10-60. The same ratio for the antler bone was only 1:2-3 and for the velvet 1:3.5. It is concluded that higher T and lower E2 concentrations found in plasma, as compared to antler bone or antler velvet, may indicate a partial metabolism of systemic androgens into estrogens xin the tissues of growing antlers.     

Internal architecture of pampas deer antlers differs in males allocated with and without females

Bone mineralization of antlers and the depth of the antler seal (the basal surface of the cast antlers) are positively related to testosterone concentrations. Pampas deer males that are in permanent contact with females have greater, heavier, and darker antlers than males that are isolated from them. The objectives were to determine if antler compact/spongy bone ratio, antler seal depth, and compact bone darkness are greater in pampas deer males permanently allocated with females than those in males isolated from them. Antlers from males permanently allocated with or without females were cut transversally in seven points and scanned, and the compact/spongy ratio was calculated. The pixel intensity of each image was determined with a software for image analysis. The coronet of the antler was cut longitudinally, and the height of the most prominent protrusion was measured. The compact/spongy ratio was greater in antlers from males that were in contact with females in the second tine (P?=?0.02) and tended to be great in the two other tines (P?=?0.06 and P?=?0.1, respectively). Compact bone pixel color was darker in males in contact with females than that in males isolated from females in two points (P?=?0.02 and P?=?0.05, respectively) and tended to do so in two more (P?=?0.055 and P?=?0.1, respectively). Antler seal convexity was also greater in antlers from males in contact with females (P?=?0.006). We concluded that permanent contact with females stimulated pampas deer males increasing compact bone portion of the antler tines, the seal depth size, and the darkness of the compact bone.     

Antler stiffness in caribou (Rangifer tarandus): testing variation in bone material properties between males and females

Vertebrate bones, including deer antler, often exhibit variation in their mechanical properties that corresponds to differences in the functional demands they encounter. Among deer, antlers are found in both males and females only in caribou (Rangifer tarandus). Several differences between male and female R. tarandus in behavior and physiology might subject their antlers to differing demands, potentially making divergence of antler material properties between the sexes advantageous. Alternatively, antler material properties might not differ between male and female R. tarandus because both sexes are members of the same species, and the properties of their antlers could, therefore, have emerged under similar pressures and constraints through evolution. To test for sexual dimorphism in antler material properties, we compared the stiffness (Young's modulus of elasticity) of antler specimens from male and female caribou using three-point bending tests. Despite behavioral and physiological differences between males and females, stiffness values did not differ significantly between the sexes in caribou, with a mean (+/-S.E.) stiffness of 5.8+/-0.4GPa across all specimens. This value differed by less than 10% from the values published for R. tarandus specimens of unknown sex, verifying the comparability of bone material property data collected across multiple studies, and lending confidence to recent analyses of the evolution of antler stiffness in deer that have drawn on literature data.     

麋鹿鹿角生长周期及影响因子

2013年8月至2014年8月,使用望远镜(SWAROVSKI 8×42 WB)和照相机(Canon 550D 70×300)观察北京南海子59只雄性麋鹿茸的生长和角脱落周期,发现麋鹿角总体趋势呈随着年龄的减少,脱角日期越迟,且等级序位高或者鹿王较年老个体先脱落。收集2012年12月至2014年2月两年度麋鹿自然脱落的角,研究表明,个体而言,左右角脱角顺序差异明显,同一天脱落两角者占17.0%,左角先脱落者占34.1%,右角先脱落者占48.8%,重量重和重量轻的角先脱落的个体各占41.5%,个体成对的左右角脱落时间间隔短,平均为1.98d;群体而言,角重量与脱角时间呈显著负相关,总体呈现角重的个体角先脱落,但不是角最重的个体角第一个脱落。2013年6月至2014年5月,对北京南海子、湖北石首、江苏大丰、天津七里海、浙江慈溪、浙江临安、海南海口、河北滦河上游、辽阳千山等9个地区麋鹿种群麋鹿角脱落起止时间进行调查,研究表明野生种群脱角时间比圈养种群早;迁入热带地区海南的麋鹿仍有规律的角周期;同一栖息地不同年份角周期存在差异,不同栖息地间麋鹿角周期存在差异;脱角起始时间与海拔、经度、纬度、年平均气温、圈养情况和气候类型等影响因子不存在统计学上显著的相关性,但光周期和激素直接主导和调节着角周期,年龄大小、角重量、营养以及与营养相关的能量、种群密度、物候等环境因子均影响角周期。     

Effects of Feed Supplementation on Mineral Composition,Mechanical Properties and Structure in Femurs of Iberian Red Deer Hinds (Cervus elaphus hispanicus)

Few studies in wild animals have assessed changes in mineral profile in long bones and their implications for mechanical properties. We examined the effect of two diets differing in mineral content on the composition and mechanical properties of femora from two groups each with 13 free-ranging red deer hinds. Contents of Ca, P, Mg, K, Na, S, Cu, Fe, Mn, Se, Zn, B and Sr, Young’s modulus of elasticity (E), bending strength and work of fracture were assessed in the proximal part of the diaphysis (PD) and the mid-diaphysis (MD). Whole body measures were also recorded on the hinds. Compared to animals on control diets, those on supplemented diets increased live weight by 6.5 kg and their kidney fat index (KFI), but not carcass weight, body or organ size, femur size or cortical thickness. Supplemental feeding increased Mn content of bone by 23%, Cu by 9% and Zn by 6%. These differences showed a mean fourfold greater content of these minerals in supplemental diet, whereas femora did not reflect a 5.4 times greater content of major minerals (Na and P) in the diet. Lower content of B and Sr in supplemented diet also reduced femur B by 14% and Sr by 5%. There was a subtle effect of diet only on E and none on other mechanical properties. Thus, greater availability of microminerals but not major minerals in the diet is reflected in bone composition even before marked body effects, bone macro-structure or its mechanical properties are affected.     

Does fluctuating asymmetry of antlers in white-tailed deer (Odocoileus virginianus) follow patterns predicted for sexually selected traits?

Secondary sexual characters have been hypothesized to signal male quality and should demonstrate a negative relationship between the size of the trait and degree of fluctuating asymmetry because they are costly to produce. We collected morphometric and antler data from 439 white-tailed deer (Odocoileus virginianus) in Oklahoma, USA, in order to determine whether measures of antler asymmetry follow the patterns predicted for sexually selected characters. Relative fluctuating asymmetry was negatively related to antler size for all deer and within age groups up to five and a half years of age. We did not detect an association between asymmetry and antler size among deer that were six and a half years or older. When categorizing deer by antler size, we found that deer with small antlers (< or = 33rd percentile) had greater levels of relative asymmetry than deer with large antlers (< or = 67th percentile). The relative asymmetry of antlers was negatively related to age and was greatest in deer that were one and a half years old. Relative asymmetry was also negatively related to carcass mass, inside spread, skull length and body length. These data suggest that asymmetry in the antlers of white-tailed deer may be a reliable signal of quality and, as such, may be important in maintaining honesty in intrasexual advertisements during the breeding season.     

White-Tailed Deer Antler Research: A Critique of Design and Analysis Methodology

ABSTRACT Debate within the popular and technical literature regarding predictability of antler size at maturity based on 1.5-year antler size in white-tailed deer (Odocoileus virginianus) has led to confusion and uncertainty within constituent groups. Koerth and Kroll (2008) provided measures of age-related antler development using recaptures of known-age males from 12 deer populations in southern Texas. Several design and analysis issues reduce the scope and validity of their conclusion that amount of growth in the first set of antlers was a poor predictor of antler growth at maturity. Although unstated, the statistical hypothesis they tested did not coincide with their specific conclusions. Using a simulation, we show that their methods were susceptible to measurement bias. Their results are applicable only to populations with similar culling and management programs. Additionally, we provide recommendations for future research projects that evaluate predictability of antler size at maturity based on antler size at younger ages.     

Antlers—a Mineral Source in Rangifer

In a calving area on western Hardangervidda, Norway, shed antlers of female reindeer, Rangifer tarandus tarandus L., have been collected for several years. Many of these antlers have been chewed on by reindeer. In the literature antler chewing has been reported to take place on both shed and unshed antlers. Our observations support these findings. Based on the present observations it is suggested that the antlers may subserve the mineral householding of the animals under special conditions. The female reindeer has to pass through a nutritionally marginal season which often coincides with calving. Bringing the bony antlers to the calving areas seems advantageous in the maintenance of calcium balance. The growth of antlers may therefore be looked upon as part of a mineral saving strategy. The mineral stored in the antler thus represents a reserve to be used during pregnancy and lactation. This is in contrast to the common view that shedding of antlers is just waste of mineral when there is a great need for calcium.     

Getting the timing right: antler growth phenology and sexual selection in a wild red deer population

There has been growing interest in the determinants of the annual timing of biological phenomena, or phenology, in wild populations, but research on vertebrate taxa has primarily focused on the phenology of reproduction. We present here analyses of the phenology of the annual growth of a secondary sexual characteristic, antlers in red deer (Cervus elaphus) males. The long-term individual-based data from a wild population of red deer on the Isle of Rum, Scotland allow us to consider ecological factors influencing variation in the phenology of growth of antlers, and the implications of variation in antler growth phenology with respect to the phenotype of antler grown (antler mass) and annual breeding success. The phenology of antler growth was influenced by local environmental conditions: higher population density delayed both the start date (during spring) and the relative end date (in late summer) of antler growth, and warmer temperatures in the September and April prior to growth advanced start and end dates, respectively. Furthermore, there was variation between individuals in this phenotypic plasticity of start date, although not in that of end date of growth. The phenology of antler growth impacted on the morphology of antlers grown, with individuals who started and ended growth earliest having the heaviest antlers. The timing of antler growth phenology was associated with breeding success in the following mating season, independently of the mass of antlers grown: an earlier start of antler growth was associated with siring a higher number of the calves born the following spring. Our results suggest that the phenology of traits that are not directly correlated with offspring survival may also regularly show correlations with fitness.     

Juvenile-to-Adult Antler Development in White-Tailed Deer in South Texas

Abstract: Past studies using penned deer provide conflicting results on the age when reliable predictions about antler growth potential in white-tailed deer (Odocoileus virginianus) can be made. We captured wild whitetail males via aerial net gun on 12 ranches in 5 counties in south Texas, USA, from 1999 to 2007 to determine if a reliable juvenile-to-adult relationship in antler development existed. We individually marked and released captured animals at the trap site after we took antler and body measurements. We recaptured marked animals as possible in subsequent years or until we obtained final measurements after legal harvest. Amount of growth in the first set of antlers in whitetail males was a poor predictor of antler growth at maturity. By 4.5 years of age there were no differences (P > 0.05) in antler measurements regardless of the amount of development of the first set of antlers at 1.5 years. We concluded culling of yearling males based on number of antler points would have little positive effect on overall antler quality in future years.     

梅花鹿(Cervus nippon hortulorum)茸角生长发育各阶段血浆睾酮、雌二醇的含量变化

本文用放射免疫测定法对3个不同年龄组的雄性东北梅花鹿茸角生长发育各阶段外周血中的睾酮、雌二醇含量进行了测定。茸期两种激素差异很大,睾酮含量低,雌二醇高,骨化期二者增高很快。对这两种激素含量的变化与茸角发育各阶段的关系进行了讨论。认为只有在二种激素同时增加时,鹿角才能骨化。