Remains of Late Pleistocene reindeer from Chmielewo,northeast Poland

Reindeer (Rangifer tarandus L.) remains in Europe are well represented in Late Pleistocene sites in France, Germany, Ukraine, Poland and other countries. In a chalk mine at Chmielewo (near niardwy Lake, NE Poland) four antlers were recovered. Radiocarbon assays date the fossils to the end of the Pleistocene. All antlers belong to male reindeer. One of the discovered specimens is an uncast antler of the tundra reindeer form, while the others are of the woodland reindeer form. The uncast antler has distinctly visible parieto-frontal fixation of the beam to the skull that confirms its parallel position in relation to the ground.     

Antlers and parasites

Development of ornamental characters exposed to directional selection may be particularly sensitive to the effect of parasitic infections. Antlers are ornamental characters of importance in intraspecific interactions, and are in reindeer (Rangifer) developed by both males and females. By antihelmintic treatment of naturally infected female reindeer we show that parasite intensities affect development of antler asymmetry, but not antler length. These results suggest that asymmetry in antlers may reflect parasite intensities and thus be of importance in intraspecific assessment of genetic resistance towards infectious organisms.     

Size and composition of the wild reindeer Rangifer tarandus platyrhynchus population in the Southeast Svalbard Nature Reserve

Alendal, E., de Bie, S. and van Wieren, S. E. 1979. Size and composition of the wild reindeer Rangifer tarandus platyrhynchus population in the Southeast Svalbard Nature Reserve. Holarct. Ecol. 2: 101-107. In the summer of 1977 we studied the reindeer population on the islands Barentsøya and Edgeøya in the eastern part of the Svalbard archipelago. A total of 1374 reindeer were observed: 326 animals in the western parts of Barentsøya and 1048 animals on Edgeøya. Considering those parts of Edgeøya which were not visited, the total number of reindeer on Edgeøya was estimated at 1300 animals. The total number of reindeer was lower than in previous years. The decline probably was due to severe winter conditions in 1975/1976 and 1976/1977 confirmed by the fact that many carcasses and few yearlings were observed. Nearly all reindeer occurred on the coastal plains and in the valleys. These areas have the relatively richest vegetation. The average recruitment of the total population (counted) was 15.9%. The adult sex ratio was in favour of females: 59% females versus 41% males. There were differences both in the recruitment and in the adult sex ratio between three distinct areas on Edgeøya and between two on Barentsøya. These differences may be due to dissimilarities in food quality and feeding conditions caused by climate, and by small exchange of reindeer between the areas. The high frequency of shed male antlers on Frankenhalvøya and Talaveraflya, north and south coast of Barentsøya respectively, indicates that these areas belong to the wintering grounds of reindeer on this island. Concentrations of shed female antlers on Barentsøya were less pronounced. The highest frequency was in the areas Sjodalen and Kvistdalen-Talaveraflya in the northwest and south respectively. Females may use these areas as late wintering grounds and possibly as calving areas. The average group size was 2.2 and the aggregation index 3.1. Seventytwo per cent of all groups, containing 48% of all reindeer, fell into group size 1 and 2. Males mostly were observed alone or together with one other animal. Females with calves most frequently occurred in groups of 2 and 4 animals.     

Origination of antlerogenesis

Antlers are unique appendages. They are shed and rebuilt at intervals, and are synapomorphic for all living Cervidae (except for the Chinese water deer, Hydropotes inermis , in which they have presumably been lost). The antlerogenic process is controlled by a complex interaction of fluctuating levels of several hormones, most importantly testosterone. The oldest antler remains are recorded from the early Miocene; these have been interpreted as non‐deciduous appendages because of supposed permanent skin coverage and the lack of a burr (a well‐developed osseous protuberance around the base of the antler, which is always present in extant cervids). The aim of this study is to test the hypothesis that antler shedding was possible in these early Miocene cervids. Antlers of all extant and eight Miocene cervid genera, including burr‐less antler fragments of the earliest cervids Procervulus , Ligeromeryx , and Lagomeryx were studied. An extensive comparative morphological analysis of external features of the antler, and of the abscission area and the base of the antler in particular, was undertaken. The results indicate that a regular, porous, and rugose abscission surface at the proximal end of the antler indicates antler shedding in both living and fossil cervids. The antler shedding mechanism must therefore have already been present in all early/mid Miocene cervid genera included in this study. On this basis, it is suggested that the presence of a burr is not prerequisite in order to shed antlers, that the presence of perpetual antlers has not yet been verified, and that the process of shedding and regeneration developed with the first appearance of these organs. This insight is particularly important for the systematic classification of early Miocene species as Cervidae, because the absence of the antler shedding and rebuilding mechanism would exclude them from the taxon Cervidae and from any relationship with extant cervids. J. Morphol. 278:182–202, 2017. © 2016 Wiley Periodicals,Inc.     

Antler mineral composition of Iberian red deer <Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis> is related to mineral profile of diet

Previous studies have suggested that antlers are costly bone structures whose mineral composition may change depending on physiological and other factors. This study examined whether nutrition variation associated with deer management influences antler mineral composition and structural characteristics of whole antler. Mineral distribution and bone structure were examined in antlers from two groups of adult Iberian red deer Cervus elaphus hispanicus Hilzheimer, 1909. They were kept under different feeding regimes at an experimental deer farm and a game estate in southeastern Spain. Protein and mineral contents differed between the diet of captive deer and that of deer in the wild. Significant differences were found for Na, Mg, K and protein. Antler composition seems to reflect the diet, as antlers of deer differed in protein, Na, Mg and K, but not in total mineral content, Ca, Fe or Zn. Thus, management conditions related to nutrition are reflected on antler composition.     

Role of oestradiol in the regulation of the seasonal antler cycle in female reindeer, Rangifer tarandus.

Reindeer (or caribou), Rangifer tarandus, is the only extant species of deer in which females as well as males normally develop antlers that are cast and regrown each year. This study investigated the role of ovarian oestradiol in the regulation of the seasonal antler cycle in female reindeer. Ovariectomized Norwegian reindeer living outdoors in northern Norway (69 degrees N) were treated with continuous-release subcutaneous Silastic implants containing oestradiol, which maintained the blood concentrations of oestradiol within the physiological range for the mating season from June to October-November. The treatment with oestradiol induced the synchronized maturation of the antlers and rapid cleaning of the velvet-like skin in August-September in the ovariectomized reindeer, a pattern very similar to that observed in ovary-intact controls living under the same conditions. The removal of the steroid implant in October-November caused the premature casting of the antlers in early winter in two of five animals, while the remainder cast at the normal time in spring; this response was seen whether the animals had received one or two oestradiol implants in autumn. The antlers developed by the ovariectomized, oestradiol-treated females were significantly heavier and carried more branches than the ovariectomized animals without oestradiol replacement, and were marginally heavier than the antlers of intact controls. These results support the view that oestradiol is the biologically active steroid secreted by the ovary in intact female reindeers that induces the normal development of the antlers. Oestradiol stimulates the growth and mineralization of the antler bone, the cleaning of the velvet, and suppresses the casting of the hard antlers. This endocrine control ensures that the hard antlers, which function as weapons, are retained throughout the autumn and winter when the females are normally pregnant and when competition between females over food in the snow is most intense; hence there is a reproductive advantage to explain the evolution of antlers in females.     

Reindeer antlers: visual indicators of individual quality?

Fluctuating asymmetry (FA) in ornamental characters may reflect developmental stability in the translation from genotype to phenotype. Antlers of reindeer show FA, are visually conspicuous ornaments and are important in intraspecific assessment. We show that there is a negative relationship between size and asymmetry in visual antler characteristics (i.e., antler length and number of tines) among free-ranging male reindeer in rut. This indicates that individuals that develop large ornaments are better able to buffer developmental stress than individuals that develop small ornaments. There is no relationship between asymmetry in antler length and asymmetry in jaw length, suggesting that symmetry in antlers does not reflect overall body symmetry. This difference may be caused by trait-specific sensitivity to developmental stress. Such stress may partly be caused by parasites, which show a positive association with asymmetry in antlers, but no relationship to asymmetry in jaws. Our results indicate that antlers may be exposed to directional selection in a visual signaler-receiver system and that information about parasite burden may be obtained from evaluation of asymmetry in antlers developed under exposure to a multitude of environmental stresses. Received: 8 October 1996 / Accepted: 13 January 1997     

Sexual dimorphism and intercorhort variation in reindeer calf antler length is associated with density and weather

We analysed intercohort variability of live weight and antler length of 5,123 reindeer calves. We further assessed the influence of climate and density on the interannual variation in antler length, and discussed sex-specific resource allocation and response to climate variability. Antler length varied significantly among years and between sexes, with interaction between year and sex. Body weight and antler length were highly positively correlated, showed similar intercohort variability, and had a strong allometrical link, suggesting that antler length could be an equally reliable measure of calf condition as live weight. We found a relative measure of antler length (i.e. antler length corrected for the allometric effect of body mass) to be positively influenced by increasing density and May–June precipitation, and also decreasing May–June temperature. We attributed the effect of early summer weather to its influence on forage availability and quality as well as the level of parasitic insect harassment. Gender difference in both the allometric exponents and the interannual variability suggest that young males and females may have different tactics for relative resource allocation towards growth of antlers as compared to body mass. Because antlers are costly to produce, they may be an honest signal of individual quality for both sexes. However, we found gender-specific allometry, as female calves more than males appear to prioritize their antler growth over body mass, especially when resources are limited. Thus, our results suggest that environmental variation may influence the extent of sexual dimorphism in antler length. Electronic Supplementary Material Supplementary material is available for this article at     

Aspects of nitrogen and mineral nutrition in Icelandic reindeer,Rangifer tarandus

Nitrogen and mineral (Fe, Mg, Na, K, and Ca) compositions of foodstuffs consumed by and dung produced by male, pregnant and lactating female adult Icelandic reindeer and calves were determined during May of 1992. Iron levels in foodstuffs are consistently above the reported toxicity level for similar-sized sheep (0.5 ppt) and may lead to iron toxicity when consumed by reindeer during periods of high lean body mass catabolism. Male and female reindeer meet nutrient requirements for all measured elements and nitrogen with the possible exception of calcium for males during antler growth.     

Development and design of a novel loading device for the investigation of bone adaptation around immediately loaded dental implants using the reindeer antler as implant bed

The assessment of the behavior of immediately loaded dental implants using biomechanical methods is of particular importance. The primary goal of this investigation is to optimize the function of the implants to serve for immediate loading. Animal experiments on reindeer antlers as a novel animal model will serve for investigation of the bone remodeling processes in the implant bed. The main interest is directed towards the time and loading-dependant behavior of the antler tissue around the implants. The aim and scope of this work was to design an autonomous loading device that has the ability to load an inserted implant in the antler with predefined occlusal forces for predetermined time protocols. The mechanical part of the device can be attached to the antler and is capable of cyclically loading the implant with forces of up to 100 N. For the calibration and testing of the loading device a biomechanical measuring system has been used. The calibration curve shows a logarithmic relationship between force and motor current and is used to control the force on the implant. A first test on a cast reindeer antler was performed successfully.     

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.     

Long or Heavy? Physiological Constraints in the Evolution of Antlers

The evolution of the investment in exaggerated secondary sexual traits is a topic of great interest for scientists. Despite antlers in the family Cervidae being one of the most interesting allometric structures, the nature of the relationships between antler and body size, and the influence of physiological factors driving the evolution of these characters, still remain unclear. In this paper, I examine these relationships in depth using the largest sample size ever studied (43 species). Under the hypothesis that antler growth may be limited by skeleton size as this process requires the allocation of huge amounts of mineral resources to the antlers, skeleton-related variables may more accurately explain these allometric relationships. The existence of physiological constraints should therefore be more clearly highlighted when studying the relationships between body size variables and the relative investment in the antler (measured as length or mass of antler per kg of skeleton). Results show that antler length is best described as being linearly related to head-body length rather than other measurements of size, and antler weight has a quadratic relationship with body mass. However, the relative investment in antler length (related to skeleton mass) is independent of body size variables, while the relative investment in antler mass has a quadratic relationship with shoulder height. The results obtained for antler mass reflect the existence of physiological constraints in the evolution of antlers, which are greater for larger sized species. On the other hand, the evolution of antler length may be linked to other factors, most probably sociobiological and biomechanical ones.     

Comparison of Effects of Shed Antler Hunting and Helicopter Surveys on Ungulate Movements and Space Use

Shed antler hunting (i.e., collecting cast cervid antlers) has increased in popularity during the past decade, but little is known about how this recreational activity affects ungulate movements and space use. We placed geographic positioning system (GPS)-collars on 133 female and male bighorn sheep (Ovis canadensis), bison (Bison bison), and mule deer (Odocoileus hemionus) to quantify their movements and space use during shed antler hunts compared with those behaviors during helicopter surveys in Utah, USA, from 2012 to 2015. For each species, we calculated means and 95% confidence intervals for distance moved during 90-minute segments (16 points/day) pre-event (control, 7 consecutive days prior to event), event (1–2 days), and post-event (7 consecutive days after event) for shed antler hunts and helicopter surveys. We also compared use of space for each species during these events. Female bighorn sheep did not increase distance moved or substantially change space use during shed antler hunts and helicopter surveys. Male bighorn sheep increased distance moved 41% on average during shed antler hunts and by 2.02 times during helicopter surveys but did not change space use during those events. Female bison increased distance moved 15% on average during shed antler hunts and 30% during helicopter surveys. Mule deer increased distance moved and altered space use the most during shed antler hunts; females increased distance moved 97%, and 54% of females moved a mean distance of 742 ± 642 (SD) m (range = 9–3,778 m) outside of their home ranges during those hunts for a mean of 9.2 ± 9.4 hours (range = 1.5 to 41 hr). Male mule deer increased distance moved by 2.10 times on average during shed antler hunts, and 82% of males moved a mean distance of 1,264 ± 732 m (range = 131–3,637 m) outside of their home ranges during those hunts for a mean of 12.6 ± 7.6 hours (range = 4.5–33 hr). Our results provide timely information about how legal shed antler hunting affects movements and space use of female and male ungulates, especially mule deer, and can guide the conservation of ungulate populations and their habitat. © 2021 The Wildlife Society.     

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.     

Annual variations in fecal androgen metabolites and antler cycle of captive red brocket bucks (<Emphasis Type="Italic">Mazama americana</Emphasis>) in southeast Brazil

Annual variations in fecal androgen metabolites (FAM) and antler cycle were analyzed in six red brocket bucks exposed to a natural photoperiod in southeast Brazil. Fecal samples and antlers status data were collected weekly during a 1-year period. Four animals were in hard antler throughout the year, whereas one individual cast antlers in September and other took several months to completely shed antler velvet. Monthly means (±SEM) of FAM concentrations exhibited no differences in all monitored bucks (P > 0.05). In addition, FAM levels from four males that kept hard antlers year-round were grouped by month and no differences among monthly means were found (P > 0.05). Comparisons between FAM levels at different antler stages were not possible because of the low number of males presenting changes in antler status. These findings indicate that red brocket bucks present no annual pattern in androgen secretion and antler changes, suggesting that testicular and antler cycles in this species are little or not affected by photoperiodic cues.     

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.     

Exploring the mechanisms regulating regeneration of deer antlers

Deer antlers are the only mammalian appendages capable of repeated rounds of regeneration; every year they are shed and regrow from a blastema into large branched structures of cartilage and bone that are used for fighting and display. Longitudinal growth is by a process of modified endochondral ossification and in some species this can exceed 2 cm per day, representing the fastest rate of organ growth in the animal kingdom. However, despite their value as a unique model of mammalian regeneration the underlying mechanisms remain poorly understood. We review what is currently known about the local and systemic regulation of antler regeneration and some of the many unsolved questions of antler physiology are discussed. Molecules that we have identified as having potentially important local roles in antlers include parathyroid hormone-related peptide and retinoic acid (RA). Both are present in the blastema and in the rapidly growing antler where they regulate the differentiation of chondrocytes, osteoblasts and osteoclasts in vitro. Recent studies have shown that blockade of RA signalling can alter cellular differentiation in the blastema in vivo. The trigger that regulates the expression of these local signals is likely to be changing levels of sex steroids because the process of antler regeneration is linked to the reproductive cycle. The natural assumption has been that the most important hormone is testosterone, however, at a cellular level oestrogen may be a more significant regulator. Our data suggest that exogenous oestrogen acts as a 'brake', inhibiting the proliferation of progenitor cells in the antler tip while stimulating their differentiation, thus inhibiting continued growth. Deciphering the mechanism(s) by which sex steroids regulate cell-cycle progression and cellular differentiation in antlers may help to address why regeneration is limited in other mammalian tissues.     

Antler extracts stimulate chondrocyte proliferation and possess potent anti-oxidative,anti-inflammatory,and immune-modulatory properties

The Sika deer antler is well known for its unique ability to regenerate repeatedly and grow rapidly. Furthermore, it is a precious traditional Chinese medicine and has been widely used for more than 20 centuries. The major bioactive components within the antlers are water-soluble proteins, polypeptides, and free amino acids. Many studies have shown that water-soluble antler extracts play pivotal roles in wound healing, immune system modulation, anti-oxidation, and anti-inflammation. However, the exact effects on chondrocytes are still largely unknown. In this study, we prepared fresh, aqueous extracts from growing deer antlers in a rapid growth stage. We isolated the chondrocytes from neonatal mouse rib cartilage and investigated the effects of antler extracts on chondrocyte viability. We also used the RNA-Seq method to analyze the gene expression pattern under antler extract treatment. We demonstrated that fresh extracts from Sika deer antlers in a rapid growth stage significantly promoted chondrocyte viability and kept chondrocytes proliferating continuously, while blocking maturation and further differentiation. Additionally, our results indicated that antler extracts might serve as a potent anti-oxidant, anti-inflammatory agent, and immune modulator to boost the abilities of chondrocytes against oxidative, inflammatory, and immune stresses. Thus, this study has greatly deepened our current knowledge of the molecular control of antler extracts on chondrocytes. It has also shed light on possible new strategies to further prevent and treat diseases of cartilage and other related diseases.     

Age-related changes in the testicular and antler cycles of reindeer, Rangifer tarandus

A total of 111 male reindeer of various ages was shot in all months of the year to study the relationship between the seasonal changes in testicular activity and the antler cycle. From the changes in testis weight, seminiferous tubular tissue area and plasma testosterone values and the occurrence of spermatogenesis, it is concluded that calves attain physiological puberty in their first year, during which they also complete an antler cycle. The amplitude of the cyclical change in testis weight and plasma testosterone values increases with age and can be correlated with the earlier onset of events in the spermatogenic and antler cycles of older animals. The duration of the spermatogenic and testosterone cycles of reindeer is short, and is inversely related to the long period spent without antlers. It is suggested that testosterone strongly influences the antler cycle of reindeer males.     

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”.