A new viewpoint on antlers reveals the evolutionary history of deer (Cervidae, Mammalia).

Recent molecular phylogeny of deer revealed that the characters of antlers previously focused on are homoplasious, and antlers tend to be considered problematic for classification. However, we think antlers are important tools and reconsidered and analysed the characters and structures to use them for classification. This study developed a method to describe the branching structure of antlers by using antler grooves, which are formed on the antlers by growth, and then projecting the position of the branching directions of tines on the burr circumference. By making diagrams, comparing the branching structure interspecifically, homologous elements (tines, beams, and processes) of the antlers of 25 species of 16 genera were determined. Subsequently, ancestral state reconstruction was performed on the fixed molecular phylogenetic tree. It was revealed that Capreolinae and Cervini gained respective three-pointed antlers independently, and their subclades gained synapomorphous tines. We found new homologous and synapomorphous characters, as the antler of Eld's deer, which has been classified in Rucervus, is structurally close to that of Elaphurus rather than that of Rucervus, consistent with molecular phylogeny. The methods of this study will contribute to the understanding of the branching structure and phylogeny of fossil species and uncover the evolutionary history of Cervidae.

Identification of working reindeer using palaeopathology and entheseal changes.

OBJECTIVE: This paper explores the potential of analyzing pathological lesions and entheseal changes in the identification of working reindeer. METHODS: The skeletons of 26 modern working reindeer from Siberia and Northern Finland are analyzed for pathological lesions and entheseal changes. RESULTS: Working results in elevated frequencies of pathological lesions, specifically joint disease in cervical and thoracic vertebrae, humeri, os coxae and proximal phalanges. Entheseal scores indicate the intensified use of shoulder flexors and extensors, and possibly elbow, hip and knee flexors and extensors in working reindeer. CONCLUSION: Patterns of skeletal changes can be used in the identification of working reindeer from the past. SIGNIFICANCE: This study provides first evidence that pathological lesions and entheseal changes can be used to assess draught and cargo use of reindeer. Therefore, the methods presented in this study provide an opportunity to scrutinize past reindeer herding practices, reindeer domestication and human-reindeer cooperation. LIMITATIONS: Methods need to be applied with caution due to the multifactorial etiologies of pathological lesions and entheseal changes. SUGGESTIONS FOR FUTURE RESEARCH: We suggest that in future studies, these methods are applied to archaeological material accompanied by osteometric and contextual analyses.

Structure, morphology and composition of fur on different parts of reindeer (Rangifer Tarandus) foot.

In the long-distance migration of reindeer in winter, furs of reindeer foot, as the part in direct contacting with the external environment, can play the role of protection and heat preservation. With furs on different parts of the right posterior foot (fibular side, tibial side and planta pedis) as research objects, the microstructure of reindeer foot furs was observed with a scanning electron microscope. The image displayed that the reindeer foot furs was divided into 3 layers, namely cuticular layer, cortical layer and medulla layer. It was observed from the fur surface that the scales of fur on tibial side had smooth edge, with the scale structure in mosaic and coronary types. The scale structure of furs on the other parts showed the irregular waves due to abrasion to different degrees. From the cross-section view of fur, there was a non-medullated segment on the medial part of fur on planta pedis. The medulla layer of fibular and tibial sides showed a porous foam structure. The medulla index (MI) of fur on fibular side and tibial side at distal part was 70.35% and 81.79%, respectively, and MI at medial part was 77.88% and 88.08%. The composition of reindeer foot fur was measured through infrared spectroscopy and energy spectrum analysis respectively. The element contents of foot fur varied on different parts. The content of sulfur of the furs on planta pedis was higher than that on other parts. The research results can provide foundations for the functional study and bionic design of reindeer foot furs during long distance migration and swimming.

Environmental and physiological influences to isotopic ratios of N and protein status in a Montane ungulate in winter.

Winter severity can influence large herbivore populations through a reduction in maternal proteins available for reproduction. Nitrogen (N) isotopes in blood fractions can be used to track the use of body proteins in northern and montane ungulates. We studied 113 adult female caribou for 13 years throughout a series of severe winters that reduced population size and offspring mass. After these severe winters, offspring mass increased but the size of the population remained low. We devised a conceptual model for routing of isotopic N in blood in the context of the severe environmental conditions experienced by this population. We measured δ15N in three blood fractions and predicted the relative mobilization of dietary and body proteins. The δ 15N of the body protein pool varied by 4‰ and 46% of the variance was associated with year. Annual variation in δ15N of body protein likely reflected the fall/early winter diet and winter locations, yet 15% of the isotopic variation in amino acid N was due to body proteins. Consistent isotopic differences among blood N pools indicated that animals tolerated fluxes in diet and body stores. Conservation of body protein in caribou is the result of active exchange among diet and body N pools. Adult females were robust to historically severe winter conditions and prioritized body condition and survival over early investment in offspring. For a vagile ungulate residing at low densities in a predator-rich environment, protein restrictions in winter may not be the primary limiting factor for reproduction.

Antlers on the Arctic Refuge: capturing multi-generational patterns of calving ground use from bones on the landscape.

Bone accumulations faithfully record historical ecological data on animal communities, and owing to millennial-scale bone survival on high-latitude landscapes, have exceptional potential for extending records on arctic ecosystems. For the Porcupine Caribou Herd, maintaining access to calving grounds on the Arctic National Wildlife Refuge (ANWR, Alaska) is a central management concern. However, variability in calving ground geography over the 30+ years of monitoring suggests establishing the impacts of climate change and potential petroleum development on future calving success could benefit from extended temporal perspectives. Using accumulations of female antlers (shed within days of calving) and neonatal skeletons, we test if caribou calving grounds develop measureable and characteristic bone accumulations and if skeletal data may be helpful in establishing a fuller, historically integrated understanding of landscape and habitat needs. Bone surveys of an important ANWR calving area reveal abundant shed antlers (reaching 10(3) km(-2)) and high proportional abundance of newborn skeletal individuals (up to 60% neonate). Openly vegetated riparian terraces, which compose less than 10 per cent of ANWR calving grounds, yield significantly higher antler concentrations than more abundant habitats traditionally viewed as primary calving terrain. Differences between habitats appear robust to potential differences in bone visibility. The distribution of antler weathering stages mirrors known multi-decadal calving histories and highlights portions of the antler accumulation that probably significantly extends records of calving activity. Death assemblages offer historically integrated ecological data valuable for the management and conservation of faunas across polar latitudes.

Why Rudolph's nose is red: observational study.

OBJECTIVE: To characterise the functional morphology of the nasal microcirculation in humans in comparison with reindeer as a means of testing the hypothesis that the luminous red nose of Rudolph, one of the most well known reindeer pulling Santa Claus's sleigh, is due to the presence of a highly dense and rich nasal microcirculation. DESIGN: Observational study. SETTING: Tromsø, Norway (near the North Pole), and Amsterdam, the Netherlands. PARTICIPANTS: Five healthy human volunteers, two adult reindeer, and a patient with grade 3 nasal polyposis. MAIN OUTCOME MEASURES: Architecture of the microvasculature of the nasal septal mucosa and head of the inferior turbinates, kinetics of red blood cells, and real time reactivity of the microcirculation to topical medicines. RESULTS: Similarities between human and reindeer nasal microcirculation were uncovered. Hairpin-like capillaries in the reindeers' nasal septal mucosa were rich in red blood cells, with a perfused vessel density of 20 (SD 0.7) mm/mm(2). Scattered crypt or gland-like structures surrounded by capillaries containing flowing red blood cells were found in human and reindeer noses. In a healthy volunteer, nasal microvascular reactivity was demonstrated by the application of a local anaesthetic with vasoconstrictor activity, which resulted in direct cessation of capillary blood flow. Abnormal microvasculature was observed in the patient with nasal polyposis. CONCLUSIONS: The nasal microcirculation of reindeer is richly vascularised, with a vascular density 25% higher than that in humans. These results highlight the intrinsic physiological properties of Rudolph's legendary luminous red nose, which help to protect it from freezing during sleigh rides and to regulate the temperature of the reindeer's brain, factors essential for flying reindeer pulling Santa Claus's sleigh under extreme temperatures.

No 'bypass' in adult ruminants: passage of fluid ingested vs. fluid inserted into the rumen in fistulated muskoxen (Ovibos moschatus), reindeer (Rangifer tarandus) and moose (Alces alces).

In young ruminants, the reticular groove ensures that ingested milk is channelled past the forestomach to avoid malfermentation. It has been speculated that some adult wild ruminants, in particular browsing species, maintain a functional oesophageal (reticular) groove, that soluble nutrients can thus bypass the rumen, and that thus the energetic gain from the diet can be increased. We inserted a fluid marker (Co-EDTA) via cannula into the rumen and simultaneously fed a diet that contained a second fluid marker (Sm-EDTA), and analysed the faecal marker excretion patterns, in muskoxen (Ovibos moschatus, n=4 in two experiments each), reindeer (Rangifer tarandus, n=4 in a total of six experiments) and moose (Alces alces, n=1 in one experiment). In no case was the orally fed marker excreted distinctively earlier than the marker inserted into the rumen, which indicates that substantial bypass did not occur in these animals. However, differences between the three species in the excretion of the two markers from the rumen are consistent with hypothetical differences in the stratification of rumen contents. We suggest that effects previously ascribed to a "rumen bypass" in wild ruminants most likely reflect differences in the passage from the rumen.

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+/-0.4 GPa 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.

[Changes of the femoral bone microstructure in the reindeer during the fetal period of ontogenesis].

The diaphysis of the femoral bone was studied morphometrically in 30 reindeer fetuses aged 2-7.5 months. The formation of the diaphyseal cavity of the femoral bone took place from 3 till 7.5 months, rapidly progressing during all the fetal period. The thickness of the periosteum in the epiphysis and the diaphysis of the bone increased from 2 to 6 months inclusive, whereupon it decreased by the time of birth. The cartilaginous tissue in the epiphyses was present from 2 till 7.5 months; its growth was registered up to 3 months in a proximal epiphysis, while it continued till 4 months in a distal epiphysis. Later on, the thinning of the cartilage was noted till the birth. The thickness of spongy substance of bone epiphyses increased with the fetal age. The osteons in the diaphysis of the femoral bone were formed in 2-month-old fetus, their numbers were found to increase with age. The compact substance of the diaphysis of the femoral bone increased in thickness till 5 months, whereupon the process of thinning of tissue till the time of birth, was noted.

Papillar morphology of the rumen of forest reindeer (Rangifer tarandus fennicus) and semidomesticated reindeer (R. t. tarandus).

The papillar morphology of the ventral and dorsal rumen of the wild forest reindeer (Rangifer tarandus fennicus Lönn.) and semidomesticated reindeer (R. t. tarandus L.) was studied in October and November 1996. The morphological measurements which were carried out were: the lengths of the papillae, the number of the papillae per square centimetre, the cross-sectional area and perimeter of sections cut from the middle of papillae. From these measurements mean papillar volume, areal papillar volume, mean papillar (epithelial) surface area, areal papillar surface and surface enlargement factor were calculated. No differences in these measurements between ventral and dorsal walls of the rumen were evident. The semidomesticated reindeer had longer papillar perimeters, larger mean and areal papillar surface areas, larger mean papillar volumes, and a larger surface enlargement factor in the ventral rumen than did forest reindeer. This may be a result of differences between feeding habits, the semidomesticated reindeer preferring a diet including more plants rich in carbohydrates e.g. lichens, which has resulted in a high production of volatile fatty acids and thus stimulation of papillar growth.

Nordic rattle: the hoarse vocalization and the inflatable laryngeal air sac of reindeer (Rangifer tarandus).

Laryngeal air sacs have evolved convergently in diverse mammalian lineages including insectivores, bats, rodents, pinnipeds, ungulates and primates, but their precise function has remained elusive. Among cervids, the vocal tract of reindeer has evolved an unpaired inflatable ventrorostral laryngeal air sac. This air sac is not present at birth but emerges during ontogenetic development. It protrudes from the laryngeal vestibulum via a short duct between the epiglottis and the thyroid cartilage. In the female the growth of the air sac stops at the age of 2-3 years, whereas in males it continues to grow up to the age of about 6 years, leading to a pronounced sexual dimorphism of the air sac. In adult females it is of moderate size (about 100 cm3), whereas in adult males it is large (3000-4000 cm3) and becomes asymmetric extending either to the left or to the right side of the neck. In both adult females and males the ventral air sac walls touch the integument. In the adult male the air sac is laterally covered by the mandibular portion of the sternocephalic muscle and the skin. Both sexes of reindeer have a double stylohyoid muscle and a thyroepiglottic muscle. Possibly these muscles assist in inflation of the air sac. Head-and-neck specimens were subjected to macroscopic anatomical dissection, computer tomographic analysis and skeletonization. In addition, isolated larynges were studied for comparison. Acoustic recordings were made during an autumn round-up of semi-domestic reindeer in Finland and in a small zoo herd. Male reindeer adopt a specific posture when emitting their serial hoarse rutting calls. Head and neck are kept low and the throat region is extended. In the ventral neck region, roughly corresponding to the position of the large air sac, there is a mane of longer hairs. Neck swelling and mane spreading during vocalization may act as an optical signal to other males and females. The air sac, as a side branch of the vocal tract, can be considered as an additional acoustic filter. Individual acoustic recognition may have been the primary function in the evolution of a size-variable air sac, and this function is retained in mother-young communication. In males sexual selection seems to have favoured a considerable size increase of the air sac and a switch to call series instead of single calls. Vocalization became restricted to the rutting period serving the attraction of females. We propose two possibilities for the acoustic function of the air sac in vocalization that do not exclude each other. The first assumes a coupling between air sac and the environment, resulting in an acoustic output that is a combination of the vocal tract resonance frequencies emitted via mouth and nostrils and the resonance frequencies of the air sac transmitted via the neck skin. The second assumes a weak coupling so that resonance frequencies of the air sac are lost to surrounding tissues by dissipation. In this case the resonance frequencies of the air sac solely influence the signal that is further filtered by the remaining vocal tract. According to our results one acoustic effect of the air sac in adult reindeer might be to mask formants of the vocal tract proper. In other cervid species, however, formants of rutting calls convey essential information on the quality of the sender, related to its potential reproductive success, to conspecifics. Further studies are required to solve this inconsistency.

The relative role of winter and spring conditions: linking climate and landscape-scale plant phenology to alpine reindeer body mass.

The relative importance of winter harshness and early summer foraging conditions are of prime interest when assessing the effect of global warming on Arctic and mountainous ecosystems. We explored how climate and vegetation onset (satellite-derived normalized difference vegetation index data) determined individual performance in three reindeer populations (data on 27814 calves sampled over 11 years). Snow conditions, spring temperatures and topography were the main determinants of the onset of the vegetation. An earlier onset positively affected the body mass of calves born the following autumn, while there was no significant direct negative impact of the previous winter. This study underlines the major impact of winter and spring climatic conditions, determining the spring and summer food availability, and the subsequent growth of calves among alpine herbivores.

Functional anatomy of the omasum in high Arctic Svalbard reindeer (Rangifer tarandus platyrhynchus) and Norwegian reindeer (Rangifer tarandus tarandus).

The structure and fill of the omasum was investigated in summer and in winter in adult female reindeer living on the polar desert and tundra of the high Arctic archipelago of Svalbard and in sub-Arctic mountain habitats in northern Norway. The mean total mass of the omasum in non-lactating adult female Svalbard reindeer was 467 g (0.65 g per 100 g live body mass (BM)) in September and 477 g (1.03 g per 100 g BM) in April. By contrast, the mean mass of the omasum in non-lactating adult reindeer in northern Norway was 534 g (0.83 g per 100 g BM) in September but only 205 g (0.35 g per 100 g BM p < 0.05) in late March, owing to a decrease in both tissue mass and the wet mass of the contents of the organ. The mean absorptive surface of the omasum in Svalbard reindeer was 2300 cm2 in September and 2023 cm2 in April. In Norwegian reindeer, by contrast, the absorptive surface area decreased from 2201 cm2 in September to 1181 cm2 (p < 0.05) in late March. The marked seasonal decline of omasal tissue and contents in Norwegian reindeer probably results from intake of highly digestible forage plants, including lichens, in winter. Svalbard reindeer, a non-migratory sub-species, survive eating poor quality fibrous vascular plants in winter. The absence of any marked seasonal change in the mass, total absorptive surface area or filling of the omasum in Svalbard reindeer in winter despite a substantial decline in body mass presumably reflects their need to maintain maximum absorption of nutrients, including volatile fatty acids, when feeding on such poorly fermentable forage.

The effects of a prolonged undernutrition on serum lipids and fatty acid composition of reindeer calves during winter and spring.

We examined the effects of undernutrition on lipid metabolism in reindeer (<1 year) during mid-winter and spring, with particular focus on the proportions of polyunsaturated fatty acids (PUFAs) in major serum lipids. The reindeer (n=8) were fed their winter feed, lichen, ad libitum for 5 weeks, followed by 40% restriction of energy for 8 weeks and refeeding to normal for 6 weeks. The concentrations of major serum lipids, cholesterol and phospholipids decreased significantly during the ad libitum period (by 50 and 44%, respectively). The proportion of major PUFA, linoleic acid in serum cholesteryl esters, decreased from 48.2 to 38.4% during the ad libitum period (P < 0.01), and to 29.2% during the restriction period (P < 0.001). The proportion of linoleic acid in phospholipids decreased from 27.9 to 15.6% during the ad libitum period (P < 0. 001), and to 13.0% during the restriction (P < 0.01). Also alpha-linolenic acid in the major lipids decreased significantly during the ad libitum and restriction periods. The decreases in the major lipids and linoleic acid were reversed during the refeeding. The control group (n=8) which was fed high-quality concentrates ad libitum gained weight most of the spring but showed similar although slower decreases in the major serum lipids and PUFAs than the lichen group. Our results indicate that feeding reindeer on lichen during winter leads to the retardation of growth and reductions in major serum lipids and their principal C18-PUFA proportions. The decreased proportions of the principal PUFAs most probably reflect their low dietary intake but may have been modified also by seasonal factors.

Adults only. Reindeer hunting at the middle palaeolithic site salzgitter lebenstedt, northern Germany.

The Middle Palaeolithic site Salzgitter Lebenstedt (northern Germany), excavated in 1952, is well known because of its well-preserved faunal remains, dominated by adult reindeer (Rangifer tarandus). The archaeological assemblage accumulated in an arctic setting in an earlier part of the last (Weichsel) glacial (OIS5-3). The site is remarkable because of the presence of unique Middle Palaeolithic bone tools and the occurrence of the northernmost Neanderthal remains, but this paper focuses on an analysis of its reindeer assemblage. The results indicate autumn hunting of reindeer by Middle Palaeolithic hominids. After the hunt, carcasses were butchered and in subsequent marrow processing of the bones a selection against young and sub-adult animals occurred. Adults were clearly preferred, and from their bones, again, poorer marrow bones were neglected. This focus on primeness of resources has been documented in other domains of Neanderthal behaviour, but Salzgitter Lebenstedt is the best example yet known in terms of systematic and routinized processing of game. The Salzgitter Lebenstedt assemblage displays some remarkable similarities to the Late Glacial reindeer assemblages from the Ahrensburg tunnel valley sites. The subsequent review of the evidence on subsistence strategies from earlier periods of the European Palaeolithic shows that hunting of large mammals may have been a part of the behavioural repertoire of the Middle Pleistocene occupants of Europe from the earliest occupation onwards. At the same time, it is suggested that these early hunting strategies were incorporated in ways of moving through landscapes ("settlement systems") which were different from what we know from the middle parts of the Upper Palaeolithic onwards.

[Character identification of 12 kinds of pilose antler medicinal materials].

In this paper, commercial medicinal materials of 12 kinds of pilose antler, Cervus nippon, C. elaphus, C. albirostris, C. unicolor, C. eldihainanus, C. timorensis C. porinus, Dama dama, Rangifer farandus, Alces alces, Elaphurus davidianus, Capreolus capreolus were compared and identified. A key and simple character illustration were listed.

The hypoglossal nuclear complex in reindeer.

The hypoglossal complex has an ovoid shape and is largest at the level of obex. Caudally to the obex is 2/3 of the complex. The complex is divided into 4 groups. Sublingual nucleus of Roller presents a narrow strip of cells located ventrally to the middle part of hypoglossal nucleus. Laterally to obex is the paramedian nucleus of Jacobson in the form of short, narrow band. Posteriorly to Jacobson's nucleus is the intercalated nucleus of Staderini. Constinuation of the hypoglossal nucleus is the nucleus prepositus of Marburg.

Morphological peculiarities of respiratory compartments of arctic animal lungs.

BACKGROUND: Morphological and ultrastructural peculiarities of interalveolar septa in endemic arctic animals (reindeer, polar fox, lemming) are compared with laboratory animals (rat,dog). METHODS: For light microscopy, tissue samples were taken from the central and peripheral sections of all lobes of the right lung. They were fixed in 10% neutral formalin and embedded in paraffin. For electron microscopy, samples were taken from subpleural sections of the caudal lobe of the right lung, fixed in 4% paraformaldehyde for 24 hours, subsequently postfixed in 2% OsO4. for 2.0 hours. Samples were dehydrated in acetone and embedded in a mixture of Epon 812 and Araldite. Ultrathin sections were photographed at a magnification of x4,000. For each interalveolar septum, lengths and diameters were recorded and the squares of septa surface, air-blood barrier surface and the number of the structures were determined. The topography of capillaries and the ultrastructure of interstitium were described. RESULTS: Acini in the arctic animals (reindeer, polar fox, lemming) are compact. In all lobes they are fully expanded and uniformly filled with air. There is no physiological atelectasis. Alveoli appear straight and homogeneous in form and size. In the polar fox, the quantity of interalveolar pores of Kohn is twice that in the dog. The number of pores in the lemming are similar to those in the rat but their size is 1.6 times greater in diameter. In arctic animals more capillaries connect with both alveolar surfaces by an air-blood barrier and simultaneously participate in the gas exchange of two adjoining alveoli. In the polar fox and lemming the thickness of the air-blood barrier is 1.3-1.4 times less than that in the dog and rat. CONCLUSIONS: The set of morpho-functional peculiarities of the acini of arctic animals allows for an increase in gas exchange in the respiratory compartments of the lungs and provides necessary oxygenation of arterial blood at a low partial pressure of oxygen in the alveolar gas.