A new species of crown‐antlered deer Stephanocemas (Artiodactyla,Cervidae) from the middle Miocene of Qaidam Basin,northern Tibetan Plateau,China, and a preliminary evaluation of its phylogeny

Stephanocemas Colbert, 1939 is one of the earliest groups of deer (Cervidae) that shows indications of shedding their antlers, and as such, occupies a special position in the transition from nondeciduous to deciduous antlers in the evolution of cervids. Despite this basal position in cervid phylogeny, however, it has a highly peculiar, palmate antler morphology that is easily recognized in the fossil records of the early to middle Miocene of Eurasia. A new species, Stephanocemas palmatus sp. nov. , is described based on three partial antlers ranging from juvenile to full adult individuals, recently collected from the late middle Miocene Lower Youshashan Formation in the Qaidam Basin of northern Tibetan Plateau. Another indeterminate species (Stephanocemas sp.) of much smaller size and primitive characteristics is described from more fragmentary materials in strata lower than the level of S. palmatus but still middle Miocene in age. To the extent that it was possible, we also attempted to place into modern stratigraphical context historical collections of Stephanocemas from Qinghai Province that were obtained during the Sino‐Swedish Expeditions in the 1930s. These materials were described by Birger Bohlin in 1937 but their stratigraphical context has been in question ever since. A species‐level phylogeny suggests that S. palmatus is the most derived among known species, with its large size, great expansion of the palm part of the antler, and horizontally radiating tines. We recognize a Paradicrocerus clade that includes P. flerovi Gabuniya, 1959, P. elegantulus (Roger, 1898), and P. brevistephanos (Baschanov & Nurumov, 1955); the latter two were traditionally regarded as belonging to Stephanocemas. As a result, the Paradicrocerus clade is largely European and western Asian, whereas the restricted Stephanocemas is now Asiatic in geographical distribution. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 680–695.     

The petrosal bone and bony labyrinth of early to middle Miocene European deer (Mammalia,Cervidae) reveal their phylogeny

Deer (Cervidae) have a long evolutionary history dating back to the Early Miocene, around 19 million years ago. The best known fossils to document this history belong to European taxa, which all bear cranial appendages more or less similar to today's deer antlers. Despite the good fossil record, relationships of the earliest stem deer and earliest crown deer are much debated. This hampers precise calibration against the independent evidence of the fossil record in molecular clock analyses. While much has been written on the Early and Middle Miocene deer, only two phylogenetic analyses have been performed on these taxa to date mostly based on cranial appendage characters. Because the petrosal bone and bony labyrinth have been shown to be relevant for phylogeny in ruminants, we describe for the first time these elements for four iconic early cervids from Europe (Procervulus dichotomus, Heteroprox larteti, Dicrocerus elegans and Euprox furcatus) and include them in a phylogenetic analysis based on the ear region exclusively. The analysis recovered E. furcatus in a sister position to the living red deer (Cervus elaphus). Further, it placed D. elegans in a sister position to Euprox + Cervus and a clade Procervulinae that includes P. dichotomus and H. larteti, in sister position to all other deer. The inclusion of E. furcatus in crown Cervidae, which was previously suggested based on antler morphology, cannot be ruled out here but needs a more comprehensive comparison to other crown deer to be confirmed. J. Morphol. 277:1329–1338, 2016. © 2016 Wiley Periodicals, Inc.     

Virtual brain endocast of Antifer (Mammalia: Cervidae), an extinct large cervid from South America

A diverse fossil record of Cervidae (Mammalia) has been documented in the South American Pleistocene, when these animals arrived during the Great American Biotic Interchange. Using computed tomography-scanning techniques, it is possible to access the endocranial morphology of extinct species. Here, we studied the brain endocast of the extinct late Pleistocene cervid Antifer ensenadensis from southern Brazil, one of the largest forms that lived on this continent, using comparative morphology, geometric morphometrics, and encephalization quotients. The analyzed endocasts demonstrate that A. ensenadensis had a gyrencephalic brain, showing a prominent longitudinal sinus (=sagittal superior sinus), which is also observed in the large South American cervid Blastocerus dichotomus. The encephalization quotient is within the variation of extant cervids, suggesting maintenance of the pattern of encephalization from at least the late Pleistocene. Geometric morphometric analysis suggested a clear and linear allometric trend between brain endocast size and shape, and highlights A. ensenadensis as an extreme form within the analyzed cervids regarding brain morphology.     

Territory Establishment and Antler Cycle in Male Roe Deer

Territory establishment and antler cycle in male roe deer, Capreolus capreolus, were studied in two areas in south-eastern Sweden during 1989–1993. Initial territorial behaviour was observed in early Mar., when overlap between male ranges decreased from 66 % to 0 % over a 3–6-wk period, as determined from telemetry data. The decrease was associated with an increased number of agonistic interactions. Most territories were established at the end of Mar. to early Apr., with a tendency for males re-establishing old territories to be first. In all years, territories were taken up 1–3 wk before males shed velvet from their antlers. The distance between geometric centres of neighbouring males in winter and after territory establishment increased from 164 to 364 m. Former territory holders moved the geometric centres of their winter range and subsequent territory a shorter distance than first-year territorials, 129 and 267 m respectively. Territory defence ceased after the rut in late Aug., and the spatial relations between neighbours reverted to the situation before the territorial season. Neither velvet shedding nor antler casting was correlated with male age (yearlings excluded), which contradicts earlier statements. Neither were there any associations between timing of antler casting and subsequent velvet shedding, nor vice versa. Hence, a male casting antlers early one year derives no advantage by early shedding the next season. Furthermore, to be in hard antlers is not a prerequisite for territory take-up, whereas antlers probably are necessary for maintaining the area throughout the season. Functional aspects on roe buck territoriality are also discussed.     

关于柄杯鹿亚科(Lagomerycinae)的系统位置

柄杯鹿亚科的系统位置争论已久。争论的焦点是它们属于鹿科中的亚科还是麒麟鹿超科。经过对头骨和头后骨骼形态特点的研究和用扫描电镜检查角的结构显出柄杯鹿具有原始的角,而这种原始的角与后期鹿类的角柄同源。因此认为柄杯鹿科是鹿类中独立的一分支。     

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.     

The Miocene differentiation of old world Pecora (Mammalia)

Dental and horn core/antler morphology is discussed for some representative lower and middle Miocene pecorans in a review of the phylogeny and early zoogeography of the modern Old World families. Giraffes and bovids are more closely related than either is to deer. Andegameryx, Walangania and Teruelia are related to giraffoids plus bovoids together; Amphimoschus and Hispanomeryx are a probable sister group of bovoids; Propalaeoryx is the sister group of the giraffids plus climacoceratids. Procervulus and Lagomeryx may be closer to modern deer than are Stephanocemas and Dicrocerus but all are included in the Cervidae. The horn structure of Palaeomeryx has resemblances to giraffids, but its teeth and postcranial bones are more like cervids and it is placed in the Cervoidea. Pecorans seem to have evolved in Eurasia and there gave rise to deer. By the early Miocene some pecorans had entered Arabo‐Africa where they gave rise to giraffids and probably bovids. The relationships and zoogeography of the early bovids Eotragus, Homoiodorcas and Caprotragoides are likely to be important for later bovid history.     

Improbable appendages: Deer antler renewal as a unique case of mammalian regeneration

Deer antlers are periodically replaced cranial appendages that develop from permanent outgrowths of the frontal bones known as pedicles. Antler re-growth is a unique regenerative event in mammals which in general are unable to replace bony appendages. Recent evidence suggests that antler regeneration is a stem cell-based process that depends on the activation of stem cells located in the pedicle periosteum which are presumed to be neural crest-derived. It has been demonstrated that several developmental pathways are involved in antler regeneration that are also known to play a role in the control of skeletal development and regeneration in other vertebrates. However, in contrast to most other natural examples of regeneration of complete body structures, antler regeneration apparently neither depends on a functional nerve supply nor involves a direct contact between wound epithelium and mesenchymal tissue. Antlers thus demonstrate that regeneration of a large bony appendage in a mammal can be achieved by a process that differs in certain aspects from epimorphic regeneration in lower vertebrates.     

Revisiting the allometry of antlers among deer species: male–male sexual competition as a driver

The positive allometry between antlers and shoulder height reported for cervids has previously been interpreted as resulting from a high male‐male competition in large species that form large breeding groups. We aim at revisiting relative antler size variation among deer species by including more species (n = 31) and by testing both direct and indirect influences of different sexual selection proxies on the relative antler length using path analysis. The absence of direct effect of mating tactic on relative antler allometry indicates that the strength of fights does not differ among mating tactics. On the other hand, the main effect of breeding group size is revealed by polygyny. Highly polygynous species have relatively longer antlers than less polygynous ones but the difference in the relative effect of breeding group size on relative antler length is weak. Strong direct effect of breeding group size and indirect effect of mating tactic shaped the observed variation in the relative antler size among deer, suggesting that the amount of male‐male competition is the main evolutionary force of antler allometry in cervids.     

An appraisal of the fossil record for the Cirolanidae (Malacostraca: Peracarida: Isopoda: Cymothoida), with a description of a new cirolanid isopod crustacean from the early Miocene of the Vienna Basin (Western Carpathians)

Abstract:  Isopod crustaceans are rarely preserved in the fossil record. Herein, an appraisal of the fossil record for the cirolanid isopods is presented. Five genera are briefly discussed, including Bathynomus, Brunnaega, Palaega, Pseudopalaega and Cirolana. A key for the cirolanid genera known to date from the fossil record is provided based mostly on pleotelson characters. From the early Miocene of the Slovak part of the Vienna Basin, Cirolana feldmanni sp. nov. is described being only the fifth fossil Cirolana species known to date and one of the few with preserved appendages. The material exhibits preservation suggesting biphasic moulting; the mode of preservation suggests a rather short time between shedding the posterior and anterior parts of the exoskeleton instead of hours or even days known in extant taxa. As no subsequent transport or physical disturbance was inferred, the specimens can be stated as in situ preservation. From the palaeoecological point of view, it is concluded that Cirolana feldmanni sp. nov. is the first unequivocal fossil deep‐water Cirolana as suggested by the accompanied fauna.     

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.     

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.     

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.     

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.     

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     

Evolution of ruminant headgear: a review

The horns, ossicones and antlers of ruminants are familiar and diverse examples of cranial appendages. We collectively term ruminant cranial appendages 'headgear'; this includes four extant forms: antlers (in cervids), horns (in bovids), pronghorns (in pronghorn antelope) and ossicones (in giraffids). Headgear evolution remains an open and intriguing question because phylogenies (molecular and morphological), adult headgear structure and headgear development (where data are available) all suggest different pictures of ruminant evolution. We discuss what is known about the evolution of headgear, including the evidence motivating previous hypotheses of single versus multiple origins, and the implications of recent phylogenetic revisions for these hypotheses. Inclusion of developmental data is critical for progress on the question of headgear evolution, and we synthesize the scattered literature on this front. The areas most in need of attention are early development in general; pronghorn and ossicone development in particular; and histological study of fossil forms of headgear. An integrative study of headgear development and evolution may have ramifications beyond the fields of systematics and evolution. Researchers in organismal biology, as well as those in biomedical fields investigating skin, bone and regenerative medicine, may all benefit from insights produced by this line of research.     

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.     

The mechanism of antler casting in the fallow deer.

The process by which antlers are detached from their pedicles was examined histologically in fallow deer castrated in the autumn to induce precocious casting. Osteoclastic erosion across an abscission line between the dead bone of the antler and the living bone of the pedicle was found to be responsible for the separation of the 2. As early as 3 days after castration, osteoclasts and associated lacunae were present on the sides of the pedicle bone. These were then found in progressively deeper locations, by 2 weeks extending across the entire width of the pedicle. Concomitant with the centripetal spread of osteoclasts was the enlargement of Haversian canals, the surfaces of which became lined with osteoclasts. These widening vascular channels within the bone were filled with connective tissue, which in precasting stages formed a mesodermal pad about 1 mm thick. In later stages, a circumferential cleft was excavated beneath the antler burr, and connective tissues from the surrounding pedicle skin invaded the space between the antler and pedicle. After casting, the ingrowing integumental tissues fused with the mesodermal tissues derived from the vascular channels of the pedicle to give rise to an incipient antler bud beneath the scab. The ingrowth of epidermis capable of de novo hair follicle formation gave rise to the future velvet skin that envelops the elongating antler.     

New fossils of Eucladoceros boulei (Artiodactyla,Mammalia) from Early Pleistocene Nihewan Beds,China

Recent excavations at Shanshenmiaozui site in Nihewan Basin of North China uncovered a rich collection of comb-antlered deer, which includes the first discovery of the yearling antler, complete upper and lower dentitions (both deciduous and permanent), associated limb bones including the longest metapodials ever recovered. Based on toothrow length and the dental characters as well as the postcranial bones, the new fossil materials can be referred to Eucladoceros boulei that is estimated to be 350 kg and represents the largest Pleistocene cervid ever recovered in China. Besides the large size, E. boulei is also characterized by the pronounced anterior cingulids and entostylid ribs on lower molars as well as the moderately pachyostosed mandibles. The present study shows that the body weight of large cervids can be estimated by the length of toothrow and metacarpal, with exception for Cervus elaphus, which has larger toothrow length, but shorter metacarpal and smaller body size. E. boulei is a typical element of the Early Pleistocene fauna in northern China. The early Villafranchian is a bottleneck period for cervid evolution in northern China, which is characterized by the following features: decrease of cervid diversity, disappearance of archaic groups, and the rise of the medium to large-sized three-tined cervini taxa. The sudden appearance of the very large and/or multi-tined cervids at the Pliocene–Pleistocene transition may represent a great migration event of mammals. The Early Pleistocene cervids from Nihewan Basin are very diverse, and are in need of more taxonomic work.     

准噶尔盆地北缘中中新世偶蹄类

本文记述了新疆准噶尔盆地北缘哈拉玛盖组中的偶蹄类七属七种(包括一个新种Eot-ragus halamagaiensis sp.nov.),奇蹄类三属三种。其中始羚在在国内尚属首次报道。该化石集群与内蒙东部的通古尔动物群比较两者成分相接近,但其中一些分子具有更原始的特征。显然哈拉玛盖组比通古尔组的沉积时期早,但仍属中中新世通古尔期。