Bull bellows and bugles: a remarkable convergence of low and high-frequency vocalizations between male domestic cattle Bos taurus and the rutting calls of Siberian and North American wapiti

Whereas low-frequency bellows (below 200 Hz) have been investigated in detail in both male and female domestic cattle (Bos taurus), male high-frequency bugle calls (over 800 Hz) have not been described so far in any large bovid species. In this study, high-frequency bugles and low-frequency bellows were recorded from three crossbred free-ranging domestic cattle bulls and analysed spectrographically. The maximum fundamental frequency of bellows was 113.6 Hz, whereas the maximum fundamental frequency of bugles was 958.4 Hz, ranging from 801 to 1125 Hz in different males. These amazingly high fundamental frequencies of bull bugles are comparable with those reported for Siberian wapiti Cervus elaphus sibiricus, but lower than reported values for some subspecies of North American wapiti Cervus canadensis. The similarity with both Siberian and North American wapiti was also observed in the production of bull biphonic bugles with two fundamental frequencies: the low and the high one. We suggest that bugles of domestic cattle bulls provide an excellent model for comparative research with cervid bugles concerning the mechanism of vocal production and the underlying anatomical and behavioural adaptations.     

Spectrographic analysis points to source–filter coupling in rutting roars of Iberian red deer

Source–filter coupling is the rarest acoustic phenomenon not only in Iberian red deer, but in any mammal. In most mammals, sound production can be well described in the framework of source–filter theory. The vocal output is the result of combined work of the larynx (the source) and of the supralaryngeal vocal tract (the filter). The source–filter theory suggests the independence of source and filter. Thus, vocal tract filtering should not affect the fundamental frequency (f0) of the sound created in the larynx. Spectrographically, the source is mostly characterized by the f0 and its harmonics, while the filter by the vocal tract resonances, i.e., formant frequencies. Nevertheless, a non-independent (coupled) source and filter can be proposed when the vocal folds start oscillating at one of the formant frequencies. Coupling between source and filter has been found in human singers and predicted for red deer Cervus elaphus by a computer modeling approach. This study describes different modes of phonation in a natural bout of rutting calls of Iberian red deer Cervus elaphus hispanicus and the transition from a chaotic mode to a probable source–filter coupling mode. This phenomenon might be involved in the production of extremely high-frequency bugles of North American and Asian subspecies of C. elaphus.     

Seasonal migration of male red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) in southern Sweden and consequences for management

Red deer Cervus elaphus is a highly appreciated and intensively managed game species throughout Europe. A common management objective is a sustainable harvest of large trophies. In southern Sweden, management has mainly aimed at preserving the nominate subspecies C. elaphus elaphus. Seasonal migration of red deer males may, however, complicate both harvest management as well as conservation efforts. I used individually identified male red deer in southern Sweden to observe distance travelled from rutting areas to areas used by males in summer and winter. Adult males were identified by antler shape and photo-documented during rut. Photos from the rut were compared to trophies of deer harvested or found dead, to found cast antlers and to stags photographed during summer. From 1969 to 2007, a distance between rutting ground and summer/winter quarters was established for 96 identified stags. An average distance of 14 km and a maximum distance of 47 km were recorded between rut and summer/winter observations. The seasonal migration of males increases the risk of overexploitation of males with harvest in both rutting areas and wintering areas. Harvest management and conservation efforts may fail if males seasonally migrate outside the management unit. The results suggest that seasonal migration must be considered in harvest management and conservation and that there is a need for a regulation of male harvest. Furthermore, the study stresses that the success in deer management of single hunting units, may be largely dependent on the harvest policies in the near surroundings as well as in areas tenths of kilometres away, suggesting that a successful management must rely on co-operation and co-ordination on a landscape scale.     

Phylogeography of red deer (Cervus elaphus): Analysis of MtDNA cytochrome b polymorphism

Analysis of MtDNA cytochrome b gene (1140 bp) polymorphism of 106 samples red deer (Cervus elaphus) from different regions of Eurasia was performed; the phylogenetic relationships of groups throughout the entire inhibiting area (including North America) were reconstructed. Totally 75 haplotypes were detected, 33 of which were found in the European and 42 in the Asian part of the area. There were no identical haplotypes for these two parts of the area found. The close relatedness between Siberian red deer (C. e. sibirica) and North American wapiti (C. e. canadensis) was confirmed. Red deer inhibiting Yakutia were close to the Siberian red deer from Altai and Tuva, whereas red deer inhibiting Krasnoyarsk and Irkutsk regions formed a separate clade. Overall, the reconstructed phylogeographic pattern of the species was significantly different from the subspecies differentiation based on morphological traits.     

A phylogenetic comparison of red deer and wapiti using mitochondrial DNA

A phylogeny was constructed for red deer/wapiti (Cervus elaphus) subspecies using sequence data from the control region of mitochondrial DNA (mtDNA). The tree was rooted using Cervus nippon (sika deer), Cervus albirostris (Thorold's white-lipped deer), and several Odocoileinae species. A division between the mtDNA haplotypes of red deer (European) and wapiti (Asian/North American) corresponds to subspecies found on opposite sides of the Himalayan Mountains and Gobi, which suggests wapiti should be reconsidered for the status of C. canadensis. Using parsimony and distance analysis, red deer and wapiti are derived from a single recent common ancestor, which is consistent with current taxonomy that recognizes the subspecies of Cervus elaphus as monophyletic group. However, maximum-likelihood analysis using weighted transitional substitutions caused red deer to form a sister group to sika deer (Cervus nippon) and wapiti. A phenetic comparison revealed wapiti also share more nucleotide similarities with sika deer, although approximately 5% sequence divergence separates wapiti, sika, and red deer. Phylogenetic evidence from the cytochrome b sequences corroborated observations from the control region. Observations from this study suggest that the species status of wapiti should be reinstated.     

Advertising individual identity by mother and adolescent contact calls in Siberian wapiti Cervus elaphus sibiricus

Individualistic contact calls facilitate mother‐offspring reunion after separation. However, in many mammals, both the acoustic structure and individuality of contact calls differ between mother and young. In contrast, in Siberian wapiti Cervus elaphus sibiricus, contact calls are similar in the acoustics between mother and young, whereas effects of this similarity on vocal individuality were not investigated. In this study, we analyzed acoustic differences between closed‐mouth (nasal) and open‐mouth (oral) contact calls and examined individuality of the most usual oral calls of 19 Siberian wapiti (9 hinds and 10 5–6‐month adolescents) emitted in response to mother‐offspring separation. In the oral calls, the values of frequency and power variables were higher than in the nasal calls. Calls of hinds and adolescents did not differ by the maximum fundamental frequency and duration, whereas the peak frequency was higher in the young. Discriminant function analysis (DFA) based on 11 acoustic variables of oral calls accurately classified to individual 92.5% of hind calls and 96.9% of adolescent calls (chi‐square test for differences between hinds and adolescents, p = 0.19). Variables mainly contributing to vocal identity (duration, start, and maximum fundamental frequency) were the same in calls of mothers and adolescents. We conclude that similarities in the acoustics calls of mothers and adolescents mean that they do not differ in their potential for encoding individual identity, suggesting a mutual process of mother‐offspring vocal recognition in Siberian wapiti.     

Do Red Deer Stags (Cervus elaphus) Use Roar Fundamental Frequency (F0) to Assess Rivals?

It is well established that in humans, male voices are disproportionately lower pitched than female voices, and recent studies suggest that this dimorphism in fundamental frequency (F0) results from both intrasexual (male competition) and intersexual (female mate choice) selection for lower pitched voices in men. However, comparative investigations indicate that sexual dimorphism in F0 is not universal in terrestrial mammals. In the highly polygynous and sexually dimorphic Scottish red deer Cervus elaphus scoticus, more successful males give sexually-selected calls (roars) with higher minimum F0s, suggesting that high, rather than low F0s advertise quality in this subspecies. While playback experiments demonstrated that oestrous females prefer higher pitched roars, the potential role of roar F0 in male competition remains untested. Here we examined the response of rutting red deer stags to playbacks of re-synthesized male roars with different median F0s. Our results show that stags’ responses (latencies and durations of attention, vocal and approach responses) were not affected by the F0 of the roar. This suggests that intrasexual selection is unlikely to strongly influence the evolution of roar F0 in Scottish red deer stags, and illustrates how the F0 of terrestrial mammal vocal sexual signals may be subject to different selection pressures across species. Further investigations on species characterized by different F0 profiles are needed to provide a comparative background for evolutionary interpretations of sex differences in mammalian vocalizations.     

Male red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) dispersal during the breeding season

Breeding dispersal can be of significant ecological and evolutionary importance. Yet, it is seldom considered in mammals. I present data on male red deer (Cervus elaphus) movements between sub-populations in southern Sweden during the rut. I investigated whether these movements could be breeding dispersal driven by mate competition. During the ruts of 1998–2009, I recorded 91 movements of males. The longest movement distance was 18.5 km. Dispersal was not restricted to yearlings or sub-adults, but also observed among adult stags. Of 91 movements observed, 7 were made by yearlings, 46 by sub-adults and 38 by adults. There was a significant move among yearlings and sub-adults towards areas with a higher ratio of females/adult males and towards areas with more females. The movements between rutting areas thereby seemed driven by sexual competition.     

The fossil fallow deer <Emphasis Type="Italic">Dama geiselana</Emphasis> (Cervidae,Mammalia, upgrade to species level) in the context of migration and local extinctions of fallow deer in the Late and Middle Pleistocene in Europe

The fossil fallow deer Dama dama geiselana Pfeiffer from Neumark-Nord (Saxony-Anhalt, Germany) is upgraded to species level and discussed within the current taxonomy of recent and fossil fallow deer. Typical antler and skeletal characteristics are discussed in comparison with the recent Dama dama, Dama mesopotamica and the fossil species Dama clactoniana, Dama nestii and Dama rhenana. The phylogenetic relationship of fallow deer can be traced back to the Late Pliocene and distinguished morphologically from Cervus elaphus, Cervus nippon and Axis axis. Late and Middle Pleistocene finds from Germany are presented and discussed in the context of the finds from the Mediterranean region and Great Britain. The differences in antler morphology and bone dimensions in West German and North-East German fallow deer from the Late Pleistocene support the hypothesis of different immigration channels, on the one hand from the eastern Mediterranean along the Danube and on the other from the west along the Rhone and Rhine. In the Middle Pleistocene, Dama mesopotamica is considered as the typical fallow deer in the eastern Mediterranean, while in the west, Dama clactoniana is widespread. The hypothesis of immigration from the eastern Mediterranean is supported by the fossil record in Germany with the fallow deer from Edesheim. Conversely, Dama geiselana probably influenced East Mediterranean populations. Special tooth characteristics of Dama geiselana occur with lower frequency in Dama mesopotamica. In the Bronze Age, the fallow deer from Kastanas (Macedonia) shares antler characteristics, the high frequency of specific features of the scapula, and the astragalus with Dama geiselana. A relict population of Dama geiselana probably reached the Eastern Mediterranean at the beginning of the last cold stage.     

四川梅花鹿发情期的几种发声行为

2006 年9 ～11 月在四川省若尔盖县铁布自然保护区,对四川梅花鹿发情期的声行为进行了初步观测,结果表明:在发情期,四川梅花鹿的发声行为可分为雌、雄鹿的报警叫声,雄鹿的吼叫声和求偶叫声。雌鹿的报警叫声持续时间257 ～ 539 ms,频率范围1409. 5 ～ 4474. 6 Hz,主频率3534. 8 ±89.12 Hz;雄鹿的报警叫声持续时间136 ～187 ms,频率范围271.8 ～3910.5 Hz,主频率3244.3 ±79.32 Hz;两者在持续时间、最低频率、最高频率上差异极显著(P < 0.01),在间隔时间上差异不显著(P = 0.624)。吼叫是雄鹿的主要发情行为之一。雄鹿每次吼叫1 声,持续时间1580 ～4972 ms,频率范围234.6 ～6171.4 Hz,主频率2264.6 ± 166.44 Hz。雄鹿吼叫声的主频率存在显著的个体差异(P <0.01)。在整个吼叫过程中,一只雄鹿的吼叫常会引起周围其它雄鹿的吼叫反应。雄鹿每日吼叫的次数与其在繁殖群中的等级序列有关,不同序列等级雄鹿的吼叫频次存在显著差异(P<0.01)。雄鹿的吼叫声在白天和夜晚均能听到,但主要发生在06:00 ～ 08:00、17:00 ～ 19:00 和01:00 ～ 03:00 三个时间段。四川梅花鹿雄鹿的求偶叫声有4 种,其生物学意义与发情炫耀、追逐、激惹和爬跨等行为有关。     

梅花鹿华南亚种和东北亚种警戒声比较

梅花鹿Cervus nippon是社群性哺乳动物,其声音通讯在社群中发挥十分重要的作用。梅花鹿目前存在3个亚种,本研究采集了浙江省清凉峰国家级自然保护区的野生华南亚种C.n.kopschi和浙江省温州康寿鹿业科技有限公司的圈养东北亚种C.n.hortulorum的警戒声,进行声音特征分析。结果显示:(1)华南亚种具有较为多态性的警戒声,具有单音节和双音节2种不同类型的叫声,明显清晰的频带有3~4个,频率分布在1 726~15 403 Hz,而东北亚种虽也有单音节和双音节2种不同类型的叫声,但明显清晰的频带只有1个,频率分布在307~4 523 Hz;(2)综合其他亚种的相关参数,认为清凉峰华南亚种的警戒声频率范围最广且频带最多,推测由其生存的环境或遗传因素导致;(3)东北亚种的警戒声频带数量少和频率范围较窄,推测与长期的人为养殖有一定的关系。本研究结果将对浙江清凉峰野生梅花鹿种群的保护、管理与利用提供一定的帮助。     

A new sexual signal in rutting male red deer: Age related chemical scent constituents in the belly black spot

Rutting behaviour of red deer stags (Cervus elaphus) includes an extensive repertoire of visual and acoustic signals directed either to rival males or to females. As in other mammals, olfactory communication is expected to play a central role in these rutting interactions too, but this has rarely been investigated. Only during the rutting season, red deer males show a conspicuous black spot area throughout most of their underbelly produced by the impregnation of substances with a strong scent. Here, we examined the origin of these compounds and their potential role as chemical signals. By using gas chromatography–mass spectrometry (GC–MS), we identified 67 compounds in the hair from the belly black spot of red deer stags, mainly heterocyclic aromatic organic compounds, such as m-cresol, benzoic acid, cyclohexanecarboxylic acid and ethylphenol, but we also found steroids, such as cholesterol and androstane-3,17-dione, carboxylic acids and their esters between n-C₆ and n-C₂₂, alcohols, squalene and other minor compounds. Many of these compounds are found in the belly black spot but not in other hair areas, and may have originated from several sources, such as the urine or the sebaceous glands of the skin, which impregnated the belly. Moreover, we found differences in chemical profiles depending on age, with older males having higher proportions of benzoic acid and androstane-3,17-dione, but lower proportions of m-cresol. Because most of these compounds are strongly odoriferous, and appear related to male characteristics, our data indicate that scent from the hairs forming the black spot of the belly may be regarded as an overlooked new sexual chemical signal in red deer in the context of competition for mates during the rutting season.     

Ungulate: vehicle collision rates are associated with the phase of the moon

The phase of the moon can affect activity patterns of nocturnal animals, and may also affect visibility for motorists. However, surprisingly little is known about whether the risk of a wildlife-vehicle collision (WVC) is associated with lunar phase. We investigated the relationship between frequency of WVC at night and lunar phase for four large ungulate species that account for a high proportion of serious WVC along roads in agricultural and forested landscapes of two continents: wild boar Sus scrofa, roe deer Capreolus capreolus, and red deer Cervus elaphus in Castile and Leon, Spain, and white-tailed deer Odocoileus virginianus in New York State, USA. Three of the four species most frequently collided with vehicles at night during the full moon phase of the lunar cycle; this pattern was evident throughout the year but was stronger during some months. For roe deer, the species for which WVC was most closely associated with the lunar cycle, the frequency of WVC was 71.3% greater during the full than new moon period. Our results indicate that rates of ungulate WVC at night cycle on a period of a lunar month, which has implications for traffic safety planning and for motor vehicle collision emergency response preparation.     

A new form of wapiti Cervus canadensis Erxleben, 1777 (Cervidae,Mammalia) from the Late Pleistocene of France

A well-preserved antlered braincase of wapiti (Cervus canadensis) from the Late Pleistocene of Saint-Hippolyte (Puy de Dôme, France) is described herein. The specific morphology of antlers suggests peculiar adaptations to the open landscapes of periglacial tundra-steppe that permitted to identify a new fossil subspecies Cervus canadensis combrayicus n. ssp. The specimen attests the occurrence of Cervus canadensis in the paleontological record of Western Europe and helps to clarify the systematical position of some disputed findings of Cervus from Western Europe. The revised systematic position of some fossil and sub-fossil cervid findings reveals the paleobiogeographic story of wapiti in Western Europe during the Last Glacial Maximum and the postglacial time. The treeless Alpine altitudes and Sweden are proposed as the glacial refugia for the last European wapiti. The medium-sized deer from Capri Island is regarded as an insular dwarfed wapiti C. canadensis tyrrhenicus closely related to the continental form C. canadensis palmidactyloceros from the alpine refugium.     

Red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis> Linnaeus, 1758) increasing density effects on species assemblage of ground beetles (Coleoptera: Carabidae) in Alpine forests

Red deer (Cervus elaphus) increasing density may trigger several modifications at the ecosystem level. Positive and negative impact of red deer on the invertebrate fauna is known in northern Europe, whereas it is currently unknown in the Alps. In this paper, we tested the impact of red deer density on ground beetles’ (Coleoptera: Carabidae) richness and species distribution in forests of the central-eastern Italian Alps (Stelvio National Park). Carabid beetles were sampled using pitfall traps in nine forest sites with different density of red deer. Carabid species richness was related to elevation, habitat type and red deer density in summer and winter using generalised linear mixed models and canonical correspondence analysis. The high winter red deer density affected positively the carabid species richness. Overgrazing seemed to increase presence of generalist species of dried and unshaded soils and decreasing of specialised species typical of well-preserved Rhododendro-Vaccinietum cover. Accordingly, the presence of patches with different red deer density seems to contribute to maintaining high levels of carabid beetles’ richness, albeit overgrazing needs to be monitored because it is likely to cause a local decrease of specialised forest species.     

Great expectations: dominance,breeding success and offspring sex ratios in red deer

Differences in dominance rank among red deer (Cervus elaphus) females (hinds) on Rhum were related to their breeding success as well as to the comparative success of male and female offspring. Males (stags) born to mothers above median rank were more successful than hinds, while hinds born to subordinate mothers were more successful than stags. The ratio of male to female calves produced by dominant mothers was significantly higher than that produced by subordinates. Since dominance rank among hinds is related to their body weight as adults and to their birth weight, these results suggest that the birth sex ratio may be affected by environmental factors operating during a female's early development.     

Estimating red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) population size in the Southern Black Forest: the role of hunting in population control

Numerous studies have addressed the question of whether hunting is capable of limiting the abundance of ungulates in the northern hemisphere. We investigated whether the hunting of red deer (Cervus elaphus) has reduced their abundance in the Southern Black Forest (area 17,500 ha), Southern Germany, since 2006. Red deer abundance was estimated using data obtained from visual counts at winter feeding sites, track counts, and bag records. An age- and sex-structured population model to estimate the winter population size was also constructed using bag records. The estimated red deer population size was evaluated according to a non-invasive genetic mark-recapture approach. The results showed that the hunting of red deer can reduce their population size if the hunting regime is part of a holistic management concept that takes into account the uncertainty of population size estimates and is implemented at scales appropriate to the management of this species.     

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”.     

Genetic variation and structure in Scandinavian red deer (Cervus elaphus): influence of ancestry,past hunting,and restoration management

In the 19^th century, the red deer (Cervus elaphus) population in Sweden experienced a rapid decline in numbers and distribution. A small population was, however, remnant in the southernmost province (Skåne) of the country, presumably corresponding to the nominate form of red deer (Cervus elaphus elaphus Linnaeus, 1758). After management, reintroductions, and supplementary release during the 20^th century the Swedish C. elaphus population recovered. The recovery was partially uncontrolled, and included introductions of C. elaphus of continental origin. In northern central Sweden (Jämtland) the current C. elaphus population may stem from natural colonization from Norway and/or from specimens of Swedish origin that have escaped from enclosures. To evaluate the status of the current, partially separated populations, we investigated variation at microsatellite markers in 157 C. elaphus specimens from ten locations in Sweden and Norway. Analyses suggest that the highest‐likelihood phylogenetic structure among the individuals sampled is described four distinct genetic clusters: (1) animals from the province of Västergötland in south‐western Sweden; (2) deer from the southernmost province of Skåne; (3) deer from the provinces Jämtland, Blekinge, and Västmanland; and (4) Norwegian deer. Cervus elaphus from a captive herd at the Skåne Zoo cluster with deer from Skåne or deer from Västergötland, depending on the method of analysis. A number of populations in Sweden may genetically match the nominate form of red deer (C. e. elaphus). The recently established C. elaphus population in Jämtland seems to stem mainly from escapees from enclosures, with a mixed ancestry from the wild remnant population in Skåne and continental deer, whereas the influx from Norway is minor, if any. Our results show the need for a detailed assessment of genetic differentiation, and emphasize the value of local management plans when planning and managing introductions. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 43–53.     

Selectivity of wolf predation on red deer in the Bieszczady Mountains, Poland

A pattern of wolfCanis lupus Linnaeus, 1758 predation on red deerCervus elaphus Linnaeus, 1758 was studied in Bieszczady Mountains in 1991–2002. In total 324 remains of red deer > 4 months old, killed by wolves throughout the year, were found. The sex, age and bone marrow fat content of wolf kills were compared with the same characteristics within the free living red deer population. The overall contribution of calves killed by wolves (24%) in October-May was higher than in the population (17%), and decreased from autumn to spring. Adult males were more vulnerable to wolf predation than adult females: stags constituted 62% and hinds 38% of adult red deer killed by wolves, whereas in the population, the percentages were 37 and 63%, respectively. Stags killed by wolves were younger ([`(x)] = 4.1\bar x = 4.1 years old) than hinds ([`(x)] = 8.9\bar x = 8.9 years old). Wolves killed more > 8 years old hinds and < 5 years old stags than available in the population. In wolf kills, the average fat content in femur marrow was higher among hinds (84.9%) than stags (69.3%) and calves (66.1%). Only 8% of hinds had < 70% femur marrow fat content, whereas 40% of calves and 38% of stags had marrow fat values below that level. Marrow fat content showed seasonal variation and was the lowest in March among all sex-age classes. The monthly share of stags in all kills, and hinds in hind-calf part of the sample was negatively correlated with their monthly average bone marrow fat content, and monthly share of calves was positively correlated with monthly average bone marrow fat content of adults. The segregation of social units (hind-calf and stag groups), except during the rutting season, and the low fat reserves of males from midwinter until spring contribute to the high overall incidence of calves and adult males and the relatively low incidence of adult females among wolf kills.