Seasonal foraging behavioural compensation in reproductive wapiti hinds (Cervus elaphus canadensis)

Captive wapiti hinds were observed during seven periods between June 1996 and November 1997. We investigated their activity and foraging behaviour on two pastures, heavily and lightly grazed, during four phases of the reproductive cycle (early gestation, late gestation, peak lactation, and late lactation). Within season, differences in forage availability between pastures had little apparent effect on wapiti feeding behaviour (p0.05). However, within each pasture, hinds tended to select locations of higher phytomass than the pasture average. Among seasons, effects of forage availability on feeding behaviour were pronounced. Hinds grazed longest in late lactation (12.8 h/day), when they spent 94% of their active time foraging, whereas during early gestation they foraged fewer hours (8.2 h/day) and less intensively (66% of active time). The longest foraging bouts also occurred in late lactation (100 min) and decreased linearly as the number of bouts per day increased (R(2)=0.88). The annual peak bite rate (BR) was in late lactation (62 bites/min), whereas the annual nadir occurred in early gestation (37 bites/min). Smallest bite sizes (BS) (mg) were recorded in late gestation (127 mg), and increased linearly with forage availability (R(2)=0.46), with largest BS occurring during peak lactation (280 mg). This study demonstrated how seasonal modifications in activity and foraging behaviour enabled gestating and lactating wapiti hinds to satisfy their changing nutritional requirements on seasonal pastures. Knowledge of behavioural compensation in response to nutritional demand and pasture conditions will be useful in designing supplementation program for farmed wapiti.     

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.     

The gestation length of wapiti (Cervus elaphus) revisited

As an ancillary activity to an artificial insemination program in farmed wapiti, the length of gestation of 28 wapiti hinds that delivered single calves of established parentage was calculated. Estrus was synchronized in 47 wapiti using progesterone impregnated devices (controlled internal drug release, CIDR) and an injection of PMSG. All hinds were artificially inseminated between 60 and 63h after CIDR removal. Pregnancy was determined between 45 and 65 days by ultrasound. A verifiable figure for gestation length was obtained based both upon timed-artificial insemination, date of parturition, and confirmation of sire identity through microsatellite DNA technology. The calculated gestational length of 247 +/- 5 days was significantly (P < 0.0001) shorter than the generally quoted figure of 255 +/- 7 days.     

Forage quality and patch choice by wapiti (Cervus elaphus)

Recent models suggest that herbivores might optimize energygain by selecting patches of intermediate vegetation biomass.We tested this hypothesis in wapiti (Cervus elaphus) by estimatingdaily rates of energy gain in relation to grass biomass andby measuring patch choice in experimental pastures in whichgrass biomass was manipulated by mowing. The digestible energycontent of grasses declined with increasing biomass due to maturationalchanges in fiber and lignin content. Daily rates of dry matterintake by wapiti increased with grass biomass at a deceleratingrate, implying a Type II functional response. Linking thesevalues to published ad libitum energy intake and energy expenditureparameters, Fryxell's (1991) model predicted that the dailyrate of energy gain should be highest when wapiti feed in grasslandswith 1000–1100 kg/ha. In trials in which grass biomasswithin a mosaic of patches was manipulated experimentally between800–2900 kg/ha, wapiti preferred patches of 1200 kg/ha,close to the value predicted by the energy gain model. Our resultssuggest that the rate of energy gain by wapiti is constrainedby both grass biomass and grass fiber content, the latter ofwhich varies inversely with grass biomass. Behavioral preferencefor grass patches of intermediate biomass and fiber contentcould help explain patterns of aggregation and seasonal migrationreported previously for wapiti.     

Microsatellite analysis of North American wapiti (Cervus elaphus) populations

Eleven populations of wapiti (Cervus elaphus) were analysed for genetic diversity using 12 microsatellite loci. Samples were taken from Vancouver Island, British Columbia; Burwash and French River herds in Ontario; Ya Ha Tinda Ranch, Alberta; and Banff, Elk Island, Jasper, Kootenay, Riding Mountain, Yellowstone and Yoho National Parks. Overall, wapiti populations have on average three to four alleles per locus and an average expected heterozygosity that ranged from 25.75 to 52.85%. The greatest genetic distances were observed between the Vancouver population and all other populations. Using the assignment test, Roosevelt wapiti (C. e. roosevelti Merriam 1897) assigned only to the Vancouver Island population. The distance and assignment values suggest a divergence of the Roosevelt wapiti from other populations and support the subspecific status for the Vancouver Island population. No evidence was found for the existence of unique Eastern wapiti (C. e. canadensis Erxleben 1777) in the Burwash or French River herds in Ontario. The overlapping distribution of genotypes from indigenous populations from Riding Mountain, Elk Island and Yellowstone National Parks suggests that wapiti were once a continuous population before settlers decimated their numbers. The lack of differentiation between these populations raises questions about the status of Manitoban (C. e.manitobensis Millais 1915) and Rocky Mountain (C. e.nelsoni Bailey 1935) subspecies.     

Induction of ovarian follicular wave emergence in wapiti (Cervus elaphus)

Two experiments were done to test the effects of treatments designed to electively induce ovarian follicular wave emergence in wapiti for the purpose of group synchronization. In Experiment 1, hinds were assigned randomly to three groups and given saline im (controls; n=5), 5mg of estadiol-17ss im (n=4), or 5mg estradiol-17ss plus 100mg progesterone im (n=5). In Experiment 2, hinds were assigned randomly to two groups and given no treatment (controls; n=6), or transvaginal ultrasound-guided follicle ablation (n=7). In both experiments, ovarian follicular dynamics were monitored by daily transrectal ultrasonography from Day 0 (day of treatment) to Day 9. In Experiment 1, blood samples were collected at each examination for measurement of serum concentrations of progesterone and FSH. Both experiments were conducted during the late anestrous period (July and August). The mean (+/-S.E.M.) day of wave emergence did not differ between the control and estradiol alone groups, but tended to be later in the estradiol plus progesterone group Day 4.0+/-0.7, Day 3.5+/-0.3, and Day 5.2+/-0.2, respectively; P=0.06). The interval from treatment to wave emergence was less variable in the estradiol plus progesterone group (P<0.05) and tended to be less variable in the estradiol-alone group (P=0.07) than in the control group. The day of wave emergence was more variable (P<0.05) and tended to be later (P=0.10) in the control group compared to the ablation group (Day 2.5+/-0.8 versus Day 1.4+/-0.2). All three treatments were effective in synchronizing ovarian follicular wave emergence among a group of wapiti hinds. Follicle ablation may be an alternative method for synchronization of follicular waves in estrus synchronization and superstimulatory protocols.     

Behavior of Elk (Cervus canadensis) during the Rut

Während zweier Monate (1965) wurden im Banff National Park (Kanada) qualitative und quantitative Daten über das Brunftverhalten des amerikanischen Wapiti (Cervus canadensis) gesammelt. Einige Verhaltensweisen sind erstmals genauer beschneben. Die ♀♀-Gruppen sowie die Zahl der mit ihnen lebenden Spießer und 21/2- und 31/2 jähngen ♂♂ wurden zu Beginn der Brunft kleiner, an ihrem Ende wieder größer. Das Zahlenverhältnis von ♀♀ und Kälbern blieb dabei gleich. Spießer blieben länger als ältere ♂♂ in den ♀♀-Gruppen und schlossen sich ihnen früher wieder an. Bei Brunftbeginn traten die über 31/2Jährigen ♂♂ zu den ♀♀-Gruppen, spalteten sie und vertrieben jüngere ♂♂. Sonogramme erläutern genauere Angaben über das Röhren und den Alarmruf, der bei ♂♂ höher ist als bei ♀♀. Durch Röhren, Geweihschlagen am Gesträuch, Graben und Suhlen scheinen andere ♂♂ aufmerksam und angelockt zu werden. Das führt zu Zweikämpfen und fördert die geschlechtliche Auslese. ♂♂ ab 31/2 Jahren können ejakulieren beim Röhren, Geweihschlagen, Graben, Stehen in der Suhle, nach dem Zusammentreiben von ♀♀ und beim Alarmrufen. Heftige Kämpfe waren selten, nur einmal kam es zu Schlägen auf den Körper. Kampfwunden scheinen unwesentlich als Todesursache. ♂♂ kämpfen bevorzugt mit Altersgenossen, ältere kämpfen mehr als junge, alle zu bestimmten Tages- und Jahreszeiten mehr als sonst. Reviere scheint es nicht zu geben. Harem-♂♂ treiben zu bestimmten Tages- und Jahreszeiten besonders häufig ♂♀ mit Kälbern zusammen. Mehr als 31/2 Jahre alte ♂♂ bestritten 94 % der heterosexuellen Kontakte, die ebenfalls tages- und jahreszeitlich verschieden häufig waren. Harem-♂♂ vertrieben am häufigsten über 31/2jährige ♂♂, selten Spießer, am seltensten 21/2jährige ♂♂. Wenn es beim Drohen blieb, war immer der Angreifer gleich alt oder älter als sein Gegner; die Häufigkeit solcher nicht zur Berührung führenden aggressiven Begegnungen änderte sich mit der Tageszeit. ♀♀-Gruppen werden offenbar von adulten ♀♀, nicht von jungen ♂♂ ?geführt”. Über 31/2jährige ♂♂ verbrauchen wohl in der Brunft Energiereserven, denn sie fressen weniger lange und sind mehr mit Laufen und anderen Bewegungen beschäftigt als jüngere ♂♂.     

Identification of dorsal-spined larvae from free-ranging wapiti (Cervus elaphus) in southwestern Manitoba, Canada

Dorsal-spined first-stage larvae recovered from feces of free-ranging wapiti (Cervus elaphus) were passaged through snails (Triodopsis multilineata) and two hand-raised white-tailed deer fawns (Odocoileus virginianus). A total of 74 adult Parelaphostrongylus tenuis were recovered from the fawns; no other protostrongylid nematodes were recovered. The study indicates that wapiti may be infected with natural infections of meningeal worm and pass larvae suitable for transmission to gastropod intermediate hosts. Wapiti from areas endemic with P. tenuis should not be translocated to areas currently free of the parasite.     

Artificial insemination of red deer (Cervus elaphus) with frozen-thawed wapiti semen

Semen collected from wapiti (Cervus elaphus) in Canada in 1983 was frozen in two extenders. In 1988, the semen was used to inseminate 200 red deer hinds on 2 farms in New Zealand. Oestrus was synchronized in the hinds with progesterone-impregnated intravaginal devices (CIDR); 200 iu pregnant mares' serum gonadotrophin was given to each hind on Day 11. The CIDRs were removed on Day 12 at 20/h, as the numbers of the hinds were recorded. On Day 14, 54-56 h after CIDR removal, the hinds were brought into the yards in the same batches and laparoscopically inseminated. Semen from three sires was used. The overall conception rate was 51%. Gestation length ranged from 239 to 247 days. One hind was lost at calving, 3 calves had to be hand raised and there were 2 neonatal calf deaths.     

Ovarian follicle development in wapiti (Cervus elaphus) during the anovulatory season

The ovaries of 12 mature wapiti hinds were studied by transrectal ultrasonography during the anovulatory season to characterize follicular dynamics and to test the hypothesis that follicle development occurs in a wave-like fashion. The hinds were examined daily, standing without sedation. Follicle size and numbers were recorded, and individual follicles were identified serially. Follicle development was considered wave-like if periodic changes in follicle numbers could be associated temporally with the development of a dominant follicle. There were non-random changes (P<0.01) in the number of follicles > or =4 mm in diameter detected per day. Each peak in follicle numbers was associated with the development of a single dominant follicle. The dominant follicle of the cohort was larger (P<0.05) than the other follicles 1 day after its emergence. Intervals between successive peaks (6.8 +/- 0.4 day) and troughs (6.8 +/- 0.4 day) in follicle numbers, and emergence of sequential dominant follicles (7.1 +/- 0.5 day) were not different (P=0.86). Results confirmed the hypothesis that ovarian follicles develop in a wave-like fashion in wapiti during the anovulatory season.     

马鹿卵母细胞体外培养与体外受精的超微结构

用FSH 对马鹿进行超数排卵,通过抽吸法和切割法采集卵泡卵母细胞,用M199 为基础的培养液在38.5℃、5%CO₂ 和饱和湿度条件下对马鹿卵母细胞进行体外培养与体外受精培养,利用透射电镜观察不同时期马鹿卵母细胞的超微结构,旨在揭示马鹿卵母细胞体外培养前、培养后及受精后超微结构的变化规律。结果表明,培养前卵丘细胞紧紧包围卵母细胞,卵母细胞表面的微绒毛细长,伸入透明带内,皮质区及细胞中心分布大量的细胞器。培养后卵丘细胞与卵母细胞结合松散,卵母细胞表面微绒毛短粗,倒伏于卵表面,第一极体无核,皮质颗粒在皮质区成层排列,细胞质中细胞器分布均匀。受精后卵母细胞表面的微绒毛由倒伏而竖起,第二极体有核,细胞质中细胞器丰富,主要分布于细胞中心。     

Das Wachstum des Geweihes von Cervus elaphus,Cervus barbarus und Cervus canadensis

Ohne Zusammenfassung     

Optimal grazing of wapiti (Cervus elaphus) on grassland: Patch and feeding station departure rules

Summary We studied factors which may shape giving-up decisions of wapiti grazing grassland patches (area where a wapiti initiates and terminates a feeding sequence) and feeding stations (area within a patch that a wapiti can reach without moving its forelegs). In grassland patches, cropping rate decreased after a critical period, whereas at feeding stations cropping rate increased with cumulative bites consumed. The number of feeding stations grazed, number of bites taken and grazing time did not dictate the termination of grazing in a patch. Wapiti gave up a patch only after the cropping rate at a feeding station dropped below the seasonal expectation during trials on lush pasture in May, but gave up after the cropping rate dropped below the seasonal expectation at two consecutive feeding stations in March/April and August when foraging conditions were less favourable. This confirmed a prediction of the marginal value theorem. Wapiti did not give up a feeding station according to bites taken, grazing time or cropping rate, but they left feeding stations when their lateral neck angle reached a critical point suggesting a biokinetic explanation. Leaving feeding stations when ungrazed forage can no longer be reached and patches when intake rate drops both appeared to be rules used by wapiti grazing grasslands of the boreal mixed wood forest.     

基于生境可获得性的完达山地区马鹿(Cervus elaphus xanthopygus)冬季生境选择

于2004～2005年冬季研究了完达山地区马鹿的生境利用和选择。在野外调查的基础上,获取了研究地区马鹿种群水平上的生境利用数据;利用地理信息系统和遥感技术,从区域尺度上测度了各类生境因子的可获得性。通过比较被利用生境与可获得生境,测度了马鹿冬季对各类生境或生态因子的选择。结果表明,马鹿对植被类型、海拔、坡向、NDVI等级、到公路距离表现出非比例利用,对坡度的利用与其可获得性则没有显著性差异。马鹿选择采伐迹地,低郁闭度生境（第2、3NDVI等级）,200～300m海拔区间,南坡,距离公路600～700m的区间。马鹿采食生境和卧息生境对各类因子具有不同的选择性,表现出不同功能生境分离的特征。对于采食生境,马鹿选择采伐迹地、阔叶疏林,低郁闭度（第2、3NDVI等级）,南坡;对于卧息生境,马鹿仅选择采伐迹地,低郁闭度（第2、3、4NDVI等级）。根据马鹿活动样点的出现与否,建立了马鹿冬季生境选择的逻辑斯谛回归模型。模型回判结果表明,该模型对马鹿活动样方与对照样方的总体正确预测率为74．4％,对马鹿活动样方的正确预测率为84．2％,对对照样方的正确预测率为62．4％,能够较好的预测生境利用概率。     

Introgression of exotic Cervus (nippon and canadensis) into red deer (Cervus elaphus) populations in Scotland and the English Lake District

Since the mid‐19th century, multiple introductions of Japanese sika deer (Cervus nippon nippon) and North American wapiti (C. canadensis) have taken place in the British Isles. While wapiti have generally been unsuccessful, sika have been very successful, especially in Scotland where they now overlap at least 40% of the range of native red deer (C. elaphus). Hybridization between these two species and red deer has been demonstrated in captivity and in the wild. Using a panel of 22 microsatellite loci that are highly diagnostic between red deer and sika, and moderately diagnostic between red deer and wapiti, we investigated the extent of introgression between these species in 2,943 deer sampled from around Scotland and from the English Lake District using the Bayesian clustering software STRUCTURE. We also used a diagnostic mitochondrial marker for red deer and sika. Our survey extends previous studies indicating little introgression of wapiti nuclear alleles into red deer, in particular in Northern Scotland, Kintyre, and the Lake District. We found a new area of extensive sika introgression in South Kintyre. In the North Highlands, we show for the first time geographically scattered evidence of past hybridization followed by extensive backcrossing, including one red‐like individual with sika introgression, two sika‐like individuals with red deer introgression, and six individuals that were apparently pure sika at the nuclear markers assessed but which carried red deer mitochondria. However, there has not been a collapse of assortative mating in this region. Similarly, in the English Lake District red deer, we found only traces of past sika introgression. No sika alleles were detected in the Central Highlands or the Hebridean red deer refugia. We make suggestions for management to prevent further spread of sika alleles into red deer and vice versa.     

基于粪便DNA的马鹿种群数量和性比

为分析黑龙江省完达山林区马鹿种群生存状态,制定科学有效地保护措施,从分子水平研究了种群数量和性比。2006、2007两年冬季跟踪马鹿足迹链,于五泡林场共搜集210份粪便,以成功提取DNA的167份作为分析样本。通过多态性较高的7个微卫星位点进行了基因分型分析,显示167份粪便DNA分属66只个体。基于非损伤性标志重捕法,统计出林场内马鹿数量2a平均47(39—60)只,密度0.302(0.251—0.386)只/km2,与2002年大样方法调查结果相比有减无增。SRY基因性别鉴定显示,种群雌雄性比1.00∶2.00(22♀,44♂),存在较多雄性个体,分析认为偷猎是导致性比失衡的最主要原因。数量的持续下降和性比失衡提示完达山林区马鹿种群数量的恢复需要更好地保护工作。