四川梅花鹿产仔换茸期声行为的研究

2009年4～9月在四川省若尔盖县铁布自然保护区,对四川梅花鹿Cervusnippon sichuanicus产仔换茸期的声行为进行了较系统的研究。四川梅花鹿产仔换茸期的声行为可以分为报警声、惊叫声、吼叫声、母子召唤声等20种鸣声。通过声谱分析获得了各种鸣声的语谱图及其频谱特征,并通过野外观察初步确定了这些声行为的生物学意义。在遇到危险或潜在危险时,成年雄鹿往往第一个发出报警鸣声,而雌鹿的报警声大约是雄鹿的两倍多。成年个体的主动防御鸣声频次要多于亚成体,而幼体未见有该类鸣声。对梅花鹿不同亚种和不同季节的几种声行为进行了比较。     

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。     

Female Sexual Preferences Toward Conspecific and Hybrid Male Mating Calls in Two Species of Polygynous Deer, <Emphasis Type="Italic">Cervus elaphu</Emphasis>s and <Emphasis Type="Italic">C. nippon</Emphasis>

The behavioral processes at the basis of hybridization and introgression are understudied in terrestrial mammals. We use a unique model to test the role of sexual signals as a reproductive barrier to introgression by investigating behavioral responses to male sexual calls in estrous females of two naturally allopatric but reproductively compatible deer species, red deer and sika deer. Previous studies demonstrated asymmetries in acoustic species discrimination between these species: most but not all female red deer prefer conspecific over sika deer male calls while female sika deer exhibit no preference differences. Here, we extend this examination of acoustic species discrimination to the role of male sexual calls in introgression between parent species and hybrids. Using two-speaker playback experiments, we compared the preference responses of estrous female red and sika deer to male sexual calls from conspecifics versus red × sika hybrids. These playbacks simulate early secondary contact between previously allopatric species after hybridization has occurred. Based on previous conspecific versus heterospecific playbacks, we predicted that most female red deer would prefer conspecific calls while female sika deer would show no difference in their preference behaviors toward conspecific and hybrid calls. However, results show that previous asymmetries did not persist as neither species exhibited more preferences for conspecific over hybrid calls. Thus, vocal behavior is not likely to deter introgression between these species during the early stages of sympatry. On a wider scale, weak discrimination against hybrid sexual signals could substantially contribute to this important evolutionary process in mammals and other taxa.     

Possible use of heterospecific food-associated calls of macaques by sika deer for foraging efficiency

Heterospecific communication signals sometimes convey relevant information for animal survival. For example, animals use or eavesdrop on heterospecific alarm calls concerning common predators. Indeed, most observations have been reported regarding anti-predator strategies. Use of heterospecific signals has rarely been observed as part of a foraging strategy. Here, I report empirical evidence, collected using playback experiments, showing that Japanese sika deer, Cevus nippon, use heterospecific food calls of Japanese macaques, Macaca fuscata yakui, for foraging efficiency. The deer and macaques both inhabit the wild forest of Yakushima Island with high population densities and share many food items. Anecdotal observations suggest that deer often wait to browse fruit falls under the tree where a macaque group is foraging. Furthermore, macaques frequently produce food calls during their foraging. If deer effectively obtain fruit from the leftovers of macaques, browsing fruit fall would provide a potential benefit to the deer, and, further, deer are likely to associate macaque food calls with feeding activity. The results showed that playback of macaque food calls under trees gathered significantly more deer than silence control periods. These results suggest that deer can associate macaque food calls with foraging activities and use heterospecific calls for foraging efficiency.     

Context-related acoustic variation in male fallow deer (Dama dama) groans

While social and behavioural contexts are known to affect the acoustic structure of vocal signals in several mammal species, few studies have investigated context-related acoustic variation during inter-sexual advertisement and/or intra-sexual competition. Here we recorded male fallow deer groans during the breeding season and investigated how key acoustic parameters (fundamental frequency and formant frequencies) vary as a function of the social context in which they are produced. We found that in the presence of females, male fallow deer produced groans with higher mean fundamental frequency when vocal males were also present than they did when no vocal males were in close vicinity. We attribute this to the increased arousal state typically associated with this context. In addition, groan minimum formant frequency spacing was slightly, but significantly lower (indicating marginally more extended vocal tracts) when males were alone than when potential mates and/or competitors were nearby. This indicates that, contrary to our predictions, male fallow deer do not exaggerate the acoustic impression of their body size by further lowering their formant frequencies in the presence of potential mating partners and competitors. Furthermore, since the magnitude of the variation in groan minimum formant frequency spacing remains small compared to documented inter-individual differences, our findings are consistent with the hypothesis that formants are reliable static cues to body size during intra- and inter-sexual advertisement that do not concurrently encode dynamic motivation-related information.     

四川梅花鹿的行为谱及PAE编码系统

2007年3-11月每月定期5 d,每天06:00-18:00在四川省铁布自然保护区的冻列乡、崇尔乡、热尔乡观察记录四川梅花鹿行为发生过程、内容、结果及其周围生境.建立了四川梅花鹿的行为谱、警戒行为谱及其PAE编码系统,填补了有蹄类警戒行为PAE编码的空白.分辨并记录了四川梅花鹿的11种姿势、83种动作及136种行为,描述了各种行为的相对发生频次与性别、年龄、季节的关系.与其他有蹄类相比,梅花鹿的行为与鹿科其他动物有相似之处.     

日本日光地区森林中梅花鹿的夏季巢区面积和内部利用

为了估算梅花鹿(Cervusnippon)巢区面积和内部利用以及雌雄个体间的比较,于1993-1997年在日本国日光国立公园使用无线电遥测定位法测定了8头成年雌鹿和4头雄鹿(包括1头亚成年)在森林生境中的活动定位点。按照当地梅花鹿离开和返回越夏地的日期,使用每年6月1日至10月25日的无线电遥测定位数据,采用定值核心法(Fixed kernel)计算了梅花鹿的夏季巢区面积和内部利用密度分布。梅花鹿雄鹿夏季巢区面积(192·5±50·5 hm2)显著大于雌鹿(115·7±56·7 hm2,P=0·017) ,其半家区面积(HRS0·50)和核心区面积(HRS0·25)也显著大于雌鹿(依次为P=0·027和P=0·042) ,但巢区内部利用集中度指标Ah(HRS0·50/HRS0·95)和Ac(HRS0·25/HRS0·50)在雌雄鹿之间没有显著性差异(依次为P=0·999和P=0·234)。在无线电遥测的12头梅花鹿中, 11头个体的巢区为单核心型,只有1头成年雄鹿的巢区为双核心型。11头梅花鹿个体的混合数据显示巢区面积和体重之间存在显著性相关(P=0·049) [动物学报52 (2) : 235 -241 , 2006]。     

Variability of Female Responses to Conspecific vs. Heterospecific Male Mating Calls in Polygynous Deer: An Open Door to Hybridization?

Males of all polygynous deer species (Cervinae) give conspicuous calls during the reproductive season. The extreme interspecific diversity that characterizes these vocalizations suggests that they play a strong role in species discrimination. However, interbreeding between several species of Cervinae indicates permeable interspecific reproductive barriers. This study examines the contribution of vocal behavior to female species discrimination and mating preferences in two closely related polygynous deer species known to hybridize in the wild after introductions. Specifically, we investigate the reaction of estrous female red deer (Cervus elaphus) to playbacks of red deer vs. sika deer (Cervus nippon) male mating calls, with the prediction that females will prefer conspecific calls. While on average female red deer preferred male red deer roars, two out of twenty females spent more time in close proximity to the speaker broadcasting male sika deer moans. We suggest that this absence of strict vocal preference for species-specific mating calls may contribute to the permeability of pre-zygotic reproductive barriers observed between these species. Our results also highlight the importance of examining inter-individual variation when studying the role of female preferences in species discrimination and intraspecific mate selection.     

Mule deer (Odocoileus hemionus) respond to yellow‐bellied marmot (Marmota flaviventris) alarm calls

Individuals may obtain valuable information about the presence of predators by listening to heterospecific alarm signals. Most playback studies have demonstrated that similarly sized and taxonomically related species may respond to the calls of each other, but less work has been carried out to define these factors influence responsiveness to alarm signals. In theory, individuals should respond to calls from any species that provide information about the presence of important predators, regardless of body size or taxonomic relationship. However, size is often associated with vulnerability. Coyotes (Canis latrans) in the Rocky Mountains prey upon both mule deer (Odocoileus hemionus) and yellow‐bellied marmots (Marmota flaviventris), which differ considerably in size, alarm vocalizations, and antipredator behavior. We conducted a playback experiment to see whether deer discriminated between marmot alarm calls and the non‐alarm song of a common sympatric bird. We found that deer increased vigilance significantly more after hearing broadcast marmot alarm calls compared with the bird song. Interestingly, deer that were studied within 0.5 km of homes showed significantly greater discrimination than those studied farther from humans. Our results suggest relative size differences do not prevent interspecific communication and that common predators should generally drive the evolution of the ability to learn to respond to meaningful risk cues. As long as two species share a predator, it should benefit the other to respond to its alarm calls.     

四川梅花鹿春季昼夜活动节律与时间分配

1997年4月,在四川铁布自然保护区用直接观察法对220头次四川梅花鹿群的昼夜活动节律和时间分配进行了观察。结果表明,鹿群春季昼夜活动的规律性较强。白昼,鹿群活动呈现明显的双峰型,2个高峰时段为08：30和19：30前后,活动频率分别为71．9％和94．13％,其他时间多处于休息状态;鹿群夜间活动强于昼间,活动频率均在6l％以上,仅在半夜01：30前后有一个相对不活跃期。同时发现,春季活动高峰期约90％的活动时间被四川梅花鹿用于采食和移动。这可能与光照、食物及人类活动的影响有关。     

四川梅花鹿繁殖行为

2006年4—12月和2007年3—11月在四川省铁布自然保护区观察和统计了野生梅花鹿的繁殖行为,包括发情交配、产仔、发情吼叫、爬跨及其昼夜节律行为等。结果表明,四川梅花鹿为季节性发情动物,发情交配行为发生在9月上旬至12月中旬,集中在10—11月(占(86.99±3.24)%)。四川梅花鹿发情交配日期最早见于9月8日,最晚为12月16日,跨度约90—100 d(±6 d,n=90)。雌鹿交配日期与其繁殖经历具有低度正相关性(Kendall's tau-b和Spearman's rho,0.3r0.5,P0.05),成体雌鹿交配日期稍微早于初次配种雌鹿。雄鹿发情吼叫和爬跨行为具有明显的昼夜节律性,各有2个高峰期(05:00—08:00和18:00—21:00),夜间有小节律的发情吼叫和爬跨时期。U-test检验表明发情吼叫频次和爬跨频率在昼夜间有极显著差异(P0.01)。雄鹿吼叫行为与其交配行为具有高度正相关性(Kendall's tau-b和Spearman's rho,0.8r1.0,P0.05),主雄、次雄和群外单身雄鹿的昼夜吼叫次数有极显著差异(P0.01)。雌鹿产仔期从4月下旬开始到7月下旬结束,集中在5—6月(占(91.51±4.96)%),产仔日期最早见于4月29日,最晚为7月28日,跨度约80—90 d(±5 d,n=130)。梅花鹿产仔日期与其分娩经历具有低度正相关性(Kendall's tau-b和Spearman's rho,0.3r0.5,P0.05),成体雌鹿产仔日期早于初次繁殖雌鹿。雌鹿每胎产1—2个幼仔,单双胎率分别为98.86%(±6.96%,n=129)和1.01%(±0.07%,n=1)。妊娠期和哺乳期梅花鹿采食行为分配占较大比率,其次是卧息和移动,哺乳期采食行为分配低于妊娠期,这与妊娠期正逢冬季,食物资源相对匮乏有关,而哺乳期恰逢夏季,植物生长旺盛,食物资源相对丰富。     

Sexual variations in food quality and gastrointestinal features of sika deer (Cervus nippon) in Japan during winter: implications for feeding strategy

We assessed sexual variation in food quality and gut macrostructure in adult male and pregnant female sika deer, Cervus nippon (Temminck, 1838), in Japan during winter. These variations might have important implications relative to sexual differences in habitat use, forage acquisition, and digestive strategy. According to the sexual dimorphism-body size hypothesis the larger males would feed on poorer forage and have heavier stomach contents and heavier intestine contents and longer intestines than smaller females. However, the food quality in rumen contents of males was higher than, or at least similar with, that of pregnant females. In correspondence to food quality, the relative weights of stomach contents and intestines with contents, the relative lengths of intestines to the lengths of body and total intestines in pregnant females were similar to adult males. The relative weights of omasum and abomasum tissues in pregnant females were greater than in males. Our findings suggest sexual differences in feeding strategy in sika deer in Japan during winter. To meet greater nutritional demands of high metabolic rate and gestation, pregnant females seemed to maintain a greater volume of digesta in guts and had more stomach tissues than expected by the sexual dimorphism-body size hypothesis to compensate for poorer forage quality.     

非人灵长类动物的警报叫声

警报叫声是非人灵长类动物社会交往中的一种重要的通讯行为。本文从警报叫声系统及其作用、发展和影响因素入手, 综述了相关研究。根据叫声包含的信息, 警报叫声可以归纳为功能性参照系统和紧迫性参照系统。警报叫声使灵长类动物能够更好地躲避危险, 增加存活机率。各物种产生、理解和使用警报叫声的能力有其发展过程。天敌的捕食方式、个体差异和学习, 在不同层次上影响着警报叫声。川金丝猴警报叫声的初步研究发现不同性别、年龄个体的警报声所体现的危险程度不同。对更多物种进行更深入的研究将有助于了解非人灵长类警报叫声的全貌。     

Singing in the Face of Danger: the Anomalous Type II Vocalization of the Splendid Fairy-Wren

Males of certain species of fairy-wrens (Aves: Maluridae) emit a unique vocalization, the Type II vocalization, in response to the calls of potential predators. We conducted field observations and playback experiments to identify the contexts in which the Type II vocalization is emitted by splendid fairy-wren ( Malurus splendens ) males, and to examine social and genetic factors that influence its occurrence. In field observations and controlled playback experiments, Type II vocalizations were elicited most consistently by calls of the predatory gray butcherbird ( Cracticus torquatus ). Some vocalizations from other avian species also elicited Type II vocalizations, and the majority of these were vocalizations from avian predators. Splendid fairy-wrens are cooperative breeders, and males that responded with Type II vocalizations to playbacks of butcherbird calls tended to be primary rather than secondary males, had larger cloacal protuberances, and were older than those that did not respond. In addition, secondary males that were sons of resident females were more likely than non-sons to respond with a Type II vocalization. In another playback experiment, females responded similarly to the Type I song and Type II vocalizations of their mates. Although the Type II vocalization is emitted primarily in response to predator calls, it is inconsistent with an alarm call explanation. Patterns of reproductive success among Type II calling males suggest that it does not function as an honest signal of male quality. At present, the function of the vocalization remains anomalous, but indirect fitness benefits may play a role in its explanation.     

Interference competition and group size effect in sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) at salt licks

Competition has long been considered as a confounding factor of group size effect but the understanding of interference competition is rudimentary for the difficulty in disentangling interference competition from scramble competition adequately. Here, we analysed remote-camera video records of wild sika deer (Cervus nippon) at salt licks in southern China from March 1, 2006 to November 30, 2008 to investigate how interference competition and predation risk interacted on vigilance behaviour. Scramble competition is negligible at salt licks; therefore, we could focus our interest in interference competition. We used linear mixed model to compare vigilance, licking and aggression behaviours among females with and without fawn as well as males with different group sizes to identify the primary role of vigilance behaviour in sika deer. In total, 168 individuals were recorded and observation time was 2,733.04 min. We found that deer spent much time on vigilance and scanned frequently in spring and winter, and females with fawn spent more time on vigilance than females without fawn, suggesting vigilance for predation risk. Aggression ratio increased first and then decreased, while scan frequency continued to decline and then slightly increased when group size increased from two to seven, implying vigilance for interference competition. Our results suggested vigilance in sika deer was influenced by both predation risk and interference competition, but was mainly driven by predation risk even at sites with intense interference competition. Our results of interference competition shed some light on finding the underlying mechanism of group size effect in wild populations.     

四川山鹧鸪繁殖期的鸣声行为特征

2005年4—12月在四川老君山自然保护区,通过计算机声谱分析技术,对四川山鹧鸪(Arborophilarufipectus)雄性的鸣声行为特征进行了研究,其结果表明,雄性单音节鸣声出现在啼叫、保护领域和求偶中,这些鸣声持续时间差异明显,主峰值差异不明显;雄性双音节鸣声出现在啼叫、保护领域、竞争雌体和求偶中,这些鸣声第一音节时间、第二音节时间、音节间隔时间和主峰值差异极显著,全句时间差异不明显;雄性多音节鸣声出现在保护领域、警戒和惊吓中。雄性亚成体多音节鸣声出现在惊吓和警戒中;雄性成体和亚成体警戒鸣叫的音节持续时间、音节间隔时间和主峰值差异均不明显,全句持续时间差异明显;雄性亚成体和雄性成体的惊吓鸣叫音节持续时间、音节间隔时间、全句持续时间和主峰值差异均不明显。四川山鹧鸪表现出的各种鸣声行为是为了保护配偶和防止天敌,使该种群更好地繁衍。     

Female call preferences in tree-hole frogs: why are there so many unattractive males?

In tree-hole frogs, Metaphrynella sundana, the fundamental call frequency varies widely between males. In field playback experiments, females strongly preferred calls from the lower range of frequencies found in the population. There was no correlation, however, between male size and call frequency, as is normally the case for anurans, so large males were not necessarily more attractive to females. Presence or absence of upper harmonics in the call had no effect on female choice. Tree holes with shallow air columns were more often used by calling frogs, and were presumably more common, than deep holes. Since male M. sundana actively exploit the resonant properties of tree holes for mate attraction, and high frequencies match comparatively shallow holes, the benefits of attaining acoustic matching probably select for high-frequency calls. In addition, males with high-frequency calls may be heard from a greater distance in the vicinity of torrent streams. Since the level of such noise in the forest varies in time and space, different frequencies may prove optimal in different contexts, thereby preserving the observed variation within the population. Having an ‘unattractive’ high-frequency call should be potentially beneficial only when calling males do not congregate, a condition that our data suggest is fulfilled in this system.     

一种鹿科动物标本的分子鉴定

中国是世界上鹿科动物资源最丰富的国家之一,共有18种,分属獐亚科、麂亚科、鹿亚科和美洲鹿亚科,占全球鹿种的41.7％(盛和林等,1992).在皖南分布的鹿科动物有梅花鹿华南亚种(Cervus nippon pseudaxis)、黑麂(Muntiacus crinifrons)、黄麂(Muntiacus reevesi)和毛冠鹿(Elaphadus cephalophus)(王岐山,1990).2001年11月从泾县获得一具标本(图1-A,标本现保存于安徽师范大学标本馆),体形及大小似梅花鹿,身体背部毛棕褐色,无明显白斑,尾背面黑色、腹面纯白,但鹿角主干表面光滑,无眉叉及其它分叉.鹿角是鹿科动物所特有的结构,是区别于其它有蹄类的标志之一.     

Anti-Predator Signals as Advertisements: Evidence in White-Throated Magpie-Jays

Calls and displays elicited by predators usually function as alarms or to inform predators of their detection. However, predator encounters may afford some individuals the opportunity to demonstrate quality or signal their availability. Here, I report on a class of vocal signals produced in predator-elicited displays that share many characteristics with sexually selected song. White-throated magpie-jays ( Calocitta formosa ) display at low-threat predators while producing 'loud display calls' (LDCs). I use this term because the calls occur primarily in two display contexts (see below) though occasionally in other contexts as well. Such calls and displays are primarily produced by males, and also occur in one other context, at dawn. Playback experiments showed that despite being elicited by predators, males were more likely than females to respond to LDCs, and more likely to respond when their mate was fertile. Over 134 different call types were produced in over 200 displays by 34 males; the largest minimum repertoire size was 67. Presentations of taxidermic raptor mounts elicited some LDCs, but fewer calls and lower diversity than at dawn or in predator approach displays. The male bias and high diversity suggest that LDCs are an outcome of intersexual selection, while their elicitation by predators suggests an alarm function. I propose that male magpie-jays use predator encounters as opportunities to advertise their presence and availability as mates; they use LDCs as songs. Such a communication system seems to have been favored by the unusual social system of magpie-jays, in which female groups defend territories and males have little opportunity to defend resources for mate attraction, forcing them to advertise when females are paying the most attention, during predator encounters.     

梅花鹿的微卫星多态性及种群的遗传结构

梅花鹿是我国极度濒危的鹿科动物,其野生种群已濒临灭绝。为了探讨我国野生梅花鹿的保护和管理对策,我们选用了16 个微卫星位点检测来自东北、四川、江西和浙江种群的122 份样品,以此分析我国野生梅花鹿种群的遗传多样性和遗传结构。费希尔确切性检验表明,四个种群中均存在偏离哈迪- 温伯格平衡的现象。与其它的濒危动物相比,中国梅花鹿有着相对较高的遗传多样性:每个位点的平均等位基因数为4 个,平均期望杂合度为0. 559。四个种群中,东北种群拥有最高的平均等位基因数(n = 3. 688),东北种群、四川种群、江西种群以及浙江种群的平均期望杂合度分别为0. 584,0. 477,0. 585 和0.589,它们之间不存在显著的差异。同时,利用逐步突变模型、双相突变模型和无限等位基因突变模型检测了种群的瓶颈效应,结果表明:除四川种群外,其他种群在近期内都经历过遗传瓶颈。费希尔确切性检验及配对样品F ST 的结果均表明:四个梅花鹿种群间存在显著的遗传分化(P < 0. 001)。因此,我们建议将我国梅花鹿的野生种群划分为4 个管理单元进行保护和
管理。