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    Please use this identifier to cite or link to this item: http://140.128.103.80:8080/handle/310901/25133


    Title: 領域性腹斑蛙生殖行為及聲音溝通之研究
    Other Titles: The reproductive behaviour and vocal communication in a territorial frog, Rana adenopleura
    Authors: 莊銘豐
    Ming-Feng Chuang
    Contributors: 關永才
    Yeong-Choy Kam
    生命科學系
    Keywords: 
    none
    Date: 2014
    Issue Date: 2015-03-06T08:29:46Z (UTC)
    Abstract: 性擇是偏好能促進繁殖成功的表型及行為特徵的一股重要演化力量。在資源捍衛的交配系統中,雄性競爭有限的生殖資源來促進生殖成功,因此雄性在資源可獲得性低及強烈雄性競爭下會採取有利的策略,包括降低求偶時沒有必要的衝突消耗,並最大化其生殖利益。我在蓮華池觀察領域性的腹斑蛙,分析其宣告叫聲中的變異性及獨特性,並研究其領域行為、社交辨識,以及繁殖行為。研究分成三個部分。在研究一,我分析腹斑蛙宣告叫聲的個體獨特性,檢視雄蛙是否可透過聲音來辨識鄰居或陌生人(親愛敵人現象)。這種行為可避免與建立領域的鄰居發生不必要的肢體衝突。來自45隻雄蛙的60個因子在個體之間都有顯著差異,其中第二諧音/主要音頻相對強度提供了高達94.2%統計上的個體獨特性,推測是可靠的聲音訊息。野外回播實驗中,證實了雄蛙有鄰居-陌生人辨識能力,也確定反應門檻方法在親愛敵人辨識系統中的可行性。在研究二,我以腹斑蛙來探究在資源捍衛系統中,雌性是否會以雄蛙的鳴叫特徵來挑選品質好的配偶。實驗中記錄了45隻雄蛙的899個錄音片段並測量其中的60個鳴叫因子。聲音分析的結果發現頻譜因子比起時態因子有較少的個體內的變異。第二諧音是雄蛙吻肛長及身體狀況的良好指標,且聲音低沈的雄蛙有較高的卵團受精率。然而,雄蛙交配成功次數與代表身體品質的第二諧音無關,顯示雌蛙不會透過鳴叫聲選擇好品質配偶及提高受精率;雌蛙偏好較長的音節長度,推測是易被察覺有提供雄蛙位置的功能。而本種宣告叫聲對交配次數的解釋不高,推測可能雌蛙選擇領域特徵而非鳴叫特徵;又或者是雄蛙彼此高度競爭,使雌性選擇受到限制。在研究三,我監測一個腹斑蛙族群的抱接時間,以檢測在體外受精物種中影響交配時間的可能原因。從72隻雄蛙觀察到了122次完整的交配過程,發現腹斑蛙的抱接時間相較於種類是較短的,平均少於11分鐘,包括約8分鐘的產卵前抱接以及約3分鐘的產卵時間。總抱接時間大致上是受到產卵前抱接時間的影響,而產卵時間是變異性比較低的。抱接時間正相關於窩卵數以及性比偏雄程度。抱接中的雄蛙與沒有配對的入侵雄蛙之間的衝突互動是頻繁的,這些互動往往造成交配失敗、抱接時間延長,以及受精率的降低。實驗結果顯示本種短促的抱接時間或許是一種適應性的繁殖策略,使雄性可以降低因為雄性競爭所帶來的交配干擾、受精失敗及領域喪失的風險。總而言之,本系列研究提供資源捍衛系統物種雄性競爭及吸引雌性上可能所採取的策略:在捍衛領域時會辨識鄰居以減少與威脅性較低的競爭者之間不必要的衝突互動,在吸引雌性時雄蛙會以明顯的聲音特徵來使雌蛙容易發現鳴叫者的位置,並在抱接時精簡所需的時間來降低受到干擾的風險。
    Sexual selection is an important evolutionary force to promote those morphological and behavioural characteristics favored by reproductive success. In resource defence mating system, males compete the limited breeding resources to enhance their reproductive success. Therefore, males should adopt strategies that are beneficial under the circumstance of low resource availability and high male-male competition, such as reducing energy loss from unnecessary aggressive interactions when attracting females, or to optimize their breeding benefits. I observed a population of territorial frog, Rana adenopleura, analyzed the variability and distinctiveness of their male advertisement call, and evaluated their behaviours of territoriality, social recognition, and reproduction. I divided this research into three parts. In study one, I investigated whether male R. adenopleura acoustically distinguish their neighbors from strangers (i.e. dear enemy phenomenon). This behaviour is thought to prevent the costly and unnecessary physical interactions with their neighbors that have established territories and to respond threats by individuals that are searching for available territories. The results of sound analysis from 45 males showed that, all the 60 call properties were significantly different among males, and the relative amplitude of second harmonic/dominant frequency, suggesting a reliable recognition signal, which explained statistically up to 94.2% of the individual distinctiveness. In the field playback experiment, I adopted response threshold instead of aggressive index to determine the dear enemy phenomenon. The warning response thresholds by resident males to stimuli from strangers were lower than those to stimuli from familiar neighbors, revealing the existence of dear phenomenon in R. adenopleura and confirming the application of response threshold to access dear enemy recognition system. In study two, I evaluated whether the females of resource defending species, R. adenopleura, selected high quality males by vocal characteristics. I recorded 899 calls and measured 60 call properties from 45 males. Results showed that spectral properties had small within-individual variability and temporal properties had obvious within-individual variability. Second harmonic is an indicator of snout-vent length and body condition. Males having low second harmonic correlated to high egg fertilization. However, male mating success was not correlated to second harmonic which represents male quality, indicating that females did not select males acoustically. Females preferred longer note durations was suggested function in locating calling males. The degree of mating frequency explained by advertisement call was low, suggesting females select males by traits of territory, or female choice is limited under high male-male competition. In study three, I monitored breeding activity in a population to identify factors that affect the duration of amplexus. Compared with most other frogs, the amplexus of R. adenopleura was short and lasted less than 11 min on average, which included about 8 min of pre-oviposition activity followed by 3 min of oviposition. I evaluated the relationship between amplexus duration and seven variables: male body size, male condition, operational sex ratio (OSR), population size, clutch size, territory size, and the coverage of submerged vegetation in a male’s territory. I also investigated the influence of these same variables, along with amplexus duration, on fertilization rate. Amplexus duration was positively related with clutch size and the degree of male-bias in the nightly OSR. Fertilization rate was directly related to male body size and inversely related to amplexus duration. Agonistic interactions between males in amplexus and intruding, unpaired males were frequent. These interactions often resulted in mating failure, prolonged amplexus duration, and reduced fertilization rates. Together, our findings suggest that short amplexus duration may be an adaptive reproductive strategy, and whereby males attempt to reduce the risks of mating and fertilization failures and territory loss resulting from heavy male-male competition.
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