龟类生境利用的生态形态适应及两种闭壳龟的实证研究

发布时间：2017-12-27 04:23

本文关键词：龟类生境利用的生态形态适应及两种闭壳龟的实证研究　出处：《海南师范大学》2017年博士论文　论文类型：学位论文

【摘要】：动物的形态结构决定行为功能,并与其生存环境相适应。龟类占据从水生到陆生的多样化生境,其形态和行为差异较大,是生态形态学研究的理想实验对象。龟类的生态形态学特征受到多种生态因素影响,其中生态位分离是导致龟类形态变异的主要生态因素。尽管龟类的生态形态学研究较为广泛,但还存在诸如生境分类主观而简单、对同域陆生物种的研究缺乏、忽视两性异形的影响等问题,导致不能深入、有效地解释龟类形态变异的生态机理。此外,国内尚无龟类生态形态学的研究,其他动物类群的研究也仅见于蜥蜴和鸟类的零星报道。为揭示世界龟类形态变化规律以及生态成因,本文根据不同生境特征对龟类的生境进行详细划分,研究龟类的个体大小和体形与生境的相关性,并以同域分布于海南岛的平顶闭壳龟(Coura mouhotii)和黄额闭壳龟(C.galbinifrons)为实证研究对象,研究两物种形态、运动能力与微生境选择之间的关系,阐述两物种形态差异的生态学原因。研究结果分述如下:龟类个体大小和体形与生境的相关性:结合目前龟类生境分类的方法,根据不同特征将龟类的生境分为海洋、淡水、大陆性陆地和岛屿性陆地4种类型,并根据水域面积和流速将淡水生境细分为大流水水域、大静水水域、小流水水域、小静水水域和所有水域5种亚类型,根据干旱程度和地势将大陆性陆地生境细分为荒漠、平地和高地3种亚类型。我们共收集了来自文献的331种龟类的个体大小,即最大背甲长数据(占总物种数335种的98.8%),并将其归入上述4种生境类型和8种亚类型中。为消除系统发育对生境的影响,统计分析时将科作为因素之一进行分析。结果表明,龟类个体大小与其生境类型存在相关性,海龟个体最大(134.86cm),岛屿性陆龟次之(80.05cm),淡水龟(32.6cm)和大陆性陆龟(27.08 cm)个体最小,且淡水龟类与大陆性陆生龟类的个体大小不存在显著的差异。此外,淡水龟类的个体大小与5种亚类型生境有关,大水域的个体大(大静水水域=50.30 cm,大流水水域=39.86cm),小水域的个体小(小静水水域=23.78 cm,小流水水域=23.79cm)。大陆性陆龟的个体大小从高地(24.86cm)到平地(26.15cm)再到荒漠(39.11cm)有逐渐变大的趋势,但不存在显著的差异。我们共收集和测量了115种(占总物种数34.33%)6664只成体的背甲长、背甲宽和背甲高数据,并计算背甲高与背甲宽的比值(R)作为体形指标(R越大体形越高拱),将其归入上述4种生境类型和8种生境亚类型,从而研究龟类生境与体形之间的关系。由于龟类雌性的R值(0.56±0.12)显著大于雄性(0.54±0.14),存在两性异形现象(Paired t-test:t=3.367,df=86,P=0.001),因此后续对雌雄单独进行分析,以消除两性异性的影响。相关分析表明水生环境(海洋与淡水)的龟类雌雄体形均与背甲长呈显著的负相关(♀r2=0.198,F1,63=15.571,P0.0001;♂r2=0.227,F1,54=15.813,P0.0001),即体形随个体的增大而更加扁平,而雄性陆生龟类(大陆和岛屿)则相反,体形则随个体的增大而更加高拱(r2=0.132,F1,40=6.072,P=0.018)。龟类体形在4种生境类型中存在显著的差异,从海洋、淡水、大陆性陆地到岛屿陆地,其拱形程度逐渐增大(雌性R值分别为0.40、0.50、0.63和0.70;雄性的R值分别为0.37、0.47、0.63和0.71)。雌雄淡水龟类体形与5种生境亚类型无关,雌雄大陆性陆龟的体形也与其3种生境亚类型无关。同域分布的两种闭壳龟形态、运动能力和微生境利用之间的关系:于2015年4月至2016年2月在海南吊罗山国家自然保护区,采用无线电遥测技术和圈围实验研究了平顶闭壳龟和黄额闭壳龟的微生境选择和偏好,并在室内测量其运动能力(爬坡能力和翻身能力)和形态特征,探究他们微生境利用、运动能力和形态之间的关系。结果表明平顶闭壳龟背甲扁平,在野外偏好多石且较陡的微生境,多选择石缝作为其隐蔽场所;而黄额闭壳龟背甲较高拱,偏好落叶盖度和厚度较大的缓坡微生境,并选择躲藏在疏松的落叶层下。半自然条件下(圈围实验),无论两物种之间是否存在竞争,它们的微生境选择与野外遥测结果一致,表明两物种稳定地占有不同的生态位。此外,平顶闭壳龟具有较强的爬坡能力和翻身能力,与其扁平的背甲和较大的头部有关,从而能适应陡而多石的微生境;而头部较小的高拱形黄额闭壳龟的爬坡能力和翻身能力较弱,限制其选择较平缓的微生境。这些结果表明,形态和功能的差异导致这两种闭壳龟有效地利用不同的微生境,从而实现同域共存。平顶闭壳龟的洞穴选择及其形态适应:测量平顶闭壳龟在野外所选择的隐蔽场所─石缝的大小和结构特征,并在室内设计人工控制实验验证其形态与石缝大小之间的关系。结果表明,平顶闭壳龟偏好中等宽度和洞高接近自身背甲高的石缝,并把身体嵌入石缝最深处使得背甲与洞顶紧密接触,在石缝内主要采取头朝内,而使具有锯齿的背甲后缘朝外的行为方式躲藏。综上所述,龟类的个体大小和体形与其生境利用相适应,同域分布的平顶闭壳龟与黄额闭壳龟的形态差异主要是为了适应不同的微生境和特殊的隐蔽场所。解释形态变异的成因是进化生物学研究的重要基础,本研究揭示了龟类形态变异的生态成因,为动物形态的进化适应研究奠定了基础。此外,本研究的开展将引领国内在这方面的探索,也将对濒危物种的保护、管理、人工种群的建立及今后的再引入与种群恢复等工作具有理论和实践指导意义。
[Abstract]:The morphological structure of an animal determines its behavioral function and adapts to its living environment. Turtles occupy a variety of habitats from aquatic to terrestrial, with large differences in morphology and behavior, which is an ideal experimental object for the study of ecologic morphology. The ecologic morphological characteristics of turtles are influenced by various ecological factors, and niche separation is the main ecological factor that leads to the morphological variation of tortoises. Although turbidities are widely studied, there are still some problems, such as habitat classification, subjective and simple, lack of research on the same terrestrial species and neglect of the influence of sexual dimorphism. In addition, there is no study on the ecologic morphology of the turtles in China, and the study of other animal groups is only seen in the sporadic reports of lizards and birds. In order to reveal the world turtle species variation and ecological causes, this paper divided according to the different habitat characteristics of habitat for the turtles, turtles correlation between body size and shape and habitat, and in the same domain distribution in Hainan Island box turtle (Coura mouhotii) flat and Cuora galbinifrons (C.galbinifrons) as an example the object of study, to study the relationship between the two species, the exercise ability and microhabitat selection, expounds two species morphological differences in ecology. The results were as follows: the correlation between body size and shape of the turtle habitat: combining the turtle habitat classification method, according to the different characteristics of the turtle's habitat will be divided into marine, freshwater, continental islands and land of land of 4 types, and according to the water area and flow velocity will be subdivided into large freshwater habitats water water, water, water, water hydrostatic small small still waters and all the waters of 5 sub types, according to the degree of drought and the mainland land habitat of desert, plains and Highlands is subdivided into 3 sub types. We collected from the individual size of 331 species of turtles, namely the maximum carapace length data (335 of the total species number 98.8%), and will be classified as the 4 habitat types and 8 sub types. In order to eliminate the impact of phylogeny on habitat, statistical analysis is used as one of the factors to be analyzed. The results show that the correlation between the size of individual turtles and habitat types, the largest individual turtle (134.86cm), the island of the tortoise (80.05cm), freshwater turtle (32.6cm) and continental tortoises (27.08 cm) and the smallest individual, freshwater turtles and continental terrestrial turtles a body size is not obvious the difference. In addition, the individual size of freshwater turtles is related to 5 subtypes of habitats, large individuals in large waters (large hydrostatic waters, =50.30 cm, large water =39.86cm), small individuals in small waters (small still waters =23.78 cm, and small water flowing waters =23.79cm). The size of individual continental tortoises from the highlands (24.86cm) to ground (26.15cm) to the desert (39.11cm) was gradually decreased, but there is no significant difference. We collected and measured 115 (total 34.33% species) of 6664 adult carapace length, carapace width and carapace height data, and calculate the carapace ratio and high dorsal carapace width (R) as the shape index (R is larger, the shape is high) in the 4 habitat types and 8 sub types of habitats, so as to study the relationship between turtle habitat and shape. Because the R value of female turtles is (0.56 + 0.12), which is significantly larger than that of males (0.54 + 0.14), there exists sexual dimorphism (Paired t-test:t=3.367, df=86, P=0.001). Therefore, the analysis of male and female separately is conducted to eliminate the influence of sexual heterosexual. The correlation analysis showed that the aquatic environment (Marine and freshwater turtles) male and female body shape and carapace length were negatively correlated (F1,63=15.571, female r2=0.198, male r2=0.227, F1,54=15.813, P0.0001; P0.0001), the body with individual increases more flat, while the male Lu Shenggui class (mainland and island) in shape with the increase of individual higher (r2=0.132, F1,40=6.072, P=0.018). The shape of turtles is significantly different in 4 habitats. From the ocean, fresh water, continental land to island land, the vaulting degree increases gradually (the female R values are 0.40, 0.50, 0.63 and 0.70 respectively, and the R values of males are 0.37, 0.47, 0.63 and 0.71). Regardless of male and female freshwater turtle shape and 5 kinds of habitat sub types, regardless of male and female continental tortoise shape is its 3 subtypes of habitat. The relationship between the distribution of the two domain of Cuora morphology, motility and microhabitat use: hanging in Luoshan National Nature Reserve in Hainan from April 2015 to February 2016, using radio telemetry and circle of flat box turtle Cuora galbinifrons and the surrounding microhabitat selection and preference, and measure its ability in sports (indoor climbing ability and turning ability) and morphological features, explore the relationship between their microhabitat use, between exercise capacity and morphology. The results show that the flat box turtle carapace flat in the wild, a lot of stone and partial steep microhabitats, choose more stones as its hidden places; and a high amount of yellow box turtle, coverage and thickness of the gentle slope of microhabitat preference leaves, and choose to hide in the leaves under the loose layer. Under semi natural conditions (circle experiment), no matter whether there is competition among two species, their microhabitat selection is consistent with the field telemetry results, indicating that two species occupy different niche niches. In addition, the flat box turtle has a strong climbing ability and turning ability, with flat carapace and large head, which can adapt to the steep and rocky microhabitats; and the head of small high arch C. galbinifrons climbing ability and turning ability is weak, limiting the choice of gentlier microhabitat. These results indicate that the morphological and functional differences lead to these two kinds of Cuora effectively use different microhabitats, so as to realize the coexistence. To select and shape of flat box turtle Cave: measuring flat Cuora selected in the wild
【学位授予单位】：海南师范大学
【学位级别】：博士
【学位授予年份】：2017
【分类号】：Q958.8

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