若尔盖湿地3种两栖类的种间竞争及畜牧业对其食性的影响

从空间、时间、食物3个方面研究了若尔盖湿地3种两栖类的种间竞争，运用生态位理论探讨了3种两栖类利用环境资源的能力以及它们之间的共存模式，研究了3种两栖类年周期食性，并考察了畜牧业对3种两栖类食性及食物竞争格局的影响；此外，还通过实验室研究对2种两栖类幼体的种间竞争策略进行了考察。主要结果如下： 1、两栖类的空间资源利用状况：在3种两栖类成体生态位宽度的比较中，岷山蟾蜍（Bufo minshanicus）成体在牧场性质(0.41)、牛粪数量(0.42)、植被盖度(0.20)、地表温度(0.50)、地表湿度(0.51) 5个维度上的生态位宽度最窄；而倭蛙（Nanorana pleskei）成体在牛粪数量(0.81)、植被高度(0.63)、植被盖度(0.47)、小水体距离(0.68) 4个维度上的生态位宽度最宽。在3种两栖类亚成体生态位宽度的比较中，岷山蟾蜍亚成体在牧场性质(0.66)、牛粪数量(0.58)、植被高度(0.64)、小水体距离(0.51)、地表湿度(0.79) 5个维度的上生态位宽度最宽；倭蛙亚成体在牧场性质(0.39)、牛粪数量(0.30)、地表温度(0.18)、地表湿度(0.33) 4个维度上的生态位宽度最窄。高原林蛙（Rana kukunoris）在地表温度（成体：0.62；亚成体：0.56）、地表湿度（成体：0.84；亚成体：0.60）两个维度上具有较大的生态位宽度值，而在小水体距离维度上（成体：0.27；亚成体：0.14）的生态位宽度值则很小。比较3种无尾两栖类在不同生长阶段（成体、亚成体）的生态位宽度，发现高原林蛙和倭蛙的亚成体对栖息环境的要求更高。3种两栖类空间资源利用的相似程度很高，高原林蛙与倭蛙之间的生态重叠度(0.87)较之它与岷山蟾蜍(0.81)的生态位重叠度更大。 2、两栖类的日活动节律：高原林蛙成体、亚成体、岷山蟾蜍亚成体活动的最低气温为0℃、2℃、8℃；岷山蟾蜍和高原林蛙亚成体出现的数量与气温成极显著的正相关（r=0.797, p<0.001；r=0.794, p<0.001），高原林蛙成体出现的数量与气温有一定相关性（r=0.456, p<0.05）；晴天时两栖类的活动性明显高于阴天（p<0.001）；多云转晴天气，高原林蛙和岷山蟾蜍亚成体出现两次日活动高峰，分别为中午12:30左右和下午15:30～16:30之间；多云天气，高原林蛙和岷山蟾蜍亚成体出现两次日活动高峰，分别为9:30～10:30之间和15:30～16:30之间。 3、两栖类的食物资源利用状况：春、秋两季，高原林蛙最主要的食物是蜉金龟科(Aphodiidae)昆虫，相对重要性指数(IRI)最高（春季：35.28%，秋季：28.57%），其次为昆虫的幼虫，以及双翅目的毛蚊科(Bibionidae)、蝇科(Muscidae)、丽蝇科(Calliphoridae)昆虫，秋季，蝗虫是高原林蛙食物组成中的重要部分；岷山蟾蜍最主要的食物是蚂蚁(IRI，春季：85.54%，秋季：49.70%)，其次为蜉金龟科、象甲科(Curculionidae)、步甲科(Carabidae)、粪金龟科(Geotrupidae) 等鞘翅目昆虫；倭蛙春季的最主要食物也是蜉金龟科昆虫(IRI，春季：13.41%)，其次为蚂蚁、毛蚊科昆虫、昆虫的幼虫以及狼蛛科(Lycosidae)。3种两栖类中，倭蛙的食性生态位宽度相对较宽（0.43），而岷山蟾蜍（0.09）和高原林蛙（0.22）的生态位宽度较窄，与春季相比，两栖类在秋季的食谱更宽。以利用食物种类为标准，春季高原林蛙与倭蛙的生态位重叠度(0.40)比它与岷山蟾蜍的生态位重叠度（0.33）更大。 4、畜牧业对两栖类食性及食物竞争格局的影响：以藏牦牛粪为食物或寄居场所的昆虫，如蜉金龟科、粪金龟科、毛蚊科、蝇科、丽蝇科昆虫和某些昆虫幼虫，是3种两栖类食物谱中最主要的组成部分，蜉金龟科昆虫在高原林蛙食谱中的比例更高，高原林蛙可能从畜牧业发展中获得更多的好处，使之在食物竞争方面处于优势地位。与无放牧样地相比，在有放牧样地的中，两栖类食谱中的蜉金龟科昆虫数量更多(有放牧：31.94%；无放牧：21.32%)、出现频率更高(有放牧：76.38%；无放牧：44%)。然而在不同样地上（有放牧/无放牧），两栖类的食物组成无显著性差异(P=0.188)，两栖类的数量(P=0.075)、肥满度(P=0.537)均没有显著差别。 5、两栖类幼体的竞争策略：实验室条件下，通过活动性水平，变态时的体重、增长率和完成变态所需时间考察自然条件下常同水塘分布的中华蟾蜍（Bufo gargarizans）和高原林蛙蝌蚪的竞争策略。结果表明：中华蟾蜍蝌蚪在不同食物资源条件下，所选择的生存策略可能不同，即食物资源充足时，增加活动性获取更多食物，食物资源有限时，降低活动性且提前完成变态；与中华蟾蜍蝌蚪相比，在食物资源有限时高原林蛙蝌蚪获取食物能力可能更强。 This paper presented the study of competition of three amphibians (Rana kukunoris, Nanorana pleskei, Bufo minshanicus) based on spatial, temporal and dietary scales in Zoige wetland. We measured coexistence patterns of three amphibians and analyzed their ability of exploiting resource. Effects of grazing on the diet composition and diet competition of amphibians were analyzed by their diet composition during spring and autumn. Furthermore, we examined the competitive ability of larval common frogs (Rana kukunoris)and common toads(Bufo gargarizans) in a laboratory experiment, and analyzed their competitive strategies respectively. The results were as follows: 1 .The status of using spatial resource Niche breadths of B. minshanicus adults on 5 dimensional axes including character of pasture(0.41), number of yaks dung(0.42), vegetation coverage(0.20), temperature (0.50)and humidity(0.51) of ground surface were narrower than adults of R. kukunoris and N. pleskei. Niche breadths of B. minshanicus subadults were broader than R.kukunoris subadults and N.pleskei subadults on 5 dimensional axes including character of pasture (0.66), number of yaks dung (0.58), vegetation height (0.64), distance to small waterbodies (0.51), humidity of ground surface (0.79). Niche breadths of N. pleskei subadults were the narrowest in three anurans subadults on 4 dimensional axes including character of pasture (0.39), number of yaks dung (0.30), temperature (0.18) and humidity (0.33) of ground surface, niche breadths of N. pleskei adults were the broadest in three anurans adults on 4 dimensional axes including number of yaks dung (0.81), vegetation height (0.63) and coverage(0.47), distance to small waterbodies(0.68).Comparatively, niche breadths of R. kukunoris were broader on the two microclimate factors including temperature(adults：0.62；subadults：0.56) and humidity (adults：0.84；subadults：0.60)of ground surface, but was narrow on distance to small waterbodies(adults：0.27；subadults：0.14). Strategies for using habitat resource of adults and subadults of the three species anuran were different. Generally, subadults of R. kukunoris and N. pleskei needs better habitat condition. It was quite similar that three anurans exploited spatial resource, Niche overlap between R. kukunoris and N. pleskei (0.87) was greater than that between R. kukunoris and B.minshanicus(0.81). 2.Daily activity rhythm R. kukunoris audlts were active when air temperatures were as low as 0℃, R. kukunoris subadults were active at 2℃, B.minshanicus subaudlts were active at 8℃. Positive correlation was found between activities of amphibians and air temperature, Subadults of R.kukunoris, (r=0.797, p<0.001), Subadults,of,B.minshanicus, (r=0.794, p<0.001), andbadults,of,R.kukunoris(r=0.456, p<0.05).Amphibians were more active during sunny days than cloudy days. In cloudy turning into sunny, R. kukunoris and B.minshanicus subadults had two active peak: at noon about 12:30 and 15:30～16:30 pm; in cloudy, R. kukunoris and B.minshanicus subadult had two active peak too : 9:30～10:30am,15:30～16:30pm. 3.Diet analysis Aphodiidae was the most commonly consumed food item by R. kukunoris based on index of relative importance (IRI) during spring (35.28%) and autumn (28.57%) in Zogie wetland. Besides Aphodiidae, larval insect, dipterans such as Bibionidae, Muscidae, Calliphoridae also were important food item for R. kukunoris, in autumn, locust was one of important food item for R. kukunoris. The most important food item for B.minshanicus during spring (IRI:85.54%) and autumn (IRI:49.70%) was ants, following, was coleopterans, such as Aphodiidae, dung beetle. Aphodiidae (IRI:13.41%) were the most important consumed food item by N. pleskei during spring too, following, was ants and Bibionidae. Dietary breadth of N. pleskei (0.43) were greater than R. kukunoris (0.22) and B. minshanicus (0.09). As a whole, Dietary breadth of amphibians during aurumn were greater than spring. Based on prey item, dietary overlap between R. kukunoris and N. pleskei (0.40) was greater than that between R. kukunoris and B.minshanicus (0.33) during spring. 4.Effects of grazing on the diet composition and diet competition of amphibians Amphibians are an important part of the pasture ecosystems as prey and predator. In Zogie wetland, major diet of amphibians was closely associated with dung of yaks, for example, Aphodiidae, Bibionidae, Muscidae, dung beetle. Dung of yaks was major diet and habitat of these insects. Proportion of Aphodiidae was higher in diet composition of R. kukunoris than N. pleskei and B.minshanicus, with development of pasturage, R. kukunoris may have a diet competitive advantage over N. pleskei and B.minshanicus. Number of Aphodiidae in diet composition of amphibians was higher in samples with grazing (31.94%) than in those without grazing (21.32%). Occurrence Frequency of Aphodiidae in diet composition of amphibians was higher in samples with grazing (76.38%) than in those without grazing (44%). However, There was not significantly different on diet composition (P=0.188), and number (P=0.075) and the relative fatness (P=0.537) of amphibians between grazing samples and without grazing. 5.Competitive strategies of amphibian larvae I examined the competitive ability of larval toads (Bufo gargarizans) and frogs (Rana kukunoris) which co-occur in the nature pond by activity level, the growth rate and mass at metamorphosis and larval period in a laboratory experiment. The results suggest: In laborary, B.gargarizans adapted himself to different food level by changing activity. At high food level, B. gargarizans increased activity to gain more diet. At low food level, B. gargarizans decreased activity and achieved early metamorphosis. When food resource was limit, R. kukunoris could gain more food than B. gargarizans.