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作者信息

Grimm Rainer

机构信息

Abt. Ökologie, II. Zoologisches Institut und Museum der Universität Göttingen, Göttingen, Deutschland.

出版信息

Oecologia. 1973 Sep;11(3):187-262. doi: 10.1007/BF01882783.

Abstract

The energy turnover of weevil populations ofRhynchaenus fagi L.,Strophosomus (Schönherr) andOtiorrhynchus singularis L. (Curculionidae) was investigated in the following stages: a) Observation of habits and life-cycle; b) investigation of abundance-dynamics; c) establishing the energy budget of average individuals; d) calculation of the energy turnover of the populations. In the case ofRhynchaenus fagi the investigations were carried out with all developmental and maturity stages, in the cases ofStrophosomus sp. andO. singularis only with the imagimes. 1. The larva ofR. fagi mines the beechleaf. The development takes 5\2-6 weeks from oviposition (beginning of May) to the emergence of the beetle. The imago lives for about 12 months. There is a first eating period lasting until the end of July, followed by about 9 months of almost no food consumption; after hibernation there is the second eating period (May to June) and reproduction (beginning of May). Strophosomus sp. emerges on the ground in September and eats in the top shoots of the trees till the leaves fall; it feeds on fallen foliage on the ground during the winter and high in the trees again from May to July.Strophosomus sp. reproduces mainly parthenogenetically and dies no later than the end of August. O. singularis emerges mainly in May and June. After an intensive feeding period these females also deposit mostly unfertilized eggs. The beetles live for several years, imagines from different years living side by side. 2. Beginning in JulyR. fagi migrates increasingly to nearby spruce forests which are its favourite winter quarters in the Solling. Microclimate and thigmotaxis between the needles protectR. fagi from higher mortality. 3. Abundance and mortality of eggs, larvae, pupae and young beetles ofR. fagi were determined by counting the larvae and pupae on a large number of leaves and by marking leaves with a numbered label. For the beetles a linear progression of mortality was assumed from emergence to reproduction. In the case ofStrophosomus sp. the numbers of beetles emerging and the winter mortality were determined by means of eclector catches. Shaking samples yielded data concerning the mortality after hibernation. WithO. singularis the numbers of beetles emerging was established by means of eclectors. 4. In the case ofR. fagi the production of an average individual is 2.654 cal till the end of the pupal stage, 6.353 cal until death. 1.103 cal are taken up for cocoon and exuviae, 0.809 cal for egg production. From emergence to the end of July of the following yearStrophosomus sp. produces body substance of a calorific value of 28.630 cal, of which 9.690 cal are taken up for eggs. 5. A new calorimetric method was applied for the first time in determining the respiration ofR. fagi during its leaf-mining stages. The results agreed with the calculation of energy losses based on manometric measurement of O-consumption. With all weevil imagines manometric measurement of O-consumption was carried out under near-natural conditions. The imagines showed seasonal changes of the respiration level and theQ , which were usually consistent with their life cycle. 6. The assimilation of an average individual (A=P+R) was calculated from production (P) and respiration (R); in the case ofR. fagi for all developmental and maturity stages, in the case ofStrophosomus sp. for the imagines at different times of the year. The horizontal course of the assimilation curve ofR. fagi during pupal stage and hibernation indicates that the measuring technique is highly accurate. The energy loss through respiration increases at exactly the same rate as the energy potential of the animal decreases. An average individual ofR. fagi assimilates 4.509 cal from its emergence until the end of the pupal stage, 9.042 cal by the end of the first eating period (end of July), and a total of 17.031 cal by the end of the reproduction and second feeding period (June of the following year). An average individual ofStrophosomus sp. (imago) assimilates 24.000 cal by the end of the first eating period (end of November), 32.006 cal by the end of hibernation (late April), and 78.330 cal before it dies. 7. Investigations of consumption (C) and defecation (FU) and the calculation of assimilation (A) asA=C-FU usually confirmed the results ofA=P+R. The ecological efficiency (P/C %) ranges from 14-20% for larvae ofR. fagi, from 7.0-8.5% for all weevil imagines. 8. With an energy turnover of about 21.9\sx10 kcal/ha/yr in a 120-year-old beech wood (B1a) and about 42.1\sx10 kcal/ha/yr in a 60-year-old beech wood (B4), the population ofR. fagi belongs to the ecologically dominant phytophagous insects. The larvae assimilated 62% of this amount, the imagines 31% before hibernation and 7% in spring. TheStrophosomus-population (imagines) assimilated about 4.4×10 kcal/ha/yr in B1a and about 4.3×10 kcal/ha/yr in B4. 9. As far as the production of imagines is concerned (2.1\sx10 in B1a and 4.0\sx10 kcal/ha/yr in B4)R. fagi takes first place among phytophagous insects \3-at least in young beech woods. The production of imagines inStrophosomus sp. amounted to about 1.3×10 in B1a and about 1.2×10 kcal/ha/yr in B4, inO. singularis to about 0.9×10 in B1a and about 1.3×10 kcal/ha/yr in B4. 10. Assuming an averageA/C-efficiency of 20\2-25% the populations ofR. fagi (all stages) and ofStrophosomus sp. (imagines) consumed 103-128\sx10 kcal/ha/yr in B1a and 183-229\sx10 kcal/ha/yr in B4. This means 14\2-17% and 24\2-31% (in B1a and B4 respectively) of the total leaf substance consumed by phytophagous insects, for which Funke (1972a) arrived at a figure of 7.5\sx10 kcal/ha/yr.

摘要

对栗象(Rhynchaenus fagi L.)、瘤胸象(Strophosomus (Schönherr))和奇异齿爪象(Otiorrhynchus singularis L.,象甲科)象虫种群的能量周转进行了如下阶段的研究:a)习性和生命周期观察;b)丰度动态调查;c)确定平均个体的能量预算;d)计算种群的能量周转。对于栗象,对其所有发育和成熟阶段都进行了研究;对于瘤胸象属物种和奇异齿爪象,仅对成虫进行了研究。1. 栗象的幼虫在山毛榉树叶上蛀食。从产卵(5月初)到甲虫羽化,发育需要5.5 - 6周。成虫寿命约为12个月。有一个持续到7月底的首次进食期,随后是约9个月几乎不进食;冬眠后是第二个进食期(5月至6月)和繁殖期(5月初)。瘤胸象属物种于9月在地面羽化,在树梢进食直至树叶落下;冬季以地面落叶为食,5月至7月又在树上高处进食。瘤胸象属物种主要孤雌生殖,最迟在8月底死亡。奇异齿爪象主要在5月和6月羽化。经过一段强烈进食期后,这些雌虫也大多产下未受精卵。甲虫能活数年,不同年份的成虫并存。2. 从7月开始,栗象越来越多地迁移到附近的云杉林,那里是其在索林根最喜欢的越冬场所。针叶间的微气候和趋触性保护栗象免受更高的死亡率。3. 通过统计大量树叶上的幼虫和蛹,并给树叶贴上编号标签,确定了栗象卵、幼虫、蛹和幼甲虫的丰度和死亡率。对于甲虫,假设从羽化到繁殖的死亡率呈线性变化。对于瘤胸象属物种,通过陷阱捕获确定羽化甲虫的数量和冬季死亡率。摇晃样本得出冬眠后的死亡率数据。对于奇异齿爪象,通过陷阱捕获确定羽化甲虫的数量。4. 对于栗象,到蛹期结束时,平均个体的产量为2.654卡路里,到死亡时为6.353卡路里。1.103卡路里用于结茧和蜕皮,0.809卡路里用于产卵。从羽化到次年7月底,瘤胸象属物种产生热值为28.630卡路里的身体物质,其中9.690卡路里用于产卵。5. 一种新的量热法首次应用于测定栗象在蛀叶阶段的呼吸作用。结果与基于耗氧量测压法计算的能量损失一致。对于所有象虫成虫,在接近自然的条件下进行耗氧量测压。成虫的呼吸水平和Q值呈现季节性变化,这通常与其生命周期一致。6. 根据产量(P)和呼吸作用(R)计算平均个体的同化量(A = P + R);对于栗象,计算所有发育和成熟阶段的同化量,对于瘤胸象属物种,计算一年中不同时间成虫的同化量。栗象在蛹期和冬眠期间同化曲线的水平走势表明测量技术非常准确。呼吸作用导致的能量损失增加的速率与动物能量潜力下降的速率完全相同。一只栗象平均个体从羽化到蛹期结束同化4.509卡路里,到第一个进食期结束(7月底)同化9.042卡路里,到繁殖和第二个进食期结束(次年6月)总共同化17.031卡路里。一只瘤胸象属物种(成虫)平均个体到第一个进食期结束(11月底)同化24.000卡路里,到冬眠结束(4月下旬)同化32.006卡路里,到死亡前同化78.330卡路里。7. 对消耗量(C)和排粪量(FU)的研究以及通过A = C - FU计算同化量(A)通常证实了A = P + R的结果。栗象幼虫的生态效率(P/C%)为14 - 20%,所有象虫成虫的生态效率为7.0 - 8.5%。8. 在一片120年树龄的山毛榉林(B1a)中,栗象种群的能量周转约为21.9×10千卡/公顷/年,在一片60年树龄的山毛榉林(B4)中约为42.1×10千卡/公顷/年,栗象种群属于生态上占主导地位的植食性昆虫。幼虫同化了其中的62%,成虫在冬眠前同化31%,春季同化7%。瘤胸象种群(成虫)在B1a中同化约4.4×10千卡/公顷/年,在B4中约为4.3×10千卡/公顷/年。9. 就成虫的产量而言(B1a中为2.1×10千卡/公顷/年,B4中为4.0×10千卡/公顷/年),栗象在植食性昆虫中至少在年轻山毛榉林中排名第一。瘤胸象属物种成虫的产量在B1a中约为1.3×10千卡/公顷/年,在B4中约为1.2×10千卡/公顷/年,奇异齿爪象在B1a中约为0.9×10千卡/公顷/年,在B4中约为1.3×10千卡/公顷/年。10. 假设平均A/C效率为20 - 25%,栗象种群(所有阶段)和瘤胸象属物种种群(成虫)在B1a中消耗103 - 128×10千卡/公顷/年,在B4中消耗183 - 229×10千卡/公顷/年。这意味着分别占植食性昆虫消耗的总叶物质的14 - 17%和24 - 31%(分别在B1a和B4中),Funke(1972a)得出的植食性昆虫消耗的总叶物质为7.5×10千卡/公顷/年。

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