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Taxonomic revision of the extant species of the Japanese Stephanidae (Hymenoptera), with discussion on distribution pattern and conservation importance 日本蝶形目现存物种的分类修订,以及关于分布模式和保护重要性的讨论
IF 0.7 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-10-17 DOI: 10.1111/ens.12588
Kyohei Watanabe

The Japanese species of the family Stephanidae Leach, 1815 are revised. A total of nine extant species are recognized from Japan, including two new species, Megischus pretiosus sp. nov. and Parastephanellus ryukyuensis sp. nov., and one species, Foenatopus ruficollis (Enderlein, 1913), is newly recorded from Japan. The female of Foenatopus cervinus Townes, 1958 and the male of Megischus baogong Ge & Tan, 2022 are newly described. A key to the Japanese genera and species of this family is proposed. The biogeographic pattern and the origin of Japanese stephanids, the role of Kuroshio Current, and the conservation importance of endemic species to the Ogasawara (Bonin) Islands are also discussed.

对 Stephanidae 科的日本种 Leach, 1815 进行了修订。其中包括两个新种:Megischus pretiosus sp. nov.和 Parastephanellus ryukyuensis sp. nov.。新描述了 Foenatopus cervinus 的雌性 Townes, 1958 和 Megischus baogong 的雄性 Ge & Tan, 2022。提出了该科日本属和种的检索表。此外,还讨论了日本阶鳉的生物地理格局和起源、黑潮的作用以及小笠原(波宁)群岛特有物种的保护重要性。
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引用次数: 0
Dual roles of terrestrial isopods in seed predation and seed dispersal in Phacellanthus tubiflorus (Orobancheaceae) 陆生等脚类动物在管花黄檀(萝苁科)种子捕食和种子传播中的双重作用
IF 0.7 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-09-25 DOI: 10.1111/ens.12586
Kenji Suetsugu, Osamu Nakanishi

Terrestrial isopods are known to play significant roles in litter decomposition and seed predation. The present study examines the function of terrestrial isopods, particularly Armadillidium vulgare and Armadillidium nasatum, in the seed dispersal of a nonphotosynthetic plant, Phacellanthus tubiflorus. These isopods were observed ingesting seeds in the natural habitat of P. tubiflorus in Hyogo Prefecture, Japan. Contrary to the prevailing belief that invertebrates do not serve as effective internal seed dispersers, our analysis demonstrated that some seeds ingested by A. vulgare and A. nasatum were excreted whole, with viability rates comparable to those of seeds directly sourced from fruits. This indicates that these isopods could act not only as seed predators but also as seed dispersers. The research highlights the importance of further exploration into the ecological contributions of isopods and other invertebrates as seed dispersers.

众所周知,陆生等脚类动物在垃圾分解和种子捕食中发挥着重要作用。本研究考察了陆生等脚类动物,尤其是Armadillidium vulgare和Armadillidium nasatum,在非光合作用植物Phacellanthus tubiflorus种子传播过程中的功能。在日本兵库县的Phacellanthus tubiflorus自然栖息地,观察到这些等脚类动物摄取种子。我们的分析表明,A. vulgare 和 A. nasatum 摄入的一些种子被完整地排出体外,其存活率与直接从果实中获取的种子相当。这表明这些等脚类动物不仅可以作为种子捕食者,还可以作为种子传播者。这项研究强调了进一步探索等脚类动物和其他无脊椎动物作为种子传播者的生态贡献的重要性。
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引用次数: 0
Life history of different giant water bug, Appasus japonicus (Belostomatidae: Heteroptera), populations throughout the Japanese Archipelago 日本列岛不同大水蝽种群的生活史
IF 0.7 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-09-25 DOI: 10.1111/ens.12585
Hinata Ohura, Miyuu Matsumoto, Aina Yoshimura, Shin-ya Ohba

As temperatures in the Japanese Archipelago vary, species widely distributed in Japan should have different active periods, ecologies, and life histories. Information on developmental zero and effective accumulated temperature is necessary to estimate life history. The giant water bug, Appasus japonicus, found in rice paddies, ponds, and other lentic waters, is distributed throughout Japan except for the Ryukyu Islands, and is a representative taxon in which the male cares for the eggs. In this study, we investigated the life history, including the length of the breeding season, of different A. japonicus populations from Hokkaido, Ibaraki, Hyogo, and Nagasaki. The developmental period, developmental zero, and effective accumulated temperature of the eggs and nymphs showed no significant differences among the populations. The life history for each population was estimated based on climate data. This showed that the Hokkaido population had a shorter and more extremely limited breeding season than the other populations. The estimated number of generations during one breeding season was approximately one in Hokkaido, two in Ibaraki and Hyogo, and three in Nagasaki. The estimated number of times males cared for eggs during the breeding season was less than one in Hokkaido, eight in Ibaraki and Hyogo, and twelve in Nagasaki. Although Belostomatidae may carry multiple egg masses during the breeding season, this study shows that the male of the Hokkaido population does not care for multiple egg masses.

由于日本列岛的气温不同,广泛分布于日本的物种应具有不同的活动期、生态和生活史。有关发育零度和有效积温的信息对于估计生活史非常必要。大水虻(Appasus japonicus)发现于稻田、池塘和其他透水水域,除琉球群岛外分布于日本各地,是雄性照顾卵的代表性类群。在本研究中,我们调查了北海道、茨城、兵库和长崎的不同日本鲤种群的生活史,包括繁殖季节的长度。结果表明,不同种群的卵和若虫的发育期、发育零度和有效积温没有显著差异。根据气候数据估算了各种群的生活史。结果表明,与其他种群相比,北海道种群的繁殖期更短、更有限。据估计,北海道在一个繁殖季节的世代数约为 1 代,茨城和兵库为 2 代,长崎为 3 代。在繁殖季节,雄性照顾卵的估计次数在北海道少于 1 次,在茨城和兵库为 8 次,在长崎为 12 次。虽然贝氏蝠科在繁殖季节可能携带多个卵块,但本研究表明,北海道种群的雄性贝氏蝠不会照顾多个卵块。
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引用次数: 0
Predation of salamander egg sacs by the water strider Gerris (Macrogerris) insularis Motschulsky (Hemiptera: Gerridae) 水黾Gerris (Macrogerris) insularis Motschulsky(半翅目:黾科)捕食蝾螈卵囊
IF 0.7 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-09-25 DOI: 10.1111/ens.12587
Shona Yasuda, Ryota Morii, Hiroshi Ikeda

Water striders (Hemiptera: Gerridae) are sucking predators that inhabit freshwater and marine environments. They prey mainly on insects that fall on the water surface, but they are also known to feed on amphibian eggs. Here, we report on the predation of egg sacs of Hynobius nigrescens (Caudata: Hynobiidae) by Gerris (Macrogerris) insularis. We observed predation on egg sacs of H. nigrescens by G. insularis at a pond in a mountainous area of Nagano City, Nagano Prefecture, Japan. We also found egg sacs with predation marks probably made by water striders at a pond in the mountainous area of Nishimeya Village, Aomori Prefecture, Japan. Five water striders collected around the preyed egg sacs were identified as G. insularis. We filled three tanks with water and an egg sac, and three water striders that were collected at the site in Nishimeya Village were placed together in each tank. In two tanks, the jelly layer of the egg sac was partially cut in a horizontal manner, while in another tank, the jelly layer was removed on the part of the egg sac exposed on the water surface. Notably, the water striders exhibited predatory behavior only when the jelly layers were removed. Considering the results of the rearing experiment and the predation marks observed in the field, G. insularis is also likely to prey on egg sacs at the site in Nishimeya Village. Our study suggested that predation by G. insularis occurs when eggs are accidentally exposed due to disturbance of the jelly layer.

水黾(半翅目:黾科)是一种栖息在淡水和海洋环境中的吸食性食肉动物。它们主要捕食落在水面上的昆虫,但也以两栖动物的卵为食。在此,我们报告了海岛鳗(Macrogerris)捕食黑线姬蛙(尾目:姬蛙科)卵囊的情况。我们在日本长野县长野市山区的一个池塘中观察到 G. insularis 捕食黑线姬蛙卵囊的情况。我们还在日本青森县 Nishimeya 村山区的一个池塘中发现了可能是水黾捕食痕迹的卵囊。在被捕食的卵囊周围采集到的五只水黾被鉴定为岛黾。我们在三个水箱中装满水和卵囊,并将在西目屋村采集到的三只水黾一起放入每个水箱中。在两个水槽中,水平切开卵囊的部分果冻层,而在另一个水槽中,去除卵囊露出水面部分的果冻层。值得注意的是,水黾只有在去掉果冻层后才表现出捕食行为。考虑到饲养实验的结果和在野外观察到的捕食痕迹,水黾也有可能在西宫村捕食卵囊。我们的研究表明,当卵因果冻层受到破坏而意外暴露时,就会遭到海龟的捕食。
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引用次数: 0
Effects of heterogenous forest environments on ground-dwelling beetles in a conifer plantation 异质森林环境对针叶树人工林中地栖甲虫的影响
IF 0.7 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-08-16 DOI: 10.1111/ens.12584
Akira Ueda, Hiroki Itô, Seiichi Kanetani

It is widely recognized that heterogeneous environments promote biodiversity. Consequently, we investigated whether heterogeneous environments influence the diversity of ground-dwelling beetles within a well-managed conifer plantation forest. We collected carabid beetles, and necrophagous silphid and dung beetles (necrophagous beetles) using pitfall traps at 37 sites with or without fish meat bait. The result of a two-dimensional principal component analysis (PCA) utilizing environmental indices measured at each site suggested that the environmental heterogeneity depended on the survival density of planted conifers, serving as an indicator of historical gap formation levels. Gaps populated by invading broadleaved trees exhibited low canopy opening percentages and understory cover degrees, while those filled by the growth of planted conifers displayed the opposite conditions. The presence of bait had a negligible impact on carabid beetles but significantly influenced necrophagous beetles. No significant relationships were found between the scores of PCA components and the beetle species richness, suggesting that a heterogeneous environment does not increase the beetle species diversities. However, significant relationships were observed with the scores on nonmetric multidimensional scaling axes, indicating that a heterogeneous environment contributes to the complexity of beetle assemblages. The most dominant carabid, Synuchus cycloderus, was abundant in areas where historical gap formation was thought to be more prevalent. Among necrophagous beetles, Pherotrupes laevistriatus and Panelus parvulus preferred habitats with darker environments, in contrast to Onthophagus fodiens and O. nitidus. This study will aid in developing forestry practices aimed at conserving specific ground-dwelling beetle species within conifer plantations.

人们普遍认为,异质环境能促进生物多样性。因此,我们研究了异质环境是否会影响管理良好的针叶林中地栖甲虫的多样性。我们在 37 个有或没有鱼肉饵料的地点使用坑式陷阱收集了食肉甲虫、食尸鞘甲虫和粪甲虫(食尸甲虫)。利用在每个地点测量的环境指数进行二维主成分分析(PCA)的结果表明,环境异质性取决于种植针叶树的存活密度,可作为历史间隙形成水平的指标。由入侵阔叶树填充的空隙显示出较低的树冠开阔度和林下覆盖度,而由种植针叶树填充的空隙则显示出相反的情况。诱饵的存在对食肉甲虫的影响微乎其微,但对食尸甲虫的影响却很大。在 PCA 成分得分与甲虫物种丰富度之间没有发现明显的关系,这表明异质环境不会增加甲虫物种的多样性。然而,在非度量多维标度轴上的得分之间却发现了明显的关系,表明异质环境有助于提高甲虫群落的复杂性。最主要的食肉甲虫--Synuchus cycloderus,在历史缺口形成被认为更为普遍的地区大量存在。在食尸甲虫中,Pherotrupes laevistriatus 和 Panelus parvulus 喜欢较暗的环境,与 Onthophagus fodiens 和 O. nitidus 形成鲜明对比。这项研究将有助于制定旨在保护针叶树种植园中特定地栖甲虫物种的林业措施。
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引用次数: 0
New genus and two new species of Cecidomyiidi (Diptera: Cecidomyiidae) inhabiting the fig wall of Ficus subpisocarpa and Ficus caulocarpa (Moraceae) in Japan and Taiwan 栖息于日本和台湾桑科无花果壁的 Cecidomyiidi(双翅目:Cecidomyiidae)新属和两个新种
IF 0.7 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-08-09 DOI: 10.1111/ens.12583
Kôichi Arimoto, Junichi Yukawa, Masako Yafuso, Ayako Sasaki, Zhi-Hui Su

There are approximately 850 species of Ficus (Moraceae). However, few species of gall midges (Diptera: Cecidomyiidae) inhabiting syconia and leaves of fig trees are known. In field surveys, gall midges were found in syconia of Ficus caulocarpa and Ficus subpisocarpa. Here, we examined adults, pupae and larvae of the gall midge species, describe the morphology, and provide information on distribution, behavior and genetic data. A new genus, Ficidiplosis Yukawa and Arimoto, gen. nov., is established in the supertribe Cecidomyiidi for two new species, Ficidiplosis subpisocarpae Yukawa and Arimoto, sp. nov. and Ficidiplosis caulocarpae Yukawa and Arimoto, sp. nov., which emerged from syconia of Ficus subpisocarpa and Ficus caulocarpa, respectively, in Japan and Taiwan. The larvae of Ficidiplosis species feed on the parenchyma of the fig wall and pupate there without making galls. The mitochondrial cytochrome c oxidase subunit I (COI)-based neighbor-joining tree using samples from Japan and Taiwan supported the existence of two separate species.

无花果(桑科)大约有 850 个品种。然而,栖息在无花果树的syconia和叶片上的瘿蚊(双翅目:Cecidomyiidae)种类却很少为人所知。在实地调查中,我们在无花果树(Ficus caulocarpa)和无花果树(Ficus subpisocarpa)的梧桐叶中发现了瘿蚊。在这里,我们研究了瘿蚊的成虫、蛹和幼虫,描述了它们的形态,并提供了有关分布、行为和遗传数据的信息。新属 Ficidiplosis Yukawa and Arimoto,gen. nov.,为超群 Cecidomyiidi 类中的两个新种,即 Ficidiplosis subpisocarpae Yukawa and Arimoto,sp. nov.和 Ficidiplosis caulocarpae Yukawa and Arimoto,sp. nov.,它们分别出现在日本和台湾的 Ficus subpisocarpa 和 Ficus caulocarpa 的鞘螨中。Ficidiplosis 物种的幼虫以无花果壁的实质为食,并在那里化蛹而不形成虫瘿。基于线粒体细胞色素 c 氧化酶亚单位 I(COI)的邻接树(样本来自日本和台湾)支持两个独立物种的存在。
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引用次数: 0
Alpha-amanitin tolerance of adult flies of mycophagous and nonmycophagous species in Drosophilidae 果蝇科噬菌性和非噬菌性物种成蝇对α-amanitin的耐受性
IF 0.7 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-08-08 DOI: 10.1111/ens.12577
Yang Zhang, Masanori J. Toda, Hide-aki Watabe, Toru Katoh

Mushrooms produce a great variety of defense chemical compounds to protect themselves from fungivores. Alpha-amanitin is one of such compounds found in some Amanita mushrooms. The present study focuses on the effect of α-amanitin on adult survival in 17 drosophilid species including mycophagous ones from the subgenus Drosophila and the genus Hirtodrosophila and nonmycophagous ones of the subgenera Drosophila, Siphlodora and Sophophora. The results showed that all the essential fungivores and some partially mycophagous species from different lineages are tolerant to this lethal toxin at a concentration of 50 μg/mL, suggesting that the α-amanitin tolerance may have been acquired more or less independently of the evolution of mycophagy in the family Drosophilidae.

蘑菇会产生多种防御化学物质,以保护自己不受食菌动物的侵害。α-amanitin是在一些天南星蘑菇中发现的此类化合物之一。本研究的重点是α-amanitin 对 17 种果蝇成虫存活率的影响,包括果蝇亚属和 Hirtodrosophila 属中的噬菌果蝇,以及果蝇亚属、Siphlodora 和 Sophophora 属中的非噬菌果蝇。结果表明,来自不同品系的所有基本食菌物种和一些部分食菌物种对浓度为 50 μg/mL 的这种致命毒素都有耐受性,这表明α-amanitin 耐受性的获得或多或少与果蝇科的食菌进化无关。
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引用次数: 0
New insights on the chromosomal inversion thermal adaptation in insects: American populations of Drosophila subobscura 昆虫染色体反转热适应的新发现亚种果蝇的美洲种群
IF 0.7 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-07-30 DOI: 10.1111/ens.12582
Concepció Arenas, Goran Zivanovic, Cinta Pegueroles, Francesc Mestres

Drosophila subobscura is an excellent model species for testing genetic variation in thermal adaptation due to its rich inversion chromosomal polymorphism. In Palearctic populations, the inversions of this species can be classified as “cold”, “warm” or “nonthermal” adapted. Based on this classification the Chromosomal Thermal Index (CTI) was developed, which allows measuring the thermal adaptation of populations and monitoring changes over time. Here, we aim to use this index in American colonizing populations of D. subobscura for the first time to obtain new information on the species thermal adaptation. Thus, thermal adapted inversions (“cold” and “warm”) were defined for the American continent and CTI was computed in South American (1981 and 1999 samples) and North American (1985 and 2004 samples) populations of D. subobscura. In general, both American populations showed an inverse relationship between CTI values and latitude, with CTI values decreasing when latitude increases. When comparing populations sampled in different years, an increase in CTI values was detected in four out of six temporal comparisons (only one was significant) of South America (1981 and 1999) and in six out of seven populations of North America (1985 and 2004). A global analysis using a one-way repeated measures anova of CTI values in both American hemispheres showed a trend of increase for “warm” adapted inversions in Chile and North America, but this increase was only significant for the latter. Overall, these results are in agreement with global warming expectations, although natural selection acted differently in the colonized hemispheres.

亚种果蝇具有丰富的倒位染色体多态性,是测试热适应遗传变异的极佳模式物种。在古北区种群中,该物种的倒位染色体可分为 "冷"、"暖 "或 "非热 "适应型。根据这种分类方法,开发出了染色体热指数(CTI),它可以测量种群的热适应性并监测随时间的变化。在此,我们首次将该指数用于亚种仓鼠(D. subobscura)的美洲定殖种群,以获得有关该物种热适应性的新信息。因此,我们定义了美洲大陆的热适应倒位("冷 "和 "暖"),并计算了南美洲(1981 年和 1999 年样本)和北美洲(1985 年和 2004 年样本)亚盘尾蜥种群的 CTI。总体而言,这两个美洲种群的 CTI 值与纬度呈反比关系,纬度越高,CTI 值越低。在比较不同年份采样的种群时,南美洲(1981 年和 1999 年)六次时间比较中有四次检测到 CTI 值增加(只有一次是显著的),北美洲(1985 年和 2004 年)七次时间比较中有六次检测到 CTI 值增加。对美洲两个半球的 CTI 值进行的单向重复测量 anova 全球分析表明,智利和北美洲的 "暖 "适应反常现象呈上升趋势,但这种上升只在北美洲显著。总体而言,这些结果与全球变暖的预期一致,尽管自然选择在殖民地半球的作用不同。
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引用次数: 0
Community structure of parasitoids attacking Schizomyia sasakii (Diptera: Cecidomyiidae) and local adaptation of the gall midge in the Izu region 伊豆地区瘿蚊(双翅目:瘿蚊科)的寄生虫群落结构和当地适应性
IF 0.9 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-05-22 DOI: 10.1111/ens.12576
Tomohisa Fujii, Kazunori Matsuo, Junichi Yukawa, Keizi Kiritani, Yoshihisa Abe, Makoto Tokuda

Insect community structures and biological interactions vary with the distance from the mainland to islands. Gall inducers are key organisms in local arthropod communities because their galls harbor diverse arthropods. We investigated the parasitoid community of a gall-inducing cecidomyiid Schizomyia sasakii on the Izu Peninsula and the Izu Islands, Japan. We examined relationships between parasitism by Inostemma sp. and Torymus hirtipennis, and gall characteristics, and analyzed the directional selection on gall characteristics induced by S. sasakii in each locality. The species richness of parasitoids on the Izu Islands decreased with the distance from the Izu Peninsula to each island, and area of respective islands. Inostemma sp. preferably attacked large galls on Ohshima Island, and T. hirtipennis tended to attack relatively small galls in the Izu Peninsula. Directional selection on gall characteristics of S. sasakii favored the induction of larger galls and thicker tissues by S. sasakii on Ohshima Island. In contrast, no directional selection was detected in the gall characteristics on Hachijojima Island. The number of alternative host species of parasitoid may affect variation in parasitoid richness of S. sasakii on distant and small islands, because the species richness of gall midges is necessary to maintain local parasitoid populations. Our study supported the ovipositor limitation hypothesis for the torymid species, related to gall size. The directional selection of parasitoid attacks on the Izu Islands may act to favor the induction of large and hypertrophic galls by S. sasakii to avoid the parasitoids.

昆虫群落结构和生物之间的相互作用随大陆到岛屿的距离而变化。虫瘿诱导体是当地节肢动物群落中的关键生物,因为它们的虫瘿中栖息着多种节肢动物。我们调查了日本伊豆半岛和伊豆诸岛上的引胆虫Schizomyia sasakii的寄生虫群落。我们研究了寄生虫Inostemma sp.和Torymus hirtipennis与虫瘿特征之间的关系,并分析了S. sasakii在各地诱导虫瘿特征的定向选择。伊豆群岛上寄生虫的物种丰富度随着伊豆半岛到各岛屿的距离和各岛屿的面积而降低。在大岛,Inostemma sp.更喜欢攻击大的虫瘿,而在伊豆半岛,T. hirtipennis则倾向于攻击相对较小的虫瘿。对 S. sasakii虫瘿特征的定向选择有利于大岛 S. sasakii 诱导更大的虫瘿和更厚的组织。相比之下,八丈岛的虫瘿特征没有发现定向选择。寄生虫可供选择的寄主种类的数量可能会影响寄生虫丰富度的变化,因为瘿蚊种类的丰富度是维持当地寄生虫种群的必要条件。我们的研究支持与虫瘿大小有关的卵柄限制假说。伊豆群岛上寄生虫攻击的定向选择可能有利于诱导 S. sasakii 形成大而肥大的虫瘿,以躲避寄生虫的攻击。
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引用次数: 0
What accounts for the difference in the emergence times of Drosophila melanogaster between the first and second eclosion days? 是什么原因导致黑腹果蝇在第一和第二个羽化日的出现时间不同?
IF 0.9 4区 农林科学 Q4 ENTOMOLOGY Pub Date : 2024-05-03 DOI: 10.1111/ens.12573
Yasuhiko Watari, Shin G. Goto, Yosuke Miyazaki, Izuru Kuroki, Kazuhiro Tanaka

One of the characteristics of the adult eclosion rhythm in Drosophila melanogaster is that adult emergence time differs greatly between the first and second eclosion days. The emergence time is in the middle of the light period on the first eclosion day, but immediately after light-on on the second day. We hypothesized that incomplete entrainment of the endogenous pacemaker to the light–dark (LD) cycle is responsible for the daily variation. Due to the very short pupal period of this species, adult emergence may occur before complete synchronization of the pacemaker with the external cycle on the first eclosion day. Therefore, the peak time on the first eclosion day may differ significantly from that on the second day. To verify this hypothesis, using pupae that had pupariated within 24 h, the time difference between the first and second peaks was compared in LD 12:12 or constant darkness at five different temperatures from 30 to 15°C. In both light regimes, the time difference decreased with decreasing temperature and extended pupal duration. The interval between the eclosion peaks approached 24 h, supporting this hypothesis. These results can be interpreted by using a two-oscillator model.

黑腹果蝇成虫羽化节律的一个特点是,成虫在羽化第一天和第二天的出羽时间差别很大。第一个羽化日的成虫出现时间在光照期的中间,而第二个羽化日的成虫出现时间则在光照开始后不久。我们推测,内源起搏器与光-暗(LD)周期的不完全协调是造成这种日变化的原因。由于该物种的蛹期很短,成虫的出现可能发生在第一个羽化日起搏器与外部周期完全同步之前。因此,第一个羽化日的峰值时间可能与第二个羽化日的峰值时间有很大不同。为了验证这一假设,我们使用在 24 小时内化蛹的蛹,在 30 至 15°C 的五个不同温度条件下,比较了 LD 12:12 或恒定黑暗条件下第一个和第二个峰值之间的时间差。在两种光照条件下,时间差都随着温度的降低和蛹期的延长而减小。蜕皮高峰之间的时间间隔接近 24 小时,支持了这一假设。这些结果可以用双振荡器模型来解释。
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