Larva Morphology of Shore Flies Ephydra riparia and Paracoenia fumosa (Diptera: Ephydridae) and Adaptation of Diptera to Increased Salinity

E. Yu. Yakovleva, E. B. Naimark, D. D. Sivunova, M. G. Krivosheina, A. V. Markov
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Abstract

Larvae of many shore fly species (family Ephydridae) are adapted to living in water with high or extremely high salinity. Little is known about the morphological and physiological foundations of such adaptation. We described the details of the morphology of third-instar larvae of two shore flies: Ephydra riparia and Paracoenia fumosa, and presented the images made using scanning electron microscopy (SEM). For the first time, by silver-staining and SEM, we proved that the larvae of both studied species had anal organs (AO)— specialized structures that serve an osmoregulatory function and are responsible for the transport of ions from the environment to the larval hemolymph (but not in the opposite direction). We compared the larvae morphology of the studied species with some other shore fly species from the genera Ephydra, Paracoenia, Hydrellia, and Coenia, as well as with larvae of the model species Drosophila melanogaster (family Drosophilidae). Special attention was paid to the morphology of AO, which contribute to the adaptation of larvae to increased salinity. Extremely halophilic species either do not have AO, or they are poorly developed, while the moderately halophilic ephydrids have more developed features connected with the permeability of the AO cuticle and active ion transport. These features are most developed in freshwater shore flies. AO activity can vary due to the shape and area of the AO, the smoothness or wrinkling of the cuticle, and the presence of nano-pits on it. The described variability of the AO structure is probably adaptive since, at high salinity, both the permeability of the AO cuticle and the active transport of ions from the environment to the hemolymph become less useful or even harmful.

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岸蝇 Ephydra riparia 和 Paracoenia fumosa(双翅目:Ephydridae)的幼虫形态以及双翅目对盐度升高的适应情况
摘要 许多岸蝇(Ephydridae科)的幼虫都适应生活在高盐度或极高盐度的水中。人们对这种适应的形态和生理基础知之甚少。我们描述了两种岸蝇三龄幼虫的形态细节:Ephydra riparia 和 Paracoenia fumosa,并展示了使用扫描电子显微镜(SEM)拍摄的图像。通过银染色和扫描电子显微镜,我们首次证明了所研究的两个物种的幼虫都有肛门器官(AO)--具有渗透调节功能的特化结构,负责将离子从环境中输送到幼虫血淋巴(但不是相反方向)。我们将所研究物种的幼虫形态与 Ephydra、Paracoenia、Hydrellia 和 Coenia 属的其他一些岸蝇物种以及模式物种黑腹果蝇(果蝇科)的幼虫进行了比较。研究人员特别关注AO的形态,这有助于幼虫对盐度升高的适应。极度嗜卤的物种要么没有AO,要么AO发育不良,而适度嗜卤的嗜盐水果蝇则具有与AO角质层的渗透性和活性离子传输有关的更发达的特征。这些特征在淡水岸蝇中最为发达。AO 活性会因 AO 的形状和面积、角质层的平滑或皱褶以及上面是否存在纳米凹坑而变化。所描述的 AO 结构的变化可能是适应性的,因为在高盐度条件下,AO 角质层的渗透性和离子从环境到血淋巴的主动运输都变得不那么有用,甚至有害。
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