This study is concerned with the Late Holocene floodplain history of the Karkheh River in Lower Khuzestan, and in particular with the role of human action upon its channel shifts. The research was conducted in a multidisciplinary way, in which resources and approaches from different research fields were combined: (1) geomorphological mapping based on the interpretation of Landsat and CORONA satellite imagery, (2) analyses of geological sequences, including the identification of sedimentary facies and radiocarbon dating of organic material, (3) an archaeological field survey of ancient settlements, and (4) consultation of historical documents, mainly Arabic texts from the 9th–14th century and European travel literature from the 16th-early 20th century. Three main channel belts of the Karkheh were identified (labelled Kh1, Kh2 and Kh3), corresponding to successive stages in the evolution of the floodplain. Two river shifts are documented in the datasets, both taking place within the last 2000 years. The first avulsion regards a shift from channel belt Kh1, once a tributary of the Karun, to the straight river bed of Kh2, taking place at least after 1240–1310 cal BP/710–640 AD. The second avulsion, from Kh2 to Kh3, is clearly documented in historical sources and happened in a single night event in the year 1837/113 cal BP. Reactivation of the Kh2 river bed and its irrigation canals can be attributed to the recent construction of an artificial canal bypassing the second avulsion point. Both river shifts were strongly influenced by human interference, whereby an artificial irrigation canal took over the entire river flow from the main channel belt. Most likely, a combination of human-induced factors, such as weakening of the river levees, high sedimentation rates and disadvantageous channel gradients, led to a situation prone to avulsion.
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Abstract The defense strategy of an insect toward natural enemies can include a trait that appears at first sight to contradict its defensive function. We explored phylogeny, chemistry, and defense efficiency of a peculiar group of hymenopteran sawfly larvae where this contradiction is obvious. Pseudodineurini larvae live in leaf mines that protect them from some enemies. Disturbed larvae also emit a clearly perceptible lemon-like odor produced by ventral glands, although the mine hampers the evaporation of the secretion. The mine could also lead to autointoxication of a larva by its own emitted volatiles. Citral was the major component in all Pseudodineurini species, and it efficiently repels ants. We conclude that full-grown larvae that leave their mine to pupate in the soil benefit from citral by avoiding attacks from ground-dwelling arthropods such as ants. In some species, we also detected biosynthetically related compounds, two 8-oxocitral diastereomers (i.e., (2E,6E)- and (2E,6Z)-2,6-dimethylocta-2,6-dienedial). Synthetic 8-oxocitral proved to be a potent fungicide, but not an ant repellent. The discrete distribution of 8-oxocitral was unrelated to species grouping in the phylogenetic tree. In contrast, we discovered that its presence was associated with species from humid and cold zones but absent in species favoring warm and dry environments. The former should be protected by 8-oxocitral when faced with a fungal infestation while crawling into the soil. Our work shows the importance of integrating knowledge about behavior, morphology, and life history stages for understanding the complex evolution of insects and especially their defense strategies.
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