Ancient roaches further exemplify ‘no land return’ in aquatic insects
Abstract Among insects, 236 families in 18 of 44 orders independently invaded water. We report living amphibiotic cockroaches from tropical streams of UNESCO BR Sumaco, Ecuador. We also describe the first fossil aquatic roach larvae (6 spp.; n = 44, 1, 1, 1, 1, 1) from the most diverse tropical Mesozoic sediments (Middle Jurassic Bakhar Fm in Mongolia, Kimmeridgian Karabastau Fm in Kazakhstan; Aptian Crato Fm in Brazil), and the Barremian Lebanese and Cenomanian Myanmar ambers. Tropic-limited occurrences are trophic- (biomass/litter-fall), structural- (diversity) and also abiotic-factor-dependent (high temperatures). Diverse Paleozoic aquatic eoblattids are here (re)described from the lower Permian sediments of Elmo, U.S.A. and Chekarda, Russia. They competed with true cockroaches to reach water prior to the Mesozoic. Due to different evolutionary rates or periodical changes in water characteristics, non-adapted terrestrial insects repeatedly invaded the aquatic realm with well adapted hydrobionts. Obscurely, most aquatic lineages still survive. In contrast with Crustacea, aquatic-terrestrial reversal is absent. A single principal lineage, namely of moths, ancestral to butterflies (origination of modern insects from ephemerans and dragonflies is questioned), possibly evolved from insects with aquatic immature stages, and none from aquatic adults. The rest of the orders are terrestrial-derived. The proposed reason for the lack of land return is the character of numerous aquatic adaptations related to reductions, which are unlikely to be resuppressed. The aquatic insect family/terrestrial insect family ratio over time reveals a sharp rise from the Late Carboniferous to Late Triassic followed by lasting stability. Diversification of aquatic insects seems consistent with a 62.05 ± 0.02 Ma periodicity.