(Labes, Lainodon, and Valentinella). Labes and Lainodon are well-supported members of Zhelestidae, a stem eutherian clade, whereas Valentinella is more problematic, being recently considered as a nomen dubium. Based on X-ray computed microtomography scan analysis of the holotype and thanks to the discovery of new specimens from the type locality (Vitrolles-La Plaine, south-eastern France, late Maastrichtian), we reassessed Valentinella. This genus is unique by the association of an enlarged and rounded jaw angle with an assumed relatively elevated angular process, a bulbous protoconid and an unbasined heel on p4, a p5 with a wide molariform talonid and a hypoflexid, a robust molar morphology with a potential specialized crushing-grinding function (bulbously constructed cusps, large talonid, and horizontal apical wear facet of the hypocone), a somewhat reduced m3 relative to m2, a premolariform ?P3 or ?P4 lacking a metacone, and a relatively large hypocone on upper molars. These characters reinstate Valentinella as a valid genus. We also describe Mistralestes arcensis gen. et sp. nov. from a newly discovered locality (La Cairanne-Highway, south-eastern France, late Campanian). Mistralestes is defined by a robust premolariform p5 with no cingulid, paraconid, or metaconid; molars with a transverse protocristid, a gradual compression of the trigonid from m1 to m3, and paracristid and protocristid probably confluent on m3. Based on comparisons and phylogenetic analyses, Valentinella and Mistralestes may belong to Zhelestidae but this systematic attribution remains poorly supported.
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BACKGROUND: Many insects are chemically defended against predatory vertebrates and invertebrates. Nevertheless, our understanding of the evolution and diversity of insect defenses remains limited, since most studies have focused on visual signaling of defenses against birds, thereby implicitly underestimating the impact of insectivorous insects. In the larvae of sawflies in the family Tenthredinidae (Hymenoptera), which feed on various plants and show diverse lifestyles, two distinct defensive strategies are found: easy bleeding of deterrent hemolymph, and emission of volatiles by ventral glands. Here, we used phylogenetic information to identify phylogenetic correlations among various ecological and defensive traits in order to estimate the relative importance of avian versus invertebrate predation. RESULTS: The mapping of 12 ecological and defensive traits on phylogenetic trees inferred from DNA sequences reveals the discrete distribution of easy bleeding that occurs, among others, in the genus Athalia and the tribe Phymatocerini. By contrast, occurrence of ventral glands is restricted to the monophyletic subfamily Nematinae, which are never easy bleeders. Both strategies are especially effective towards insectivorous insects such as ants, while only Nematinae species are frequently brightly colored and truly gregarious. Among ten tests of phylogenetic correlation between traits, only a few are significant. None of these involves morphological traits enhancing visual signals, but easy bleeding is associated with the absence of defensive body movements and with toxins occurring in the host plant. Easy bleeding functions through a combination of attributes, which is corroborated by an independent contrasts test indicating a statistically significant negative correlation between species-level integument mechanical resistance and hemolymph feeding deterrence against ants. CONCLUSIONS: Our analyses evidence a repeated occurrence of easy bleeding, and no phylogenetic correlation including specific visual signals is significant. We conclude that the evolution of chemically-based defenses in tenthredinids may have been driven by invertebrate as much as by avian predation. The clear-cut visual signaling often encountered in the Nematinae would be linked to differential trends of habitat use by prey and predators. Further studies on (prey) insect groups should include visual signals and other traits, as well as several groups of natural enemies, to better interpret their relative significance and to refine our understanding of insect chemical defenses.
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