B Leschen, editors. We have supplied a higher resolution image of this tree in the revised paper. Convergence was judged to have occurred when the standard deviation of split frequencies dropped below 0.005, and by ESS values higher than 200 in Tracer v1.7.0 (Rambaut et al., 2018), indicating adequate estimation of the posterior. We are aware that molecular dating studies imply that crown-group ants may have existed at this time, but because they are simply not known in thousands of ant inclusions in Burmese amber, we felt it highly unlikely that their myrmecophiles would make an appearance but they would not. Biological sources of discordance include incomplete lineage sorting (ILS) and introgression, but only ILS has received theoretical consideration in the context of hemiplasy. [Editors’ note: the author responses to the first round of peer review follow.]. Check out our clown beetle selection for the very best in unique or custom, handmade pieces from our shops. Their main argument is that extant crown-group ants are not represented in Burmese amber. For the latter three characters, Caterino and Tishechkin, 2015, said the modified gland openings involved occur only in some species groups of Operclipygus (Exosternini). Ventrites 2‒four equal in length, without posterior marginal striae. Martins-Neto, R. G. (1991). (I admit I did not examine Caterino and Tishechkin, 2015's long character list in detail to search for those.). Hind tibia flat, paddle-like, about as broad as femur (Figure 1C), with a row of strong spines along external edge, double rows of stiff and apically modified setae along inner edge, inner surface with transverse ridges and row of pointed spines before; apical spur small. For a more robust representation of the putuative sister genus of Promyrmister, we have added a Palaearctic species of Haeterius (H. ferrugineus) the type species of the genus, and scored it for all the characters. However, based on such analyses for other myrmecophile taxa, rapid evolution of specialized symbiotic phenotypes appears to be a common feature to clades of social insect symbionts (Moore and Robertson, 2014; Parker and Grimaldi, 2014), and presumably results from intense selection pressures inside colonies (Kistner, 1979; Parker, 2016). Species with such adaptations are found among the scarabs, ground beetles, and clown beetles (family Histeridae). Furthermore, the most recent estimates based on molecular data date the origin of crown-group ants to ca. The ages of crown-group ants as a whole, New World (NW) army ants, and other specific subfamilies and genera that are known hosts of Haeteriinae are also presented. Our fossil provides a minimum age, but future molecular dating of groups like Haeteriinae will be illuminating. We apologize, but the specific apomorphies shown in Figure 2—figure supplement 2, were not easily visible, e.g. We posit that the overwhelming prevalence of stem-group ants in Burmese amber implies that they were potential hosts of Promyrmister (Figure 2F). Whether haeteriines evolved in stem- or crown-group ant colonies, their original hosts are presumably long-extinct. 0 0 1 0 0 0 0. Find high-quality stock photos that you won't find anywhere else. That such an organism could have existed in the earliest-known ant societies in the Cretaceous is remarkable. However, I couldn't escape the impression that the discovery is being oversold. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix ‘http://zoobank.org/’. Classic comparative methods do not account for discordance, resulting in incorrect inferences about the number, timing, and direction of trait transitions. Molecular dating implies crown-group ants existed at the same time as Promyrmister, but stem-group ants are the only ants so far known in Burmese and other contemporaneous ambers. Morphological terminology follows Ślipiński and Mazur (1999), Zhou et al. Of all animal groups, however, the complex societies of ants play host to the greatest diversity of behavioral symbionts. They pass through 4 different body forms in their complete life cycle. However, trait evolution along discordant gene trees can also generate these incongruences – a phenomenon known as hemiplasy. Fossil ants (Hymenoptera: Formicidae): ancient diversity and the rise of modern lineages, Corrections of: Barden, P. 2017: Fossil ants (Hymenoptera: Formicidae): ancient diversity and the rise of modern lineages, Biological diversity of ants and their role in ecosystem function. Beetle fossils could in principle be associated with ants in three ways I guess. First, the fossil record is incomplete, and absence of extant ants in the fossil record therefore doesn't mean that they weren't there. The hind legs of some beetles, such as Flea Beetles (within Chrysomelidae) and flea weevils (within Curculionidae), are enlarged and designed for jumping. We also included representatives of four other tribes: Omalodini, Histerini, Hololeptini and Platysomatini, and assigned Hister unicolor as the primary outgroup, following Caterino and Tishechkin (2015). The metallic green color is what sets this clown beetle apart from other clown beetles. From the original matrix, we selected all taxa from the nearest sister clades of Haeteriinae, including 35 taxa belonging to Exosternini, including all species of Yarmister (apparently the closest genus to Haeteriinae; Caterino and Tishechkin, 2015). Scutellum obscured dorsally by secretion (Figure 1A). Complex interspecies relationships are widespread among metazoans, but the evolutionary history of these lifestyles is poorly understood. My only comments for the authors to consider are minor ones: Results and Discussion: 'mid-Cretaceous' as anon formal date is here confusing as the age of the deposit make sit Cenomanian (technically "Upper Cretaceous"), and absolute age place sits in the top 1/3 of the Cretaceous age span, so Upper cretaceous is more preferable than using' mid-Cretaceous'. Mesquite: A Modular System for Evolutionary Analysis. 100 million year old amber from Myanmar. Similarly, Caterino and Tishechkin, 2015, illustrated state 12-2 with figures of two species of Baconia (Exosternini), so again this is not truly a unique apomorphy of Sphecomyrmister. Several major radiations of myrmecophiles are known, each containing hundreds of symbiotic species, including the lycaenid butterflies (Pierce et al., 2002), eucharitid wasps (Murray et al., 2013), paussine ground beetles (Moore and Robertson, 2014) and multiple lineages of rove beetles (Kistner, 1979; Kistner, 1982; Parker, 2016; Parker and Grimaldi, 2014; Seevers, 1965; Maruyama and Parker, 2017). -The body size is usually between 1/8 - 3/8'' long. The beetles have been recorded in colonies of ant species scattered across the subfamilies Dolichoderinae, Dorylinae, Formicinae, Myrmicinae and Ponerinae (Helava et al., 1985; Tishechkin, 2007) (Figure 3). (112-2) and (113-2) and were misread by the reviewer because of the low quality of the file. We and others have previously described rove beetles (Staphylinidae) in the Burmese palaeofauna that were putative symbionts of termite colonies (Yamamoto et al., 2016; Cai et al., 2017). We hope this addresses your comment adequately. The evolutionary scenario described (subsection “Sphecomyrmister and deep time persistence of a social symbiosis", fourth paragraph) posits that these beetles originally evolved with stem-group ants and then switched to crown-group ants later. We scored Promyrmister for 259 external morphological characters used by Caterino and Tishechkin (2015) in a study investigating relationships among the tribe Exosternini, which is closely related to Haeteriinae. Stag Beetle / Fliegender Hirschkäfer! The name refers to the likely symbiotic habits of the fossil taxon inside early ant colonies. Additionally, Promyrmister possesses clear postcoxal secretory glands (Figure 1H; Figure 1—figure supplement 2A,B,D), with a globule of possible exudate emanating from the postcoxal gland opening on the right side of the body (Figure 1H; Figure 1—figure supplement 2B,D). Lesser silver water beetle (Hydrochara caraboides) Lesser silver water beetle (Hydrochara caraboides) larvae . The environments in which many hister beetles species are known to live. The implications are broad for the evolution of social interactions, early colony formation in ants, and symbiotic relationships between animals more generally. It then enters into the ‘pupal stage’ which can take up to 9 months and usually happens over the winter period. Such a scenario entails that haeteriines may not have originated with the modern ant groups that host them today; instead myrmecophily evolved first in stem-group ant colonies, with the beetles later switching to crown-group ants. Unlike reviewers 1 and 3, reviewer 2 also argues that previously-described ‘myrmecophile-looking’ histerids reduce our paper’s novelty. They also suggest new avenues for improving influenza control. The earliest examples of ancestral ants are found encapsulated in 99-million-year-old amber from a mine in northern Myanmar (Burma). The first 25% of trees were discarded as burn-in. 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