Improved Phylogenetic Analyses Corroborate a Plausible Position of Martialis heureka in the Ant Tree of Life

Abstract

Martialinae are pale, eyeless and probably hypogaeic predatory ants. Morphological character sets suggest a close relationship to the ant subfamily Leptanillinae. Recent analyses based on molecular sequence data suggest that Martialinae are the sister group to all extant ants. However, by comparing molecular studies and different reconstruction methods, the position of Martialinae remains ambiguous. While this sister group relationship was well supported by Bayesian partitioned analyses, Maximum Likelihood approaches could not unequivocally resolve the position of Martialinae. By re-analysing a previous published molecular data set, we show that the Maximum Likelihood approach is highly appropriate to resolve deep ant relationships, especially between Leptanillinae, Martialinae and the remaining ant subfamilies. Based on improved alignments, alignment masking, and tree reconstructions with a sufficient number of bootstrap replicates, our results strongly reject a placement of Martialinae at the first split within the ant tree of life. Instead, we suggest that Leptanillinae are a sister group to all other extant ant subfamilies, whereas Martialinae branch off as a second lineage. This assumption is backed by approximately unbiased (AU) tests, additional Bayesian analyses and split networks. Our results demonstrate clear effects of improved alignment approaches, alignment masking and data partitioning. We hope that our study illustrates the importance of thorough, comprehensible phylogenetic analyses using the example of ant relationships.

Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): a new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages

“…displaying a bizarre mixture of pleisiomorphic and autapomorphic traits, the species was attributed to its own subfamily, the Martialinae. this decision was supported by further morphological study (Brandão et al 2010) and multi-locus molecular phylogenetic reconstruction ; although see Kück et al 2011). Rabeling et al (2008) recovered Martialis as the sister to all remaining extant ants including the old World subfamily Leptanillinae, while a reanalysis by Kück et al (2011) found the converse.…”

“…Specimen was cleared by non-destructive extraction of dnA, which allowed for examination of internal characters. All males were examined from the following collecting events at the Fazenda Esteio study area of the Biological dynamics of Forest European Journal of Taxonomy 120: 1-62 (2015) Shared apomorphies of the basal ants of particular interest for the "basal ant" problem (Brady et al 2006;Rabeling et al 2008;Kück et al 2011) is the relationship of the Leptanillinae, Martialinae, and the Amblyoponinae. While the relationships within the formicoid clade (dorylinae, myrmeciomorphs, dolichoderomorphs, Formicinae, ectaheteromorphs, Myrmicinae) have crystallized in the past decade (Moreau et al 2006;Brady et al 2006Brady et al , 2014Ward et al 2015), the relationships of the "poneroids" (Agroecomyrmecinae, Amblyoponinae, Paraponerinae, Ponerinae, and Proceratiinae) are still unresolved (Ward 2014).…”

The Phylogeny and Evolution of Ants

“…The members of the subfamily Ponerinae have amassed considerable diversity, however, and some species are conspicuous and aggressive predators (Schmidt & Shattuck 2014). In molecular studies the poneroids appear as either a clade, sister to the formicoids, or a paraphyletic group within which the formicoids arise (Brady et al 2006, Moreau et al 2006, Rabeling et al 2008, Kück et al 2011, Moreau & Bell 2013.…”

“…These two lineages have vied for the position of sister group to all other ants. Different analyses of the same molecular data set have come to opposite conclusions (Rabeling et al 2008, Kück et al 2011, and analysis of a larger data set yielded an ambiguous outcome (Moreau & Bell 2013).…”

“…In the first comprehensive molecular The ant tree of life: a summary of well-supported relationships among the ant subfamilies, based on molecular phylogenetic analyses by Brady et al (2006Brady et al ( , 2014, Moreau et al (2006), Rabeling et al (2008), Kück et al (2011), Moreau & Bell (2013), and Ward et al (2014). Relationships among extant taxa are supported by Bayesian posterior probabilities in the range of 0.91-1.00 (mostly 1.00).…”