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Research Article the Hindwings of Ants: a Phylogenetic Analysis

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Research Article the Hindwings of Ants: a Phylogenetic Analysis Hindawi Psyche Volume 2019, Article ID 7929717, 11 pages https://doi.org/10.1155/2019/7929717 Research Article The Hindwings of Ants: A Phylogenetic Analysis Stefano Cantone and Claudio José Von Zuben Department of Zoology, Sao˜ Paulo State University, Rio Claro, SP, Brazil Correspondence should be addressed to Stefano Cantone; [email protected] Received 11 February 2019; Accepted 27 March 2019; Published 14 April 2019 Academic Editor: Jan Klimaszewski Copyright © 2019 Stefano Cantone and Claudio Jos´e Von Zuben. Tis is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. In this study, we compare and analyze diferent ant taxa hindwing morphologies with phylogenetic hypotheses of the Family Formicidae (Hymenoptera). Te hindwings are classifed into three Typologies based on progressive veins reduction. Tis analysis follows a revision of the hindwing morphology in 291 extant and eight fossil genera. Te distribution of diferent Typologies was analyzed in the two Clades: Formicoid and Poneroid. Te results show a diferent distribution of Typologies, with a higher genera percentage of hindwings of Typology I in the Clade Poneroid. A further analysis, based on genetic afnities, was performed by dividing the Clades into Subclades, showing a constant presence of hindwings of Typology I in almost all the Subclades, albeit with a diferent percentage. Te presence of hindwings of Typology I (hypothesized as more ancestral) in the Subclades, indicates the genera that could be morphologically more similar to their ancestral ones. Tis study represents the frst revision of the ants’ hindwings, showing an overview of the distribution of diferent Typologies. 1. Introduction in the Eocene epoch, most of the extinct species described have been classifed in extant genera or subfamilies, and wing Te wings in ants are present only in the winged reproductive morphology is similar or equal, representing an important caste and have the important function of promoting the characteristic in the identifcation of fossil winged forms and meeting between two sexes for mating. Wings are used widely used by various scientists. exclusively for the nuptial fight and afer mating winged In comparative studies on ant wings, more attention was Queens lose these structures and the winged male dies. given to the forewings, assuming an evolutionary history Te knowledge about the distance traveled in this fight is based on wing morphology [3–7, 9, 10]. Te diferent mor- unknown, but the loss of wings in the queens suggests a short phology of the wing veins is an important characteristic in distance of dispersion [1, 2]. the evolutionary history of the genera within the Subfamilies, Wing dimensions are directly proportional to body size, Tribe, or Group-genera identifed by phylogenetic hypotheses but vein structure has no relation to body size. In fact, wings based on molecular genetic analysis and sophisticated statis- with “complete” vein morphology are described in small tical analysis. body species and wings with reduced vein morphology in Te frst brief and incomplete analysis of the hindwings largeants[3–5].Tus,theevolutionarypathwayofthewing of ants was made by Kusnezov [11], who shows the diferent vein structure is independent of body size, representing an morphologies present in some genera. Recently, a broader important character in phylogenetic studies. In addition, in review was made by Cantone [3, 4], which classifes ant some genera described vein structure varies between species hindwings into three Typologies based on progressive wing and in some species in the forewing between males and vein reduction. Te objective of this study is to analyze queens, showing a venational evolution in progress [3, 4, 6– and compare the hindwing vein structure with phylogenetic 8]. Te stability of wing vein structure is also confrmed by the hypotheses, in the most ant genera, in order to present a constancy and permanence for millions of years as described distribution overview of the diferent hindwing morphologies in fossils from the Cretaceous and Eocene ants. In particular, in the family Formicidae. 2 Psyche Hindwing of Typology I Hindwing of Typology I with Jugal lobe 1 radial sector-media Radial sector 2 Radial sector 1 Costa Subcosta+Radius Radius 1 Basal Odontomachus bauri Subbasal Jugal lobe Media 2 Anal Media 1 Media+Cubitus cubitus-anal Cubitus 2 Anal 2 Anochetus sp. 62 Pachycondyla striata Ectatomma sp. 395 Paraponera clavata Hindwing of Typology I without Jugal lobe Eciton quadriglume Neivamyrmex sp. R1 Labidus coecus Nomamyrmex esenbeckii Acanthoponera sp. 405 Leptogenys sp. Pseudomyrmex gracilis sp. Hindwing of Typology I without Jugal lobe “azteca type” Azteca instabilis sp. Media 1+2 Figure 1: Hindwing of Typology I with Cells and Veins terminology. Te photos represent some examples of hindwings of Typology I; the dimensions are not comparable (photos by personal Cantone collection of Winged Ants). 2. Materials and Methods wing vein reduction. Te terminology of hindwings venation follows Yoshimura and Fisher [12] and Serna et al. [13]. Hindwings morphology was analyzed in 299 genera, of which Fossil specimen hindwings were examined from deposits 291 were extant and eight fossils. Tis analysis is based on the or ambers in Cretaceous and Eocene epochs and have study and revision of extant genera of winged males in 260 been based on genera belonging to the extinct Subfamilies genera [3] and winged Queens in 244 genera [4]. Hindwings Sphecomyrminae, Formiciinae, and on some extinct genera were classifed into three Typologies, based on progressive currently included in Incertae sedis. Psyche 3 Hindwing of Typology II Radial sector 1 Radial sector 2 Costa Subcosta+Radius Radius 1 Subbasal Basal Anal 1 radial sector-media Media+Cubitus Pseudomyrmex sp. 174 cubitus-anal Anal 2 Cubitus 2 Camponotus sp. 42 Nylanderia sp. 8 Formica exsecta Dolichoderus lamellosus Linepithema fuscum Linepithema humile Gnamptogenys sp. 336 Dorymyrmex pyramycus Anillidris bruchi Solenopsis sp. 209 Pogonomyrmex sp. 280 Hypoponera sp. 97 Pheidole sp. 199 Tranopelta gilva Wasmannia sp. 99 Acromyrmex sp. Procryptocerus sp. 382 Atta sexdens Figure 2: Hindwing of Typology II with Cells and Veins terminology. Te photos represent some examples of hindwings of Typology II; the dimensions are not comparable (photos by personal Cantone collection of Winged Ants). Te study and description of hindwing morphology 3. Results were based on the study of photos available on the AntWeb website and personal Cantone collection of 3.1. Hindwings Typologies Classifcation Winged Ants, as well as being based on the review of scientifc articles related to species descriptions. A Leica 3.1.1. Hindwings of Typology I (Figure 1). In this Typology, MZ8 stereoscope was utilized and hindwings photos hindwings have a more complete venation within the Family were taken in order to show some examples of diferent Formicidae. Basal and subbasal cells and media 2 vein are Typologies. always present. Tey difer in the presence/absence of the Te phylogenetic analysis is based on the comparison jugal lobe. Alternative morphology with media 1+2 vein, between hindwing Typologies and phylogenetic hypotheses that is, for the frst time, described and denominated as are based on molecular genetic analysis that divides the “azteca type”. Te hindwings of Typology I are present in family Formicidae into two Clade and into groups of genera some genera of Subfamilies Amblyoponinae, Dolichoderinae, [14–16]. Dorylinae, Ectatomminae, Heteroponerinae, Myrmeciinae, 4 Psyche Hindwing of Typology III 1 radial sector-media Subcosta+Radius Radius 1 Basal Radial sector2 Media+Cubitus Leptanilla Myrmicocrypta sp. Crematogaster sp. 6 Cyphomyrmex sp. 3 Figure 3: Hindwing of Typology III with Cells and Veins terminology. Te photos represent some examples of hindwings of Typology III; the dimensions are not comparable (photos by personal Cantone collection of Winged Ants). Ponerinae, and Paraponerinae. In the Subfamilies Myrmici- Amblyoponinae, Apomyrminae, Dolichoderinae, Dorylinae, nae and Pseudomyrmicinae only some species present hind- Leptanillinae, Martialinae, Myrmicinae, and Proceratiinae. wings of Typology I: Solenopsis bicolor Emery, described with Te hindwings of Typology III are described in 41 genera hindwings of Typology I “azteca type” [11, 17], representing [3, 4]. a rare and unique case in the subfamily Myrmicinae; Pseu- domyrmex gracilis, described with hindwings of Typology I 3.2. Hindwings of Extinct Ants in the Cretaceous and Eocene. without jugal lobe by Kusnezov [11], and some species of the Te oldest fossil ants were found in ambers or sediments of genus Tetraponera representing rare case in the subfamily the Late Cretaceous, between 110 and 75 million years ago Pseudomyrmecinae (see Figure 1). (Ma) in North America (Canadian amber ca. 78-79 Ma; New Te jugal lobe is present in some genera of the Subfamilies Jersey amber, ca. 94-90 Ma); in Botswana (Orapa, Tutorian Ponerinae, Ectatomminae, Myrmeciinae, and Paraponerinae. deposit ca. 90 Ma); in Russia (Siberia ca. 85 Ma); in France Te hindwings of Typology I with jugal lobe are described (Charentes, ca. 100 Ma); and in Myanmar (Burmese amber, in 38 genera, and the hindwings of Typology I without ca. 99 Ma). Te species in which the hindwings are described jugal lobe are described in 43 genera belonging to 10 were classifed in the extinct Subfamily Sphecomyrminae or Subfamilies (see Table 1). In seven genera of the Subfam- in some genera classifed as Incertae Sedis in the Cretaceous. ily Ponerinae the hindwings are not described; therefore, Hindwings are known from 10 species, of seven genera assuming to be of Typology I, they will be analyzed in both of Cretaceous ants. In nine species the hindwings are of genera with or without jugal lobe (see notes in Table 1), Typology I and the anal area is not visible, not showing the [3, 4]. presence/absence of the jugal lobe; the only case described by Perflieva [5, 18] is Armania robusta hindwing with jugal 3.1.2. Hindwings of Typology II (Figure 2).
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