Phylogeny, classification, and biogeography of Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 the cycloteline Therevinae (Insecta: Diptera: Therevidae) STEPHEN D. GAIMARI* AND MICHAEL E. IRWIN Universig of Illinois and Illinois Natural History Suruty, 1101 West Peabod3, Drive, Urbana, Illinois 61801, U.S.A. Received September 1998: acceptedfor publication March 1999 Twenty-one members of the Laurasian ,group of Therevinae (Diptera: Therevidae) are compared using 65 adult morphological characters. Cladistic analysis using parsimony on the 17 ingroup and 4 outgroup taxa provides a well-supported hJpothesis of relationships among taxa within the Cyclotelini, tribe nov. The Cyclotelini is a monophyletic assemblage of mostly New World genera, including Atiolitga, gen. nov., Brtoiprrna Irwin, Coltiana, gen. nov., Crebraxta, gen. nov., Cyclotelus M'alker, .&Jonana, gen.nov., and Oeodireronlyia Bigot. In addition, three Old World genera, dtnniotherem Lyneborg, Bugulaverpa, gen. nov., and Aocyrlotelus Nagatomi & Lyneborg, are included in the tribe. These ten genera are divided into two monophyletic genus-groups, the Breoipenia-group and the Qclotelus-group. Keys are provided for the genera of Cyclotelini. The tribe, the two informal genus-groups, and all genera are diagnosed; five new genera and six new species are proposed. The biogeographical histories of the genera are discussed in terms of their cladistic relationships using methods of cladistic biogeography. Two major vicariant events account for the current distribution of the tribe. The first relates to the Beringian land bridge connecting western North America and eastern Asia. Second, New World cyclotelines were profoundly affected by the Early Eocene breakup of the archipelagic bridge between North and South America, and the distributions support the hypotheses favouring the continental origin of the Greater Antilles. 0 ?OOO The Linnean Society of London ADDITIONAL KEY WORDS:-Asiloidea ~ cladistics ~ systematics ~ morpholoLg - Laurasia - Beringia. CONTENTS Introduction ....................... 130 Historical perspective .................. 131 Methods ........................ 133 Terminolog. ..................... 134 Terminal taxa .................... 13-1 Character descriptions ................... 136 Head ....................... 136 * Corresponding author. Current address: Department of Entomology, National Museum of Natural History. Smithsonian Institution, Washington, DC 20560-0 169, U.S.A. E-mail: gaimari. [email protected] 129 0024-4082/00/060129+112 $35.00/0 0 2000 The Ihnean Socier) of London Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 132 S. D. GAIMARI AND M. E. IRWIN Macquart and Agapophytus GuCrin-Menkville; Cyclotelini lack, or have a small, inner gonocoxal process); male distiphallus often curved, extending posteroventrally beyond gonocoxites (widespread in Therevidae). Lyneborg (1984) suggested that attention be paid to the possible relationship of Ammothereva Lyneborg to Ozodiceromyia, due to many species sharing a combination Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 of male genitalic characteristics, e.g. aedeagus attached to subepandrial plate; aedeagus not attached to gonocoxal apodeme or ventral section of gonocoxite; gonocoxite without a free distal inner gonocoxal process; and ventral lobe strongly sclerotized and in an upright position, forming a guide for the distiphallus (an aedeagal guide, in this case). Although his assessment of the relationships of Ammothereva is consistent with the current analysis, in effect placing the genus within the Cyclotelini, most of the characters he used are not synapomorphic for the inclusive group, except for the lack of a free, distal inner gonocoxal process. In the description of Procyclotelus elegans Nagatomi & Lyneborg (1 987), the new genus ProgJclotelus Nagatomi & Lyneborg was suggested to be most closely related to Cyclotelus, a member of the New World group of genera including Cyclotelus, Ozodiceromyia, and Breviperna Irwin. They supported this inclusion based on the following features which they said characterize the group: male genitalia telescopically concealed within the abdomen, and the epandrium consequently shortened and with sharply projecting posterolateral corners; gonocoxites broadly fused ventrally; and midcoxa without pile on posterior surface. Of these characters, the fusion of the gonocoxites and the lack of pile on the posterior surface of the midcoxa proved to be synapomorphic for the Cyclotelini in the current analysis. An epandrium with sharply projecting posterolateral corners is rare in Ozodiceromyia, and is not characteristic of Cyclotelini. Nagatomi & Lyneborg (1989) later state that Procyclotelus has gonocoxites that are divided into two pieces by a median broad ventral ditch and the hypandrium is present, while in Breviperna, Cyclotelus, and Ozodiceromyia the gonocoxites are fused ventrally at least over the anterior third and the hypandrium is absent or is part of the ‘ventral synsclerite’. Our examination of three males of Procyclotelus elegans suggests that the gonocoxites are truly fused and are not divided into two pieces. The degree of gonocoxal fusion within Cyclotelus and Ozodiceromyia is highly variable among species, and there are many species of both genera that match the degree of fusion found in Procyclotelus elegans. In addition, cyclotelines rarely have the hypandrium so fully fused with the gonocoxites as to be visibly absent, and the level of hypandrial fusion with the gonocoxites found in Proyclotelus elegans is similar to that found in all other members. Lyneborg (1989) also suggested that the genus Iberotelus Lyneborg from Spain may be closely related to Cyclotelus and Ozodiceromyia. However, he acknowledged that the characteristic used in implying this relationship, the presence of a sclerotized ‘parameral-aedeagal bridge,’ is likely a part of the groundplan for the family. He noted that Iberotelus is the only therevine genus possessing this characteristic in the Palearctic, Afrotropical, and Oriental Regions. It should also be noted that not all members of Cyclotelus or Ozodiceromyia (or the other cycloteline genera) possess this plesiomorphic condition, and, in fact, Lyneborg (1984) considers the lack of this bridge a synapomorphy that aligns Ammothereva with Ozodiceromyia. Superficial resemblances among different therevid groups seem somewhat com- mon, despite very distant relationships. For example, three genera of Phycinae, Phycus, Pherocera Cole, and Salentia A. Costa, superficially resemble more apomorphic members of the Cyclotelini. In fact, the genus Ozodiceromyia was considered by Becker PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 133 (19 12) as a junior synonym of Phycus. Members of other genera of Therevinae can also be easily misplaced into the Cyclotelini due to superficial similarity. For example, species of Euptycus can be confused with members of Orodiceromyia, and one species, Orodiceromyia mexicana Bigot, was also described as Euphycus setosus Krober (19 12g). Species of Psilocephala such as Psilocephala munda Loew, species of Nebiitus Coquillett Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 such as Nebritus pellucidus Coquillett, and most species of the predominantly African genus Schoutedenomyia Krober, externally appear closely related to Orodiceromyia, with the most obvious similarities being the shiny black frons and abdominal tergites of the females in these genera. In all of these cases, examination of internal male and female genitalia and of informative external characters provides strong evidence that they are not closely related. Lyneborg (1976) noted the superficial similarity between Schoutedenomyia and Furcijira ( = Cyclotelus), pointing to various characteristics held in common, including body shape, antennal shape and vestiture, similar placement and appearance of the antennal style, lack of dorsocentral setae, wings without structural differences, and reddish-brown male terminalia telescoped into the abdomen. However, he also noted numerous fundamental differences, especially in the male genitalia. He noted this as an excellent example of parallelism, with the genera occupying the same ecological niches in tropical and subtropical environments of the Old and New World, respectively. It also seems likely that many of these genera are mimics of sphecoid wasps, further explaining the observed parallelism. METHODS Specimens were examined using a Wild/Leica MZ8 binocular dissecting micro- scope with a range of mapification between 6.3 and 50 x . A minimum of two male specimens and two female specimens was examined for each species, except for the single pair (male and female) of Bugulaverpa rebeccae, sp. nov., the single males of Crebraseta crassicornis (Bellardi) and Coleiana nigricopis, sp. nov., and the single specimen (male) of Iberotelus cinereus Lyneborg. Additional specimens were studied for several taxa to clarify states of certain characters. For each species, at least one male abdomen and one female abdomen were macerated in 20% KOH at room temperature for 8 hours. After the initial three hours, most of the pleural membrane of one