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 side of each abdomen was teased apart to allow internal tissues direct contact with KOH. After maceration, the specimens were soaked in distilled water for 10 minutes and placed into 75% ethanol. Male terminalia were placed into glycerin for dissection of genitalia and microscopic examination. Female abdomens were carefully opened laterally by teasing apart all remaining pleural membrane, including that of segment 8. Several drops of a 40% solution of chlorazol black (chlorazol black powder dissolved in 75% ethanol) were introduced into the opened abdomen using the tips of a fine forceps. The abdomens were then rinsed with 75% ethanol and checked to determine if spermathecal ducts and sacs were dyed blue. If not, the specimen was remacerated for several hours or the tissues were teased apart to expose the ducts and sacs, after which another several drops of chlorazol black were placed in the apical part of the abdomen. When the ducts were sufficiently stained, the specimen was rinsed in 75% ethanol and placed into glycerin for microscopic examination. Further dissection of these fragde structures consisted only of teasing tissues with minuten probes and fine 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 I36 S. D. GAIMARI AND M. E. IRWIN synapomorphy supports the clade of these Laurasian groups as separate from the Gondwanan Anabarhynchus- and ,4gapophytus-groups: inner gonocoxal process of males not fully articulated (full articulation of this process is considered plesiomorphic). Species were chosen to represent an assemblage of groups within the Laurasian
Therevinae thought to be close to the Cyclotelini, but the relationships among these Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 genera are as yet speculative and under investigation.
CHARACTER DESCRIPTIONS
In this section, we describe the morphology and taxon distribution of states for the 65 characters used in this analysis. Of these, 15 are in the head, 18 are in the thorax, 26 are in the male terminalia, and 6 are in the female terminalia. Most are binary, with few showing three states: 11, 15, 48, 51, 52, and 60, and four states: 8 and 9. Character distributions are discussed relative to the presented cladogram (Fig. 1).
Head
1. Antennal length 0 shorter than or subequal to head length 1 longer than head Antennal length was measured from dorsal view, and the length of the head was measured from lateral view. Plesiomorphically, the antenna is the same length or shorter than the head (e.g. Fig. 33), as is the case for all outgroup taxa and most Therevidae. A specimen with antennae >20% longer than the head (e.g. Fig. 27) was scored as having the apomorphic state. This character and the following two supported no clades in the current analysis. However, the three characters may be useful when considering species level relationships. For example, several species with elongate antennae within Ozodiceromyia are closely related to Ozodiceromyia argenttjra (Gaimari & Irwin, 2000) (e.g. Figs 26 and 27). The same holds true for several species within Qrlotelus being related to Cyclotelus beckeri (e.g. Figs 36 and 37). The apomorphically elongate antenna was only found in these two terminal ingroup taxa, but has arisen independently in many genera of Therevidae.
2. Antennal scape width 0 same thickness or thinner than first flagellomere 1 distinctly thicker than first flagellomere The scape and first flagellomere are measured through their broadest widths in dorsal or lateral view. In the plesiomorphic state, as represented by 3 of 4 outgroup taxa and most other Therevidae, the scape is the same thickness as (e.g. Fig. 29) or thinner than (e.g. Fig. 13) the first flagellomere. To be scored as possessing the apomorphic state, the scape is >20% wider than the first flagellomere (Fig. 23), which is autapomorphic for Crebraseta.
3. Medial surface of pedicel 0 without setae 1 setose 2334665 Dialineura gorodkovi ndivirilia limata
Brachylinga cinerea 2 Ammothereva lehri c Ammothereva laticornis 5 Nesonana youngi % U
Breviperna placida 0 W 0 Anolinga psilofrons inga melanothrix E L Cyclotelus kroeberi Figure 1. Single most parsimonious cladogram for the Cyclotelini, showing character state changes under ACCTRAN character optimization. Characters are - numbered as in the tent; hash marks are as follows: black=fonvard change with no homoplasy; white=change with homoplasy. w< Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 June 24 on guest by https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 from Downloaded 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 140 S. D. CAIMARI AND M. E. IRWIN Plesiomorphically, the upper frons of the female has scattered setae, at most with a bare area along the midline (e.g. Figs 8 and 15), as is the case for all outgroup taxa and the Breviperna-group. As a synapomorphy for the Cyclotelus-group, setae are absent or are in defined patches or patterns, usually in one or more vertical lines (e.g. Fig. 23) or in a small cluster medial to the eye margin (e.g. Fig. 27). Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 11. Setae of 0 lower3ons 0 . scattered 1 in defined patches or patterns 2 absent Usually, the lower frons bulges forward beyond the upper frons in lateral view; otherwise, the lower frons is delimited above by a change in direction of the inner eye margin. This character is scored on the basis of presence of setae just above the antennal base margin. As in character 10, the lower frons of the female has scattered setae (e.g. Figs 15 and 23) plesiomorphically, as is the case for 3 of 4 outgroup taxa. Apomorphically, there are setae in defined patches or patterns (state l), usually a small cluster dorsolateral of the antennal base margin (e.g. Fig. 27), as is synapomorphic for the Ozodicemmyia-subgroup. The lack of setae or pile above the antennal base margin (state 2) (e.g. Figs 8, 19, and 33) is widespread within the Cyclotelini, in both the Breviperna-group and the Cyclotelus-subgroup, and likely arose at the base of the tribe. Brevipenza placida and Crebraseta crassicornis each display scattered setae. 12. Maxillary pa& colour 0 dark brown or black 1 orange Plesiomorphically, the maxillary palp is dark, as is the case for all outgroup taxa and for the Cyclotelus-group. The palp is apomorphically orange, as is synapomorphic for the Breviperna-group. Members of the Cyclotelus-group rarely have an orange palp, including few species of Ozodiceromyia not included in these analyses. 13. Gena 0 not protruding 1 protruding anteriorly beyond edge of eye Plesiomorphically, the gena is not visible from lateral view, as is the case for all outgroup taxa. This character was scored as protruding if the gena was visible below the eye margin from the lateral view as a raised convexity, ranging from small (e.g. Fig. 10) as in Ammothereva to large (e.g. Fig. 23) as in the Ozodiceromyia-group. The protruding gena is synapomorphic for the Cyclotelini, with a subsequent reduction in Coleiana (e.g. Figs 32 and 33). 14. lip ofgena 0 same texture and colour as remainder of gena, postgena, occiput, and parafacial 1 different texture or colour than remainder of gena, postgena, occiput, or parafacial As is the case for all outgroup taxa and most Therevidae, the genal apex is PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 141 plesiomorphically of the same texture and colour as the remainder of the gena, postgena, occiput and/or parafacial (e.g. Figs 14 and 15). In the apomorphic state, a patch of different texture or colour (e.g. Fig. 35) is present, usually as distinctly darker pruinescence, but occasionally consisting of velutinous pruinescence. This is synapomorphic for the Cyclotelus-group, with a subsequent change to state 0 in Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Coleiana. 15. Pile Ofgenal apex 0 not different from pile of postgena or lower occiput 1 forming distinctive patch of shortened pile 2 absent As is the case for 3 of 4 outgroup taxa, the genal apex plesiomorphically has setae or pile that is similar to that of the postgena or lower occiput. A dense patch of short, darkened setae on the genal apex (state 1) (e.g. Figs 20 and 23) is synapomorphic for the Ozodicmmyia-subgroup and autapomorphic for Anolinga me- lanothrix, while having the apex without pile (state 2) (e.g. Fig. 8) is otherwise synapomorphic for the Cyclotelini. Thorax 16. Clej ofthe prosternum 0 pilose 1 lacking pile Pile on the cleft of the prosternum has been used as a key character for sorting genera of therevids (Irwin & Lyneborg, 1981a,b). Plesiomorphically, as is the case for 3 of 4 outgroup taxa, the Ozodicmmyia-subgroup, and most of the Brevipaa- group, prosternal pile is present in the cleft. Apomorphically, the prosternum lacks pile in the cleft, as in the Cyclotelus-subgroup, the Ammothreua gussakovs,$+group, and Anolinga psi10jbn.s. 17. Proepimmn 0 pilose 1 lackingpile The upper half of the proepimeron is pilose plesiomorphically, as in 2 of 4 outgroup taxa and many other therevid genera. The lack of pile on the proepimeron (e.g. Figs 40 and 41) occurs throughout the Cyclotelini, except Brevipema, and is herein considered apomorphic. 18. Katepisternum 0 pilose 1 lacking pile As in 3 of 4 outgroup taxa, the katepisternum is pilose plesiomorphically. Apomorphically, this pleurite lacks pile (e.g. Fig. 40), as. is the case for the Cyclotelini, with subsequent changes to state 0 in Brevipaa, part of Anolinga, and the Ozodiceromyia- subgroup. In the Ozodiceromyia-subgroup, this pile is located along the median vertical convex crest of the pleurite (e.g. Fig. 41). 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 144 S. D. GAIMARI AND M. E. IRWIN pale macrosetae is synapomorphic for Ammothereva, although a few species within the genus, such as Ammothereva mongolica (Zaitsev) and Ammothereva splendida (Krober), have darker, nearly black macrosetae. In addition, a few species of Ozodiceromyia that were not included in the analyses also have pale macrosetae, as do other unrelated genera, such as Arenigena Irwin & Lyneborg and Ammonaios Irwin & Lyneborg. This Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 state is quite uncommon among therevids, and is certainly apomorphic within specific groups. 28. .Notal texture 0 fully pruinose 1 with reduced pruinescence As is found in all outgroup taxa and the Breviperna-group, the notum is plesio- morphically fully pruinose, with no areas that are reduced to the point of exposing ground colour. Apomorphically, a reduction of pruinescence on the notum exposes some ground colour. Most often, the lateral notum or the postpronotal lobe displays this reduced pruinescence, although the reduction can be much more extensive. This is a synapomorphy for the Cvclotelus-group. 29. Patches of matte pruinescence on notum 0 absent 1 present As is found in all outgroup taxa and most Therevidae, the notum plesiomorphically lacks patches of matte pruinescence. There are patches of matte pruinescence on the notum as a synapomorphy for Coleiana+ Cyclotelus. These patches are dull black or brown in Cyclotelus and dark brown in Coleiana, and are usually located on the mesonotum between the midline and lateral edge. 30. Scutellar setae 0 2 pairs 1 0-1 pair As in all outgroup taxa and most Cyclotelini, two pairs of scutellar setae are present plesiomorphically. Apomophically, 0-1 (usually 1) pair of scutellar setae are present, as is autapomorphic for Bugulaverpa. In addition, the South American clade of Cyclotelus beckeri+ Cyclotelus kroeberi is supported by the apomorphic condition, and this may be an important state in the generic revision of Cyclotelus underway. 31. Supra-alar setae 0 2 pairs 1 1 pair As is found in all outgroup taxa and most Cyclotelini, two pairs of supra-alar setae are present. Apomorphically, one pair of supra-alar setae is present, as is autapomorphic for Bugulaverpa. In addition, the South American clade of Cvclotelus beckeri+ Cyclotelus kroeberi is supported by the apomorphic condition, and this may be an important state in the generic revision of Cyclotelus underway. 32. Ventral surface of basal lobe ofcosta 0 with pile over entire ventral surface 1 bare, or with short pile only basally PHIZOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 145 As is the case for all examined outgroup taxa, the entire ventral surface of the basal lobe of the costa is densely covered with black or white pile (e.g. Fig. 48). Apomorphically, pile is absent or short pile is sparsely distributed only at the base of this lobe (e.g. Fig. 49). All Cyclotelini except Breviperna display this apomorphic condition, and it is considered a synapomorphy of the tribe. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 33. Basal lobe of the costa with dorsal setae 0 scattered over entire surface 1 in two distinct rows The dorsal surface of the basal lobe of the costa has short, black setae that extend into the inner and outer rows of costal setae. In the plesiomorphic condition, as is the case for 3 of 4 outgroup taxa and the Breviperna-group, these setae are scattered over the entire dorsal surface (e.g. Fig. 50) and do not form distinct lines leading into the inner and outer setal rows of the costa. In the apomorphic condition, as found in most members of the Cyclotelus-group (except Procyclotelus and Crebraseta), there is an upper row along the outer edge of the lobe that leads to the anterior row of costal setae, and a lower row along the base of the lobe that leads to the posterior row of costal setae (e.g. Fig. 51). A distinct linear bare area exists between these upper and lower rows. Male teminalia 34. Lateral ejaculatory process 0 incomplete dorsally 1 notched or complete dorsally Plesiomorphically, the lateral ejaculatory process is incomplete dorsally and is composed of two distinct sclerites, as is found in all outgroup taxa. Apomorphically, the process is complete dorsally (e.g. Fig. 135) or is at most notched along the anterior edge (e.g. Fig. 133). This notch can be deep, but the lateral ejaculatory process is distinctly a single structure. This is a synapomorphy for the Cyclotelini, but some members of the Ammothereva poecilopa-group display state 0. 35. Lateral ejaculatory process 0 set into aedeagus 1 separated and external Plesiomorphically, the lateral ejaculatory process is sunken into the aedeagus (e.g. Fig. 132), as in all outgroup taxa. Apomorphically, the lateral ejaculatory process is large and appears completely separated from the aedeagus anteriorly, as in Breviperna (Figs 118 and 1 19), Procyclotelus (Figs 136 and 137), and some Ammothereva such as Ammothereva nigrofmorata (Krober), although it is held in place with muscle and membrane. 36. Relative orientation of lateral margins of dorsal apodme 0 parallel, or slightly convergent anteriorly 1 divergent anteriorly All outgroup taxa display the plesiomorphic condition of having parallel (e.g. Fig. 133) or slightly converging (e.g. Figs 11 1 and 113) lateral margins of the dorsal 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 148 S. D. GAIMARI AND M. E. IRWIN 45. Postemlateral comers of epandrium 0 flange-like and without setae, at least distally 1 flange absent, setose over posterolateral corners As in all outgroup taxa and most genera of Cyclotelini, the epandrium is plesiomorphically dorsoventrally flattened and flange-like at the posterolateral cor- Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 ners, which lack setae (e.g. Fig. 69). Apomorphically, the posterolateral corners are not flattened flanges, and setae are present over the entire posterior epandrium, including the posterolateral corners (e.g. Fig. 65); this is synapomorphic for the Ozodiceromyia-subgroup. Even when the flange is absent, the posterolateral corners of the epandrium can be pointed (e.g. Fig. 65). 46. Anterior e&e of epandrium 0 emarginate 1 not emarginate According to 3 of 4 outgroup taxa, the plesiomorphic condition is to have an epandrium with an emarginate anterior edge in dorsal view (e.g. Fig. 62). Apomorphically, this anterior edge is not emarginate in dorsal view (e.g. Figs 63 and 68), as is the case for much of the Cyclobhs-group, but notably not for the Old World members. 47. Epandrial setae 0 covering more than two-thirds of epandrium 1 covering only posterior half of epandrium As in all outgroup taxa and most therevids, setae covering more than two-thirds of the epandrium (e.g. Fig. 68) is plesiomorphic. The condition of having setae covering at most the posterior half of the epandrium (e.g. Fig. 64) is a synapomorphy for Ozodicemmyiu. 48. Fusion ofgonocoxites 0 completely free 1 fused by sclerotized membrane between discernible gonocoxites 2 strongly fused Plesiomorphically, the gonocoxites are free in Therevidae, including all outgroup taxa. Although gonocoxal fusion occurs in other groups of Therevidae, this condition is synapomorphic for the Cyclotelini. Most commonly, the gonocoxites are completely fused (state 2) (e.g. Fig. 93). The length of the area of fusion between the two halves ranges from long, as in Coleiana (Fig. 99), to short, as in Aocyclotelus (Fig. 97). The gonocoxites can also be seen as distinct, but with a sclerotized sheet between (state 1) (e.g. Figs 72, 73, and 75), as in Ammotheva. 49. Hypandrium 0 free or absent 1 fused to gonocoxites The plesiomorphic condition, as is found in all outgroup taxa, is for the hypandrium to be free and separate from the gonocoxites or absent. The condition of having the hypandrium fdly fused to the gonocoxites (e.g. Figs 93 and 103)is a synapomorphy for the Cyclotelini. Although the hypandrium is fused to the gonocoxites, it is rarely PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 149 indistinguishable. In addition, examination of male genitalic musculature (following terminology of Ovtshinnikova, 1989), and (Ovtshinnikova & Yeates, 1998) of Ozodiceromyia nanella verified that the hypandrial muscle M 18, (and therefore the hypandrium itself) is present despite full fusion with the gonocoxites. In some unrelated groups (e.g. Chromolepida), the hypandrium can form a sclerotized bridge Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 attaching the two otherwise unfused gonocoxites. 50. Outer gonocoxal process 0 strongly everted 1 small fold, flattened flange, or absent The outer gonocoxal process is considered large and everted if it projects posteriorly by at least half the length of the gonocoxite itself (e.g. Figs 77, 78, 103, and 104). The plesiomorphic condition of having this strongly everted outer gonocoxal process is present in 2 of 4 outgroup taxa and nearly all genera of cyclotelines. The apomorphic state of lacking an outer gonocoxal process or having a small fold or flattened flange (e.g. Figs 91 and 92) is characterized by a more delicate appearance (if present) and a length shorter than half that of the gonocoxite (e.g. Fig. 94). The reduced outer gonocoxal process is synapomorphic for Ozodiceromyia. 51. Set& of the outer gonocoxal process 0 similar to that of gonocoxites 1 reduced density and size, or absent 2 dense and fine The outer gonocoxal process (or the corresponding edge of the gonocoxite in those lacking the process) having setae similar to those of the gonocoxites (e.g. Figs 76, 82, and 104) represents the plesiomorphic condition, which is found in 2 of 4 outgroup taxa and most cycloteline genera. As a synapomorphy for the Ozodiceromyia- subgroup, the process or edge has, at most, only a few short setae that are smaller and finer than other setae of the gonocoxites (state 1) (e.g. Fig. 94). As a synapomorphy for Anolinga, the gonocoxal process is densely covered with fine setae, which are finer and more densely distributed than other gonocoxal setae (state 2) (e.g. Fig. 85). 52. Inner gonocoxal process 0 strongly projecting and articulated 1 strongly projecting and not articulated 2 neither strongly projecting nor articulated The inner gonocoxal process (see Ovtshinnikova & Yeates, 1998) is strongly projecting and articulated in 1 of 4 outgroup taxa and in the Gondwanan Therevinae not included in this analysis (Anabarhyuhus-group and Agapophytus-group). The ar- ticulation is apomorphically lost, but the process is still strongly projecting in 2 other members of the outgroup. The inner gonocoxal process is reduced to a knob (e.g. Fig. 96) or is absent (e.g. Fig. 74) as is the case for the Cyclotelini and for the nearest outgroup taxon, Brachylinga cinerea. 53. Apical setue of inner gonocoxal pmcess 0 present 1 absent 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 152 S. D. GAIMARI AND M. E. IRWIN of the spermathecal and central sac ducts (e.g. Figs 155 and 152), as is the case in 3 of 4 outgroup taxa. Having the origin of these ducts at the base of the central sac (e.g. Fig. 148) is synapomorphic for Nesonana +Anolinga + Breviperna, and is present within Ozodiceromyia (e.g. Fig. 153). Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 63. Basal portion of individual spemnathecal ducts 0 same width as, or narrower than, common spermathecal duct 1 wider than common spermathecal duct As in 3 of 4 outgroup taxa and most Cyclotelini, the basal portion of the spermathecal duct is plesiomorphically the same width or narrower than the common spermathecal duct (e.g. Fig. 153). Apomorphically, the basal portion of the spermathecal duct is wider that the common spermathecal duct (e.g. Fig. 156). This widened spermathecal duct is a synapomorphy for Cyclotelus. 64. Anterior furcal membrane 0 lacking a furcal bulla 1 with a furcal bulla Nearly all Therevidae considered except Cyclotelini plesiomorphically lack a furcal bulla on the anterior part of the furcal membrane, which is instead flattened. The two exceptions (Lysilinga aurantiuca (Coquillett) and an undescribed genus from Mexico) have a dorsal extension of the furca that appears directly associated with a dorsal lobe on the 8 distiphallus. The presence of a furcal bulla is synapomorphic for the Cyclotelini, with a subsequent change to state 0 in Bugulaverpa (Fig. 150). This structure, which can be lightly to heavily sclerotized, ranges in size from large and distinct (e.g. Fig. 148) to a small convexity along the anterior furcal edge (e.g. Fig. 147). 65. Dorsocentral connection oftergite 8 with tergite 9 0 setose 1 lacking setae Among Laurasian Therevinae, the ribbon-like dorsocentral connection between tergites 8 and 9 is plesiomorphically setose, as in 3 of 4 outgroup taxa. These setae are absent, leaving the structure bare, as a synapomorphy for the Cyclotelini. This apomorphy is very consistent within the Cyclotelini, but is widespread in more distantly related therevids, indicating a possible reversed polarity when considering higher level phylogenetic relationships. CLADISTIC ANALYSIS AND RESULTS Phylogenetic analyses follow the cladistic philosophy of parsimony promoted by Hennig (1966) and other workers (e.g. Wiley, 198 1; Farris, 1983). Following Kluge (1989), the relative informativeness of characters is not prejudged by excluding characters that seem to be in opposition, and so all evidence available to us was analysed. In addition, following Yeates & Irwin (1996), characters were not excluded due to their perceived level of homoplasy, agreeing that this would amount to an a priori exclusion of evidence for some possible hypotheses of relationship as a method PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE I53 for protecting a preconceived hypothesis prior to parsimony analysis. Following Yeates (1992), autapomorphies were left in for analyses, as these character states may be found to be more widespread and may support clades in undiscovered or not included relatives of the terminal taxa. Outgroups were used to determine character polarities (Watrous & Wheeler, Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 1981; Farris, 1982; Maddison, Donoghue & Maddison, 1984; Nixon & Carpenter, 1993). Cladistic analyses were performed using PAUP version 3.1.1 (Swofford, 1993) and HENNIG86 version 1.5 (Farris, 1988) using the data matrix presented in Appendix 2. All characters were treated as non-additive, agreeing with Hauser & Presch (1991) and Hauser (1992) that congruent characters in a matrix are the evidence for the order of transformation series for multistate characters. A priori rejection or down-weighting of other possible transformation series can be used as an ad hoc assumption to protect a preconceived hypothesis, instead of using congruence with other characters to test ordering hypotheses with all possibilities equally available. PAUP analysis followed a random addition sequence order of 100 iterations, utilizing the non-exhaustive (heuristic) tree bisection-reconnection (TBR) branch swapping algorithm. HENNIG86 analysis was performed using the ‘m*; bb*’; commands (i.e. branch breaking algorithm). Cladogram figures displaying character state changes were prepared using CLADOS version 1.6.1 (Nixon, 1998) under ACCTRAN character optimization (Swofford & Maddison, 1987). Although Yeates & Irwin (1 996) considered DELTRAN optimization to be more conservative due to its biases against single origins of characters (Wheeler, Cartwright & Hayashi, 1993), we consider ACCTRAN to be more conservative, based, in part, on the arguments of Pinna (199 1) that primary homologies should be considered valid until parsimony positively falsifies them. Both analyses produced one most parsimonious tree (Figs 1 and 2) of length 166, Consistency Index (Kluge & Farris, 1969) of 0.45 (0.44, excluding autapomorphies), Retention Index (Farris, 1989) of 0.69, and Rescaled CQnsistency Index (Farris, 1989) of 0.3 1. The topology remained stable through succkive weighting (Farris, 1969; Carpenter, 1988) as implemented by HENNIG86 (i.e. m*; bb*; xs w; cc;) and PAUP (by the maximum value of the Rescaled Consistency Index, with a base weight of 1000), levelling out at a single iteration. Because the expected amount of homoplasy increases with the number of taxa in a parsimony analysis, the regression equation of Sanderson & Donoghue (1989: 1785) was used to calculate an expected value for the Consistency Index. The calculated value of 0.53 for 21 taxa is greater than the value indicated in the current analysis (0.45), so there is more homoplasy than expected for an analysis with this number of taxa under the 1989 model. The relatively high Retention Index indicates that a high proportion of the potential synapomorphies in the data matrix are present as homologies on the cladogram. Xode support Assessing cladogram or node support is a difficult endeavour, especially with the variety of available, standard methods which do not necessarily reflect cladogram or node support. Of the methods discussed below, only Bremer support (Bremer, 1988, 1994; Donoghue et al., 1992; Kallersjo et al., 1992) (decay index, branch support) is founded upon utilizing the entire original data set throughout, rather 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 156 S. D. GAIMARI AND M. E. IRWIN should provide a better estimate for the amount of additional realistic character data that would be necessary to refute a node on the cladogram. The rescaled Bremer support indices are displayed on the cladogram (Fig. 2) along with the unweighted Bremer support indices. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 CLASSIFICATION The structuring of the internal classification of the Therevidae is in its infancy. Not until recently (Irwin & Lyneborg, 198 1a) was the family even divided into two subfamilies, Therevinae and Phycinae. Irwin & Lyneborg (198 la) also provided the first classification of known Nearctic species, delimiting genera and changing the combinations of species based on putative synapomorphies, but gave no insight into the suprageneric groupings below subfamily. As presented in the historical perspective, other workers have published opinions on relationships of various genera to Ozodicmmyia and Cyclotelus, but have not attempted to reflect these opinions through formal classification. For example, Cole (1923) stated that the genera Furcijira and Epomyia (both currently junior synonyms of Cyclotelus) are very near to Psilocephala. The genus Psilocephala was broadly defined, and prior to Irwin & Lyneborg (1981a), was a polyphyletic assemblage of groups lacking pile on the parafacial. However, the most speciose and abundant group within Psilocephala was Cole’s ‘haemorrhoidalis-group’, all species of which are now members of Orodiceromyia. The fact that Cole formally described this species-group in his treatment of Psilocephala, and included it in his keys, indicated that Cole considered this a monophyletic taxon. However, he had not associated the group with the monotypic Ozodiceromyia and he did not officially separate it from Psilocephala. In addition, Cole (1923) recognized several species of lhereva to appear closer to Psilocephala, but again, did not change their generic status nor recognize the association with Ozodiceromyia, where they are currently placed. Malloch (1932) also had notions of the relationship of “the generally accepted forms of Psilocephala” ( = Ozodiceromyia) with Furcijira ( = Cyclotelus). Cole (1960b) agreed with this assessment and pointed to the possible relationship of Furcfia to Ozodiceromyia in the strict sense. A few recent workers have recognized this group as a distinct entity. Ozodiceromyia, Cyclotelus, and Breviperna were cited as forming a distinct New World group (Nagatomi & Lyneborg, 1987, 1989), with the addition of Procyclotelus from the Far East as a close relative of Cyclotelus itself. Irwin & Lyneborg (1981b) also mentioned the ‘Cyclotelus-group’ as containing Cyclotelus and Ozodiceromyia, but also included Chro- molepida, which the current analysis rejects. They noted that the genera aligned with Cyclotelus have a strong Central American base, with strong ties to genera in North and South America. The hypotheses being formulated in this study are among the first to be rigorously tested in an attempt to reflect the phylogeny of groups within the Therevidae below the subfamilial level, so it is important to erect a phylogenetically-based classification which is more biologically informative than other currently construed classification schemes (Hennig, 1966; Farris, 1979; Wiley, 1980). The classification presented in Appendix 3 follows phyletic sequencing of taxa (Nelson, 1969, 1972, 1973), rather than employing the system of strictly giving sister-groups equivalent ranks. This PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 157 study is an attempt to establish a firm foundation for organizing higher-level groupings of Laurasian Therevinae, and to aid further study of genera within the C yclotelini. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Cyclotelini, tribe nov. The resulting cladogram (Figs 1 and 2), the unweighted Bremer support index of 8 additional steps, and the rescaled Bremer support index of 20.3 steps, demonstrate that the Cyclotelini is a monophyletic group supported by synapomorphies. The hypothesized Late Cretaceous origin for this group, as discussed in the bio- geographical section of the current study, justifies the assignment of tribal rank. The two major monophyletic groups within the Cyclotelini are placed into informal genus-groups. These genus-groups are: the Brevipema-group, including Ammothereva, Anolinga, Brevipema, and Nesonana; and the Cyclotelus-group, which is divided into two monophyletic clades, or subgroups, the Cyclotelus-subgroup (Culeiana, Cyclotelus, and Procyclotelus), and the Ozodiceromyia-subgroup (Bugulaverpa, Crebraseta, and Ozodiceromyia). The genus-groups are diagnosed along with the included genera. The formal classification of the Cyclotelini, with the informal groupings included, is presented in Appendix 3. Diagnosis. The Cyclotelini can be recognized using a suite of external and internal characters. Externally the Cyclotelini lack pile on the posterior surface of the midcoxa, although it is present in the poecilupa-group of Ammuthereva, and is sparsely present in some specimens of Breznperna. The gena usually protrudes beyond the edge of the eye, at least slightly, and the gena often has a distinctive patch that differs in texture, colour, or pile from the rest of the lower occiput. Usually, the male ommatidia are larger on the dorsal than the ventral halves of the compound eye. The ventral surface of the basal lobe of the costa usually lacks pile, or has sparse short pile basally. Internally, males of all Cyclotelini have fused gonocoxites, including a fused hypandrium. The gonostylus is hinged on a gonostylar condyle rather than being hinged on the inner edge of the gonocoxite as in other therevids. The inner gonocoxal process is small and nonarticulated. In the female, the anterior portion of the furca is characterized by the presence of a furcal bulla, and the ribbon-like dorsocentral connection between tergites 8 and 9 lacks setae or pile. As in all Therevidae, spermathecae are not sclerotized. Cyclotelines have 2 spermathecae and a central sac. Synapomorphies. 9 lower frons lacking setae (character 11, state 2) (setae present within the Ozodiceromyia-subgroup, and in Brevipema); gena protruding forward beyond lower edge of eye (character 13) (gena not protruding in Coleiana); genal apex bare of setae (character 15, state 2) batch of distinctive setae (state 1) present in Ozodicmmyia-subgroup and in part of Anolinga); posterior surface of midcoxa lacking pile (character 20); ventral surface of basal costal lobe lacking pile, or with sparse short pile basally (character 32) (fully pilose in Brevzpema); lateral ejaculatory process complete or notched dorsally (character 34); ejaculatory apodeme of uniform width in dorsal view (character 40) (expanded anteriorly in part of Ozodicmmyia and in the clade of Coleiuna + Cyclotelus); gonocoxites strongly fused (character 48, state 2) (fused through sclerotized sheet (state 1) in Ammothereva); hypandrium fused to gonocoxites (character 49); gonostylus hinged to gonocoxite on gonostylar condyle (character 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 160 S. D. GAIMARI AND M. E. IRWIN Lower frons not bulging (e.g. Figs 12 and 13), usually without setae or pile (e.g. Fig. 7), or with short black setae (e.g. Fig. 9), or rarely with white pile. Scape shorter than first flagellomere, and first flagellomere thicker than scape (e.g. Fig. 18). Scape and pedicel with short black setae, but few are thickened. Proepimeron lacking pile (e.g. Fig. 40). Basal lobe of costa Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 not densely pilose on ventral surface, at most with sparse short, white pile basally (as in Fig. 49). Posterolateral epandrial flange of 6 unidentate (e.g. Fig. 61). Lateral ejaculatory process usually small, set into aedeagus (e.g. Figs 116 and 122) (although approaching condition above in some Ammotherevu) ...... 13 13(12). Macrosetae of head, notum, coxae, and legs black. 1 pair of dorsocentral setae. Legs with dark femora. Outer gonocoxal process long and thin, covered with black setae (Figs 77, 78, 79, and 80). Ventral gonocoxal process present, covered with black setae, parallel with outer gonocoxal process (Figs 77 and 79). Posterolateral epandrial flange of 6 long, thin, and pointed distally (Fig. 59), but point may be curved under epandrium and not visible from dorsal view (Fig. 58). Strong sclerotized bridge between parameral sheath and gonocoxal apodeme present. Ventral apodeme of aedeagus expanded anteriorly, and longer than dorsal apodeme (Fig. 116) ...... Nesonunu, gen. nov...... 14 Macrosetae of head, notum, coxae, and legs pale or black. 0-2 pairs of dorsocentral setae. Legs usually entirely orange. Outer gonocoxal process long, but usually laterally flattened, covered with white setae (e.g. Figs 72 and 85). Ventral gonocoxal process absent (e.g. Figs 74 and 85). Posterolateral epandrial flange of 6 not elongated or thin (e.g. Fig. 57), only rarely pointed (e.g. Fig. 61). Parameral sheath and gonocoxal apodeme not attached by a sclerotized bridge. Dorsal apodeme of aedeagus longer than ventral apodeme (e.g. Fig. 120), which is not expanded distally ...... 15 14(13). Single row of postocular setae. Median occipital sclerite with some silver pruinescence; gold pruinescence along upper edge of postocular area. 6 epandrium with posterolateral flange pointed and directed posteriorly beside cerci (Fig. 59) ...... Nesonana youngi, sp. nov. - Second row of setae beneath row of postocular setae. Median occipital sclerite with gold pruinescence. S epandrium with posterolateral flange folded under cerci (Fig. 58) ...... Nesonana xenoverpa, sp. nov. 15(13). Macrosetae of head, notum, coxae, and legs black. 2 pairs of dorsocentral setae. Dorsoapical third of femora with brown, appressed, scale-like pile on 9 (and usually on S),in addition to short, black setae. 9 abdomen with tergites dark brown dorsally. Basal distiphallus with ventrolateral wing-like processes (Fig. 12 1). Distiphallus oriented upward from posterior portion of dorsal apodeme before curving downward (Fig. 120). In 9, common spermathecal duct >2 furcal lengths (Fig. 149) ...... Anolingu, gen. nov...... 16 Macrosetae of head, notum, coxae, and/or legs pale. 0-2 pairs of dorso- central setae. Femora with appressed, scale-like pile white throughout; short, black setae present or absent. 9 abdomen with tergites pale or orange, as pruinescence or exposed ground colour. Distiphallus variable, but never with ventrolateral wing-like processes, and always oriented posteriorly from dorsal apodeme (e.g. Fig. 112). In 9, the common PHYLOGENY AND BIOGEOGR4PHY OF CYCLUI'ELINE 'I'HERE\INAE 161 spermathecal duct is <2 furcal lengths ...... Amnzothereua Lyneborg (see key to species in Lyneborg, 1984) 16(15). 8 frons lacking setae (Fig. 18); 9 frons with few setae (Fig. 19). Kate- pisternum and cleft of prosternum lacking pile. Epandrial setae white ..... ...... Anolinga psilofrons, sp. nov. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 - 8 frons with 2 patches of short setae (Fig. 16); 0 from with many scattered setae (Fig. 17). Katepisternuni and cleft of prosternum pilose. Epandrial setae black ...... Anolinga melanothrix, sp. nov. 17(10). One pair of supra-alar setae. 1 pair of scutellar setae, oriented posteriorly. Ommatidia of d the same size. In 9 , lacking furcal bulla; medially oriented pegs separate furca into anterior and posterior halves (Fig. 150) ...... Bugulauerpa, gen. nov. (rebeccae, sp. nov.) - Two pairs of supra-alar setae (1 pair in some Cjdotelus). 2 pairs of scutellar setae (0-1 in some Cjcloteh); if only 1 pair, usually oriented upward. Ommatidia of 8 with larger facets on upper half than lower half of eye. In 9,furcal bulla present; lacking medially oriented pegs on furca ... 18 18(19). Prosternum and katepisternum (e.g. Fig. 41) pilose. ? frons lacking patches of black or brown matte pruinescence, usually shiny black (e.g. Fig. 27), at least in part. Posterolateral corners of epandrium lacking flattened, bare flange; with setae encompassing edge (e.g. Fig. 65). Outer gonocoxal process small or absent, with reduced setal covering (e.g. Figs 91, 92, 94, and 96). Inner gonocoxal process usually a small knob with apical setae (e.g. Figs 92, 94, and 96). Gonostylus usually turned up at tip by 90" and ventrally expanded into lobe (e.g. Fig. 109). Foretibia sometimes expanded distally, clavate (as in Fig. 45) ...... Ozodiceromvia Bigot Prosternum and katepisternum lacking pile. 9 frons often with patches of black or brown matte pruinescence (e.g. Figs 33, 35, and 39), but if shiny black (Fig. 31) (Pro@otelus), then other characters not as above. Posterolateral corners of epandrium with flattened, bare flange; with setae only to base of flange (e.g. Figs 67, 68, and 70). Outer gonocoxal process large and distinct, covered with vestiture similar to remainder of gonocoxite (e.g. Figs 98, 99, 101, and 103). Inner gonocoxal process often indistinct from outer (e.g. Figs 98 and loo), with or without (e.g. Fig. 106) setae. Gonostylus extended posteriorly at tip, not upcurved; not expanded ventally into lobe (e.g. Fig. 107). Foretibia and forebasitarsus variable ...... 19 19(18). Lower frons bulging and shiny black (Figs 30 and 31). Upper frons of with velutinous pruinescence (Fig. 3 1). Flange of posterolateral corner of epandrium bidentate (Fig. 67). Lateral ejaculatory process enlarged, sep- arated from aedeagus anteriorly (Figs 136 and 137). Common spermathecal duct of 9 longer than 3 furcal lengths ...... Procvclotelus Nagatomi & Lyneborg (elegans Nagatomi & Lyneborg) Lower frons not bulging. Frons usually with patches of black or brown matte pruinescence. Upper frons of 0 occasionally shiny black above patch of matte pruinescence (e.g. Fig. 35). Flange of posterolateral corner of epandrium usually unidentate (e.g. Figs 68, 70 and 7 1). Lateral ejaculatory process small, set into aedeagus (e.g. Fig. 143) ...... 20 20(19). Katatergite covered with black pile, and with stiff black setae posteriorly. 162 S. D. GAIMARI AND M. E. IRWIN Cervical sclerite, anepisternum, coxae, and tergite 1 with black pile. Gena not protruding forward (Figs 32 and 33). Lower frons with brown matte pruinescence, upper frons of 9 dark brown (Fig. 33). Foretibia and forebasitarsus normal. Midbasitarsus longer than midtarsomere 2-5 in- clusive (Fig. 47). Style apical on first flagellomere (Figs 32 and 33). Inner Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 gonocoxal process indistinct from outer, with apical setae (Fig. 100). Aedeagus with dorsal shelf over distal part of distiphallus (Fig. 138). Common spermathecal duct of 9 longer than 2 furcal lengths (e.g. Fig. 155) ...... Coleiana, gen. nov. (nigricopis, sp. nov.) Katatergite, cervical sclerite, anepisternum, coxae, and tergite 1 with white pile. Gena protruding forward (Figs 34 and 35). Lower frons silver pruinose; upper frons often with patches of brown or black matte pruinescence (Figs 35, 36 and 39), and occasionally with shiny black area above patch (e.g. Fig. 35). Foretibia expanded distally, clavate (e.g. Fig. 45). Forebasitarsus swollen relative to other basitarsi (Fig. 45). Midbasitarsus subequal or shorter than tarsomeres 2-5 (e.g. Fig. 45). Style subapical and usually set into first flagellomere. Inner gonocoxal process usually a small bump lacking apical setae (e.g. Fig. 106). Distal distiphallus without dorsal shelf. Common spermathecal duct of 0 shorter than 2 furcal lengths (e.g. Fig. 156) ...... Cyclotelus Walker (Cole, 1960b, has the most current key to species) Genus-groups, genera, and specie5 To reflect the two monophyletic clades within the tribe, the genera are divided into two informal genus-groups. Within each genus-group, the genera are presented alphabetically, without regard to subgroup. Species within each genus annotated with ** were used in the phylogenetic analyses of the current study. An exhaustive list of references for each previously described taxon is provided by Gaimari (1998), but only taxonomically relevant references are listed herein. Within species descriptions, values in [ ] represent the values taken from paratypes of the same sex. Abbreviations for repositories follow Arnett, Samuelson & Nishida (1993), and are listed in Appendix 4. This manuscript was prepared and submitted long before the 4th edition of the International Code of Zoological Nomenclature was available (see Art. 86.1). All new nomenclatural acts except the lectotype designations conform to this edition. While each lectotype designation is specific for one nominal taxon, we here state that each and every one of these designations is for the sole purpose of fixing and stabilizing the current concept of the taxon. Breviperna-group Diagnosis. Members of this group display full pruinescence on the frons, pleura, and notum, with no areas of shiny black, velutinous, or matte pruinescence. They typically have an antenna1 first flagellomere that is onion-shaped, or bulbous basally, tapering to the tip. In addition, the style is terminal on the first flagellomere. The PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREL’INAE 163 maxillary palps are orange. The 9 upper frons possesses scattered setae, or setae are occasionally in defined patches. The genal apex usually lacks any patch of distinctive colour or texture and usually lacks pile. The foreleg is normal, with no swollen or enlarged parts. On the wings, the setae of the basal lobe of the costa are scattered over the entire surface, extending into both outer and inner costal rows. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Wing-like processes are present on the ventrolateral portion of the basal distiphallus in New World members. Most of the epandrium is covered with setae. The inner gonocoxal process lacks apical setae. The gonostylus usually has a subapical, lateral spur. Synapomoqhies. Maxillary palpus colour orange (character 12); gonostylus with sub- apical lateral spur (character 56) (absent in Breviperna). Synapomorphies for Nesonana + Breviperna + Anolinga: Ventral apodeme of aedeagus longer than dorsal apodeme (character 37) (dorsal apodeme longer in Anolinga); ejaculatory apodeme expanded anteriorly in dorsal view (character 40, state 0); basal distiphallus with wing-like processes ventrolaterally (character 42); gonostylus expanded ventrally into lobe (character 57) (strap-like in part of Nesonana); spermathecal ducts originate on base of central sac (character 62). Synapomorphies for Breviperna +Anolinga: Katepisternum with pile (character 18, state 0) (pile lacking in part of Anolinga); ventral apodeme laterally flattened, thinned, and sword-like (character 38); dis- tiphallus extended upwards, oriented in line with ventral apodeme (character 4 1); common spermathecal duct >2 furcal lengths (character 6 1). Ammothereva Lyneborg Ammothereva Lyneborg, 1984: 206 (incl. revision, illust.: numerous, e.g. antennae, heads, 8 genit.). Type species, Psilocephala gussakovskyi Zaitsev, by original designation. Majer, 1997: 522 (key ref., key does not work for Ammothereva), 523 (illust.: 9 head). Diagnosis. The best diagnosis, with the most complete consideration of the species is found in Lyneborg (1984). An attempt to diagnose this genus here, without all of the necessary specimens, would be incomplete, especially when the genus was revised so recently and no new species or combinations have since been recognized. A few comments are in order, however. Within the Cyclotelini, this is the only genus that regularly displays the state of having pale macrosetae of the notum, head, coxae, and legs. Also, the epandrium is consistently longer than wide (e.g. Figs 56 and 57). Interestingly, this genus seems to be variable in certain characters which are not variable in other groups of Cyclotelini. Ammothereva lepida Lyneborg and Ammothereua salentioides Lyneborg have a conical first flagellomere rather than the more typical basally-bulbous first flagellomere. Ammothereva mesasiatica (Zaitsev) and Ammothereva salentioides have males with the eyes separated by a distance greater than the anterior ocellus. Species in the poecilopa-group have a pilose prosternal cleft while others lack pile, and Ammothereva salentioides has pile on the posterior surface of the midcoxa, a state shared by no other member of the tribe except atypical specimens of Breviperna. In the male genitalia, Ammothereva n&gemorata and Ammothereva gussakovskyi have the lateral ejaculatory process enlarged and anteriorly separated from the aedeagus as is found in Breviperna and Procyclotelus. Many of these differences may be attributable 16.1 S. D. GAIMARI AND M. E. IRWIN to the ancient divergence and geographical separation (and therefore open niches available) of this genus from the remainder of the tribe. Synapomorphies. Prosternum lacking pile (character 16) (pile present in poecilopa-group); macrosetae of head, notum, coxae, and/or legs pale (character 27) (dark, or nearly black in few species); epandrium longer than wide (character 44); gonocoxites fused Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 by sclerotized membrane between discernible gonocoxites (character 48, state 1). Distribution. This genus is found throughout the steppes, semideserts, and deserts of the southeastern Palearctic Region from Hungary through Mongolia (Fig. 3A). Specimens examined. Ammothereva laticornis (Loew): AUSTRIA: Thalhammer, Keczel (1 8,ME1 109643, OSUC). HUNGARY: (as) Ungarn, leg. Kalocsa (1 8,ME1 103402, 1 9,ME1 104436, USNM), leg. Keczel (18, ME1 105181, USNM), leg. Vadkert (1 8, ME1 109639, MEIC); Hung.[-ary] (1 8,ME1 109646, UMSP); Hungaria, Keczel, V-4 (1 8, ME1 109642, OSUC); Gyon [ = Dabas], KertCsz, collection C.W. Johnson (18, ME1 109647, 10, ME1 109644, MCZC); Deliblat, KertCsz, collection C. W. Johnson (1 8, ME1 080876, MCZC); SzCkesvehkr-vhr- Sost6,15-VI-1928 (288, ME1 079725,109641,l O,ME1 109645, CNCI).RUSSIA: Omsk Region: Troickoe, Slavgorodsk (c.e.), 27-VI-1922, Reichardt (ME1 105 180, 1 8,SDGC); Terekli-Mekteb, Karanogajskaya Steppe [between Caspian and Black Seas], 16-V-1925, Kirichenko (1 9,ME1 105183, SDGC). UNKNOWN: 7/Sajo (1 8 , ME1 105182, 1 0,ME1 109640, EMEC), 8/Sajo (1 8,ME1 109638, USNM). Ammothereva lehri (Zaitsev): TADJIKISTAN: Kazalkum, GA, Tamdy, town of Ak- Tau [=Gory, Ak-Tau] , 9-V- 1965, V. Zaitsev (1 paratype 0,ME1 105324, SDGC). UZBEKISTAN: Kazalkum, Aznek, 70 km S. Tamdy-Bulak [ = near Tashkent], 2- V-1965, V. Zaitsev (1 paratype 8,ME1 105326, SDGC; 1 paratype 8, ME1 105325, 1 paratype 0,ME1 105327, MEIC). Included species. A revision of this genus is not necessary at this time, as Lyneborg (1984) treated the species of this group and no new species or combinations have been recognized. Lyneborg's (1984) treatment included synonymies and a justification for generic rank. The following list of species, with their original generic placements and synonymies, is presented to complete the picture of what currently constitutes the Cyclotelini. Refer to Lyneborg (1984) for complete taxonomic information, including information on types, diagnoses, distributions, and a key to species. Ammothereva albula (Zaitsev), 1974: 3 16 (Psilocephala). A. gussakovs@i (Zaitsev), 1973: 704 (Psilocephala). ** A. laticornis (Loew), 1856: 32 (7hereva). ** A. lehri (Zaitsev), 1973: 707 (Psilocephala). A. lepida Lyneborg, 1984: 2 19. A. lobata Lyneborg, 1984: 2 15 (replacement name for Psilocephala mesasiatica Zaitsev) Psilocephala mesasiatica Zaitsev, 1973: 7 10 (preoccupied by Neothereva mesas- iatica Zaitsev, 1970: 795). A. mesasiatica (Zaitsev), 1970: 795 (Neothereva). A. mongolica (Zaitsev), 1970: 797 (Neothereva). A. n&gemorata (Krober), 19 12i: 1 19 (Psilocephala). A. poecilopa (Loew), 1871a: 320 (nereva) (replacement name for 7hereva hebes Loew) PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE I65 Thereua hebes Loew, 1869a: 123 (preoccupied by Thereua hebes Walker, 1852: 160). A. salentioides Lyneborg, 1984: 22 1 (replacement name for Psilocephala mongolica Zaitsev) Psilocephala mongolica Zaitsev, 1976: 488 (preoccupied by Neothereua mongolica Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Zaitsev, 1970: 797). A. splendida (Krober), 19 12i: 1 18 (Psilocephala). Psilocephala caucasica Zaitsev, 1973: 7 1 1. A. uzbekorum (Zaitsev), 1973: 7 12 (Psilocephala). Anolinga, gen. nov. ljpe species. Anolinga melanothrix, sp. nov. Epmology. Feminine. Gr., ano-: upward + Skrt., f., linga: phallic symbol, referring to the upwards orientation of the basal distiphallus. Diagnosis. The from are covered with gold pruinescence. The scutum is grey or brown pruinose, with erect black and appressed pale or gold pile. Two pairs of dorsocentral setae are present. Legs are orange. The dorsal apodeme of the aedeagus is parallel sided and elongate, >2 x length of ventral apodeme (e.g. Figs 121 and 123). The ventral apodeme is thin and sword-like (e.g. Fig. 120). The distiphallus is oriented upwards, in line with the ventral apodeme (e.g. Fig. 120). The outer gonocoxal process has setae around the outer edges that are denser and finer than setae on the remainder of the gonocoxite (e.g. Fig. 85). In 0, tergite 6 is silver pruinose with a medial brown stripe. The common spermathecal duct is >2 x the length of the furca and originates on the posterior surface of the furcal bulla (e.g. Fig. 149). The furcal bulla is sclerotized anteriorly (e.g. Fig. 149). Synapomorphies. Dorsal apodeme of aedeagus longer than ventral apodeme (character 37, state 0); outer gonocoxal process with dense, fine setae (character 51, state 2); common spermathecal duct originates on posterior part of furcal bulla (character 60, state 2). Distribution. This genus occurs in the Brazilian states of S5o Paulo, Rio de Janeiro, and Minas Gerais (Fig. 3A). Included species. Anolinga melanothrix, sp. nov., and Anolinga psilofons, sp. nov. **Anolinga melanothrix, sp. nov. Epmology. Gr., melano: black + f., thrix: hair, referring to the black epandrial setae; to be treated as a noun in apposition. Dzagnosis. This species is larger than its congener. In 8,the frons is setose; ommatidia are larger in the upper half than the lower. The genal apex has a small patch of short black setae. The katepisternum and the cleft of the prosternum are pilose. The setae of the epandrium are black. I66 S. I). GAIhIARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figure 3A-C. Distributions of cycloteline genera. A, Breuiperna-group; B, Ozodiceronyia-subgroup; C, Cyclotelus-subgroup. Autupomorphy. Genal apex with patch of short, black pile (character 15, state 1). Descrzption ofholoppe 8 (ME1 1002 15): length [8.9-] 9.4 mm. Heud(Fig. 16). 1.25 [1.13-1.261 mm long, [2.10-] 2.34mm wide, [1.52-] 1.62mm high. Distance between eyes at antenna1 level 0.68 [0.60-0.691 mm; at genal level [0.81-] 0.95 mm. Scape 0.26 mm long, 0.15 mm wide; silver pruinose; bare medially; with fine black setae otherwise evenly distributed, and few thick setae concentrated distally. Pedicel 0.1 1 mm long; bare medially. First flagellomere 0.44 mm long, PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 167 0.20 mm wide; orange; bulbous basally, tapering distally. Style 0.20 mm long; terminal. Antenna1 base to nearest edge of eye 0.18 mm. Frons bulging 0.09 mm beyond eye in lateral view. Parafacial lacking pile, covered with gold pruinescence extending dorsally through frons, which is also gold pruinose; upper frons with two distinct patches of black setae; lower frons bare. Genal pile short and black. Palpus Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 orange, covered with white pile. Postgena and occiput silver pruinose; pile white; occiput with several black setae. Median occipital sclerite flattened; lacking setae or pile; covered with gold pruinescence. Postocular setae black, arranged in single row. Ocellar tubercle gold pruinose; with fine, forward-directed, black setae. irhorux. Scutum [2.73-] 2.88 mm long, [2. lo-] 2.22 mm wide; grey-brown pruinose; covered with erect black and erect and recumbent pale gold pile; median vitta and dorsocentral vittae faintly present as thin lines of darker brown pruinescence; 2 pairs dorsocentral setae; 4 [usually 31 pairs notopleural setae; 2 pairs supra-alar setae; 1 pair postalar setae. Scutellum brown pruinose, with white pile. Prosternum with white pile. Pleura entirely silver pruinose. Katatergite with dense, long, white pile. Anepisternum and katepisternum with white pile. Proepimeron lacking pile. Legs. Coxae with silver pruinescence, posterior surfaces less so; fore- and midcoxae each with 2 black macrosetae and long, white pile on anterior surfaces; hindcoxa with 1 black macroseta and long, white pile on lateral surface. Legs entirely orange below coxae. Femora with appressed, scale-like white pile dorsally; and with scattered, short, black setae. Forefemur with erect white pile. Tibiae and tarsi covered with short, erect, black setae. Basitarsi orange basally, becoming dark brown through tarsomeres 2-5. Wng. [7.84-] 8.64 mm long. Basal lobe of costa with scattered setae extending into inner and outer rows of costal setae. Cell m3 closed at edge of wing. Membrane unmarked. Halter light brown. Abdomen. Tergites silver pruinose; covered with recumbent white and brown pile; tergites 2-4 lacking pruinescence on anterior edges. Sternites covered with silver pruinescence and erect white pile. Tminuliu (paratype, ME1 100224). Sternite 8 with white setae restricted to posterior edge, which is emarginate medially. Tergite 8 dumbell-shaped, with white setae restricted to posterior edge. Epandrium (Fig. 61) 0.68mm long, 0.90mm wide at widest point; orange; emarginate anteriorly; dorsal surface entirely covered with black setae; lateral edges tapering in posterior third; posterolateral flange 0.17 mm long, bare. Cerci 0.29 mm long; not fused medially; extending posteriorly beyond posterolateral epandrial flange; subequal in length to hypoproct. Subepandrial plate with setae on ventral surface. Gonocoxites (Fig. 83) 0.96mm wide; orange; with white setae. Outer gonocoxal process 0.47 mm long; laterally flattened; with fine, dense setae along outer edge. Gonocoxal apodeme entirely within anterior edge of gonocoxite; lacking sclerotized bridge to parameral sheath of phallus. Ventral gonocoxal process absent; setae of corresponding edge absent. Ventral lobes distinct, fused basally. Gonostylus strap-like, tip extending posteriorly; with lateral wing-like process and subapical, lateral spur. Aedeagus in Figures 120 and 121. Dorsal apodeme of aedeagus parallel sided; 2 x longer than ventral apodeme. Ventral apodeme thin, sword-like. Ejaculatory apodeme 0.56 mm long; stick-like, with uniform width. Lateral ejaculatory process complete dorsally, set into aedeagus; 168 S. D. GAIRL\RI AND M. E. IRWIN lightly sclerotized. Basal distiphallus with ventrolateral wings. Distiphallus basally oriented upwards; long, recurved. DeJcription ofallo&pe 0 (ME1 100157): 9.6 mm long. Similar to 6 except as follows. Head (Fig. 17). [ 1.13-1 1.20 mm long, [2.00-] 2.26 mm wide, [ 1.38-1 1.50 mm high. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Eyes separated below anterior ocellus by [0.29-] 0.30 mm. Distance between eyes at antennal level [0.65-] 0.7 1 mni; at genal level [0.83-] 0.90 mm. Upper from with scattered setae. Second row of postocular setae present ventrally to those of upper edge. nzorux. Scutum [2.50-] 2.73 mm long, [1.7t] 1.89 mm wide. Whg. [8.08-] 8.48 mm long. Abdomen. Tergites with brown pruinescence and areas of silver pruinescence; with brown recumbent pile dorsally. Laterally, tergites mostly silver pruinose, with erect and recumbent white pile; middle of tergite 2 and anterior of tergite 3 brown pruinose. Sternites covered with silver pruinescence. Zrminalia (as in Fig. 149) (paratype, ME1 100211). Furca 0.54mm long, 0.35mm wide; entire; anterolateral wings dorsoventrally flattened; furcal bulla sclerotized anteriorly. Gonopore on posterior part of furcal bulla. Common spermathecal duct 2.40 mm long. Spermathecal ducts originate on base of central sac. Diytribution. Known from Minas Gerais and S%oPaulo, Brazil. Speriiizens examined. Holotype: BRAZIL: Minas Gerais: Sapucai-mirim, Cidade Azul, 1400 m, 6-IX- 1953, F. Pd. Pereira and Medeiros (6,ME1 1002 15, MZSP). Paratypes: same data as holotype (allotype 9,ME1 100157, MZSP; 1 d, ME1 100224, SDGC; 1 9,ME1 1002 1 I, SDGC); Sao Paulo: Campos de JordZo Eug. Lefevre, 1200 m, 4- IX to 8-IX- 1953, F. Pd. Pereira and Medeiros (1 8,ME1 1002 13, MZSP). **Anolinga psilofrons, sp. nov. E&molog)1. Noun. Gr., psilo: naked, bare + L., f., frons: front, forehead, referring to the lack of setae on the frons of the male; to be treated as a noun in apposition. DiugnosiJ. This species is smaller than its known congener. In 8,the from lacks setae; ommatidia are all the same size. The genal apex is bare of setae. The katepisternum and the cleft of the prosternum lack pile. The setae of the epandrium are white. Autapomoqhies. d ommatidia all the same size (character 7); prosternum lacking pile (character 16); katepisternum lacking pile (character 18). Description ofholoppe 6 (ME1 077737): length 7.0 mm. Head(Fig. 18). [0.93-] 0.96mm long, [1.60-] 1.70nim wide, [1.26-] 1.29mm high. Distance between eyes at antennal level [0.53-] 0.54mm; at genal level [0.66-] 0.7 1 mm. Scape 0.17 mm long, 0.14 mm wide; silver pruinose; bare medially; with few thick and fine, black setae, otherwise evenly distributed. Pedicel 0.09 mm long; bare medially. First flagellomere 0.36 mm long, 0.17 mm wide; orange; bulbous PHYLOGENY AN11 BIOGEOGRAPHY OF CYCLUIELINE THEREL’INAE 169 basally, tapering distally. Style 0.1 7 mm long; terminal. Antenna1 base to nearest edge of eye 0.14 mm. Frons bulging 0.03 mm beyond eye in lateral view. Parafacial lacking pile, covered with silver pruinescence extending dorsally along eye margin halfway up frons; middle of frons with gold pruinescence; frons bare of setae. Genal pile absent. Palpus orange, covered with white pile. Postgena and occiput silver Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 pruinose; pile white; occiput with several black setae. Median occipital sclerite flattened; lacking setae or pile; covered with silver pruinescence. Postocular setae black, arranged in single row. Ocellar tubercle with fine, forward-directed, black setae. Thorax. Scutum [2.04-] 2.20 mm long, [1.34-] 1.50 mm wide; silver-grey pruinose; covered with erect black and erect and recumbent white pile; median vitta and dorsocentral vittae faintly present as thin lines of light brown pruinescence; 2 pairs dorsocentral setae; 3 pairs notopleural setae; 2 pairs supra-alar setae; 1 pair postalar setae. Scutellum silver-grey pruinose, with white pile. Prosternum lacking pile. Pleura entirely silver pruinose. Katatergite with dense, long, white pile. Anepisternum with white pile. Katepisternum and proepimeron lacking pile. Legs. Coxae with silver pruinescence, posterior surfaces less so; fore- and midcoxae each with 2 black macrosetae and long, white pile on anterior surfaces; hindcoxa with 1 black macroseta and long, white pile on lateral surface. Legs entirely orange below coxae. Femora with appressed, scale-like white pile dorsally. Forefemur with erect white pile. Tibiae and tarsi covered with short, erect, black setae. Tarsi orange, becoming dark brown at tips. ll4ng. 5.76 [-5.801 mm long. Basal lobe of costa with scattered setae extending into inner and outer rows of costal setae. Cell m3 closed and stalked. Membrane unmarked. Halter with knob yellow and stalk light brown. Abdomen. Tergites entirely silver pruinose; covered with recumbent white pile. Sternites covered with silver pruinescence and erect white pile. Tminalia (paratype, ME1 077736). Sternite 8 with white setae restricted to posterior edge, which is emarginate medially. Tergite 8 dumbell-shaped, with white setae restricted to posterior edge. Epandrium (as in Fig. 61) 0.48 mm long, 0.75 mm wide at widest point; orange; not emarginate anteriorly; dorsal surface entirely covered with white setae; lateral edges tapering in posterior third; posterolateral flange 0.1 1 mm long, bare. Cerci 0.23 mm long; not fused medially; extending posteriorly beyond posterolateral epandrial flange; subequal in length to hypoproct. Subepandrial plate with setae on ventral surface. Gonocoxites (Figs 84 and 85) 0.66mm wide; orange; with white setae. Outer gonocoxal process 0.41 mm long; laterally flattened; with fine, dense setae along outer edge. Gonocoxal apodeme entirely within anterior edge of gonocoxite; lacking sclerotized bridge to parameral sheath of phallus. Ventral gonocoxal process absent; setae of corresponding edge absent. Ventral lobes distinct, fused basally. Gonostylus strap-like, tip extending posteriorly; with lateral wing-like process and subapical, lateral spur. Aedeagus in Figures 122 and 123. Dorsal apodeme of aedeagus parallel sided; 2 x longer than ventral apodeme. Ventral apodeme thin, sword-like. Ejaculatory apodeme 0.38 mm long; stick-like, with uniform width. Lateral ejaculatory process complete dorsally, set into aedeagus; lightly sclerotized. Basal distiphallus with ventrolateral wings. Distiphallus basally oriented upwards; long, recurved. 170 S. D. GAIMARI AND M. E. IRWIN Description ofallo&pe ? (ME1 077734): 8.5 mm long. Similar to 8 except as follows. Head (Fig. 19). [0.92-] 1.02 mm long, [ 1.62-1 1.88 mm wide, [ 1.17-1 1.35 mm high. Eyes separated below anterior ocellus by [0.2 1-3 0.24 mm. Distance between eyes at antenna1 level [0.56-] 0.62mm; at genal level [0.66-] 0.75mm. Upper frons setose. Second row of postocular setae present ventrally to those of upper edge. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 norax. Scutum [2.14-] 2.55 mm long, [1.43-] 1.61 mm wide; brown pruinose, but more silver-grey laterally; covered with erect black and recumbent pale gold pile; dorsocentral vittae present as silver-grey pruinescence. Scutellum light brown anteriorly, silver-grey posteriorly. Mng. [6.00-] 6.60 mm long. Abdomen. Tergites with brown pruinescence and brown recumbent pile dorsally; posterior edges of tergites 2 and 3 with silver pruinescence; anterior three-fourths of tergite 5 silver pruinose. Laterally, tergites mostly silver pruinose, with erect white pile; middle of tergite 2 brown pruinose. Sternites covered with silver pruinescence. Zmzinalia (Fig. 149) (paratype, ME1 077735). Furca 0.45 mm long, 0.33 mm wide; entire; anterolateral wings absent; furcal bulla sclerotized anteriorly. Gonopore on posterior part of furcal bulla. Common spermathecal duct 1.65 mm long. Sperm- athecal ducts originate on base of central sac. Distribution. Known from the Brazilian state of Rio de Janeiro. Specimens examined. Holotype: BRAZIL, Rio de Janeiro: Grajahu, 20-VIII- 1939, S. Lopes (d,ME1 077737, MZSP). Paratypes: same data as holotype (allotype 9,ME1 077734, MZSP; 1 d, ME1 077736, 1 0,ME1 077735, SDGC). Brevipema Irwin Breviperna Irwin, 1977a: 288. Type species, Psilocephala placida Coquillett, 1894, by original designation. Irwin & Lyneborg, 1981a: 203 (key ref.), 247 (redescr.). Included species. This genus was only recently treated by Irwin (1977a) and Irwin & Lyneborg (1981a). Since that time, no new species have become apparent, so we only provide additional morphological and distributional information for the de- scribed species. **Breviperna placida (Coquillett) Psilocephalaplacidu Coquillett, 1894: 99. Type locality: in literature as Florida (likely an error-label reads “Fla.”), more likely Flagstaff, Arizona. Type: HT? in USNM, catalog # 10420. Cole, 1923: 64 (annot. listing, misident.), Plate 6 (misident., illust.: 0 head). Brevipema placida: Irwin, 1977a: 289 (illust.: head, wing, ? termin.), 291 (redescr., comb. change), 292 (illust.: c3 genit.), Irwin & Lyneborg, 1981a: 248 (illust.: 8 genit.), 249 (listing). PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 171 Diagnosis. This large bodied species has an antenna1 scape that tapers distally and is as thick as the first flagellomere. This is one of the few groups known within the Cyclotelini where the male compound eyes are dichoptic, separated by a width greater than the anterior ocellus (Fig. 14) (certain members of Ammothereva also share this condition). The same is true for the condition of having scattered setae on the Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 female lower frons, which bulges in Brevz$erna placida (Fig. 15). There is one pair of dorsocentral setae. The prosternum has white pile. The proepimeron is pilose on the upper half. The katepisternum is also pilose, as is the case for part of the related genus Anolinga. The posterior surface of the midcoxa occasionally has sparse pile, but this is thought to be an aberration and not a characteristic for the species. On the wing, cell m3 is open, or occasionally closed at the edge of the wing. Of the Cyclotelini, this is the only member with pile covering the ventral surface of the basal lobe of the costa (Fig. 48). The gonostylus lacks a subapical, lateral spur. The aedeagus (Figs 118 and 119) is characterized by having a short dorsal apodeme with a slightly longer ventral apodeme, and an ejaculatory apodeme with a large posterior tip. The lateral ejaculatory process is greatly enlarged and appears separated anteriorly from the remainder of the aedeagus. The furcal bulla in this species is quite large and distinct (Fig. 148), with the long common spermathecal duct originating on the furca basal to the furcal bulla. The spermathecal ducts originate from the base of the large, round, central sac. Autapomorphies. 8 compound eyes separated by width greater than width of anterior ocellus (character 6); 9 lower frons with scattered setae (character 11, state 0); proepimeron with pile (character 17, state 0); ventral surface of basal costal lobe pilose (character 32, state 0); lateral ejaculatory process enlarged and separated from aedeagus anteriorly (character 35); gonostylus lacking a subapical lateral spur (character 56, state 0); setae of tergite 8 widespread (character 58, state 0); sternite 8 with setae widely distributed (character 59). Description of 9 temzinalia (ME1 028316) (Fig. 148). Furca 0.63 mm long, 0.39 mm wide; anterior edge not sclerotized; anterolateral wings dorsoventrally flattened; furcal bulla large, lightly sclerotized. Gonopore basal to but not on furcal bulla. Common spermathecal duct 6.60 mm long; wider than spermathecal ducts. Central sac 1.50 mm in diameter; round. Spermathecal ducts originate from base of central sac. Distribution. This species was previously recorded only from the montane regions of the southeastern quarter of Arizona, most extensively collected in the Chiricahua and Santa Rita Mountains. Its known range is extended to include the Pinos Altos Mountains in the Gila National Forest in New Mexico, the southwestern part of the Sierra Madre Occidental mountains in Nayarit, Mexico, and the northwestern part of the Sierra Madre del Sur in Jalisco, Mexico. It seems likely that this species covers the low altitude montane regions throughout this range (Fig. 3A), but is undercollected. Specimens examined. MEXICO: Jalisco: Nevada de Colima, 14.5 km from hwy.jct., 13-IV-1980, J. Schaffner, J. Cuda, ex Quercus spp. (1 8, ME1 076525, 1 9,ME1 076526, TAMU); Nayarit: 14.5 km NW Santa Isabella, 10-111-1972 (1 8, ME1 004052, 1 9,ME1 004053, collected in copula, USNM). UNITED STATES: Arizona: Cochise Co., Portal, Southwestern Research Station, 1600 m, 28-IV- 1969, V.D. Roth (1 0,ME1 026944, AMNH); Gila Co., Pinal Mts., VI, D.K. Duncan (1 9, I72 S. D. CAIMARI AND M. E. IRWIN ME1 105319, AMNH), 6-V-1940, F.D. Parker (1 0,ME1 078517, UAIC), Sierra Ancha Mts., VI, D.K. Duncan (2 0 ? , ME1 105300, 105320, AMNH), 17-VI-1938, L.M. Jackson (1 8, ME1 105268, AMNH); Pima Co., Tucson, 15-V-1937, R.H. Crandall (1 8, ME1 105269, AMNH); Santa Cruz Co., Santa Rita Mountains, Madera Canyon, 1500 m, 3-V- 1963,J.G. Franclemont (1 8, ME1 004027, CUIC), Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 8-V-1963 (1 8 , ME1 004025, 1 0,ME1 004028, CUIC), 2 1-VI to 27-VI-1940, L. M. Martin (1 0, ME1 0283 16, LACM); New Mexico: Grant Co., Gila National Forest, Lake Roberts, 28-V to 5-VI-1991, J.E. Swann, FIT in dry stream bed near lake (1 8,ME1 076419, DEBU). Nesonana, gen. nov. 7jpe species. Nesonana young, sp. nov. EpmoloQ. Feminine. Gr., f., nesos: island + L., m., nanus: dwarf, referring to the small body size and Antillean island distribution. Diagnosis. These small-bodied flies have frons with gold pruinescence. This is one of the few cycloteline genera where the 8 ommatidia are of uniform size on the upper and lower halves. The notum is brown with lighter dorsocentral vittae and one pair of dorsocentral setae. The ventral and outer gonocoxal processes are parallel and oriented posteriorly (e.g. Fig. 80). The ventral gonocoxal process is weakly sclerotized and has smaller setae than the remainder of the gonocoxite. A strong sclerotized bridge between the gonocoxal apodeme and the parameral sheath of the aedeagus is present. The lateral edges of the dorsal apodeme of the aedeagus are divergent anteriorly (e.g. Fig. 117). Both ventral and ejaculatory apodemes (e.g. Fig. 117) are distinctly expanded anteriorly. The distiphallus is short. Synapomorphies. 8 ommatidia all the same size (character 7); lateral edges of dorsal apodeme divergent anteriorly (character 36); ventral apodeme expanded into dorso- ventrally flattened bulb anteriorly (character 39). Distribution. This genus is only known from the island of Hispaniola (Fig. 3A). Included species. Nesonana xenoverpa, sp. nov., and Nesonana youngi, sp. nov. **Nesonana xenoverpa, sp. nov. Etymology. Gr., m., xenos: stranger + L., f., verpa: penis; to be treated as a noun in apposition. Diagnosis. There is a second row of setae crossing the median occipital sclerite below the row of postocular setae. The epandrium is more or less parallel sided, with the posterolateral flanges folded medially under the hypoproct and cerci, which are both subequal in length. The dorsal apodeme of the aedeagus is longer than the ventral apodeme. The phallus is considerably more complex than in Nesonanayoungi, with dorsolateral wings folding over the top of the distiphallus. In the 0,the central sac is round. Autapomorphy. Dorsolateral wings folding over top of distiphallus (not included in analysis, but clearly autapomorphic). PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 173 Description ofholoppe 8 (ME1 030366): length 6.0 mm. Head (Fig. 1 1). 0.80 mm long, 1.34 mm wide, 1.13 mm high. Distance between eyes at antennal level 0.41 mm; at genal level 0.56 mm. Antennae brown. Scape 0.17 mm long, 0.1 1 mm wide; bare medially; with fine black setae otherwise evenly distributed, and thick black setae concentrated distally. Pedicel 0.08 mm long. First flagellomere Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 0.24 mm long, 0.15 mm wide; bulbous basally, tapering distally. Style 0.14 mm long; terminal. Antenna1 base to nearest edge of eye 0.09 mm. Frons bulging 0.05 mm beyond eye in lateral view. Parafacial lacking pile, covered with silver pruinescence extending dorsally along eye margin halfway up frons; middle of frons with gold pruinescence; with scattered, fine, black setae (up to 0.26 mm long) on upper frons, except medially; lower frons bare. Genal pile absent. Palpus orange, covered with white pile. Postgenal and occipital pile white; occiput with several black setae. Median occipital sclerite flattened; lacking setae or pile; gold pruinose. Postocular setae black, arranged in single row; with second row below, across median occipital sclerite. Ocellar tubercle with fine, forward-directed, black setae. Ihorux. Scutum 1.47 mm long, 1.25 mm wide; brown pruinose, lighter laterally; covered with recumbent black and pale pile; dorsocentral vittae faintly present as lines of lighter brown pruinescence; 1 pair dorsocentral setae; 3 pairs notopleural setae; 2 pairs supra-alar setae; 1 pair postalar setae. Scutellum brown pruinose with recumbent pale pile. Prosternum with white pile. Pleura entirely silver pruinose. Katatergite with dense, long, white pile. Anepisternum with silver pruinescence and white pile. Katepisternum with silver pruinescence, lacking pile. Proepimeron with silver pruinescence, lacking pile. Lgs. Coxae with silver pruinescence, posterior surfaces less so; fore- and midcoxae each with 2 black macrosetae and long, white pile on anterior surfaces; hindcoxa with 1 black macroseta and long, white pile on lateral surface. Femora dark brown; with appressed, scale-like white pile dorsally; distally, this pile is dark brown. Fore- and midfemora with sparse, erect, white pile. Tibiae orange, becoming dark brown at distal tip; covered with short, erect, black setae. Tarsi dark brown, with basal halves of basitarsi orange. Wng. 5.06 mm long. Basal lobe of costa with scattered setae extending into inner and outer rows of costal setae. Cell m3 closed and stalked. Membrane slightly darkened from stigma through distal r, and r2 cells. Halter brown. Abdomen. Tergites covered with recumbent white and black pile, more erect laterally; tergite 1 also with erect white pile. Tergites dorsally covered with silver pruinescence; laterally, similarly pruinose, except middle of tergite 2, and anterior of tergites 3-5. Sternites with very little silver pruinescence. Tminuliu (paratype, ME1 030356). Sternite 8 with black setae restricted to posterior edge, which is emarginate medially. Tergite 8 dumbell-shaped, with black setae restricted to posterior edge. Epandrium (Fig. 58) 0.51 mm long, 0.62 mm wide at widest point; dark brown; emarginate anteriorly; dorsal surface entirely covered with black setae; lateral edges tapering in posterior half; posterolateral flange 0.20 mm long, bare, tips oriented medially uder hypoproct. Cerci 0.27 mm long; not fused medially; extending posteriorly beyond hypoproct and posterolateral epandrial flange. Subepandrial plate bare ventrally. Gonocoxites (Figs 77 and 78) 0.68mm wide; dark brown; with black setae. Outer gonocoxal process 0.39mm long; with 174 S. D. GAIMARI AND M. E. IRWIN setae of same density and size as gonocoxites. Gonocoxal apodeme contained within anterior edge of gonocoxite; with strong sclerotized bridge to parameral sheath of phallus. Ventral gonocoxal process 0.35 mm long; lightly sclerotized; parallel with outer gonocoxal process; with shorter setae than remainder of gonocoxite. Ventral lobes fused medially at base; reduced. Gonostylus expanded ventrally into lobe; tip Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 extending posteriorly; wider basally than distally; with subapical, lateral spur. Aedeagus in Figures 114 and 115. Lateral edges of dorsal apodeme of aedeagus divergent anteriorly; longer than ventral apodeme. Ventral apodeme distinctly widened anteriorly. Ejaculatory apodeme 0.45 mm long; with distinctly enlarged bulb anteriorly. Lateral ejaculatory process complete dorsally, set into aedeagus. Basal distiphallus with ventrolateral extensions and dorsolateral, medially oriented wings. Distiphallus short, recurved. Description ofalloppe 0 (ME1 102256): 6.8 mm long. Similar to 8 except as follows. Head (similar to Fig. 13). 0.80 mm long, 1.35 mm wide, 1.08 mm high. Eyes separated below anterior ocellus by 0.14 mm. Distance between eyes at antenna1 level 0.42 mm; at genal level 0.56 mm. irhorax. Scutum 1.50 mm long, 1.19 mm wide. Pile on katatergite and anepisternum slightly shorter than on 8. Mng. 5.25 mm long. Abdomen. Tergites brown, dorsally without pruinescence, but tergite 5 with silver pruinescence except for median brown stripe; tergites covered with short, appressed black and sparse white pile. Eminalia (paratype, ME1 030434). Furca 0.33 mm long, 0.29 mm wide; entire, but anterior edge lightly sclerotized; anterolateral wings thin and pointed; furcal bulla lightly sclerotized. Gonopore basal to furcal bulla. Common spermathecal duct 0.27 mm long. Central sac 0.33 mm in diameter; round. Spermathecal ducts originate from equal trifurcation with central sac duct. Dzstribution. Dominican Republic. Specimens examined. Holotype: DOMINICAN REPUBLIC: Pedernales: 3.3 km NE Los Arroyos, 1450m, 16-VII to 18-VII-1990, L. Mavier, J.E. Rawlins, C.W. Young, sweep sample in wet montane forest (8,ME1 030366, CMNH). Paratypes: DOMINICAN REPUBLIC: Pedernales: 3.3 km NE Los Arroyos, 1450 m, 16-VII to 18-VII-1990, L. Masner, sweep sample in wet montane forest (1 8,ME1 030356, MEIC; 1 S,ME1 030360, SDGC; 3 S S, ME1 030350,030361,030368, CMNH), 14.5 km N Cab0 Rojo, 165 m, 18'03" 7 1"39'W, 26-IX to 27-IX-1991, C. Young, S. Thompson, R. Davidson, J. Rawlins, arid thornscrub (1 d, ME1 100164, 1 0, ME1 100191, SDGC; 1 d, ME1 100221, 1 0,ME1 100166, CMNH; 1 8, ME1 100231, USNM), 23.5 km N Cab0 Rojo, 540 m, 13-VII to 25-VII- 1990, L. Masner, J.E. Rawlins, C.W. Young, flight intercept trap in deciduous forest (1 8,ME1 030359, CNCI), 26 km W Cab0 Rojo, 730 m, 13-VII to 25-VII-1990, L. Masner, J.E. Rawlins, C.W. Young, flight intercept trap in wet deciduous forest (1 6,ME1 030435, CMNH); 31-VII-1990, J.E. Rawlins, C.W. Young, S. Thompson (1 0, ME1 030434, CMNH), nr. Cab0 Rojo, 26 km Alcoa Rd, 17-1 to 20-1-1989, J. Swann, S.A. Marshall, in pan trap, (allotype 0,ME1 102256, 1 0, ME1 102255, DEBU); Monte Cristi: 5 km NNE Botoncillo, 50m, 29-XI to 30-XI-1992, R. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 175 Davidson, M. Klingler, S. Thompson, J. Rawlins, arid thornscrub (1 9, ME1 106339, MEIC). **Nesonanayoungi, sp. nov. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Epmolou. Patronym for Chen Young, one of the collectors of this interesting species; to be treated as a noun in apposition. Diagnosis. There is only one row of postocular setae present, with the remainder of the median occipital sclerite bare of setae. The epandrium is distinctly converging posteriorly, and the posterolateral flanges are long, thin, and posteriorly oriented. The cerci are notably longer than the hypoproct. The ventral apodeme of the aedeagus is longer than the dorsal apodeme. In the 9, the central sac is more elongate than in Nesonana xenoperna. Autaponorphies. Epandrium longer than wide (character 44); gonostylus strap-like, not expanded ventrally into lobe (character 57, state 0). Description ofholobpe 8 (ME1 030376): length 5.4 mm. Head (Fig. 12). 0.69 mm long, 1.22 mm wide, 0.98 mm high. Distance between eyes at antennal level 0.32 mm; at genal level 0.44 mm. Antennae brown. Scape 0.17 mm long, 0.09 mm wide; bare medially; with fine black setae otherwise evenly distributed, and thick black setae concentrated distally. Pedicel 0.06 mm long. First flagellomere 0.26 mm long, 0.12 mm wide; bulbous basally, tapering distally. Style 0.1 1 mm long; terminal. Antenna1 base to nearest edge of eye 0.08mm. Frons bulging 0.03mm beyond eye in lateral view. Parafacial lacking pile, covered with silver pruinescence extending dorsally along eye margin halfway up frons; middle of frons with gold pruinescence; with scattered, fine, black setae (up to 0.17 mm long) on upper frons, except medially; lower frons bare. Genal pile absent. Palpus orange, covered with white pile. Postgenal and occipital pile white; occiput with several black setae. Median occipital sclerite flattened; lacking setae or pile; silver pruinose; upper edge and ocellar tubercle with gold pruinescence. Postocular setae black, arranged in single row. Ocellar tubercle with fine, fonvard-directed, black setae. norax. Scutum 1.35 mm long, 1.02 mm wide; brown pruinose, lighter laterally; covered with recumbent black and gold pile; dorsocentral vittae faintly present as lines of lighter brown pruinescence; 1 pair dorsocentral setae; 3 pairs notopleural setae; 2 pairs supra-alar setae; 1 pair postalar setae. Scutellum brown pruinose with recumbent gold pile. Prosternum with white pile. Pleura entirely silver pruinose, as in Figure 40. Katatergite with dense, long, white pile. Anepisternum with silver pruinescence and white pile. Katepisternum with silver pruinescence, lacking pile. Proepimeron with silver pruinescence, lacking pile. hgs. Coxae with silver pruinescence, posterior surfaces less so; fore- and midcoxae each with 2 black macrosetae and long, white pile on anterior surfaces; hindcoxa with 1 black macroseta and long, white pile on lateral surface. Femora dark brown; with appressed, scale-like white pile dorsally; distally, this pile is dark brown. Fore- and midfemora with sparse, erect, white pile. Tibiae orange, becoming dark brown at distal tip; covered with short, erect, black setae. Tarsi dark brown, with basal halves of basitarsi orange. 176 S. D. GAIMARI AND M. E. IRWIN Mng. 3.98mm long. Basal lobe of costa with scattered setae extending into inner and outer rows of costal setae. Cell m3 closed and stalked. Membrane slightly darkened from stigma through distal rl and r2 cells. Halter brown. Abdomen. Tergites covered with recumbent white and black pile, more erect laterally; tergite 1 also with erect white pile, except mediodorsal posterior edge with recumbent, Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 brown pile. Tergites dorsally covered with silver pruinescence; laterally, similarly pruinose, except middle of tergite 2, anterior of tergite 3, and tergite 4. Sternites with little silver pruinescence. Zminalia (paratype, ME1 030417). Sternite 8 with black setae restricted to posterior edge, which is emarginate medially. Tergite 8 dumbell-shaped, with black setae restricted to posterior edge. Epandrium (Fig. 59) 0.53 mm long, 0.45 mm wide at widest point; dark brown; emarginate anteriorly; dorsal surface entirely covered with black setae; lateral edges tapering in posterior hale posterolateral flange 0.20 mm long, bare, thin, and pointed. Cerci 0.23 mm long; not fused medially; extending posteriorly beyond hypoproct and posterolateral epandrial flange. Subepandrial plate bare ventrally. Gonocoxites (Figs 79 and 80) 0.51 mm wide; dark brown; with black setae. Outer gonocoxal process 0.33 mm long; with setae of same density and size as gonocoxites; >2 x length of ventral gonocoxal process. Gonocoxal apodeme extends slightly beyond anterior edge of gonocoxite; with strong sclerotized bridge to parameral sheath of phallus. Ventral gonocoxal process 0.17 mm long; parallel with outer gonocoxal process; with shorter setae than remainder of gonocoxite. Ventral lobes completely fused medially and reduced. Gonostylus strap-like, tip extending posteriorly; wider basally than distally; with subapical, lateral spur. Aedeagus in Figures 116 and 1 17. Lateral edges of dorsal apodeme of aedeagus divergent anteriorly; shorter than ventral apodeme. Ventral apodeme distinctly widened anteriorly. Ejaculatory apodeme 0.30 mm long; with distinctly enlarged bulb anteriorly. Lateral ejaculatory process complete dorsally, set into aedeagus. Basal distiphallus with ventrolateral wings. Distiphallus short, recurved. Description ofalloQpe 0 (ME1 030367): 5.9mm long. Similar to 8 except as follows. Head (Fig. 13). 0.77 mm long, 1.32 mm wide, 1.04 mm high. Eyes separated below anterior ocellus by 0.12 mm. Distance between eyes at antenna1 level 0.39 mm; at genal level 0.5 1 mm. Thorax. Scutum 1.47 mm long, 1.17 mm wide. Pile on katatergite and anepisternum shorter than on 8. I4ng. 4.43 mm long. Abdomen. Tergites brown, dorsally without pruinescence, but tergite 5 with silver pruinescence except for median brown stripe; tergites covered with short, appressed black and sparse white pile. Zminalia (Fig. 147) (paratype, ME1 030436). Furca 0.35 mm long, 0.18 mm wide; entire, but anterior edge lightly sclerotized; anterolateral wings thin and pointed. Gonopore basal to furcal bulla. Common spermathecal duct 0.53 mm long. Central sac 0.77 mm long, 0.23 mm wide. Spermathecal ducts originate on base of central sac. Spermatheca 0.17 mm in diameter; round. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 177 Distribution. Dominican Republic. Specimens examined. Holotype: DOMINICAN REPUBLIC: Pedernales: 26 km W Cab0 Rojo, 730 m, 31-VII-1990, J.E. Rawlins, C.W. Young, S. Thompson (8,ME1 030376, CMNH). Paratypes: same data as holotype (allotype 0, ME1 030367, CMNH), 13-VII to 25-VII- 1990, flight intercept trap in wet deciduous forest (1 d , Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 ME1 030417, MEIC); 16-VII-1992, C.W. Young, R. Davidson, S. Thompson, J. E. Rawlins, mesic deciduous forest with scattered pines (1 8, ME1 030375, SDGC), 13-VII to 20-VII-1990, L. Masner, J.E. Rawlins, C.W. Young, flight intercept trap in wet deciduous forest (1 9, ME1 030354, CMNH; 1 ? , ME1 030355, CASC), 13-VII to 25-VII-1990 (1 9,ME1 030436, MEIC, 2 ? $2, ME1 030409, 030437, CMNH), 18-VII to 25-VII-1990 (1 8, ME1 100182, CMNH), 19-VII to 25-VII- 1990 (1 0,ME1 030369, SDGC; 1 $2, ME1 030363, USNM), 14.5 km N Cab0 Rojo, 165m, 13-VII-1990, J.E. Rawlins, C.W. Young, S. Thompson (1 ?, ME1 030439, CMNH; 1 0,ME1 030438, SDGC). Cyclotelus-group Diagnosis. The female frons usually has reduced or modified pruinescence, ranging from matte or velutinous pruinescence to absence, fully exposing ground colour, as in many Ozodiceromyia. Most members of this group have an antennal first flagellomere that is cylindrical, with a subapical style set into a ventral pit on the first flagellomere. The maxillary palps vary in coloration and vestiture, but are rarely orange. On the upper and lower frons of the 9,setae are usually either lacking or are distributed in defined patches, but are only rarely scattered. The genal apex usually has a patch of different texture or colour from the remainder of the postgena and parafacial. Most members of this group (except Crebraseta and the basal members of Ozodiceromyia) also have a vertical line of reduced pruinescence from the wing base through the meron, encompassing the anepimeron, katepimeron, and meron. In the two largest genera within this group, Ozodiceromyia and Cyclotelus, the foretibia is often swollen relative to the other tibiae, and in Cyclotelus, the forebasitarsus is usually swollen relative the remainder of the foretarsus or other basitarsi. The notum usually has areas of reduced (exposing ground colour) or modified (matte or velutinous) pruinescence, especially on the lateral edges. The basal lobe of the costa usually has two distinct lines of setae on the dorsal surface leading into the inner and outer rows of costal setae, rather than being covered with scattered setae. The epandrium is either only covered with setae on the posterior half (e.g. Cyclotelus and Ozodiceromyia) or is almost entirely covered. The inner gonocoxal process usually (except for Cyclotelus) has apical setae (e.g. Figs 98, 100, and 92). The gonostylus usually lacks a subapical, lateral spur. Synapomorphies. Antenna1 first flagellomere cylindrical (character 4) (onion-shaped in Crebraseta and basal members of Ozodiceromyia); antennal style subapical, set into ventral pit on first flagellomere (character 5) (apical in Coleiana, Crebraseta, and basal members of Ozodiceromyia); 9 upper frons with velutinous pruinescence (character 8, state 3) (shiny black (state 2) in Ozodiceromyia and part of Cyclotelus; with matte pruinescence (state 1) in part of Cyclotelus); ? lower frons shiny black 178 S. D. GAIMARI AND M. E. IRWIN (character 9, state 2) (with normal pruinescence (state 0) in Crebraseta; with velutinous pruinescence (state 3) in Coleiana; with matte pruinescence (state 1) in Cyclotelus); setae of 9 upper frons in defined patches (character 10); genal apex with a patch of different texture or colour (character 14) (lacking patch in Coleiana); vertical stripe of reduced pruinescence from wing base through meron Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 (character 19) (stripe lacking in Crebraseta and part of Ozodiceromyia); notum with areas of reduced pruinescence (character 28); basal lobe of costa with dorsal setae in two rows extending into inner and outer rows of costal setae (character 33) (dorsal setae scattered in Procyclotelus and Crebraseta); apical setae of inner gonocoxal process present (character 53, state 0) (setae absent in Cyclotelus). Cyclotelus-subgroup synapomorphies: Prosternum lacking pile (character 16); midtarsomere 1 longer than midtarsomeres 2-5 inclusive (character 25) (subequal in Cyclolelus); sternite 8 with setae widely distributed (character 59) (restricted to posterior edge in some Cyclotelus); common spermathecal duct origin on posterior part of furcal bulla (character 60, state 2); common spermathecal duct >2 furcal lengths long (character 6 1) (<2 furcal lengths in Cyclotelus). Synapomorphies for Coleiana + Cyclotelus: Notum with patches of matte pruinescence (character 29); lateral edges of dorsal apodeme divergent anteriorly (character 36); ventral apodeme laterally flattened, thinned, and sword-like (character 38) (widened distally in part of Cyclotelus); ejaculatory apodeme expanded distally (character 40, state 0); anterior margin of epandrium not emarginate, straight across (character 46); setae of tergite 8 widespread (character 58, state 0). Ozodiceromyia- subgroup synapomorphies: ? lower frons with setae in defined patches (character 11, state 1) (scattered (state 0) in Crebraseta); genal apex with a patch of short, black pile (character 15, state 1); katepisternum with pile (character 18, state 0); posterolateral corners of epandrium lacking flattened flange (character 45); outer gonocoxal process with setae reduced in density and/or size relative to those of gonocoxites (character 5 1, state 1). Synapomorphies for Crebraseta + Ozodiceromyia: Antenna1 first flagellomere tear-drop or onion-shaped (character 4, state 0) (cylindrical in the more derived members of Ozodiceromyia); antennal style apical on flagellum (character 5, state 0) (subapical in the more derived members of Ozodiceromyia); vertical stripe of reduced pruinescence from wing base through meron lacking (character 19, state 0) (stripe present in more derived members of Ozodiceromyia); anterior margin of epandrium not emarginate, straight across (character 46) (emarginate in some members of Ozodiceromyia); gonostylus curved upward distally by 90" or more (character 55) (extended out distally in few members of Ozodiceromyia). Bugulaverpa, gen. nov. irype species. Bugulaverpa rebeccae, sp. nov. Epmology. Feminine. L. prefix, bu-: large + L., f., gula: gullet, throat + L., f., verpa: penis, referring to the large dorsal lobe on the basal distiphallus. Included species. Bugulaverpa rebeccae, sp. nov. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 179 **Bugulaverpa rebeccae, sp. nov. Epmology. Patronym for Rebecca Marie Gaimari, daughter of the first author, born during this study; to be treated as a noun in apposition. Dzugnosis. The genal apex is enlarged and covered with short black setae and a patch Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 of darkened pruinescence. The frons of the d is silver pruinose, with short setae dorsolateral to the antennal margin. The parafacial is silver pruinose and lacks pile. The cleft of the prosternum lacks pile. Dorsocentral setae are absent. Only 1 pair of supra-alar and 1 pair of scutellar setae are present. The katepisternum has short white pile only on the upper half. The aedeagus is small, with dorsal and ventral apodemes parallel and close together. The distiphallus has a dorsal lobe basally, but is short distally. This is the only cycloteline that lacks a furcal bulla. The lateral edges of the furca are flattened and folded over dorsally. The common spermathecal duct is very short. Autupomorphies. Ommatidia of d the same size on upper and lower halves (character 7); 1 pair of scutellar setae present (character 30); 1 pair of supra-alar setae present (character 31); epandrium longer than wide (character 44); common duct origin on membrane without furcal bulla (character 60, state 0); lacking furcal bulla (character 64, state 0). Description ofholoppe d (ME1 0899 12): length 6.5 mm. Head (Fig. 20). 0.92 mm long, 1.47 mm wide, 1.22 mm high. Distance between eyes at antennal level 0.47 mm; at genal level 0.75 mm. Antennae with grey pruinescence. Scape 0.45 mm long, 0.12 mm wide; bare medially; with fine black setae otherwise sparsely distributed from dorsal view; with dense, longer setae on ventral surface. Pedicel 0.09 mm long. First flagellomere 0.39 mm long, 0.12 mm wide; cylindrical, slightly tapering distally. Style 0.05 mm long; subapical. Antenna1 base to nearest edge of eye 0.17 mm. Ommatidia of same size on upper and lower halves. Frons bulging 0.14 mm beyond eye in lateral view. Parafacial silver pruinose; lacking pile. Frons with silver pruinescence; lower frons with few, short (up to 0.05mm long) black setae dorsolateral to antennal base. Gena enlarged and darkened; with short black setae. Mouthparts broken OK Postgena and occiput with silver pruinescence; pile white; occiput with several black setae. Median occipital sclerite flattened; with fine white pile; with silver pruinescence, except darker near vertex. Postocular setae black, arranged in single row, with shorter black setae near vertex. Ocellar tubercle with gold pruinescence, and short, fine, forward-directed, black setae. norax. Scutum 1.58 mm long, 1.35 mm wide; with silver pruinescence; covered with short, erect, black setae and recumbent, gold pile; median vitta present as line of reduced pruinescence, exposing black ground colour; dorsocentral vittae absent; dorsocentral setae absent; 3 pairs (4 on left side) notopleural setae; 1 pair supra-alar setae; 1 pair postalar setae. Scutellum silver pruinose, with appressed gold pile; 1 pair scutellar setae. Prosternum with white pile. Anepimeron, katepimeron, and meron lacking pile, and with reduced pruinescence, appearing as vertical line from wing base through meron. Katatergite with dense, long, white pile. Anepisternum with silver pruinescence and white pile. Katepisternum with silver pruinescence, and short, white pile along upper edge. Proepimeron with silver pruinescence, lacking pile. 180 S. D. GAIMARI AND M. E. IRWIN Legs. Coxae with silver pruinescence, posterior surfaces less so; fore- and midcoxae each with 2 black macrosetae and white pile on anterior surfaces; hindcoxa with 1 black macroseta and short, white pile on lateral surface. Femora dark brown; with appressed, scale-like white and brown pile dorsally. Forefemur also with erect brown pile. Tibiae and tarsi covered with short, erect, black setae. Tibiae orange; foretibia Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 dark brown on distal half. Tarsi dark brown, with some orange basally. Wing. 5.3 mm long. Basal lobe of costa with setae arranged in line along outer edge extending into outer row of costal setae; second line of setae along base of lobe extending into inner row of costal setae. Membrane hyaline. Cell m3 closed before wing margin and stalked. Halter stalk light brown; knobs broken OK Abdomen. Tergites covered with recumbent, white pile, except recumbent, brown pile in median patches; dorsally and laterally covered with silver pruinescence, except anterior edges showing dark brown ground colour. Sternites with pruinescence similar to tergites. Temzinalia. Sternite 8 with black setae dense along posterior edge; posterior edge emarginate medially. Tergite 8 dumbell-shaped, with black setae dense along posterior edge. Epandrium (Fig. 62) 0.41 mm long, 0.51 mm wide at widest point; light brown; emarginate anteriorly; dorsal surface covered with black setae; lateral edges parallel; posterolateral flange 0.06 mm long, bare. Cerci 0.23 mm long; not fused medially; extending posteriorly slightly beyond posterolateral epandrial flange; subequal in length to hypoproct. Subepandrial plate bare ventrally. Gonocoxites (Figs 86, 87, and 88) 0.56 mm wide; dark brown; with black setae. Outer gonocoxal process 0.20 mm long; with setae along outer edge. Inner gonocoxal process indistinct from outer, with both fine and thick setae at the tip. Gonocoxal apodeme extending slightly beyond anterior edge of gonocoxite; lacking sclerotized bridge to parameral sheath of phallus. Ventral gonocoxal process broad; setae along edge thicker than those of gonocoxite. Ventral lobes enlarged and distinct, fused basally. Gonostylus expanded ventrally into lobe, tip extending posteriorly and with dull subapical spur. Aedeagus in Figures 124 and 125. Dorsal apodeme of aedeagus parallel sided; slightly longer than ventral apodeme. Ventral apodeme nearly as wide as dorsal apodeme, uniformly wide. Ejaculatory apodeme 0.2 1 mm long; thin, stick-like. Lateral ejaculatory process complete dorsally, set into aedeagus. Basal distiphallus expanded into lobe dorsally; distiphallus short. Description ofallogpe 0 (ME1 106349): 9.5 mm long. Similar to d except as follows. Head (Fig. 2 1). 0.95 mm long, 1.58 mm wide, 1.17 mm high. Eyes separated below anterior ocellus by 0.20 mm. Distance between eyes at antennal level 0.60 mm; at genal level 0.78 mm. Antenna1 base to nearest edge of eye 0.23 mm. Palpus brown, covered with white pile. 7horax. Scutum 1.88 mm long, 1.44 mm wide; with short, fine, erect, pale setae and recumbent, gold pile. Wing. 5.9 mm long. Cell m3 closed at wing margin. Halter light brown. Abdomen. Tergites and sternites mostly lacking pruinescence, showing shiny, dark brown ground colour, except posterior edges silver pruinose. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 181 Zminalia (Fig. 150). Furca 0.47 mm long, 0.27 mm wide; medially oriented pegs separate furca into anterior and posterior halves (anterior half 0.23 mm wide); lateral edges dorsoventrally flattened and wing-like, folded over top of furca; anterior edge sclerotized; furcal bulla absent. Common spermathecal duct 0.04 mm long. Spermathecal ducts originate from equal trifurcation with central sac duct. Central Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 sac duct 0.32mm long. Central sac 0.23mm in diameter, round. Spermathecae 0.17 mm in diameter, round. Distribution. This species is known only from Luzon, Philippines (Fig. 3B). Specimen examined. Holotype: PHILIPPINES: Luzon: Manila, 3-IV- 1932, leg. Banks (d, ME1 089912, USNM [originally from CSUC, deposited in USNM with permission]). Paratype: PHILIPPINES: Luzon: Pampanga Prov., Clark Airforce Base, 21-1 to 23-1-1980, C.W. Mills I11 (allotype 9,ME1 106349, BPBM). Coleiana, gen. nov. Irype species. Coleiana nigricopis, sp, nov. Epmology. Feminine. Patronym for Frank R. Cole, one of the great early workers on Nearctic Therevidae, and the first to recognize that part of this tribe forms a natural group. Included species. Coleiana nigricopis, sp. nov. Coleiana nigricopis, sp.nov. Epmology. L., nigos: black+f., copis: knife, referring to the colour and shape of the body; to be treated as a noun in apposition. Diagnosis. This is one of the few Cyclotelus-subgroup members with an apical antenna1 style set on the first flagellomere. This is the only known member of the tribe with a gena that does not protrude at least slightly in lateral view. The midbasitarsus is longer than tarsomeres 2-5 inclusive. This is the darkest coloured of all known Cyclotelini. The patches of matte pruinescence characteristic of most of the Cyclotelus- subgroup are dark brown in this species. Dorsocentral setae are absent. The wing membrane is darkened. This species is sexually dimorphic, more so than most therevids examined: in 9,the halteres and basal part of the wing are dark, while these are yellow in the d. The setae covering the gonocoxites and outer gonocoxal process are dense and uniform, as are the setae covering the epandrium, sternite 8, and tergite 8. The ventral apodeme of the aedeagus is thinned and sword-like. There is a dorsal, shelf-like projection over the distal distiphallus. Autapomorphies. Antenna1 style apical on first flagellomere (character 5, state 0); 9 lower frons with velutinous pruinescence (character 9, state 3); gena not protruding (character 13, state 0); tip of gena with same texture and colour as remainder of gena, postgena, occiput, and parafacial (character 14, state 0); basal distiphallus with dorsal shelf over distal distiphallus (character 43). 182 S. D. GAIMARI AND M. E. IRWIN Description ofholobpe 8 (ME1 028449): length 10.2 mm. Head (Fig. 32). 1.13 mm long, 2.22 mm wide, 1.58 mm high. Distance between eyes at antennal level 0.60 mm; at genal level 0.83 mm. Antennae brown. Scape 0.26 mm long, 0.14 mm wide; bare medially; with fine black setae otherwise distributed evenly, and few thick black setae concentrated distally. Pedicel 0.09 mm long. First Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 flagellomere 0.47 mm long, 0.17 mm wide; bulbous basally, tapering distally. Style 0.17 mm long; terminal. Antenna1 base to nearest edge of eye 0.17 mm. Frons bulging 0.08 mm beyond eye in lateral view. Parafacial silver pruinose; lacking pile. Frons lacking setae; with gold pruinescence; lower frons with patch of brown matte pruinescence lateral to antennal base; upper frons with median diamond-shaped patch of brown matte pruinescence. Genal pile absent. Palpus brown, with clubbed tip; covered with black pile, densest at tip. Postgena and occiput with silver pruinescence and black pile; occiput with several black setae. Median occipital sclerite flattened; with few heavy black setae; covered with gdd pruinescence, except black triangle of matte pruinescence from base of ocellar tubercle. Postocular setae black, arranged in single row. Ocellar tubercle with gold pruinescence, and short, fine, forward-directed, black setae. Thorax. Scutum 2.78 mm long, 1.80 mm wide; dark brown pruinose; covered with short, recumbent, brown pile; median vitta wide, darker brown; dorsocentral vittae present as lines of lighter brown pruinescence; patches of dark brown matte pruinescence on shoulders, except postpronotal lobes with normal pruinescence; dorsocentral setae absent; 3 pairs notopleural setae; 2 pairs supra-alar setae; 1 pair postalar setae. Scutellum dark brown pruinose, with sparse short brown pile. Prosternum lacking pile. Katatergite with thick black setae posteriorly, and dense, long, black pile. Anepimeron, katepimeron, and meron lacking pile, and lacking pruinescence. Anepisternum with light covering of brown pruinescence and black pile. Katepisternum and proepimeron with light brown pruinescence, lacking pile. Legs. Coxae with brown pruinescence, posterior surfaces less so; fore- and midcoxae each with 2 black macrosetae and short dark pile on anterior surfaces; hindcoxa with 1 black macroseta and short dark pile on lateral surface. Femora dark brown; with appressed, scale-like brown pile dorsally. Forefemur with erect and recumbent, brown pile. Tibiae and tarsi dark brown; covered with short, erect, black setae. Midbasitarsus longer than midtarsomeres 2-5 inclusive (Fig. 47). Wng. 9.20mm long. Basal lobe of costa with setae arranged in line along outer edge extending into outer row of costal setae; second line of setae along base of lobe extending into inner row of costal setae. Veins and membrane yellow basally, darkened in distal two-thirds. Cell m3 wide open. Halter yellow. Abdomen. Tergites covered with recumbent, brown pile; dorsally covered with silver pruinescence, except tergite 1 and anterior half of tergite 2; laterally with reduced pruinescence (showing dark brown ground colour). Sternites with very little silver pruinescence. Terminalia. Sternite 8 with black setae widely distributed; posterior edge emarginate medially. Tergite 8 dumbell-shaped, with black setae widely distributed. Epandrium (Fig. 68) 0.83 mm long, 1.13 mm wide at widest point; dark brown; emarginate anteriorly; dorsal surface densely covered with black setae; lateral edges tapering in posterior two-thirds; posterolateral flange 0.17 mm long, bare. Cerci 0.35 mm long; PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 183 not fused medially; extending posteriorly slightly beyond posterolateral epandrial flange; subequal in length to hypoproct. Subepandrial plate bare ventrally. Gono- coxites (Figs 99 and 100) 1.08 mm wide; dark brown; densely covered with black setae. Outer gonocoxal process 0.56 mm long; appearing bifed distally; with setae of same density and size as gonocoxites. Inner gonocoxal process (upper part of Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 bifed outer gonocoxal process tip) indistinct from outer, with apical setae. Gonocoxal apodeme extending slightly beyond anterior edge of gonocoxite; lacking sclerotized bridge to parameral sheath of phallus. Ventral gonocoxal process absent; setae of corresponding edge same as those of remainder of gonocoxite. Ventral lobes distinct, fused basally. Gonostylus expanded ventrally into lobe; tip extending posteriorly, lacking subapical spur. Aedeagus in Figures 138 and 139. Lateral edges of dorsal apodeme of aedeagus slightly divergent anteriorly; slightly longer than ventral apodeme. Ventral apodeme thin, sword-like. Ejaculatory apodeme 0.62 mm long; widened anteriorly. Lateral ejaculatory process complete dorsally, slightly enlarged. Basal distiphallus with dorsal shelf over distal distiphallus. Distiphallus short. Description ofalloppe 9 (ME1 028450): 10.6 mm long. Similar to 8 except as follows. Head. (Fig. 33) [0.98-] 1.02mm long, 1.80 [1.77-2.101 mm wide, 1.32 [-1.381 mm high. Eyes separated below anterior ocellus by [0.24 -1 0.29 mm. Distance between eyes at antenna1 level 0.56 [0.53-0.571 mm; at genal level [0.72 -1 0.75 mm. Upper frons entirely matte brown. [Frons often with row of short black setae between eye margin and midline.] irhorax. Scutum 2.50 mm long, 1.59 mm wide; entirely matte brown dorsally, except for wide. light brown median vitta with thinner dark brown median vitta through middle. Halter dark brown. Midbasitarsus slightly longer than remainder of midtarsus. Wng. 9.20 [8.32-9.441 mm long. Bristles of basal lobe of the costa more scattered than in 8. Veins and membrane darkest basally, becoming lighter in distal two- thirds. Veins slightly darkened distally. Abdomen. Tergites and sternites entirely lacking pruinescence. Tminalia (Fig. 155) (paratype, ME1 088220). Furca 0.47 mm long, 0.33 mm wide; anterior edge not sclerotized; anterolateral wings dorsoventrally flattened; furcal bulla not sclerotized. Gonopore on posterior part of furcal bulla. Common sperma- thecal duct 0.98 mm long. Spermathecal ducts originate from equal trifurcation of central sac duct. Spermatheca 0.20 mm in diameter; round. kriution. One 9 (ME1 088219, in USNM, with the same labels as ME1 028450) similar to the allotype and 2 paratypes, but notum and scutellum entirely orange, except for brown pruinescence on postpronotal lobe and patches of matte pruin- escence on anterior part of notum. Also, mediotergite, anatergite, anepimeron, and posterior edge of anepisternum orange. This specimen was collected with the type series, but represents either a different species or a large amount of intraspecific variation. Distribution. This species is known only from southeastern Brazilian states of Rio de Janiero and Santa Catarina, from Rio de Janeiro (city) southwest beyond Curitiba (Fig. 3C). 184 S. D. C:AIMAKI AND M. E. IRWIN Specimens examined. Holotype: BRAZIL: Santa Catarina State: Nova Teutonia (27’1 l’S, 52’23’W), 300-500 m, X-1966, F. Plaumann (6,ME1 028449, CNCI). Paratypes: BRAZIL: Rio de Janeiro State: Rio de Janeiro, VIII-1938 (allotype 9, ME1 028450, 1 9,ME1 088220, USNM), VII-1938 (1 0,ME1 028451, SDGC). Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Crebraseta, gen.nov. ljpe species Ziereva crassicornis. Bellardi, 186 1. Epmology. Feminine. L., crebra: thick, close, numerous + L., f., seta: bristle, referring to the thick cluster of heavy setae on the posteroventral surface of the gonostylus. Included species As the single species of this genus has not been treated since Lyneborg (1969), a redescription of the holotype 6 and, for the first time, a description of the 9,is presented. * * Crebraseta crassiconiis (Bellardi), comb. nov. 7hereva crassicornis Bellardi, 1861: 88. Type locality: ‘Messico.’ Type: LTd (desig- nated by Lyneborg, 1969: 398, by assumption of HT status) in MRSN. “lhereva’ crassicornis: Lyneborg, 1969: 398 (redescr., LT designation), 400 (illust.: d genit.). Osodiceromya [sic] crassiconzis: Irwin & Lyneborg, 198 la: 257 (listing, comb. change). Diagnosis. This large-bodied therevid is characterized by having a thick scape. The lower frons bulges and is densely covered with setae, although they do not extend down the parafacial. This genus and the basal members of Osodiceromyia are the only members of the Qclotelus-group that lack the vertical stripe of reduced pruin- escence from wing base through meron. Dorsocentral setae are absent. The legs are entirely orange. The gonostylar condyle is large and distinct (Fig. 89). The ventral apodeme of the aedeagus is thinned and sword-like, and the ejaculatory apodeme is stick-like with uniform width (Fig. 127). The gonostylus is unique with its dense cluster of thickened setae on the ventral surface (Fig. 89). In the 9,the ducts at the trifurcation of the spermathecal and central sac ducts are all the same size basally. Synapomorphies. Antennal scape wider than first flagellomere (character 2); median surface of pedicel setose (character 3); 9 lower frons with scattered setae (character 11, state 0); basal lobe of costa with dorsal setae scattered over surface (character 33, state 0); ventral apodeme laterally flattened, thinned, and sword-like (character 38). Redescription ofholoppe 8 (ME1 070536): length 9.9 mm. Head (Fig. 22). 1.22 mm long, 2.42 mm wide, 1.56 mm high. Distance between eyes at antenna1 level 1.14 mm; at genal level 1.34 mm. Scape 0.80 mm long, 0.32 mm wide; pruinose; bare medially; with numerous, otherwise evenly distributed, thick setae (up to 0.54 mm long). Pedicel 0.12 mm long; covered with small setae, including medial surface. First flagellomere broken of€. Antennal base to nearest edge of eye 0.35 mm. Frons bulging 0.17 mm beyond eye in lateral view. Parafacial lacking pile, covered with silver or gold pruinescence extending dorsally through frons, which is PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 185 also silver or gold pruinose; upper from bare; lower from bulging, densely covered with long (up to 0.53 mni) setae. Gena slightly darkened; pile short and black. Palpi silver-grey, covered with yellow pile. Postgena and occiput silver pruinose; pile yellow; occiput with several black setae. Median occipital sclerite flattened; with several black setae; covered with silver or gold pruinescence. Postocular setae black, Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 arranged in single row. Ocellar tubercle pruinose; with long, fine, forward-directed, black setae. irhorax. Scutum and scutellum covered with erect black and recumbent gold pile; pruinescence and vittae not visible due to greasy condition of specimen. Scutum 2.63 mm long, 1.80 mm wide; dorsocentral setae absent; 3 pairs notopleural setae; 2 pairs supra-alar setae; 1 pair postalar setae. Prosternum with white pile. Pleura entirely pruinose. Katatergite, anepisternum, and katepisternum with dense, long, yellow pile. Proepimeron lacking pile. figs. Coxae with silver or gold pruinescence, posterior surfaces less so; fore- and midcoxae each with 2 black macrosetae and long, yellow pile on anterior surfaces; hindcoxa with 1 black macroseta and long, yellow pile on lateral surface. Femora orange, with appressed, scale-like white and black pile dorsally. Fore- and midfemora with erect white and black pile. Tibiae and tarsi covered with short, erect, black setae. Tibiae orange. Forebasitarsus orange basally, becoming dark brown through tarsomeres 2-5. Midtarsus as in foretarsus; all other tarsi broken OK Wng. 9.04mm long. Basal lobe of costa with scattered setae extending into inner and outer rows of costal setae. Cell m, closed and stalked. Membrane unmarked. Halter brown, stalk lighter. Abdomen. Tergites orange, except dorsally with large, posteriorly-oriented, brown triangle on tergites 2-5, and entire tergite 1 brown. Sternites entirely orange. Tminalia. Sternite 8 with white setae restricted to posterior edge, which is emarginate medially. Tergite 8 dumbell-shaped, with white setae restricted to posterior edge. Epandrium (Fig. 63) 0.68 mm long, 0.90 mm wide at widest point; orange; emarginate anteriorly; dorsal surface entirely covered with black setae; lateral edges tapering in posterior third; posterolateral flange 0.17 mm long, bare. Cerci 0.29 mm long; not fused medially; extending posteriorly beyond posterolaterd epandrial flange; subequal in length to hypoproct. Subepandrial plate with setae on ventral surface. Gonocoxites (Figs 89 and 90) 0.96 mm wide; orange; with white setae. Outer gonocoxal process 0.47 mm long; laterally flattened; with setae along outer edge. Inner gonocoxal process indistinct from outer, with short distal setae. Gonocoxal apodeme entirely within anterior edge of gonocoxite; lacking sclerotized bridge to parameral sheath of phallus. Ventral gonocoxal process absent; setae of corresponding edge absent. Ventral lobes distinct, fused basally. Gonostylus strap-like, with cluster of thick setae ventrally; tip upcurved by 90'; with lateral wing-like process, and lacking subapical, lateral spur. Aedeagus in Figures 126 and 127. Dorsal apodeme of aedeagus parallel sided; 2 x longer than ventral apodeme. Ventral apodeme thin, sword- like. Ejaculatory apodeme 0.56 mm long; stick-like, with uniform width. Lateral ejaculatory process complete dorsally, set into aedeagus; lightly sclerotized. Basal distiphallus enlarged dorsally, with median dorsal keel. Description of 9 (ME1 100439, ME1 010766): 10.8 mm long. Owing to the poor condition of the holotype 8,a full description of the female follows. 186 S D. GAIMAKI AND M. E. LRU’IN Head (Fig. 23). 1.20 mm long, 2.42 mm wide, 1.44 mm high. Eyes separated below anterior ocellus by 0.45 mm. Distance between eyes at antennal level 1.07 mm; at genal level 1.22 mm. Scape 0.83 mm long, 0.33 mm wide; brown pruinose; bare medially; otherwise with numerous long, fine black setae, and with thick setae (up to 0.30 mm long) concentrated distally. Pedicel 0.12 mm long; covered with small Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 setae, including medial surface. First flagellomere 0.42 mm long, 0.2 1 mm wide; onion-shaped, tapering to tip. Style 0.17 mm long; terminal. Antenna1 base to nearest edge of eye 0.33 mm. Frons bulging 0.12 mm beyond eye in lateral view. Parafacial lacking pile, covered with silver pruinescence; line of darker brown knap from ventral part of antennal insertion to lateral edge of eye. Upper frons gold pruinose, with scattered long, fine setae, but bare medially; lower frons bulging, densely covered with long (up to 0.27mm) setae. Gena slightly darkened; with patch of short, black pile. Palpi silver pruinose, covered with white pile. Postgena and occiput silver pruinose; pile white; occiput with several black setae. Median occipital sclerite with several black setae; covered with silver pruinescence. Postocular setae black, arranged ‘in single row. Ocellar tubercle gold pruinose; with long, fine, forward- directed, black setae. Thorax. Scutum 2.78mm long, 2.24mm wide; brown pruinose; covered with re- cumbent black and gold pile; median vitta present as thin, darker brown line; shoulders dark brown to nearly black from dorsal view; dorsocentral setae absent; 3 pairs notopleural setae; 2 pairs supra-alar setae; 1 pair postalar setae. Scutellum brown pruinose; covered with recumbent gold pile. Prosternum with white pile. Pleura entirely pruinose. Katatergite with dense, long, white pile. Anepisternum with dark brown pruinescence and dark brown pile on upper half, and silver pruinescence and white pile on lower half. Katepisternum with white pile only along vertical crest of pleurite. Proepimeron lacking pile. Lpgs. Coxae with silver or gold pruinescence, posterior surfaces less so; fore- and midcoxae each with 2 black macrosetae and long, white pile on anterior surfaces; hindcoxa with 1 black macroseta and long, white pile on lateral surface. Femora orange, with appressed, scale-like white and black pile dorsally. Fore- and midfemora with erect white and black pile. Tibiae and tarsi covered with short, erect, black setae. Tibiae orange. Forebasitarsus orange, becoming dark brown through tarsomeres 2-5. Other tarsi entirely orange. Wng. 10.13 mm long. Basal lobe of costa with scattered setae extending into inner and outer rows of costal setae (Fig. 50). Cell m3 closed and stalked. Membrane unmarked. Halter brown, stalk lighter. Abdomen. Tergites orange, except dorsally with large, posteriorly-oriented, brown triangle on tergites 2-5, and entire tergite 1 brown. Sternites entirely orange. Zrminalia (Fig. 151) (ME1 100439). Furca 0.62mm long, 0.39mm wide; anterior edge not sclerotized; anterolateral wings dorsoventrally flattened; furcal bulla not sclerotized. Gonopore basal to furcal bulla. Common spermathecal duct 0.35 mm long. Spermathecal ducts originate from equal trifurcation with central sac duct; all three ducts with same diameter basally. Spermatheca 0.23 mm in diameter; round. Distribution. The type locality, ‘Messico’, is too broadly defined to know its true distribution. However, the female specimens are from Mexico City and Amecameca (about 60 km southeast of Mexico City) (Fig. 3B). PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 187 Specimens examined: MEXICO: ‘165’ [Messico, =Mexico] (holotype 8,ME1 070536, MRSN); District0 Federal: Amecameca (de Juarez), VIII-1900, O.W. Barrett (1 9, ME1 100439, USNM), Mexico, 27-VIII-1922, E.G. Smyth (1 9,ME1 010766, USNM). Nomenclatural bookkeeping. The species Thereva crassicornis Williston ( 1886) was described Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 as a junior, primary homonym of the species of Bellardi (186 1). Cole (1965) erected the new name Dialineura willistoni Cole for the species of Williston, apparently not recognizing the fact that Williston himself (1908) had recognized his own error and erected the new name Thereva pachyceras Williston for his homonymous species. Therefore the name of Cole (1965) is considered an unjustified replacement name (but is the next available name for the species), and the name of Williston (1908) is formally recognized as the valid name, notwithstanding combination changes. Following is the current status for the species. Pallicephala pachyceras (Williston) Thereva crassicornis Williston, 1886: 293 (preoccupied by Thereva crassicornis Bellardi, 186 1: 88). Type locality: California. Type LTd (designated by Webb & Irwin, 1991a) in SEMC. Psilocephala crassicornk Coquillett, 1893b: 197 (comb. change). Dzalineura crassicornk Krober, 19 12g: 2 16 (comb. change); Cole, 1923: 79-80 (redescr.), 1927: 473 (illust.: 8 genit.). Thereva pachyceras Williston, 1908: 206 (replacement name for Thereva crassicornis Williston, nec Bellardi). Dzalineura willistoni Cole, 1965: 352 (unjustified replacement name for Thereva crassicornis Williston). Thereva willistoni: Lyneborg, 1968: 148, 166 (comment on uncertain generic placement), 167-168 (8 genit. descr., comb. change), 169 (illust.: d genit.). Pallicephala willistoni: Irwin & Lyneborg, 1981 a: 206 (illust.: antenna, d genit.), 208 (listing, comb. change), 1981b: 5 15 (illust.: antenna); Webb & Irwin, 1991a: 870 (phylogeny), 876 (key ref.), 893-896 (lectotype design., redescr., gen. revision), 895 (distrib. map). Cyclotelus Walker Cyclotelus Walker, 1850: 4. Type species, Cyclotelus pruinosus Walker, by subsequent designation (Becker, 19 12: 3 15). Roder, 1885: 138 (comment on rel. to Agapophytus); Krober, 1913a: 6 (key ref.), 8 (annot. listing), 9 (key to spp.); Irwin & Lyneborg, 1981a: 203 (key ref.), 25 1 (illust.: d genitalia), 25 1-254 (redescr.). Furcij2r-a Krober, 19 1 1: 524. Type species, Furcijira fascipennis Krober, by subsequent designation (Cole, 1960b: 16 1) ( = Cyclotelus socius Walker, 1850: 6). Krober, 1912g: 220 (redescr.), 1913a: 8 (key ref.), 27 (redescr., key to spp.); Cole, 1960b (revision, illust.: numerous, e.g. heads, antennae, d genit.); Oldroyd, 1968: 377 (comment on rel. to Entesia). Epomyia Cole, 1923: 26. Type species, Thereva pictipennis Wiedemann, by original designation. Cole, 1960b: 161 (prop. syn. of Furcijira). 188 S. D. GAIMARI AND M. E. IRWIN Diagnosis. The genal apex is enlarged, with a patch of darkened pruinescence, lacking pile (e.g. Fig. 35). The upper frons of the 6 usually has a diamond of brown or black matte pruinescence, and the lower frons is silver pruinose (e.g. Figs 34 and 38). The frons of the 9 usually has a well defined central patch of brown or black matte pruinescence (e.g. Fig. 35), and often has smaller patches laterally. The area Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 above this patch of matte pruinescence and below the ocellar tubercle may be shiny black and without pruinescence (e.g. Fig. 35), or can have matte pruinescence (e.g. Fig. 37). The area below the patch is silver pruinose. Parafacial pile is absent. The cleft of the prosternum lacks pile. The mesonotum usually has patches of black matte pruinescence dorsolaterally. Dorsocentral setae are absent. There are 1 or 2 supra-alar setae and 0-2 scutellar setae. The scutellum often has matte pruinescence. The anterolateral surface of the midcoxa has 1-2 macrosetae, and the lateral surface of the hindcoxa has 0-1 macrosetae. The foretibia is swollen and clavate (e.g. Fig. 45). The forebasitarsus is usually swollen relative to other tarsomeres (e.g. Fig. 45). Setae are widely distributed on tergite 8 (e.g. Fig. 53). This is the only member of the Cyclotelus-group that lacks apical setae on the inner gonocoxal process (e.g. Figs 102 and 106). A strong sclerotized bridge often attaches the gonocoxal apodeme and the parameral sheath of the aedeagus. Aedeagus as in Figures 140, 141, 142, 143, 144, and 145. The lateral edges of the dorsal apodeme of the aedeagus are divergent anteriorly. Synupomorphies. 9 upper frons with matte pruinescence (character 8, state 1); 9 lower frons with matte pruinescence (character 9, state 1); foretibia thickened distally and clavate (character 23); forebasitarsus swollen (character 24); midtarsomere 1 subequal in length to midtarsomeres 2-5 (character 25, state 0); common spermathecal duct <2 furcal lengths long (character 61, state 0); basal spermathecal duct wider than common spermathecal duct (character 63). Synapomorphies for Cyclotelus beckeri + Cyclotelus kroeberi: Midcoxa with one macroseta (character 2 1); hindcoxa lacking macrosetae (character 22); 1 pair of scutellar setae present (character 30); 1 pair of supra-alar setae present (character 3 1); ventral apodeme dorsoventrally flattened distally (character 38, state 0); inner gonocoxal process lacking apical setae (character 53). Distribution. Species of the genus Cyclotelus are widespread throughout the Americas (including the Bahama Islands and the Greater Antilles), with notable absence from western North America (Fig. 3C). Specimens examined. Cyclotelus beckeri (Krober): ARGENTINA: Cordoba Province: San Martin, 6-X-1948, F. Lopez (1 6,ME1 028372, IMLA); Salta: Rosario de Lerma, 3- XI to 8-XI-1983, M. Wasbauer (1 9,ME1 028375, CDAE); Tucuman: 23-VI to 28- VI-1951, M.AczelandBach(286,MEI 105272,105273, 19,MEI028374,IMLA), San P. Colalay, 1-1948, leg. Arnay (1 9, ME1 105274, IMLA). BRAZIL: S%oPaulo State: Porto Cabral, Rio Parana, 1-XI to 10-XI-1941, L. Travassos Filho Co. (1 6, ME1 028373, 1 9,ME1 105275, MZSP). Cyclotelus kroeberi (Cole): URUGUAY: Rio Negro: 15 km S. Paysandu, Arroyo Negro, 27-XI1 to 31-XII-1962, R.G. Van Gelder (2 9 9,ME1 076072,076074, AMNH), 1-1 to 6-1-1963 (36 6, ME1 076059,076062, 076063, 1 9,ME1 076081, AMNH). Cyclotelus pictipennis (Wiedemann): UNITED STATES: Florida: Highlands Co., Archbold Biological Station, 12-V to 13-V-1979, H.V. Weems Jr. and Sylvia Halkin, insect flight trap (1 6,ME1 105 184, FSCA), 16-V- 1979 (18,ME1 103276, FSCA), 18-V-1979 (19, ME1 103277, FSCA), 25-V-1979 PHYLOGENY :WD BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE I89 (1 ? , ME1 103274, FSCA), 26-V-1978, H.V. WeemsJr. and Lisa K. Klein, insect flight trap (1 6,ME1 104444, FSCA), 8-X-1979, H.V. Weems Jr. and Thomas A. Webber, insect flight trap (1 0,ME1 105 185, FSCA). Included species. This genus is currently under investigation at the species level, and it is beyond the scope of the current study to present a complete revision, despite Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 the fact that the last consideration of the group was by Cole (1960b), as Furczfera. Below is a list of species known to belong to this genus, including synonymy listings and one new combination. Additional information for many species can be found in Cole (1960b), Irwin & Lyneborg (198 1a), and Irwin & Webb (1992). A full revision of this genus, including redescriptions, descriptions of new species, diagnoses, and keys, will be forthcoming by the authors. Cyclotelus achuetus (Malloch), 1932: 256 (Furr$ra arhaeta). C. badicrasus Irwin & Webb, 1992: 94. ** C. beckeri (Krober), 191 1: 481 (Phyrus). C. bellus (Cole), 1923: 32 (Epomyia bella). C. brazilianus (Cole), 1960b: 166 (Furczjka braziliana). C. colei Irwin & Lyneborg, 1981a: 254 (replacement name for Psilocephala scutellaris Loew) Psilocephala scutellaris Loew, 186913: 17 1 (preoccupied by niereva scutellaris Walker, 1857: 133). C. diversipes (Krober), 19 1 1: 493 (nereria). C. Javipes (Krober), 1928b: 1 13 (Furczfera). Furczfera divprsipes Krober, 1928b: 1 16. C. hardyi (Cole), 1960b: 167 (Furrzfera) (replacement name for Epomyia jlaziipes Hardy) Eponpiajavipes Hardy, 1943: 26 @reoccupied by FurctjiraJlavipes Krober, 1928b: 113). ** C. kroeberi (Cole), 1960b: 169 (Furrzferu kriiberz). C. laetus Walker, 1850: 6. C. longiconzis (Krober), 191 1: 526 (Furczfern). C. n@r@ammus Walker, 1850: 4. Furc@ra$lvipennis Krober, 192813: 114. ** C. pictipennis (Wiedemann), 182 1: 63 (nereva). Psilocephala evthrura Loew, 1869c: 172. C. politus (Krober), 19 1 1: 524 (Furczjka polita). C. pruinosus Walker, 1850: 5. C. rujcornis (Macquart), 1840: 25 (nereva), comb. nov. C. rujiventris (Loew), 1869b: 17 (Psilocephala). Psilocephala lacteipennis Krober, 19 14: 53. C. scutellaris (Walker), 1857: 133 (irhereva). C. silacrasus Irwin & Webb, 1992: 99. C. socius Walker, 1850: 6. Furc@ra fascipennis Krober, 19 1 1 : 526. C. sumichrasti (Bellardi), 186 1 : 9 1 (Psilocephala). Ozodiceromyia Bigot Ozodiceromyia Bigot, 1890: 323. Type species, Ozodiceromvia mexicana Bigot, 1890, by original designation. 190 S. D. GAIMARI AND M. E. IRWIN Krober, 1912g: 210 (key ref.), 21 1 (repr. orig. descr.), 1913a: 6 (key ref.), 8 (repr. orig. descr., cat. cit.); Cole, 1923: 19 (trans. orig. descr., annot. listing); Sabrosky, 1978: 143-1 44 (formally fixed spelling as ‘first reviser’). Ozodiceromya (original misspelling): Bigot, 1890: 32 1 (original descr.); Irwin & Lyneborg, 1981 a: 203 (key ref.), 254 (redescr.). Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Ozodiceronyma (misspelling): Godman, 1901 : 378 (listing). Phycus Walker, 1850: Becker, 1912: 294 (prop. as senior syn.). Psilocephala ‘haemorrhoidalis-group’:Cole, 1923: 34-37 (key ref.), 37 (descr.). Diagnosis. This genus is highly variable and diverse. The genal apex is enlarged and has a patch of darkened pruinescence and shortened, dark setae (e.g. Figs 25, 26, 27, 28, and 29). The frons, especially that of the 9, usually has some shiny black areas (e.g. Figs 25, 27, 28, and 29), but never has patches of matte pruinescence. The lower frons is sometimes bulging at the antenna1 insertion. Parafacial pile is sometimes present (e.g. Figs 24 and 25). The cleft of the prosternum has white pile. The scutum usually displays at least some ground colour, at least on the lateral edges; often with dorsocentral and/or median vittae. The katepisternum is pilose along the vertical median convex crest of the pleurite (e.g. Fig. 41). The foretibia is occasionally swollen and clavate. The epandrium (e.g. Figs 64,65, and 66) is typically parallel sided and lacks a dorsoventrally flattened posterolateral flange, but may be pointed. Setae cover only the posterior half of the epandrium, including the posterolateral corners. The gonostylus is usually expanded ventrally into a lobe, and the tip is usually curved up by 90’ (e.g. Fig. 109). The inner gonocoxal process is knob-like with distal setae (e.g. Figs 92, 94, and 96). The ventral gonocoxal process and the outer gonocoxal process are often present, and are variably setose or bare. The aedeagus and distiphallus are highly variable (e.g. Figs 128, 129, 130, 131, 132, 133, 134, 135). Synapomorphies. 9 upper frons with shiny black, pruinescence-free area (character 8, state 2) (subshiny in few species); setae covering less than posterior half of epandrium (character 47); outer gonocoxal process small or absent (character 50); gonostylus expanded ventrally into lobe (character 57). Distribution. This large and diverse genus has its greatest diversity in the southwestern U.S.A., California, and northern Mexico. Species range from Canada and throughout North America south into Venezuela and Ecuador (Fig. 3B). Specimens examined. Ozodiceromyia argentiJera (Krober): MEXICO: Durango: 1 1.3 km W Durango, 2100 m, 26-VII-1964, W.C. McGuffin (1 9,ME1 052230, SDGC); Hi- dalgo: Guanajuato, 19.3 km W Dolores, 7-IX-1970, E.M. Fisher, J.L. Fisher (1 d, ME1 037789, SDGC);Jalisco: 40.3 km SE Acotomilco, 23-VII-1952,J.W. MacSwain (1 8,ME1 038790, SDGC); MichoacAn: 29 km NW of Quiroga, 2100 m, 22-VIII- 1962, R.H. Painter, E.M. Painter (1 9, ME1 052183, KSUC). Ozodiceromyia nanella (Cole): UNITED STATES: Colorado: Rio Grande Co., South Fork, 2400m, 20- VI-1972, W.W. Wirth, malaise trap (2dd, ME1 054136, 054140, 29 9, ME1 054122, 0541 24, USNM). Ozodiceromyia notata (Wiedemann): UNITED STATES: Illinois: Champaign Co., Urbana, 3-IX-1914, in yard (1 9, ME1 082483, INHS); Pope Co., 1.9 km S Glendale, 8-IX to 10-1X-1978, E.A. Lisowski (1 9,ME1 082597, INHS); New York: Madison Co., Chittenango, 25-VI-1970, D.J. Peckham (26 d, ME1 037564, 037570, USNM). PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 191 Included species. This large and diverse genus is currently undergoing intensive study by the authors. It is beyond the scope of the current work to fully revise the genus despite the fact that it has never been properly revised. Although many new species are recognized and being described, the current work only lists described species, including new synonymies, new combinations, and lectotype designations where Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 necessary. The remaining lectotype designations, redescriptions, descriptions of new species, diagnoses, and keys will be forthcoming in the phylogenetic revision of the genus by the authors. Otodiceromyia anomala (Adams), 1904: 444 (irhereva). 0. algentata (Bellardi), 186 1: 89 (irhereva). Psilocephalafrontalis Cole, 1923: 40, syn. nov. Psilocephala univittata, sensu Williston, 190 1 : 279, nec Bellardi, 186 1 : 90, misident. ** 0. aEentzjra (Krober), 1929: 418 (Phycus algentzjir). 0. arizonensis (Cole), 1923: 45 (Psilocephala). 0. breviventris (Krober), 1928a: 19 (Psilocephala),comb. nov. 0. calgornica (Krober), 19 12g: 259 (7hereva). 0. costalis (Loew), 186913: 11 (Psilocephala). 0. jlavipennis (Cole), 1923: 42 (Psilocephala). 0. gennana (Walker), 1848: 222 (irhereva). Psilocephala johnsoni Coquillett, 1893a: 228, syn. nov. Psilocephala davisi Johnson, 1926: 300, syn. nov. Psilocephala notata, sensu Krober, 1914: 38, and Cole, 1923: 43, nec (Wiedemann), 182 1: 1 14, misident. 0. lateralis (Adams), 1904: 444 (Psilocephala), nom. rev. 0.frommeri Irwin & Lyneborg, 198 1a: 257 (unjustified replacement name for Psilocephala lateralis Adams, nec irhereva lateralis Eschscholtz, 1822: 112.) 0. levipzlta (Loew), 1876: 3 19 (Psilocephala). Psilocephala aldrichii Coquillett, 1893a: 227, syn. nov. Lectotype designation: (ME1 070538) pinned 8 (in USNM) with the following labels: 7-22/Los Angeles Co., Cal./Collection Co- quillett/Cotype No. 10414 U.S.N.M. (orange label). Paralectotypes (examined; neither are the same species as the lec- totype): (ME1 084837) pinned 8 (in MCZC) with the following labels: Jamesburg, NJ, 7.4. (1 8)91/Cotype No. (red label)/P. aldrichi Coq./Paratype (yellow label, referring to Psilocephala jlavipennis Cole)/Type 14443 (red label)/Collection C.W.Johnson/ this specimen is: a syntype of Psilocephala aldrichi Coq. AND a paratype of P. flavipennis Cole (red label); (ME1 033098) pinned Q (in USNM) with the following label data: Mon(tana)/Collection Coquillett/?Cotype No. 10414 U.S.N.M. (orange label). Paralectotypes (not examined): 8, Wyoming, Aldrich collection; 8, locality unknown, but either California, New Jersey, Montana, or Wyoming. 0. melanoneura (Loew), 187 lb: 74 (irhereua, alternate spelling of irhereva). 0. metallica (Krober), 19 14: 68 (irhereva). 'Ihereva melanoneura, sensu Cole, 1923: 91, nec Loew, 1871b: 74, misident. 192 S. D. GAIMARI AND M. E. IRWIN 0. mexicana Bigot, 1890: 321. Euphycus setosus Krober, 19 12g: 2 1 1. 0. milleri (Irwin), 1977a: 294 (Breviperna). ** 0. nanella (Cole), 1960a: 118 (Ziereva; replacement name for irhereva nana Cole) irhereva nana Cole, 1959: 148 (replacement name for irhereva fpgmaea Cole; Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 preoccupied by Ziereva nana Fallkn, 1816: 233). irhereva pygmaea Cole, 1923: 89 (preoccupied by irhereva pygmaea Fallen, 1816: 234) irhereva nigra, sensu Krober, 1914: 69, nec Say, 1823: 40, misident. 0. nigrimana (Krober), 1912g: 238 (Psilocephala). Psilocephala coloradensisJames, 1936: 341, syn. nov. Psilocephalaplatancala, sensu Cole, 1923: 48, nec Loew, 1876: 32 1, misident. ** 0. notata (Wiedemann), 182 1 : 1 14 (irhereva). Ziereva nigra Say, 1823: 40, syn. nov. Lectotype designation: (ME1 084101) pinned 2 (in NHMW) with the following labels: Pensylvania (sic)/nigra, coll. Winthem/nigra Say, Pensylvan. (Wiedemann handwritten)/Psilocephala haem- orrhoidalis (sic) Mcq., det. Krober 191 (sic). irherezia hoemorrhoidalis Macquart, 1840: 26, syn. nov. 0. obliquefasciata (Krober), 19 1 1 : 504 (Psilocephala). 0. platancala (Loew), 1876: 32 1 (Psilocephala). Psilocephala albertensis Cole, 1925: 86, syn. nov. Psilocephala montiradicis James, 1949: 10, syn. nov. 0. proxima (Schiner), 1868: 147 (Psilocephala), comb. nov. Psilocephala stigmaticalis, sensu Schiner, 1868: 146, partim (1 paralectotype), misident . 0. rug$ons (Krober), 19 14: 54 (Psilocephala). 0. schroederi (Krober), 19 1 1: 503 (Psilocephala schmdem). 0. signatipennis (Cole), 1923: 47 (Psilocephala). 0. subnotata (Johnson), 1926: 299 (Psilocephala). 0. univittata (Bellardi), 186 1: 90 (Psilocephala). 0. xanthobasis (James), 1949: 12 (Ziereva). Nomenclatural bookkeeping. The species Psilocephala quadrimaculata Krober was described from one of the five all female syntypes of Psilocephala stigmaticalis Schiner (Krober, 191 1). Since that time, no lectotype has been designated to fix the identity of Psilocephala stigmaticalis. Because neither species has been extensively treated in the literature, and to provide nomenclatural stability, it seems prudent to designate the holotype of Psilocephala quadrimaculata as the lectotype of Psilocephala stigmaticalis, fixing Psilocephala quadrimaculata as a junior objective synonym of Psilocephala stigmaticalis. Below is the lectotype designation, the bibliographic information, and the new combination for Psilocephala stigmaticalis. Penniverpa stigmaticalis (Schiner), comb. nov. Psilocephala stigmaticalis Schiner, 1868: 146. Type locality: Venezuela. Type: LTQ (designated herein) in NHMW. Krober, 191 1: 5 12-5 13 (redescr.). Lectotype designation: (ME1 084125) pinned 0 (in NHMW) with the following labels: Lindig, 1864, Venezuela/stigmaticalis, Alte Sammlung/Type @ink label)/Psilocephala quadrimaculata Krob., PHYLOGENY AND BIOGEOGFUPHY OF CYCLOTELINE THEREVINAE 193 det. Krober 19 1 1/Holotype 0, Psilocephala quadrimaculata Krober, det. S.D. Gaimari 1996. Paralectotypes (series consists of specimens of Ozodiceromyia proxima and an undescribed species of Ozodiceromyia): (ME1 109322) pinned ? (in NHMW) with the following labels: Lindig, 1864, Venezuela/ Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 stigmaticalis, Alte Sammlung/ Psilocephala stigmaticalis Schin., det. Krober 19 1 1/Syntypus 9,Psilocephala stigmaticalis Schiner, det. S.D. Gaimari 1996/0zodiceromyia proxima (Schiner), det. S.D. Gaimari 1996; (ME1 0841 23, ME1 0841 24) pinned 9 9 (in NHMW) with same first four labels as above, with the following additional label: Paralectotype, Psilocephala stigmaticalis Schiner 0 , det. S.D. Gaimari 1996. The fifth syntype was not located, but a d specimen (ME1 0841 22) was present with the same first three labels that could have been the fifth specimen. Psilocephala quadrimaculata Krober, 19 1 1 : 5 14, syn. nov. Type locality: Venezuela. Type: HT 0 ( = LT of Psilocephala stigmaticalis Schiner) in NHMW. Procyclotelus Nagatomi & Lyneborg Procyclotelus Nagatomi & Lyneborg, 1987: 1 16. Type species, Procyclotelus elegans Nagatomi & Lyneborg, by original designation. Included species. The single species in this genus was described thoroughly by Nagatomi and Lyneborg (1987, 1989), so we only provide additional morphological and distributional information for the species, rather than a full redescription. * * Procyclotelus elegans Nagatomi & Lyneborg Procyclotelus elegans Nagatomi & Lyneborg, 1987: 1 17. Type locality: Kyushu, Japan. Type: HTd in KUIC. Nagatomi & Lyneborg, 1989: 349, 350 (key ref.), 351 (illust.: coxa, head), 358 (annot. listing); Majer, 1997: 521 (illust.: antenna), 523 (illust.: d head, coxa). Diagnosis. This species has a shiny black lower frons (Figs 30 and 31), as in most Ozodict7omyia. The notum has dense, appressed, gold pile. Dorsocentral setae are absent. The prosternum lacks pile. The midbasitarsus is longer than midtarsomeres 2-5 inclusive. The ventral surface of the subepandrial plate is setose (Fig. 67). The inner gonocoxal process is indistinct from the outer, with short, apical setae (Fig. 98). The aedeagus is in Figures 136 and 137. The lateral ejaculatory process is greatly enlarged and separated from the aedeagus anteriorly. The ventral apodeme of the aedeagus is expanded anteriorly, and the ejaculatory apodeme is stick-like with uniform width. In 9,the common spermathecal duct is very long, >5 x length of furca. At the trifurcation of the spermathecal and central sac ducts, the ducts are all the same diameter basally. The furcal bulla is large and sclerotized anteriorly (Fig. 154). 194 S. D. GAIMARI AND M. E. IRWIN Autupomorphies. Basal lobe of costa with scattered dorsal setae (character 33, state 0); lateral ejaculatory process enlarged and separated from aedeagus anteriorly (character 35). Description of 9 temzinuliu (Fig. 154) (ME1 076054). Furca 0.53 mm long, 0.35 mm wide; not sclerotized anteriorly; anterolateral wings dorsoventrally flattened; furcal Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 bulla large, not sclerotized. Gonopore on posterior part of furcal bulla. Common spermathecal duct long and coiled (>5 x length of furca). Central sac 0.38 mm in diameter; round. Spermathecal ducts originate from equal trifurcation with central sac duct. Spermathecae 0.17 mm in diameter; round. Distribution. This species has been recorded from Japan and Vladivostok, and its known range is herein recognized to include Korea (Fig. 3C). Specimens examined.JAPAN: Higo: Gokano-Sho, 2 1-VII-1966, A. Nagatomi (1 8,ME1 103265, ZMUC); Hitakatsu: Tsushima Island, 27-V-1968, H. Shima (1 8, ME1 084354, MEIC) Tsushima Island, Mt. Tatsura, 23-V-1965, M. Iga (1 paratype 9, ME1 105277, KUIC); Iwawakisak: 19-VII-1951, Synsirolto (1 8, ME1 105276, ZMUC); Kyushu: Oita Pref., Mt. Sobo, 6-VII to 7-VII- 1980, A. Yonetsu (1 paratype 8,ME1 105278, KUIC). KOREA: 11.3km E Uijongbu, National Forest, 200m, 3-VI- 1954, T.A. Derse (1 0,ME1 076054, AMNH). RUSSIAN FAR EAST: Primo- rskij kraj (region): Dubnjak, 15 km SW Slavjanki, 16-VI-1993, coll. Belokobylskij (1 8, ME1 105279, SDGC); Vladisvostok: Sedanska, 24-VI- 1950, R. Malaise (1 0, ME1 084353, SDGC). BIOGEOGRAPHY Methods of cladistic biogeography (Rosen, 1978; Nelson & Platnick, 198 1; Humphries & Parenti, 1986)were used to compare patterns of cycloteline cladogenesis and the vicariant events in geological history. Congruence between geological and phylogenetic histories provides corroboration that earth history has influenced cycloteline cladogenesis. The areas occupied by Cyclotelini (Fig. 3A-C) and used in this biogeographic analysis are as follows: Asia, North America, Central America, South America, Greater Antilles. Phylogenetic analysis resulted in several well defined genera with > 1 species being analysed, but for the most part, these biogeographic analyses were performed at the genus level. The only exception is the separation of the North and South American members of the genus Cyclotelus. Morphological evidence from more detailed studies in progress suggest that the two faunas are each monophyletic and warrant separate consideration. A resolved area cladogram was derived from the taxon-area cladogram (Fig. 4) using component analysis (Nelson & Platnick, 1981) via COMPONENT version 2.0 (Page, 1993), using the nearest-neighbour interchange algorithm in an heuristic search. Unfortunately, COMPONENT is limited in its ability to implement as- sumption 2 of Nelson & Platnick (198 1) (Page, 1993), although this currently seems to be the most realistic approach for dealing with dispersal (Morrone & Carpenter, 1994; Humphries, 1989) as it treats each geographical occurrence of a taxon separately. As Enghoff (1998) pointed out, by turning off the ‘map widespread associates’ option in COMPONENT, either assumption 1 (Nelson & Platnick, 198 1) PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 195 Ammothereva - Asia Nesonana - Greater Antilles ~~~~i~~~~~ - North America Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Central America Anolinga - South America Bugulauerpa - Asia Crebraseta - Central America Ozodiceromyia - CentralNorth America America Procyclotelus - Asia IColeiana - South America North America rCyclotelus pictipennis group - Central America Greater Antilles € Cyclotelus beckeri group - South America Figure 4. Taxon-area cladogram for the genera of Cyclotelini, including the division of Cycloklus into two species-groups. or 2 will be implemented, depending on whether the widespread taxa are in a terminal or basal position, repectively, on the taxon-area cladogram. In our case, all widespread taxa were in terminal postions (Fig. 4), and thus assumption 1 was effectively followed, allowing area relationships to be either mono- or paraphyletic relative to their widespread taxa. We minimized the optimality criterion ‘leaves added’, which is the number of branches added to reconcile both the taxon and area cladograms. Because counting the number of leaves added to reconcile a taxon cladogram with an area cladogram may overestimate the number of actual events (Page, 1989), we separately analysed the data minimizing the optimality criterion ‘losses,’ which takes into account, for example, that a single loss can account for absence of a larger taxon clade from an area or area clade, potentially resulting in several leaves added. To test for significant congruence between area and phylogenetic relationships (i.e. could the observed pattern have arisen by chance alone), the number of ‘leaves added’ was compared between a random set of 10 000 area cladograms (with labelled, rooted topologies) reconciled with the taxon cladogram (Page, 1993). The percentage of random trees that matched or had less than the number of ‘leaves added’ in the resolved area cladogram provided a measure of probability that the topology did not arise by chance alone. The random area cladograms were calculated using the ‘equiprobable’ model of random tree topology (Page, 1993). I96 S. D. GAIMAKI AND M. E. IRWIN In assessing the probable ancestral area inhabited by early cyclotelines, several quantitative methods were employed. Bremer’s method of ancestral area analysis (Bremer, 1992) was used to compare possible gains versus losses of taxa in each area, respectively, within a ‘gain, no loss’ scenario starting from the root of the taxon-area cladoLgram with no areas represented initially, and a ‘loss, no gain’ Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 scenario starting from the root of the taxon-area cladogram with all areas represented initially. In addition, we used Fitch optimization, according to the protocol of Liebherr (1988a) to deduce the ancestral area for the tribe. As per EnghoFs (1995) assesment, the descending phase (or downpass) of Fitch optimization is sufficient for estimating ancestral areas. Ronquist’s ( 1997) dispersal-vicariance analysis, as implemented by DIVA version 1.1 (Ronquist, 1996), was also an essential tool for quantitatively assessing the probable ancestral area for the tribe. Earb histoly of Tiierevidae Although still under analysis, Therevidae are thought to have a Jurassic, Pangaean origin, primarily based upon fossil evidence and present-day distributions of putatively primitive and derived members of the family. This follows the likely Triassic origin of the Brachycera (Woodley, 1989; Krzeminski, 1992), and the Jurassic origins of Asiloidea as displayed by fossil species representing a variety of asiloids (see Evenhuis, 1994), and by phylogenetic evidence, including that for Mydidae (Yeates & Irwin, 1996; Papavero & Wilcox, 1974) and Apioceridae (Yeates & Irwin, 1996). The Upper Jurassic species Rhagiophvrne bianalis Rohdendorf, 1964 (translation in 1974; illustration in Rohdendorf, 1991: 488, fig. 1091), which is represented by a series of well preserved specimens from the flyscoid deposits of the Karatau mountain ridge in southern Kazakhastan, is the earliest known fossil member of the Therevidae (Mostovski, 1998). In addition, the species Protomphrale marbnovi Rohdendorf, 1938 (illustration in Rohdendorf, 1991 : 49 1, fig. 1094), from the same deposits, represents the earliest known fossil member of the Scenopinidae (Rohdendorf, 1938; Mostovski, 1998), indicating that these two families were likely already differentiated by the Upper Jurassic. Geological histoly The geological history presented herein is a synthesis of pertinent portions of Briggs’ (1 995) review of biogeographical events, Smith, Smith, & Funnell’s (1 994) atlas of ancient coastlines, and Rosen’s (1976, 1985) hypotheses on the origins of the Greater Antilles. Despite the earlier origin of the family, biogeographical evidence herein suggests a Late Cretaceous origin for the Cyclotelini. By this time, 65-90 million years before present (Myr), high sea levels resulted in extensive epicontinental seas. Potentially important to cycloteline cladogenesis was the Mid-Continental Sea (Cox, 1974) separating Euramerica (i.e. eastern North America +western Europe) from West- america which formed a single continent with eastern Asia via the Beringian land bridge. This sea dried up in the Paleocene, but a north-south oriented mountain range (the present-day Rocky Mountains) through western North America remained an effective physical and climatic barrier to biotic movement (Frakes, 1979). In the PHYLOGENY AND BIOGEOGRAPHY OF CYCI,OTEI.INE THEREVINAE 197 late Cretaceous and early Paleogene, these northern latitudes were characterized by boreotropical to paratropical climates (Wolfe, 1975, 1977; Hsii, 1983). The Beringian land bridge was a route common to many groups, including insects (e.g. Matthews, 1979, 1980; Nalepa et al., 1997), plants, mammals, and lizards. Notable examples of insect-plant congruence include several groups of Cerambycidae (Coleoptera) and Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 their host trees displaying this Asian-Westamerican pattern (Linsley, 1963). From the middle to late Paleogene (Eocene into Oligocene), the northern latitudes encountered a steady increase in humidity (Frakes, 1979) and precipitation (Wolfe & Hopkins, 1967), and a drop in temperatures (Frakes, 1979), all of which could have acted as vicariant barriers separating the North American and Asian faunas and floras. Further south, at the juncture of North and South America, Rosen (1976, 1985) proposed an archipelagic bridge, and Savage (1982) and Guyer & Savage (1986) proposed a solid land connection, linking the two continents through the Cretaceous- Paleogene boundary. This proto-Antillean archipelago, or isthmanian link, was on the eastern edge of the Caribbean plate which moved eastward in the early Eocene, to eventually become the present-day Greater Antilles. A second land mass followed on the western edge of the plate, becoming attached to southern North America. This Central American isthmus only reattached North and South America as recently as the Pliocene (less than 5 Myr). Perfit & Williams (1989) review geological events in the tectonic history of the Caribbean, noting that most evidence supports the continental origins of the Greater Antilles, which is also supported by the numerous biological examples in Woods (1989) and Liebherr (1988~).This differs from Briggs’ (1987, 1995) hypothesis that the archipelagic bridge remained in place to later form Central America, concluding that the Greater Antilles were oceanic in origin with subsequent colonization by over-water dispersal. Taxon and area cladogram congruence Early in cycloteline cladogenesis, there was a land bridge connecting Asia and Westamerica. At this time, western North America extended south through the proto-Antillean arc into South America. Five million years after the drying of the Mid-Continental Sea, the next major vicariant event was the eastward movement of the Caribbean plate. This resulted in the separation of the Greater Antilles from the mainland, the resulting separation of North from South America, and the subsequent attachment of present-day Central America into southern North America. This isthmus reattached North and South America less than 5 Myr. The resulting area cladogram of reviewed geological history is as follows: (Asia, (Greater Antilles, (South America, (Central America, North America)))), which is identical to the resolved area cladogram (Fig. 5) revealed by component analysis of the areas currently inhabited by members of the Cyclotelini, following assumption 1, and minimizing either optimality criterion, ‘leaves added’ or ‘losses’. In addition, the area cladograms of Rosen’s (1976) North American-Caribbean track and South American-Caribbean track (see Liebherr, 1988b, fig. 2a-b) are identical to the New World portion of the resolved area cladogram in Figure 5. The randomization tests strongly suggested that the area cladogram topology did not arise by chance alone. Of 10 000 random trees, only 99 trees (i.e. Greater Antilles South America Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Central America North America Figure 5. Resolved area cladogram from component analysis, which is identical to the hypothesized area cladogram from geological evidence. the number of ‘leaves added’ found in the resolved area cladogram after reconciling with the phylogenetic cladogram (5 leaves added, with 5 area leaves). All three methods used for assessing the probable ancestral area for the Cyclotelini pointed to Asia as being their place of origin, with subsequent movement into the New World. Dispersal-vicariance analysis suggested 10 possible dispersal events to account for this Asian origin for the tribe. In comparing each major clade of the Cyclotelini with the area cladogram (Fig. 6A-C), all branches are explainable relative to their distributions (with some duplications), except for the unexplained presence of the North American contingent of Cyclotelus in the Greater Antilles (Fig. 6C). This points to potential problems associated with widespread taxa in biogeographical analyses. Biogeographical synthesis The Cyclotelini represents a diverse assemblage of genera with an apparently Laurasian distribution. Given the current distributions of taxa, the place of origin is likely to be in Asia. The plesiomorphic members of each major clade (i.e. Brevipema- group, Cyclotelus-subgroup, Ozodiceromyia-subgroup) are Asian, so it is likely that each of the groups independently moved from Asia and diversified more fully in the New World. The other alternative is that the tribe originated in the New World, with the plesiomorphic members each moving to Asia through Beringia. This latter possibility is not supported by the analytical methods invoked for assessing ancestral areas. Despite the likely Asian origin for the tribe, there is a relative scarceness of Asian taxa, and more complex distributional patterns are displayed by the New World cycloteline fauna. The lack of cycloteline taxa in Africa, Australia, or Chile suggests that their origin was not Gondwanan, as could be otherwise suggested by the current distribution of the genera Anolinga and Coleiana in southeastern Brazil. The general absence of even primitive members of the tribe from western Europe, and the paucity of taxa and the lack of more plesiomorphic taxa from eastern North America suggests that the cyclotelines diversified in the New World after the late Cretaceous separation of Euramerica and Westamerica by the Mid-Continental Sea. After the Mid-Continental Sea dried up in the Paleocene, few groups (e.g. part of Cyclotelus and few more derived members of Ozodiceromyia) moved eastward; others Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 200 S. D. GAIMARI AND M.E. IRWIN were likely prevented by the north-south oriented mountain range running through western North America. The Beringian land bridge linked North America and Asia. Climatic characteristics, which varied considerably through this geological time, were likely the vicariance inducing factors separating the North American from Asian cycloteline faunas. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Current New World distributions were profoundly affected by the early Eocene eastward migration of the archipelagic or isthmanian bridge between North and South America, providing support for Rosen’s (1976, 1985) hypothesis favouring the continental origin of the Greater Antilles. The widespread North American contingent of Qclotelus occurring on these islands, however, is difficult to explain in the context of vicariance. Many other groups of insects, and numerous non-insect examples, share some of the vicariant patterns found in the present study, including several groups of Antillean Trichoptera (Flint, 1976, 1978; Hamilton, 1988), the dipteran families Drosophilidae (Grimaldi, 1988) and Keroplatidae (Matile, 1990), and the coleopterous subfamily Scarabaeinae (Scarabaedidae) (Halffter, 1974). CONCLUSIONS This study provides the first phylogenetic classification of a monophyletic, supra- generic grouping within the subfamily Therevinae. Using the long hypothesized relationship between Ozodiceromyia and Cyclotelus as a starting point, numerous generic level taxa, both described and undescribed, were considered for inclusion within this putatively monophyletic group. In the end, a large set of synapomorphic characteristic were found to support a monophyletic Cyclotelini, which is currently an assemblage of 10 genera spanning the Americas and extending into Middle, Far East, and Southeast Asia. With time and further collecting efforts, the faunas of South America (especially the underexplored parts of Brazil) and eastern Asia will probably yield additional cycloteline genera, and will likely add further species to the existing genera. ACKNOWLEDGEMENTS This study is part of the doctoral research of SDG, supported by NSF DEB- PEET grant # 95-2 1925 to MEI, in addition to funding provided from the Schlinger Foundation and a graduate research assistantship to SDG through the Illinois Natural History Survey. Heartfelt thanks are extended to Jill D. Mullett, for her superb illustrations; Gail E. Kampmeier (and our cadre of lab assistants, including Mona Abou-Batnin, Jessica Adams, Jamie Bender, Jessica Bickham, Amanda Buck, Shelly Cook, Andrea Gavurnik, Mary Grabowski, Dyanna Gregory, Traci Gustafson, Sarah Kastelic, Sandy Kawanaka, Erin Leslie, Leslie Marsh, Robert Pollok, Ann Ramsey, Stephanie Strothoff, Johnson Zeledon, and Anjun Zhou), for making this research much easier through databasing; Kevin C. Holston, the late Ellis G. MacLeod, Mark A. Metz, Evert I. Schlinger, Donald W. Webb, Brian M. Wiegmann, Shaun L. Winterton, Long-Long Yang, and David K. Yeates, for many fruitful discussions about systematic theory, therevid and asiloid evolution and characters, PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 20 1 distributions and biogeography, and for numerous shared experiences in the field. We also thank Neal L. Evenhuis and F. Christian Thompson for providing helpful nomenclatural and bibliographic insight, Mark A. Metz for translating German primary literature, and Mikhail B. Mostovski for access to unpublished information about Upper Jurassic dipteran fossils in the Rohdendorf collection. The comments Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 of three anonymous reviewers are also deeply appreciated. In addition, we wish to thank the following curators for loans of specimens and for their help during the travels of SDG studying type materials: John E. Chainey (BMNH); Loic Matile (MNHN), Ruth Contreras-Lichtenberg (NHMW), and Vadim F. Zaitsev and Vitali N. Tanasijtshuk (ZISP); and for loans of specimens (some including types), we thank the following curators, in order by collection: David A. Grimaldi (AMNH), Neal L. Evenhuis and Keith Arakaki (BPBM), Paul H. Arnaud and Norman D. Penny (CASC), Eric M. Fisher (CDFA), Chen W. Young (CMNH),Jeffrey M. Cummings (CNCI), Boris C. Kondratieff and Scott J. Fitzgerald (CSUC), E. Richard Hoebeke (CUIC), Stephen A. Marshall (DEBU),Joachim Ziegler (DEIC), Cheryl B. Barr, John A. Chemsak, and Jerry Powell (EMEC), Howard V. Weems, Jr. and Gary J. Steck (FSCA), Guillermo L. Claps (IMLA), Donald W. Webb (INHS), Francisco Fernandez Ykpez (IZAV), H. Derrick Blocker (KSUC), Akira Nagatomi (KUIC), Brian V. Brown (LACM), Philip D. Perkins (MCZC), Mauro Daccordi (MRSN), Nelson Papavero (MZSP),Andrey Sharkov and Norman F. Johnson (OSUC),Robert W. Brooks (SEMC),Wade C. Sherbrooke (SWRS), Edward G. Riley (TAMU), Carl A. Olson (UAIC), Saul I. Frommer (UCRC), Philip J. Clausen (UMSP), Atilano Contreras-Ramos and Harry Brailovsky (UNAMJ, F. Christian Thompson (USNM), Richard S. Zack (WSUC), H. Wendt (ZMHB), and Leif Lyneborg and Verner Michelsen (ZMUC). The maps for Fig. 3 were modified from an-output of RANGEMAPPER, version 2.0, by Kenelm W. Philip. Special thanks are due to our wives, Helen G. Gaimari and Bonnie J. Irwin, for their unending patience and understanding. REFERENCES Adams CF. 1904. Notes on and descriptions of North American Diptera. Kansas Universip Science Bulletin 2: 433455. Arnett RH Jr.,, Samuelson GA, Nishida GM. 1993. The Insect and Spider Collections ofthe World, 2nd edition. Flora and Fauna Handbook No. 1 1. Gainesville: Sandhill Crane Press. Becker T. 1912. Beitrag zur Kenntnis der Thereviden. Kaiserlich-koniglichen Verhandlungen der Carpenter JM. 1988. Choosing among equally parsimonious cladograms. Cladistics 4: 29 1-296. Carpenter JM. 1992. Random cladistics. Cladistics 8: 147-153. Carpenter JM. 1996. Uninformative bootstrapping. Cladistics 12: 177-181. Carpenter JM, Goloboff PA, Farris JS. 1998. PTP is meaningless, T-PTP is contradictory: a reply to Trueman. Cladistics 14: 105-1 16. Cole FR. 1923. A revision of the North American two-winged flies of the family Therevidae. Proceedings Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 ofthe US. National Museum 62(4), 140 pp. Cole FR. 1925. Notes on the dipterous family Therevidae. Canadian Entomologzst 57: 84-88. Cole FR. 1927. A study of the terminal abdominal structures of male Diptera (two-winged flies). Aoceedirgs ofthe Calijhia Acadmy of Sciences 16: 397-499. Cole FR. 1959. A new name proposed in the genus 7lzereva (Diptera: Therevidae). Pan-Puc$c Entomologist 35: 148. Cole FR. 1960a. New names in Therevidae and Bombyliidae (Diptera). Pan-Pac$c Entomologist 36: 118. Cole FR. 1960b. Stiletto-flies of the genus Furctjira Krober (Diptera: Therevidae). Annals $ the Entomological Socieg of America 53: 160- 169. Cole FR. 1965. Family Therevidae. In: Stone A, Sabrosky CW, Wirth WW, Foote RH, Coulson JR, eds. A Catalog ofthe aptera ofAmerica North OfMexico. U.S. Department of Agriculture Agricultural Handbook No. 276. Washington, DC, 348-354. Coquillett DW. 1893a. Synopsis of the dipterous genus Psilocephala. Canadian Entomologist 25: 222-229. Coquillett DW. 1893b. Synopsis of the dipterous genus Thereva. Canadian Entomologist 25: 197-201. Coquillett DW. 1894. Revision of the dipterous family Therevidae. Journal ofthe New rOrk Entomological Sock& 2: 97-101. Cox CC. 1974. Vertebrate palaeodistributional patterns and continental drift. Journal of Biogeogra~~ 1: 75-94. Donoghue MJ, Olmstead RG, SmithJF, Palmer JD. 1992. Phylogeneticrelationships of Dipsacales based on rbcL seouences. Annals of the Missouri Botanical Garden 79 333-345. Domes JA. 1969. The swarming and mating flights of Diptera. Annual Re&w of Entomology 14: 27 1-298. Enghoff H. 1995. Historical biogeography of the Holarctic: area relationships, ancestral areas, and dispersal of non-marine animals. Cladistics 11: 223-263. Enghoff H. 1998. Widespread taxa and COMPONENT 2.0. Cladistics 14: 383-386. Eriksson T. 1997. AUTODECAT, version 2.9.9 (Hypercard stack). Distributed by author: Botaniska institutionen, Stockholm University, Sweden. Eschscholtz JF. 1822. Entomagraphien. Erster Lieferung. Berlin: G. Reimer. Evenhuis NL. 1994. Catalogue ofthe Fossil Flies ofthe WorM (Insecta: Diptera). Leiden: Backhuys. 600 pp. Faith DP. 1991. Cladistic permutation tests of monophyly and nonmonophyly. $stmutic Farris JS. 1995. Conjectures and refutations. Cladistics 11: 105-1 18. Far& JS, KiSllersjii My Kluge AG, Bult C. 1994. Permutations. Cladistics 10: 65-76. Felsenstein J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39 783-79 1. Flint OS, Jr. 1976. The Greater Antilles species of Pobcentropus (Trichoptera: Polycentropidae). Pmceedings ofthe Biological Socieg of Washington 89: 233-246. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Flint OS, Jr. 1978. Probable origins of the West Indian Trichoptera and Odonata faunas. In: Crichton MI, ed. Pmceedings of the 2d Intematwnal Symposium on Tihoptera, 1977. The Hague: Dr W. Junk, 2 15-228. Frakes LA. 1979. Climata Throughout Geologic Zme. Amsterdam: Elsevier. Gaimari SD. 1998. Phylogeny, classification, and biogeography of the cycloteline Therevinae (Diptera: Therevidae). Unpublished Ph.D. Thesis, University of Illinois, Urbana. Gaimari SD, Irwin ME. 2000. Revision of the mexicana-group of the cycloteline genus Otodicmmyia Bigot (Diptera: Therevidae). Aoceedings ofthe Entomological So&& of Wahington 102: 56 1-600. Godman FD. 1901. List of species recorded from Mexico or Central America since 1887, not enumerated in the Supplement. In: Godman FD, Salvin 0, eds. Biologia Centrali-Americana. Class Insecta. Order Diptera. Volume 1 (1886-1901). London: Taylor and Francis, 377-378. Goloboff PA. 1991. Random data, homoplasy, and information. Cladistics 7: 395406. Goloboff PA. 1993. Estimating character weights during tree search. Cladistics 9: 83-9 1. Grimaldi DA. 1988. Relicts in the Drosophilidae (Diptera). In: Liebherr JK, ed. zoogeografihy of Caribbean Insects, Ithaca: Cornell University Press, 183-2 13. Guyer C, Savage JM. 1986. Cladistic relationships among anoles (Sauria: Iguanidae). Systematic James MT. 1936. New species and records of Colorado Diptera. Journal ofthe New Tbrk Entomologcal Sociep 44: 341-344. James MT. 1949. Some new and poorly known Therevidae (Diptera) from Colorado. Annals of the Entomologual Sociep of America 42: 10-1 3. Johnson CW. 1926. New species of Diptera from North Carolina and Florida. Pgche (1925) 32: 299-302. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Kitllersjo M, Farris JS, Kluge AG, Bult C. 1992. Skewness and permutation. Cladistics 8: 275-287. Kluge AG. 1989. A concern for evidence and a phylogenetic hypothesis of relationships among Epicrates (Boidae, Serpentes). Systematic Liebherr JK. 1988b. The Caribbean: Fertile Ground for Zoogeography. In: Liebherr JK, ed. zoogeography of Caribbean Insects. Ithaca: Cornell University Press, 1- 14. Liebherr JK. ed. 1988c. ~oogeographyOJ Caribbean Insects. Ithaca: Cornell University Press. Linsley EG. 1963. Bering arc relationships of Cerambycidae and their host plants. In: Gressitt JL, ed. Pacijic Basin Biogeographv. Honolulu: Bishop Museum, 159-1 78. Loew H. 1856. Neue Beitree tur k‘enntnis der Dzpteren. Vierter Beitrag. Programm K. Realschule Meseritz Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 (= Miedzyrzecz, Poland). Loew H. 1869a. Systmatisrhe Beschreibung der bekannten eumpaischen Zwe$tigel&n Insecten, Lion Johann M.%lhelm Meken. Achter Theil oder zweiter Supplementband. Beschreibung europaischer Dipteren. Erster Band. Halle: H. W. Schmidt. Loew H. 1869b. Diptera Americae septentrionalis indigena. Centuria Octava. Berliner Entomologische zeitschnz 13: 1-52 (in German). Loew H. 1869c. Diptera Americae septentrionalis indigena. Centuria Nona. Berliner Entomologische zeitschnz 13: 130-186 (in German). Loew H. 1871a. Systmatische Beschreibung der bekannten europaischen zwezjltgel~enInsecten, van Johann Mlhelm Aleken. Neunter Theil oder dritter Supplementband. Beschreibung europaischer Dipteren. Zweiter Band. Halle: HW Schmidt. Loew H. 1871b. Diptera Americae septentrionalis indigena. Centuria Decima. Berliner Entomologische zeitxhnj 16: 49-1 15 (in German). Loew H. 1876. Beschreibungen neuer amerikanischer Dipteren. Zeitschnz $r die Gesummten Na- tumiissenschajen 48: 3 17-340. Lyneborg L. 1968. On the genus Balineura Rondani, 1856 (Diptera, Therevidae). Entornologisk Ttdskrift 89: 147-172. Lyneborg L. 1969. Redescriptions of six Therevidae from the Americas, described by J. C. Fabricius and L. Bellardi (Diptera). Entornologiskt Meddelelser 37: 389412. Lyneborg L. 1972. A revision of the Xestomyza-group of Therevidae (Diptera). Annals of the Natal hfuseum 21: 297-376. Lyneborg L. 1976. A revision of the therevine stiletto-flies (Diptera: Therevidae) of the Ethiopian Region. Bulletin ofthe British Museum (Natural Histoy), Entomology 33: 189-346. Lyneborg L. 1980. The South African species of Neotubuda Krober (Diptera: Therevidae). Entomologica Scandinavica 11: 313-342. Lyneborg L. 1983. A review of the Palaearctic genera of Phycinae (Insecta, Diptera, Therevidae). Steenstrupia 9: 18 1-205. Lyneborg L. 1984. Ammothereva, a new Palearctic genus of Therevidae, with a review of the 13 included species (Insecta, Diptera). Stpenstrupia 10: 205-222. Lyneborg L. 1986a. Genera of Therevidae new to the Palaearctic region (Insecta, Diptera). Steenstrupia 12: 61-71. Lyneborg L. 1986b. The genus Acrosathe Irwin & Lyneborg, 1981, in the Old World (Insecta, Diptera, Therevidae). Steenstrupia 12: 10 1- 1 13. Lyneborg L. 1986c. The Palaearctic species of Pandioirilia Irwin & Lyneborg, 1981 (Insecta, Diptera, Therevidae). Steenstrupia 12: 85-98. Lyneborg L. 1988. Revision of Orthartia Krober, 1912, with descriptions of six new species (Diptera: Therevidae: Phycinae). Annals ofthe Natal Museum 29: 537-555. Lyneborg L. 1989. Iberotelus, a new genus of Therevidae (Diptera) from Spain. Eos 64 89-94. Lyneborg L. 1992. Therevidae (Insecta: Diptera). Fauna of New zealand 24. Macquart J. 1840. Dzpt2re.s exotigues nourxaux ou peu ronnus. Tome deuxitme, premitre partie. Paris: NE Roret. Maddison WP, Donoghue MJ, Maddison DR. 1984. Outgroup analysis and parsimony. Systematic ~OO~OQ33: 83-103. Majer JM. 1997. Family Therevidae. In: Papp L, Darvas B, eds. Contributions to a Manual ofPalaearctic Diptera, Volume 2, Nematocera and Lower Brachycera. Budapest: Science Herald, 5 19-529. Malloch JR. 1932. Rhagionidae (Leptidae), Therevidae, Scenopinidae, Mydaidae, Asilidae, Lon- chopteridae. Diptera of Patagonia and South Chile, Volume 5, Fascicle 3: 199-293. Mann JS. 1928. Revisional notes on Australian Therevidae, part 1. Australian ~oologist5: 151-194. Mann JS. 1929. Revisional notes on Australian Therevidae, part 2. Australian MatthewsJV, Jr. 1979. Tertiary and Quaternary environments: historical background for an analysis of the Canadian insect fauna. In: Danks HV, ed. Canada and its Insect Fauna. Memoirs ofthe Entomological SocieQ of Canada 108: 3 1-86. Matthews JV,Jr. 1980. Tertiary land bridges and their climate: backdrop for development of the present Canadian insect fauna. Canadian Entomologist 112: 1089-1 103. McAlpine JF. 1981. Morphology and Terminolow- Adults. In: McAlpine JF, Peterson BV, Shewell Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 GE, Teskey HJ, Vockeroth JR, Wood DM, eds. Manual OfNearctic Dipbra, Volume 1. Research Branch, Agriculture Canada. Monograph 27, 9-63. Morrone JJ, Carpenter JM. 1994. In search of a method for cladistic biogeography: an empirical comparison of component analysis, Brooks parsimony analysis, and three-area statements. Cladbtics 10 99-153. Mostovski MB. 1998. The ear4 stages ofthe evolution of brachyceran Jlies (Diptera Brachycera). Unpublished PhD Thesis, Paleontological Institute of the Russian Academy of Sciences, Moscow (in Russian). Nagatomi A, Lyneborg L. 1987. A new genus and species of Therevidae from Japan (Diptera). Xontyri 55: 116-122. Nagacomi A, Lyneborg L. 1989. The Therevidae (Diptera) of Japan. Japanese Journal of Entomology 57: 347-373. Nalepa CAYByers GW, Bandi C, Sironi M. 1997. Description of Cyptocercus clevelandi (Dictyoptera: Cryptocercidae) from the northwestern United States, molecular analysis of bacterial symbionts in fat body, and notes on biology, distribution, and biogeography. Annals ofthe Entomological SocieQ of America 90 416-424. Nelson GJ. 1969. Gill arches and the phylogeny of fishes, with notes on the classification ofvertebrates. Bulletin ofthe American Museum ofNatural Histov 141: 475-552. Nelson GJ. 1972. Phylogenetic relationships and classification. systematic Ronquist F. 1996. DIVA, version 1.1. Uppsala, Sweden: Uppsala University (computer program and manual available by anonymous FI’P from ftp.uu.se or ftp.systbot.uu.se). Ronquist F. 1997. Dispersal-vicariance analysis: a new approach to the quantification of historical biogeography. Systmatic Biology 46: 195-203. Rosen DE. 1976. A vicariance model of Caribbean biogeography. Systematic 431464. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Rosen DE. 1978. Vicariant patterns and historical explanation in biogeography. Systematic Wheeler WC, Cartwright P, Hayashi CY. 1993. Arthropod phylogeny: a combined approach. Cladistics 9 1-39. White A. 1916. The Diptera-Brachycera of Tasmania. Part 11. Families Tabanidae & Therevidae. Papers and Proceedings of the Royal Sock& of Tismania (19 15). Wiedemann CRW. 1821. Diptera exotica. Pars I. Tabulis aeneis duabus. Kiliae (=Kid). Wiley EO. 1980. Phylogenetic systematics and vicariance biogeography. Systematic Botany 5: 194-220. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Wiley EO. 1981. Phylogenetics: The Theoly and Practice of Phylogenetic Systematics. New York: Wdey Interscience. Williston SW. 1886. Dipterological notes and descriptions. Transactions of the American Entomological Society and Proceedings of the Entomological Section of the Academy of Natural Sciences of Philadelphia 13: 287-307. Williston SW. 1901. Supplement. In: Godman FD, Salvin 0, eds. Biologia Centrali-Americana. Class Insecta. Order Diptera. Volume 1 (1886-1901). London: Taylor and Francis, 273-296. Williston SW. 1908. Manual ofNorth Ameriian Diptera, Third edition. New Haven: J. T. Hathaway. 405 pp. Winterton SLYIrwin ME, Yeates DK. 1999. Systematics of Nanexila Winterton & Irwin, gen. nov. (Diptera: Therevidae) from Australia. Invertebrate Tmonomy 13: 237-308. Wolfe JA. 1975. Some aspects of the plant geography of the northern hemisphere during the Late Cretaceous and Tertiary. Annals ofthe Missouri Botanical Garden 62: 264-279. Wolfe JA. 1977. Paleogene floras from the Gulf of Alaska region. U.S. Geological Sumg Professional Papers 997. Wolfe JAYHopkins DM. 1967. Climatic changes recorded by Tertiary land floras in northwestern North America: Tertiary correlations and climatic changes in the Pacific. In: Hatai K, ed. Ertialy Correlations and Climatic Changes in the Paczjc, Symposium 25. Tokyo: Eleventh Pacific Science Congress, 67-76. Woodley NE. 1989. Phylogeny and classification of the ‘Orthorrhaphous’ Brachycera. In: McAlpine JF ed. Manual ofNearctic Diptera, Volume 3. Research Branch, Agriculture Canada. Monograph 32: 1371-1395. Woods CA (ed.). 1989. APPENDIX 1 List of taxa coded for analysis of the Cyclotelini, and their distributions ~~~ Outgroup Distribution Generic distribution Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Brachvlinga rinena Central America New World halineura gomdkooi Holarct ic Holarctic Pandiiirilia limata western Nearctic Holarctic T?ienna nebulosa western Nearctic Holarctic Ammothhrreua latzromu eastern Palearctic Middle Asia Aniniohrez~alehn Uzbekistan, Kazhakstan .4nnlinga melanolhnr southeastern Brazil southeastern Brazil Anohnga psdojons southeastern Brazil Brezxpema placuia southwestern Nearctic southwestern Nearctic Bugulaiserpa rebeccae Philippines Philippines CoLiana nigncoprc southeastern Brazil southeastern Brazil Crebrasela rrassicomiJ southern Mexico southern Mexico Cyclolplus becken Neotropical New World Qrlolplus kmebm Neotropical cyrlotelus pict2pennz.r eastern Nearctic NeJonana renouerpa Hispaniola Hispaniola Nesonana-voury Hispaniola Ozohcemmyia axent&a Mexico Nearctic & northern Neotropics Ozosimeonyza nanella western Nearctic Ozodicemmyza notata eastern Nearctic Aogdotplus elegans Asian Far East Asian Far East APPENDIX 2 Data matrix used in analysis of the Cyclotelini I 10 20 30 40 50 60 ~ ~ ~~ Brarhplinga cinerea 0000000000 0000001100 0000000000 0000000100 0000000000 0210101000 00000 halineura gomdkoui 0100000000 0000000000 0000000000 0010001000 0001010000 0000010000 00001 Pandizirilia limata 0000001000 2000210000 0000000000 0000001001 0001000001 1100010000 01100 ?;hema npbulosa 0001000100 0000001000 0000000000 0000000000 0000000001 1100000110 00000 Amniothereua laticomis 0000000000 2 I I02 I I I0 I 000000 1000 0 I0 I00000 I 000 I000 I20 02 I I0 10 I0 I 000 I 1 Ammolhereua lehn 0000000000 2110211101 0000001000 0101000001 0001000120 0211010101 0001I Anohnga melanothrir 0000000000 2110101001 0000000000 0101000100 1100000220 2211011102 11011 Anolinga psilofins 0000001000 2110211101 0000000000 0101000100 1100000220 2211011102 11011 Breuipenia placida 0000010000 0110200001 0000000000 0001101100 1100000220 0211001011 11011 Bugulaveqa rebeccae 000110132I 1011101011 0000000101 1111000001 0001100220 I201000100 00001 Coleiana nigricopis 0001000331 2000211111 0000100110 0111010100 0010010220 0201000012 10011 Crebraseta erassicomis 0110000301 0011101001 0000000100 0101000101 0000110220 1201100101 00011 Cyclolplrcs bechi 1011100111 2011211111 111101011I 1111011010 0000010220 0211001012 00111 Qclolplus kroebm’ 0001100111 2011211111 11110001II 1111010000 0000010220 0211000012 00111 Civclotelus pictipmnir 0001100211 2011211111 0011000110 0111010100 0100010220 0211000002 00111 .Nesonana uozoverpa 0000001000 2110201101 0000000000 0101011010 0100000220 0211011101 01011 Nesonanayoui2gi 0000001000 21 10201 101 0000000000 010101 1010 0101000220 021 1010101 0101 1 Ozodicemmyia axentiJia 1011100221 1011101011 0010000100 0111000000 0000101221 1201111101 00011 Ozodiceromvia nanella 0000000221 1011101001 0000000100 0111010001 0000111221 1201101101 00011 Ozodicemtnyia notata 0001100221 1011 10101I 0010000100 0111000001 0000111221 1201111101 OIOII Ao(-vrln!elus elegans 00011003’21 201 121 I1 11 0000100100 0101 100001 0000000220 02010001 12 1001 1 210 S. D. GAIMARI AND M. E. IRWIN APPENDIX 3 Classification for the genera of Cyclotelini, including informal genus-groups and subgroups Family Therevidae Subfamily Therevinae Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Tribe Cyclotelini, tibe nov. Brevipema-grou p Genus Ammothereva Lynehorg Genus Nesonana, gen. nov. Genus Brevipema Irwin Genus Anolinga, gen. nov. Cyclotelus-group Cyclotelus-subgroup Genus Procyclotelus Nagatomi and Lyneborg Genus Coleiana, gen. nov. Genus Cyclotelus Walker Ozodicemmy ia-su bgroup Genus Bugulaoerpa, gen. nov. Genus Crebrateta, gen. nov. Genus Ozodicemmyia Bigot PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 21 1 APPENDIX 4 List of repositories and their abbreviations. AMNH American Museum of Natural History, New York, New York. BMNH The Natural History Museum, London, United Kingdom. Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 BPBM Bernice P. Bishop Museum, Honolulu, Hawaii. CASC California Academy of Sciences, San Francisco, California. CDFA California State Collection ofArthropods, California Department of Food and Agriculture, Sacramento, California. CMNH Carnegie Museum of Natural History, Pittsburgh, Pennsylvania. CNCI Canadian National Collection of Insects, Ottawa, Ontario, Canada. CSUC C.P. Gillette Arthropod Biodiversity Museum, Colorado State University, Fort Collins, Colorado. CUIC Cornell University Insect Collection, Ithaca, New York. DEBU University of Guelph Insect Collection, Guelph, Ontario, Canada. DEIC Deutsches Entomologisches Institut, Eberswalde, Germany. EMEC Essig Museum of Entomology, University of California, Berkeley, California. FSCA Florida State Collection of Arthropods, Gainesville, Florida. IMLA Instituto Miguel Ldo, Universidad Nacional de Tucuman, Tucuman, Argentina. INHS Illinois Natural History Survey, Champaign, Illinois. IZAV Instituto de Zoologia Agricola, Universidad Central de Venezuela, Maracay, Venezuela. KSUC Kansas State University Insect Collection, Manhatten, Kansas. KUIC Kagoshima University Insect Collection, Kagoshima, Japan. LACM Los Angeles County Museum of Natural History, Los Angeles, California. MCZC Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts. MEIC Michael E. Irwin private collection, for eventual deposit with CASC. MNHN Museum National #Histoire Naturelle, Paris, France. MRSN Spinola Collection, Museu Regionale di Scienze Naturali, Torino (=Turin), Italy. MZSP Museu de Zoologia, Universidade de SPo Paulo, SPo Paulo, Brazil. NHMW Naturhistorisches Museum Wien, Wien (=Vienna), Austria. OSUC Ohio State University Collection of Insects and Spiders, Columbus, Ohio. SDGC Stephen D. Gaimari private collection. SEMC Snow Entomological Collection, University of Kansas, Lawrence, Kansas. SWRS Southwestern Research Station of the American Museum of Natural History, Portal, Arizona. TAMU Texas A&M University Insect Collection, College Station, Texas. UAIC University of Arizona Insect Collection, Tucson, Arizona. UCRC UCR Entomological Research and Teaching Collection, University of California, River- side, California. UMSP University of Minnesota Insect Collection, St. Paul, Minnesota. UNAM Coleccion Nacional de Insectos, Universidad Nacional Autonoma de Mexico, Mexico, Distrito Federal, Mexico. USNM United States National Insect Collection, United States National Museum, Washington, District of Columbia. WSUC Maurice T. James Entomological Collection, Washington State University, Pullman, Washington. ZMHB Museum fur Naturkunde, Humboldt Universtitat, Berlin, Germany. ZMUC Zoological Museum, University of Copenhagen, Copenhagen, Denmark. ZISP Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia. 212 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 7-1 1. Head (left antenna removed). 7-8. Ammotherem Laticomis: 7. 6 (ME1 105180), 8. 9 (ME1 104436). 9-10. Ammotherevu Lehn 9. d (ME1 105326), 10. 9 (ME1 105324). 11. Nesonana xenoverpa: d (ME1 030366). Scale bar = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLO‘TELINE THEREk’INAE 213 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 12-17. Head (left antenna removed). 12-13. .Nesonanayoungi 12. d (ME1 030376), 13. ? (ME1 030367). 14-15. Brevipernaplacida: 14. d (ME1004027), 15. P (ME1004028). 16-17. Anolinga rnelanothrix: 16. d (ME1 100224), arrow points to small patch of black setae on gena, 17. ? (ME1 10021 1). Scale bars=0.2 mm. 214 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 18-23. Head (left antenna removed). 18-19. Anolingu psilgmns: 18. 8 (ME1 077736), 19. ? (ME1 077735). 2Ck-21. Bugulaverpa rebeccae: 20. 6 (ME1 089912), arrow points to patch of black setae on gena, 21. ? (ME1 106349). 22-23. Crebrueta crcrrsicornis: 22. 6 (ME1 070536) (note, flagellum missing), 23. 0 (ME1 100439). Scale bars = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 215 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 24-27,Head (left antenna removed). 2G25. Ozodiceromyia nanella: 24. 6 (ME1 054140), 25. 9 (ME1 054124). 26-27. Ozodzcemmyia argent$mu: 26. 6 (ME1 038790), 27. 0 (ME1 052183). Scale bar= 0.2 mm. 216 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 28-33. Head (left antenna removed). 28-29. O~odiceromyzu notutu: 28. 6 (ME1 037564), 25. ? (ME1 082483). 30-31. Procyclotelus &guns: 30. d (ME1 103265), 31. ? (ME1 105277). 32-33. Coleiunu nz@copix 32. d (ME1 028449), 33. 0 (ME1 088220). Scale bar = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 217 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 34-39. Head (left antenna removed). 34-35. Cyclotelus pictipennix 34. d (ME1 104444), 35. 0 (ME1 105185),arrow points to textured patch on gena. 36-37. Cyclotelu beck 36. 6 (ME1 028372), 37. 0 (ME1 028374). 38-39. Cyclotelus kmeben’: 38. 6 (ME1 076059), 39. ? (ME1 076074). Scale bar = 0.2 mm. 218 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 40, 41. Lateral thorax. 40. Neeonanagoung: 6 (ME1 030376). 41. Ozodiceronyia argentfira: 8 (ME1 037789). Labels: as = anepisternum, am = anepimeron, p = proepimeron, k = katepisternum, m = meron. Scale bars=0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 219 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 42-47, Antenna, foreleg, and midtarsus. 42-43. Antenna, medial view: 42. Ozodicemmyia cal$rnica 8 (ME1 038294), 43. Ozodiceromyia rostalis 8 (ME1 037741). 44-45. Foreleg: 44. Ozodicemmyia nanella ? (ME1 054122), 45. ~_vclolpluspirtipennis ? (ME1 105185). 46-47. Midtarsus: 46. Ozodiceromyia nanella ? (ME1 054122), 47. Coleiana nigncopis Q (hlE1 028450). Scale hars=0.2 mm. S. D. GAIhlARI AND hI. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 48-5 1. Basal lobe of' costal vein on wing. 48-49. \'entral view: 48. Brevzpenza placzda 9 (hlEI 004028), 49. Ozodzcerovyza algentlfPra 8 (ME1 038790). 50-5 I. Dorsal view: 50. CrebraJeta craszcomzr P (ME1 100439), 51. O~odiceromyzaagnt@ra P (hlEI 052 183) Scale bar = 0.2 mm. PHYI.OGENY AND BIOGEOGRkPHY OF CYCLOTELINE THEREVINAE 22 1 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 52-57. 8 pregenitalia (tergite 8 lobe, sternite 8) and epandrium. 52-53. Lobe of tergite 8: 52. Oiodicemmyia argeiitijira (ME1 037789), 53. Cyrloklus becken' (ME1 028372). 5.1-55. Sternite 8: 54. Ozodicemmyia argentijira (ME1 037789), 55. Cvrlotelus becken' (ME1 028372). 56-57. Epandrium: 56. Ammothereva laticomis (ME1 105 180), dorsal view, 57. Anzmolhereru lehri (ME1 105326), split view (left, dorsal; right, ventral). Scale bar = 0.2 mm. 222 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 58-63. 6 epandrium. 58. Nesonana xenoverpa (ME1 030356), dorsal view. 59. Nesonana youngi (ME1 03041 7), dorsal view. 60. Breviperna placida (ME1 004027), split view (left, dorsal; right, ventral), arrow points to posterolateral flange. 61. Anolinga mrlanothrix (ME1 100224), dorsal view. 62. Bugulaverpa rebeccae (ME1 089912), dorsal view. 63. Crebraseta crauicomis (ME1 070536), split view (left, dorsal; right, ventral), arrow points to subepandrial plate. Scale bars=0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE 'I'HEREVINAE 223 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 64-69. 6 epandrium. 64. Ozodiceromyia natiella (hlEI 0541 36), dorsal view. 65. Ozodiceromyia apnt@a (ME1 037789), split view (left, dorsal; right, ventral). 66. Ozodiceromyia notata (ME1 037570), ventral view, arrow points to subepandrial plate. 67. Proyclotelus degans (hIEI 084354), split view (left, dorsal; right, ventral). 68. Coleiana nigricopis (ME1 028449), dorsal view. 69. Cjclotdus pictipennis (ME1 103276), split view (left, dorsal; right, ventral). Scale bars = 0.2 inin. 224 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 70-74. 8 epandrium and gonocoxites. 70-7 1. Epandrium: 70. Cyclotelus becka' (ME1 028372), split view (left, dorsal; right, ventral), 71. Cyclotelus kmeba' (ME1 076063), split view (left, dorsal; right, ventral). 72-74. Gonocoxites of Aminolhereua laticomis (ME1 105180): 72. split, ventral view, 73. split, dorsal view, 74. lateral view (gonostyli and ventral lobes not shown). Scale bars = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 225 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 75-80. 6 gonocoxites. 75, 76. Ammotherezlu lehri (ME1 105326): 75. split view (left, ventral; right, dorsal), 76. lateral view (gonostyliand ventral lobes not shown). 77, 78. Nesonunu xenoverpu (ME1 030356): 77. split view (left, ventral; right, dorsal), arrow points to outer gonocoxal process, 78. lateral view (gonostyli and ventral lobes not shown). 79, 80. Nesonunu youngi (ME1 03041 7): 79. split view (left, ventral; right, dorsal), arrow points to ventral gonocoxal process, 80. lateral view (gonostyli and ventral lobes not shown), arrow points to ventral gonocoxal process. Scale bar = 0.2 mm. 226 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 81-85. d gonocoxites. 81, 82. Brcvipernu plucidu (ME1 004027): 81. split view (left, ventral; right, dorsal), arrow points to gonostylar condyle, 82. lateral view (gonostyli and ventral lobes not shown). 83. Anolingu melunothrix (ME1 100224), split view (left, ventral; right, dorsal). 84, 85. Anolingu psilofins (ME1 077736): 84. ventral view, arrow points to subapical lateral spur of gonostylus, 85. lateral view (gonostyli and ventral lobes not shown). Scale bars = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 227 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 86-90. 8 gonocoxites. 86-88. Bugulaverpa rebeccae (ME1 089912): 86. ventral view, 87. split dorsal view, 88. lateral view (gonostyli and ventral lobes not shown). 89-90. Crebraseta cra.ssicornzC (ME1 070536): 89. split view (left, ventral; right, dorsal), arrow points to gonostylar condyle, 90. lateral view (gonostyli not shown), arrow points to ventral lobe. Scale bars = 0.2 mm. 228 S. D. GAIhIARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 91-96. 8 gonocoxites. 91-92. O~odzceronyiunanellu (ME1 054136): 91. split view (left, ventral; right, dorsal), 92. lateral view (gonostyli not shown), arrow points to outer gonocoxal process. 93-94. Oiodiceromyia agnt$ra (ME1 037789): 93. split view (left, ventral; right, dorsal), arrow points to hypandrium, 94. lateral view (gonostyli and ventral lobes not shown), arrow points to outer gonocoxal process. 95-96. Ozodiceromyiu not& (ME1 037570): 95. split view (left, ventral; right, dorsal), 96. lateral view (gonostyli and ventral lobes not shown), arrow points to inner gonocoxal process. Scale bars = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 229 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 97-102. 6 gonocoxites. 97-98. Aocyclotelus &guns (ME1 084354): 97. split view (left, ventral; right, dorsal), 98. lateral view (gonostyli and ventral lobes not shown), arrow points to inner gonocoxal process, fused with outer gonocoxal process. 99, 100. Coleiuna nigricopis (ME1 028449): 99. split view (left, ventral; right, dorsal), 100. lateral view (gonostyli and ventral lobes not shown), arrow points to inner gonocoxal process. 101, 102. Qclotelzls pirtipennk (ME1 103276): 101. split view (left, ventral; right, dorsal), 102. lateral view (gonostyli and ventral lobes not shown). Scale bars=0.2 mm. 230 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 103-109. 0 gonocoxites and gonostylus. 103-106. Gonocoxites. 103, 104. Cyclotelus beck& (ME1 028372): split view (left, ventral; right, dorsal), arrow points to hypandrium, 104. lateral view (gonostyli and ventral lobes not shown). 105, 106. Cyclotelus kroeben' (ME1 076063): 105. split view (left, ventral; right, dorsal), 106. lateral view (gonostyli and ventral lobes not shown). 107-109. Gonostylus, lateral view: 107. Coleiana nigricopk (ME1 028449), 108. O~odicemmyzanigrimana (ME1 044582), 109. Ozodicemmyia agent@ra (ME1 037789). Scale bars =0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 23 I Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 110-1 17. 6 aedeagal complex. 110, 1 11. Ammohrma laticornu (ME1 1051 80): 1 10. lateral view, 111. dorsal view. 1 12, 113. Ammothema lehn (ME1 105326): 112. lateral view, 113. dorsal view. 1 14, 115. Ntmnana xenoverpa (ME1 030356): 114. lateral view, 115. dorsal view. 116, 117. Ntsanana yourgz (ME1 030417): 116. lateral view, arrow points to ventrolateral wing-like process, 117. dorsal view. Scale bars = 0.2 mm. 232 S. D. GAIMARI AND hf. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 118-125. 6 aedeagal complex. 1 18, 1 19. Brevipemu plucidu (ME1 004027): I 18. lateral view, arrow points to ventrolateral wing-like process, 119. dorsal view, arrow points to lateral ejaculatory process. 120, 12 1. Anolingu melunothrix (ME1 100224): 120. lateral view, 12 1. dorsal view. 122, 123. dnolingu psilofions (ME1 077736): 122. lateral view, 123. dorsal view, arrow points to ventrolateral wing- like process. 124, 125. Buguluverpu rebeccae (ME1 089912): 124. lateral view, 125. dorsal view. Scale bars = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 233 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 126-133. d aedeagal complex. 126, 127. Crebrmb urnsicomis (ME1 070536): 126. lateral view, 127. dorsal view. 128, 129. OzodiCemmyiu nunella (ME1 054136): 128. lateral view, 129. dorsal view. 130, 131. Orodzremmyia anomulu (ME1067707): 130. lateral view, 13 1. dorsal view. 132, 133. Ozodimmyia ugmtfiu (ME1 037789): 132. lateral view, arrow points to ventral apodeme, 133. dorsal view, arrow points to dorsal apodeme. Scale bars = 0.2 mm. 234 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 134-141. 6 aedeagal complex. 134, 135. Ozodicemmyiu notutu (ME1 037570): 134. lateral view, 135. dorsal view, arrow points to lateral ejaculatory process. 136, 137. hcyclotelus elegum (ME1 084354): 136. lateral view, 137. dorsal view. 138, 139. Coleianu ntgncopis (ME1 028449): 138. lateral view, 139. dorsal view. 140, 141. Cyclotelu piclipennk (ME1 103276): 140. lateral view, 141. dorsal view. Scale bars = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 235 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 142-147. 6 aedeagal complex and 9 furcal complex. 142-1 45. Aedeagal complex. 142-143. Cyclotelus becken (ME1 028372): 142. lateral view, 143. dorsal view, arrow points to ejaculatory apodeme. 144-145. Cyclotelus kroeben (ME1 076063): 144. lateral view, 145. dorsal view. 146-147. Furcal complex (accessory glands removed). 146. Ammotherem lehn (ME1 105327), furca (dorsal view) and basally truncated common spermathecal duct, 147. Nesonana young. (ME1 030436), furca (dorsal view) through central sac including basally truncated spermathecal ducts, arrow points to central sac. Scale bars= 0.2 mm. 236 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figure 148. ? furcal complex: Brevipma phcida (ME1 028316), furca (dorsal view) through central sac including truncated spermathecal ducts and truncated accessory glands, arrow points to furcal bulla. Scale bar = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 237 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 149-15 1. ? furcal complex (accessory glands removed). 149. Anolingu psiln~ns(ME1 077735), furca (dorsal view) through central sac including basally truncated spermathecal ducts. 150. BupluvgFu rebeccue (ME1 106349), furca (dorsal view) through central sac and spermatheca (left spermathecal duct is truncated basally). 15 1. Crebraietu craisicomk (ME1 100439), furca (dorsal view) and basally truncated common spermathecal duct, arrow points to medioventral, posterolateral process. Scale bars = 0.2 mm. 238 S. D. GAIMARI AND M. E. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 152, 153. 0 furcal complex. 152. O~odicmmyiuugenttjiiu (ME1 052230), furca (dorsal view) through central sac including truncated basal spermathecal ducts (accessory glands removed), arrow points to common spermathecal duct. 153. 0t.odicmmyia notutu (ME1 082597), furca (dorsal view) through central sac and spermatheca (right spermathecal duct and accessory glands truncated basally), arrow points to spermathecal duct. Scale bar = 0.2 mm. PHYLOGENY AND BIOGEOGRAPHY OF CYCLOTELINE THEREVINAE 239 Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figures 154, 155. 0 furcal complex (accessory glands removed). 154. Aocycloklus ehgunr (ME1 084353), furca (dorsal view) and basally truncated central sac duct. 155. Coleianu nigncopk (ME1 088220), furca (dorsal view) through basally truncated central sac duct and spermathecal ducts. Scale bar =0.2 mm. 240 S. D. GAIMARI AND M, F. IRWIN Downloaded from https://academic.oup.com/zoolinnean/article-abstract/129/2/129/2682484 by guest on 24 June 2020 Figure 156. 9 furcal complex: Qclotelus becken' (ME1 028375), furca (dorsal view) through central sac and spermatheca (left spermathecal duct and accessory glands truncated basally). Measure bar = 0.2 mm.