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FIGURE 1 in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 1. Lixini (A, B), Rhinocyllini (C) and Cleonini (D–F), live adult beetles. A: Lachnaeus crinitus (Russia, Rostov-on- Don); B: Lixus (Phillixus) subtilis (Russia, Rostov-on-Don); C: Rhinocyllus conicus (Russia, Adygea); D: Adosomus (Adosomus) roridus (Russia, Volgograd); E: Ammocleonus aschabadensis (Iran, Hormozgan); F: Apleurus saginatus (USA, Arizona).
FIGURE 7 in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 7. World distribution of Cleonini. The presence of the tribe in Indochina is poorly documented and shown as striped. Northern limits in North America are from Bouchard et al. (2013). Text boxes list genus-group taxa in five main "peripheral" regions; those endemic have the number of species indicated. Asterisk (*) denotes non-native taxa introduced by humans. Point on the map denotes the Baskunchak-Elton locality on the Kazakhstan-Russia border with the highest documented diversity of Cleonini containing at least the following 52 species: Adosomus (Adosomus) roridus, Asproparthenis carinata, A. carinicollis, A. foveocollis, A. obsoletefasciata, A. punctiventris, A. vexata, Bothynoderes affinis, B. declivis, Chromonotus (Chevrolatius) bipunctatus, Ch. (Chevrolatius) hirsutulus, Ch. (Chevrolatius) pilosellus, Ch. (Chromonotus) pictus, Ch. (Chromonotus) vittatus, Cleonis pigra, Coniocleonus (Augustecleonus) hollbergi, C. (Plagiographus) crinipes, C. (Plagiographus) nigrosuturatus, Conorhynchus excavatus, C. lacerta, C. nigrivittis, C. pulverulentus, Cyphocleonus achates, C. adumbratus, C. cenchrus, C. dealbatus, C. trisulcatus, Epexochus lehmani, Eumecops kittaryi, Eurysternus limis, Leucomigus candidatus, Leucophyes pedestris, Maximus strabus, M. verrucosus, Mecaspis alternans, Pachycerus segnis, Pleurocleonus quadrivittatus, P. sollicitus, Pseudocleonus cinereus, P. grammicus, P. marginicollis, Rhabdorrhynchus echii, Scaphomorphus vibex, Stephanocleonus (SanZia) microgrammus, S. (SanZia) tetragrammus, Temnorhinus (Massimocleonus) elongatus, T. (Massimocleonus) nasutus, T. (Temnorhinus) hololeucus, T. (Temnorhinus) kirghisicus, T. (Temnorhinus) verecundus, Terminasiania granosa, Xanthochelus nomas.
FIGURE 10 in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 10. Unambiguously optimized synapomorphies of Lixinae, Cleonini and 24 genus-group Cleonini taxa from a randomly selected (=first) among the overflow of 100 most parsimonious trees. Solid circles indicate unique synapomorphies, open circles indicate reversals or homoplasies. Numbers above and below circles are character numbers and states, respectively. Note that the genera Pseudocleonus, Stephanocleonus and the subgenus Eremocleonus recovered as monophyletic on the MP consensus tree (Fig. 9) were not recovered as such on the selected most parsimonious tree.
FIGURE 4. Cleonini, live adult beetles. A in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 4. Cleonini, live adult beetles. A: Cyphocleonus trisulcatus (Russia, Volgograd reg.); B: Entymetopus (Entymetopus) lineolatus (Kazakhstan, Kendirli); C: Eurycleonus talamellii (Morocco, Tan-Tan prov.); D: Glebius confluens (Kazakhstan, Karatau); E: Leucochromus imperialis (Kazakhstan, Akkuduk); F: Leucomigus candidatus (Kazakhstan, Balkhash); G: Maximus granulatus (Kazakhstan, Ustyurt); H: Maximus strabus (Russia, Volgograd reg.); I: Mecaspis alternans (Ukraine, Kharkiv).
FIGURE 3. Cleonini, live adult beetles. A in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 3. Cleonini, live adult beetles. A: Chromonotus (Chromonotus) vittatus (Russia, Astrakhan reg.); B: Chromosomus fischeri (Kasakhstan, Onere); C: Cleonis pigra (Russia, Rostov reg.); D: Coniocleonus (Augustecleonus) turbatus (Russia, Rostov reg.); E: Coniocleonus (Plagiographus) nigrosuturatus (Russia, Rostov reg.); F: Conorhynchus lacerta (Kazakhstan, Karynzharyk); G: Conorhynchus nigrivittis (Russia, Volgograd reg.); H: Cyphocleonus cenchrus (Kazakhstan, Karynzharyk); I: Cyphocleonus dealbatus (Russia, Rostov-on-Don).
FIGURE 2. Cleonini, live adult beetles. A in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 2. Cleonini, live adult beetles. A: Asproparthenis carinata (Russia, Rostov-on-Don); B: Asproparthenis carinicollis (Russia, Rostov-on-Don); C: Asproparthenis obsoletefasciata (Kazakhstan, Karynzharyk); D: Asproparthenis punctiventris (Russia, Rostov-on-Don); E: Bothynoderes affinis (Russia, Volgograd reg.); F: Bothynoderes declivis (Russia, Volgograd); G: Brachycleonus fronto (Kazakhstan, Priaral Karakum); H: Calodemas errans (Namibia, Khomas); I: Chromonotus (Chromonotus) pictus (Kazakhstan, Koksem).
FIGURE 6. Cleonini, immature stages. A-F in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 6. Cleonini, immature stages. A-F: Rhabdorrhynchus sp.: habitat in Kazakhstan (Kegen) with host plant Lappula sp. (Boraginaceae) flowering blue (A); roots of Lappula sp. with gall formed by immature beetle (B-D); immature stages (E-F). G- I: Adosomus (Adosomus) roridus: egg (G), larva (H) and pupa (I); from Trnka et al. (2015). J-L: Coniocleonus (Plagiographus) nigrosuturatus; larva (J, K) and pupa (L), from Stejskal et al. (2014).
FIGURE 5. Cleonini, live adult beetles. A in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 5. Cleonini, live adult beetles. A: Menecleonus virgatus (Kazakhstan, Mametkazgan); B: Pleurocleonus quadrivittatus (Russia, Rostov-on-Don); C: Pleurocleonus sollicitus (Russia, Rostov reg.); D: Pseudocleonus cinereus (Russia, Rostov reg.); E: Pseudocleonus marginicollis (Kyrgyzstan, Osh reg.); F: Rhabdorhynchus karelini (Kazakhstan, Kendirli); G: Scaphomorphus quadrilineatus (USA, Arizona); H: Scaphomorphus vibex (Kazakhstan, Malye Barsuki); I: Trichocleonus leucophyllus (Kazakhstan, Khantau).
FIGURE 9 in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 9. Maximum Parsimony majority rule bootstrap consensus phylogram of Cleonini. Numbers on branches are bootstrap values. Terminal names are truncated to the first 32 symbols; see Appendix 3 for the full names. Wide branches are those leading to monophyletic taxa, either Lixinae or Cleonini, or the genera or subgenera of Cleonini.
FIGURE 8 in Cleonini (Coleoptera: Curculionidae: Lixinae) are monophyletic and flightless: tribe overview, rampant adult homoplasy and illustrated global diversity
FIGURE 8. Bayesian inference majority rule consensus phylogram of Cleonini. Numbers on branches are posterior probabilities. Terminal names are truncated to the first 32 symbols; see Appendix 3 for the full names.
FIGURE 7 in A new species of Ungla (Neuroptera: Chrysopidae) that exhibits remarkable homoplasy in male secondary sexual characters
FIGURE 7. Ungla pseudomeleoma, sp. nov., Paratype, cleared abdomen (female, Peru: La Libertad, TRC). A. Terminal segments of abdomen (lateral); B. Fused ninth tergite and ectoproct (lateral); C, D. Spermathecal complex; E, F. Spermathecal complex and subgenitale. c.c., callus cerci; g.l. gonapophysis lateralis; inv, spermathecal invagination; pouch, pouch at base of subgenitale; sg, subgenitale; sm, spermatheca; sm.d., spermathecal duct; sm.d.(b), sm.d.(d), basal, distal sections of spermathecal duct; S7, seventh sternite; T7, T8, seventh, eighth tergites; T9+e, fused ninth tergite and ectoproct; vel, spermathecal vellum.
FIGURE 8 in A new species of Ungla (Neuroptera: Chrysopidae) that exhibits remarkable homoplasy in male secondary sexual characters
FIGURE 8. Similarity in the antennal features of Ungla pseudomeleoma with those of three species of Meleoma. A. Ungla pseudomeleoma, sp. nov., Holotype (Peru: La Libertad, FSCA); B. Meleoma emuncta Fitch (USA: California, TRC); C. Meleoma undescribed sp. #1 (USA: Arizona, TRC); D. Meleoma stangei Penny (Mexico: Baja California, San Diego County Museum of Natural History). f1, first flagellomere; h, horn extending from scape; ped, pedicel; sc, scape.
FIGURE 2 in A new species of Ungla (Neuroptera: Chrysopidae) that exhibits remarkable homoplasy in male secondary sexual characters
FIGURE 2. Ungla pseudomeleoma, sp. nov., Paratypes (female, Peru: La Libertad). A. Head, frontal (FSCA); B. Head, prothorax (dorsal, FSCA); C. Head, prothorax, mesothorax (dorsal, TRC); D. Antenna (FSCA). f1, first flagellomere; in.fu., interantennal furrow; in.h., interantennal horn; ped, pedicel; sc, scape; tor, torulus..
FIGURE 6 in A new species of Ungla (Neuroptera: Chrysopidae) that exhibits remarkable homoplasy in male secondary sexual characters
FIGURE 6. Ungla pseudomeleoma, sp. nov., Holotype, genitalia (male, Peru: La Libertad, FSCA). A. Gonarcal complex (anterolateral); B. Gonarcus (posterodorsal); C. Gonarcal complex (frontal); D. gonarcus (caudal); E. Gonarcal complex (frontoventral). c, comes; gsac, gonosaccus; g.ap., gonarcal apodeme; g.br., gonarcal bridge; h.i., hypandrium internum; mu, mediuncus; pr, unarticulated process on frontal margin of gonarcal apodeme; s.p., setose subanal plate; s.pl., paired setose plate of gonosaccus.
FIGURE 5 in A new species of Ungla (Neuroptera: Chrysopidae) that exhibits remarkable homoplasy in male secondary sexual characters
FIGURE 5. Ungla pseudomeleoma, sp. nov., Holotype, cleared abdomen (male, Peru: La Libertad, FSCA). A. Abdomen (lateral); B. Abdominal terminus (lateral); C. Terminal abdominal segments and genitalia (lateral); D. Fused ninth tergite and ectoproct (dorsal); E. Fused eighth and ninth sternites (ventral, suture on dorsal integument between T8 and T9 visible). c.c., callus cerci; g.ap., gonarcal apodeme; g.br., gonarcal bridge; inv, invaginated dorsal cleft in T9+e, above anus; S3, third sternite; S8+9, fused eighth and ninth sternites, s.p., setose subanal plate; sp, spiracular opening; T4, T6, T8, T9, fourth, sixth, eighth, ninth tergites; T9+e, fused ninth tergite and ectoproct.
FIGURE 4 in A new species of Ungla (Neuroptera: Chrysopidae) that exhibits remarkable homoplasy in male secondary sexual characters
FIGURE 4. Ungla pseudomeleoma, sp. nov., Paratype (female, Peru: La Libertad, TRC). Wings. A. Forewing B. Hindwing. b2, b3, second, third upper Banksian cells (cells below RP that do not include a gradate vein); b'2, b'4, second, fourth lower Banksian cells (cells between Psm and Psc; gc3, third gradate cell (cell with inner and outer gradates at the two ends); icu1, first intracubital cell; 1ig, first inner gradate vein, im1, first intramedian cell; Psc, Pseudocubitus; Psm, Pseudomedius; rarp1, rarp9, first, ninth closed cells between RA and RP; RA, anterior branch of Radius; RP, posterior branch of Radius.
FIGURE 9 in A new species of Ungla (Neuroptera: Chrysopidae) that exhibits remarkable homoplasy in male secondary sexual characters
FIGURE 9. Comparison of the male cranial features of Ungla pseudomeleoma with those of selected Meleoma species, A. Ungla pseudomeleoma, sp. nov., Holotype (Peru: La Libertad, FSCA). A1. Dorsal aspect (base) of interantennal horn (arrow indicates dorsal origin of mesal trough), A2. Frontolateral aspect of interantennal and lower horns (The arrows indicate paired distal arms of the interantennal horn; the lower horns are brown, triangular, below and anterior to the interantennal horns), A3. Frontal close-up of interantennal trough, interantennal horn, and lower horn (upper arrows indicate tips of paired interantennal horns; lower arrows indicate paired lower horns); B-D. Dorsal aspect of interantennal horn (lower arrows on each figure indicate base of the horn; upper arrows indicate tip of interantennal horn. B. Meleoma emuncta Fitch (USA: California, TRC); C. Meleoma hageni Banks (USA: Arizona, TRC); D. Meleoma unidentified sp. (Argentina: Jujuy, TRC).
FIGURE 1 in A new species of Ungla (Neuroptera: Chrysopidae) that exhibits remarkable homoplasy in male secondary sexual characters
FIGURE 1. Ungla pseudomeleoma, sp. nov., Holotype (male, Peru: La Libertad, FSCA). A. Head, prothorax (frontolateral); B. Head, prothorax (lateral); C. Head (frontal); D. Head (lateral); E. Face (frontal); F. Head, prothorax (dorsal). f1, first flagellomere; in.fu., interantennal furrow; in.h., interantennal horn; lo.h., lower horn; ped, pedicel.
Fig. 4 in Phenotypical plasticity and homoplasy complicate species delimitation in the Cladonia gracilis group (Cladoniaceae, Ascomycota)
Fig. 4 Principal component analysis (PCA) based on seven morphological and anatomical variables of C. coniocraea and C. ochrochlora
Fig. 2 in Phenotypical plasticity and homoplasy complicate species delimitation in the Cladonia gracilis group (Cladoniaceae, Ascomycota)
Fig. 2 Phylogeny of the Cladonia coniocraea and C. ochrochlora complex. 50% Majority rule Bayesian tree based on ITS rDNA, IGS, RPB2 and EF1-α. Branches supported with posterior probability ≥0.95 and bootstrap>70% are indicated in bold. Bootstrap value>70% for MP/ Bootstrap value>70% for ML/posterior probability>0.95 for Bayesian analysis at branches
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