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Zeitschrift/Journal: Systematics and Phylogeny

Jahr/Year: 2020

Band/Volume: 78

Autor(en)/Author(s): Korneyev Severyn V., Smit John T., Hulbert Daniel L., Norrbom Allen L., Gaimari Stephen, Korneyev Valery A., Smith James J.

Artikel/Article: Phylogeny of the Latreille, 1804 (Diptera: ) 111-132 78 (1): 111 – 132 2020

© Senckenberg Gesellschaft für Naturforschung, 2020.

Phylogeny of the genus Tephritis Latreille, 1804 (Diptera: Tephritidae)

Severyn V. Korneyev *, 1, John T. Smit 2, Daniel L. Hulbert 3, Allen L. Norrbom 4, Stephen D. Gaimari 5, Valery A. Korneyev 1 & James J. Smith 3

1 I.I. Schmalhausen Institute of Zoology, NAS of , Kiev, Ukraine; Severyn V. Korneyev * [[email protected]] — 2 European Invertebrate Survey – the , Naturalis Center, Leiden, the Netherlands — 3 Michigan State University, Department of Entomology, East Lansing, USA — 4 Systematic Entomology Laboratory, ARS, USDA, c/o National Museum of Natural History, Smith- sonian Institution, , DC, USA — 5 Pest Diagnostics Center, Department of Food and Agriculture, Sacramento, USA — * Corresponding author

Accepted on April 30, 2020. Published online at www.senckenberg.de/arthropod-systematics on May 26, 2020. Editors in charge: Rudolf Meier & Klaus-Dieter Klass

Abstract. The phylogenetic relationships of Tephritis Latreille, one of the largest genera of true fruit fies, are poorly understood due to the large number of , lack of reliable morphological characters and diffculties in identifcation. In the present study we used two datasets to address these problems: one with 35 species of the genus Tephritis (28 from the Palaearctic and seven from the Nearctic Re- gion) and seven species from closely related genera of the tribe (one species each of Rondani, Capitites Foote & Freidberg, Loew, Hendel, Schrank and two species of Heringina Aczél). For this dataset, we inferred the phylogenetic relationships using fve genes (2914 bp): mitochondrial COI (687 bp), mitochondrial 16S rRNA (561 bp), and the nuclear genes Period (481 bp), AATS (444 bp), and the 28S rRNA (741 bp). The second dataset consisted of 64 species of the genus Tephritis, two species of Heringina and one Capitites as an ougtroup and phylogenetic relationships were only inferred using COI (687 bp). The results of both analyses advance the understanding of species relationships, composition of species groups, and existing concepts of homology, re- lationships and divergence of the genus. Our results based on molecular phylogenetic relationships show that morphological characters are occasionally uninformative for delimiting monophyletic species groups; however, 10 species groups are moderately well supported based on their association with certain host plant taxa (genera, subtribes or tribes). Furthermore, we discuss monophyly of the genus Tephritis, and its relationships with the closely related genera Heringina and Capitites.

Key words. Diptera, Tephritidae, Tephritis, , species groups, host , phylogeny, distribution.

1. Introduction

The genus Tephritis Latreille 1804 is one of the largest Anthemideae, , Cardueae, , , genera of Tephritidae, with over 160 described species, Senecioneae and Calenduleae, occasionally inducing mainly occurring in the Holarctic and, to a lesser degree, non-lignifed galls in fower heads, shoots and rosettes. Afrotropical, Oriental, and Australasian Regions (Norr- Most Tephritis species are monophagous or stenophagous bom et al. 1999), but entirely lacking in the Neotropical (feed on several species of the same genus: S. Korneyev Region. Recently most of Australian species of Tephri­ 2016). Tephritis are potential control agents for invasive tis and were reassigned to the genus Aus­ species of asteraceous plants, like species from Cecido­ trotephritis (Hancock & Drew 2003). Tephritis species chares Bezzi, Hendel, Munro, inhabit most climatic zones and altitudes from dry and Robineau-Desvoidy, Robineau-Des- hot semidesert to circumpolar and alpine tundra in voidy (White & Elson-Harris 1992; Foote et al. 1993; mountainous regions. Larvae of Tephritis species feed in Edwards et al. 2009; Day et al. 2013; Day et al. 2016; fower heads of a wide range of Asteraceae of the tribes Randall et al. 2017; Aigbedion-Atalor et al. 2018). For

ISSN 1863-7221 (print) | eISSN 1864-8312 (online) | DOI: 10.26049/ASP78-1-2020-05 111 Korneyev et al.: Phylogeny of Tephritis

example, T. neesii (Meigen 1830) is considered as a po- Mohamadzade & S. Korneyev 2012; S. Korneyev & tential biocontrol agent of Oxeye daisy, Leucanthemum Mohamadzade 2013; Mohamadzade et al. 2015; S. Kor­ vulgare Lam. (Asteraceae), but little is known about neyev et al. 2015a,b; Mohamadzade & S. Korneyev host-specifcity (McClay et al. 2013). There are several 2017; S. Korneyev & V. Korneyev 2019). These studies examples of unsuccessful introduction: T. dilacerata, a also included new host plant records, a review of the Palearctic species, has been released in existing information, and a DNA barcoding analysis as the biocontrol agent for arvensis, but popula- based on mitochondrial COI sequences, which yielded tion has not survived (Foote et al. 1993); T. postica was preliminary hypotheses about phylogenetic relationships released in in 1995 and 1997 but also has not within the genus. The present study is based on analyses established (Turner 1996). For many Tephritis species, which include additional genetic markers as well as new host plant relationships remain unknown. interpretation of their host plant associations and mor- Identifcation of Tephritis species is based on the keys phological characters in order to test the monophyly of by Hendel (1927) and Hering (1944) or more recent keys the genus Tephritis, as well as delimitation of species for particular countries such as the ones by White (1988) groups proposed. (), Freidberg & Kugler (1989) (), The most reliable data on Tephritis host plants were Foote et al. (1993) (US & Canada), Merz (1994) (Swit- provided by Merz (1994) based on his reared material. zerland), V. Korneyev & Ovchinnikova (2004) (Far East Other data were previously provided and summarized by of Russia), and Smit (2010) (The Netherlands). The ge- Frauenfeld (1857), Freidberg & Kugler (1989), and V. nus is in need of thorough taxonomic revision, for there Korneyev & Kameneva (1993) and many published re- are still numerous undescribed species, and a number of cords are compiled on the CoFFHI website (https://coff- names are likely synonyms. There is also a lack morpho- hi.cphst.org/), but these may represent unreliable records logical information for inferring phylogenetic relation- due to sweeping or rearing from samples contaminated ships in Tephritis, because the species have very little by other plants, or misidentifcation of host plants. In any variation in the male genitalia while the same structures case, use of these data requires thorough reviews of these provide characters of phylogenetic signifcance in many papers and subsequent critical analysis. other tephritid genera (V. Korneyev 1985, 1990; V. Kor- In the Palearctic Region. Tephritis species are asso- neyev & Ovchinnikova 2004). ciated with tribes of Asteraceae: Anthemideae, Astereae, Only a few morphologically delimited (but not nec- Cardueae, Cichorieae, Gnaphalieae, Inuleae, Senecio- essarily monophyletic) species groups have been reco­ neae, and Calenduleae. In some cases, both associations gnized in the past: the group (Jenkins & Turn- with certain host plant tribe or genus correlate with unique er 1989), the leontodontis and hyoscyami groups (Merz morphological characters (e.g., aculeus tip shape), which 1995), the maccus group (V. Korneyev & Dirlbek 2000), possibly support the monophyly of some species groups and the pulchra group (Merz 1993; Freidberg & Kütük (for example, the pulchra group: see Merz 1993; Freid­ 2002). Furthermore, a number of poorly morphologically berg & Kütük 2002; and the formosa group: see S. Kor­ differentiated “cryptic species” or “host races” are recog- neyev & V. Korneyev 2019). A major goal of the present nized in collections based on reared material from differ- study was the reconstruction of phylogenetic relation- ent plants, thus complicating the systematic understand- ships within the genus Tephritis based both on molecular ing of the genus (S. Korneyev, personal observation). and morphological data. In addition to the COI barcode Association with certain plant taxa may be used as an region, we sequenced four additional genetic mark- informative character, but host plant associations remain ers (mitochondrial 16S and nuclear Period, AATS, and unknown for many species. Therefore, a reconstruc- 28S). tion of phylogenetic relationships with molecular char- The genus Tephritis belongs to the tribe Tephritini of acters is particularly attractive in Tephritis. Previously, the subfamily . The subfamily Tephritinae is a sequences of Tephritis species were scarcely available, diverse taxon containing more than 200 genera and 1,950 and collection material was too old or stored under inap- species from all zoogeographical regions (Norrbom et al. propriate conditions for successful DNA extraction and 1999). Its monophyly is supported both by morphology sequencing. (V. Korneyev 1999) and molecular data (Han et al. 2006). A frst step towards understanding the relationships The tribe Tephritini contains almost 1000 described spe- was Korneyev’s (2016) phylogenetic analyis of the genus cies of more than 80 genera including Tephritis, the larg- Tephritis based on morphological characters (52 species, est of them. Many genera are similar to Tephritis in wing 30 characters), host plant preferences (host plant tribe/ patterns and habitus. A phylogenetic analysis based on genera used as character state for 52 species, 4 charac- sequences of the 16S mitochondrial gene for 38 genera ters), and mitochondrial COI sequences (41 species, 271 of different Tephritinae tribes (Han et al. 2006) does not specimens). The analysis was used to delimit preliminary support the current tribal classifcation of the Tephritinae species groups for Tephritis. Additional morphological based on morphology; e.g., Euaresta and , studies of almost 9,000 Tephritis specimens representing which are morphologically similar to Tephritis, are not more than 90 species from the Western Palaearctic have supported as closely related and are placed in a distant served as a base to describe several new species and re- related group of Nearctic genera. On the other hand, Han describe many little-known species (S. Kor­neyev 2013; et al. (2006) proposed that Tephritomyia and Trupanea

112 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (1) 2020

are closely related to Tephritis. Additionally, relationships between several different Tephritis species were hypothesized for the frst time as a large number of European fruit fies were barcoded (Smit et al. 2013). For example, some of the morphologically defned species (°C) (min:sec) groups such as pulchra and leontodontis were supported, and Herin­ ext. period Final gina Aczél appeared as an in-group within Tephritis, showing a pos- sibility that Heringina might be an aberrant member of Tephritis or its sister-group (Smit et al. 2013). Both studies provided a necessity to use Euaresta, Actinoptera Rondani, Goniurellia Hendel, Trupanea, (°C) (min:sec)

Capitites Foote & Freidberg and Heringina as outgroup taxa for the Extension time present study.

2. Material and methods (°C) (min:sec) Annealing temp.

Taxon sampling. Specimens were collected from May to October 2004 – 2017 in eastern and Middle (Severyn Korneyev and Valery Korneyev; deposited in I.I. Schmalhausen Institute of Zo-

ology NAS of Ukraine, SIZK), in western and and (°C) (min:sec)

Israel (J. Smit; deposited in Naturalis Biodiversity Center, Leiden, short Denaturation The Netherlands, RMNH, except for one specimen collected by Al- len Norrbom, now in SIZK collection), and in North America (Allen No. of Norrbom and colleagues, material deposited in SIZK, with additional cycles voucher specimens at National Museum of Natural History, Washing- ton, DC, USA; and Dr. Martin Hauser and Dr. Steve Gaimari, material deposited in California Department of Food and Agriculture, Sacra- 95°C, 5:00 34 95°C, 0:30 50°C, 1:00 72°C, 1:00 72°C, 5:00 95°C, 5:00 35 95°C, 1:00 62.7°C, 1:00 72°C, 1:30 72°C, 5:00 mento, California, USA, CSCA) (Table 3). 95°C, 10:00 45 95°C, 0:30 50°C, 1:00 72°C. 1:00 72°C, 4:00 (°C) (min:sec)

All material was stored in 96% ethanol at –18 – 20°C. DNA was long Denaturation extracted from more than 170 specimens. In the present study, for the COI analyses 67 specimens of different Tephritis species were used, while for the fve-gene analysis used 35 specimens of the Tephritis species (28 from the Palaearctic Region and 7 from the Nearctic Re- Reference gion) and 7 species from the other genera of tribe Tephritini (one spe- Hebert et al. 2004 94°C, 5:00 34 94°C, 1:00 46°C, 1:30 72°C, 1:00 72°C, 5:00 Han et al. 2006 Barr et al. 2005 95°C, 2:00 35 95°C, 0:30 57°C, 1:00 72°C, 1:00 72°C, 4:00 Morgulis 2017 with modifcations cies each of Actinoptera, Capitites, Euaresta, Goniurellia and Trupa­ Han et al. 2002 with modifcations nea and two species of Heringina) (Table 3). After testing primers and excluding samples that failed to extract or resulted in poor quality or limited DNA, we selected Actinoptera, Capitites, Goniurellia, Her­ ingina, Trupanea for inclusion in the present study because of their different degrees of relatedness to Tephritis, and fnally, Euaresta ae­ qualis (Loew, 1862) was selected as the outgroup. The selection of the outgroup was based on the phylogenetic analysis of Tephritinae by Han et al. (2006) as the distantly related representatives from the tribe

Tephritini. Primer sequence

Abbreviations of depositories. CSCA – California State Collection of , Plant Pest Diagnostics Center, California Department of Food and Agriculture, Sacramento, California, USA; RMNH – Naturalis Biodiversity Centre, Leiden, the Netherlands; SIZK – I.I. 5’-ATTCAACCAATCATAAAGATAT-3’ 5’-TAAACTTCTGGATGTCCAAAAA-3’ 5’-CGCCTGTTTATCAAAAACAT-3’ 5’-CTCCGGTTTGAACTCAGATC-3’ 5’-ATTCWTGGGARGGRGATGCC-3’ 5’-AABGACATGGGTTGGTACATC-3’ 5’-TGTAAAACGACGGCCAGTTAYCAYCAYACNTTYTTYGARATG-3’ 5’-CAGGAAACAGCTATGACCATNCCRCARTCNATRTGYTT-3’ 5’-GTGCAAATCGATTGTCAGAA-3’ 5’-AGCAGGACGGTGGACATGGA-3’ 5’-ACTTAAGCGCCATCCATTT-3’ 5’-GAAGTGGAGAAGGGTTTCGT-3’ 5’-TGGAACCGTATTCCCTTTCG-3’ 5’-CTTACCTACATTATTCTATCGACT-3’ Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Kyiv, Ukraine.

Microscopy. This work is partly based on morphological studies giv- LepF1 LepR1 LR-N-13398 LR-J-12883 Per2612Fdeg Per2612Fdeg Per3105R M13A1x92F M13rA1x244R en in previous papers (S. Korneyev 2013; Mohamadzade et al. 2015; S28C (forward) S28E (forward) A28D (reverse) S28G (forward) A28F (reverse) A28HL (reverse) S. Korneyev et al. 2015a,b; S. Korneyev 2016; S. Korneyev & V. Kor­neyev 2019). Studied morphological characters show differences of wing patterns and aculeus among different species and are presented

on the tree fgures (Figs. 2, 3). Genitalia were prepared for study us- Locus (size) Primer pair COI (687 bp) 16S (561 bp) Period (481 bp) Period AATS (444 bp) AATS 28S (1280 bp) ing the following procedure: the abdomen was excised from a relaxed 1. Primer sequences. Table

113 Korneyev et al.: Phylogeny of Tephritis

Table 2. List of specimens and GenBank accession numbers.

Taxon COI 16S Period AATS 28S Tephritis cf. dioscurea MN312021 MN331953 MN311979 MN394180 MN331891 Tephritis multiguttata MN312022 MN331954 MN311980 MN394181 MN331892 Tephritis sp. nr. tanaceti MN312023 MN331955 MN311981 MN394182 MN331893 Tephritis araneosa MN312024 MN331956 MN311982 MN394183 MN331894 Tephritis cf. ovatipennis MN312025 MN331957 MN311983 MN394184 MN331895 Tephritis spn. 1 MN312026 MN331958 MN311984 MN394185 MN331896 Tephritis spn. 2 MN312027 MN331959 MN311985 MN394186 MN331897 Tephritis spn. 4 MN312028 MN331960 MN311986 MN394187 MN331898 Tephritis spn. 3 MN312029 MN331961 MN311987 MN394188 MN331899 Tephritis cf. tanaceti MN312030 MN331962 MN311988 MN394189 MN331900 Tephritis cf. nigricauda MN312031 MN331963 MN311989 MN394190 MN331901 Tephritis stigmatica MN312032 MN331964 MN311990 MN394191 MN331902 Tephritis labecula MN312033 MN331965 MN311991 MN394192 MN331903 Tephritis baccharis MN312034 MN331966 MN311992 MN394193 MN331904 Tephritis arizonaensis MN312035 MN331967 MN311993 MN394194 MN331905 Tephritis sahandi MN312036 MN331968 MN311994 MN394195 MN331906 Tephritis hurvitzi MN312037 MN331969 MN311995 MN394196 MN331907 Tephritis cf. conyzifoliae MN312038 MN331970 MN311996 MN394197 MN331908 Tephritis cf. separata MN312039 MN331971 MN311997 MN394198 MN331909 Tephritis kovalevi MN312040 MN331972 MN311998 MN394199 MN331910 Tephritis youngiana MN312041 MN331973 MN311999 MN394200 MN331911 MN312042 MN331974 MN312000 MN394201 MN331912 Tephritis cometa MN312043 MN331975 MN312001 MN394202 MN331913 MN312044 MN331976 MN312002 MN394203 MN331914 Tephritis arsenii MN312045 MN331977 MN312003 MN394204 MN331915 Tephritis kogardtauica MN312046 MN331978 MN312004 MN394205 MN331916 Tephritis hyoscyami MN312047 MN331979 MN312005 MN394206 MN331917 Tephritis cf. maccus MN312048 MN331980 MN312006 MN394207 MN331918 MN312049 MN331981 MN312007 MN394208 MN331919 Tephritis tatarica MN312050 MN331982 MN312008 MN394209 MN331920 Tephritis nozarii MN312051 MN331983 MN312009 MN394210 MN331921 Tephritis oedipus MN312052 MN331984 MN312010 MN394211 MN331922 Tephritis praecox MN312053 MN331985 MN312011 MN394212 MN331923 Heringina arezoana MN312054 MN331986 MN312012 MN394213 MN331924 Heringina guttata MN312055 MN331987 MN312013 MN394214 MN331925 MN312056 MN331988 MN312014 MN394215 MN331926 Tephritis cf. leavittensis MN312057 MN331989 MN312015 MN394216 MN331927 Trupanea cf. jonesi MN312058 MN331990 MN312016 MN394217 MN331928 Actinoptera discoidea MN312059 MN331991 MN312017 MN394218 MN331929 Goniurellia tridens MN312060 MN331992 MN312018 MN394219 MN331930 Capitites ramulosa MN312061 MN331993 MN312019 MN394220 MN331931 Euaresta aequalis MN312062 MN331994 MN312020 MN394221 MN331932 Tephritis dudichi MN312063 Tephritis angustipennis MN312064 MN312065 Tephritis valida MN312066 Tephritis sauterina MN312067 Tephritis subpura MN312068 Tephritis alamutensis MN312069 Tephritis robusta MN312070 Tephritis crinita MN312071 MN312072 MN312073 MN312074 MN312075 Tephritis crepidis MN312076 Tephritis mutabilis MN312077 Tephritis cf. cameo MN312078 Tephritis divisa MN312079 Tephritis pallescens MN312080

114 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (1) 2020

Table 2 continued.

Taxon COI 16S Period AATS 28S Tephritis pulchra MN312081 Tephritis recurrens MN312082 Tephritis scorzonerae MN312083 Tephritis ruralis MN312084 Tephritis cf. gladius MN312085 Tephritis cf. admissa MN312086 Tephritis baccharis MN312087 Tephritis palmeri MN312088 Tephritis subpura MN312089 Tephritis signatipennis MN312090 Tephritis heliophila MN312091 specimen, cleared in NaOH solution (10%) for 2 hours at PCR Purifcation Kit protocol (QIAGEN). Sanger se- 90 – 95°C, and then washed in distilled water. Genitalia quencing was performed at the Michigan State Uni- were examined in a drop of glycerin on a depression mi- versity Research Technology Support Facility via Big- croscope slide, covered with a glass cover slip. Dissected Dye Terminator Sequencing on an Applied Biosystems structures are stored in polypropylene microvials contain- 3730xl DNA Analyzer (Foster City, CA, USA) using the ing glycerin pinned together with a specimen. PCR primers as sequencing primers. Sequences are de- Photographs of genitalia were taken using a Nikon posited in GenBank (Table 2). Coolpix P50 through the ocular of a Wild M11 light mi- croscope; photographs of wing and habitus were taken us- Phylogenetic analyses. The alignments of the protein- ing a Canon PowerShot A640 connected to a Zeiss Stemi coding DNA were performed using MEGA 7 (Kumar et C-2000 microscope. Digitized photographs were stacked al. 2016) using MUSCLE to align translated amino acids using CombineZM® (Hadley 2007). Specimens depos- (Edgar 2004). Non-coding rRNA genes (16S and 28S) ited in the collection of the Museum für Naturkunde Ber- were aligned using MAFFT (Katoh et al. 2002). To fnd lin were taken under supervision of Dr. Joachim Ziegler optimal partitioning schemes for the Bayesian and maxi- and Bernhard Schurian, and were stacked using Helicon mum likelihood analyses, we used PartitionFinder 2.1.1 Focus®. Pictures of all Nearctic specimens were taken (Lanfear et al. 2016) with the “greedy” search algorithm at the National Museum of Natural History with the as- (Lanfear et al. 2012). Before running PartitionFinder sistance of Dr. Allen Norrbom and Lucrecia Rodriguez the data were divided into 11 biologically relevant parti- using a Systematic Entomology Laboratory Visionary tions, which included frst, second and third nucleotide Digital camera system. positions for each protein-coding gene (9 partitions) and two partitions for non-coding genes 16S and 28S. The DNA isolation and PCR amplifcation. All DNA was following partitions were defned (with models): GTR + extracted using the DNeasy Blood & Tissue Kit protocol I + G for COI, Period (frst positions) and 16S; HKY + (Qiagen, Valencia, CA). For most specimens, a single leg I + G for 28S; HKY + I for COI (second positions); or head was removed, ground in a microcentrifuge tube GTR + G for COI and Period (third positions); HKY + with proteinase K and lysing buffer and then incubated G for Period (second positions) and AATS (third posi- for 12 hours. A non-destructive method was used for rare tions); K2P + G for AATS (frst positions); F81 for AATS specimens: a whole abdomen was placed in the buffer (second positions). Phylogenetic analyses using Bayes- and incubated for 12 hours. We PCR amplifed each lo- ian inference were performed on the concatenated ge- cus (COI, 16S, period, AATS, and 28S) using GotaqFlexi netic dataset in MrBayes 3.2.6 (Ronquist et al. 2012). DNA polymerase (Promega, Madison, WI) in 25 μl re- The Bayesian analysis used four independent runs each actions with the following concentrations of reagents: with four Metropolis-coupled chains with default heating reaction buffer (5 mM), MgCl2 (8 mM), dNTP (1.25 parameters (one cold and three heated). The chains were mM each), forward and reverse primers (1.25 mM each), sampled once every thousand generations for 10 million DNA polymerase (0.25 mM), water (4.5 mM) and DNA generations and the frst 25% of samples were discarded template (3.5 mM). Primer sequences and PCR condi- as burn-in. All analyses converged to an average standard tions for every gene were taken from literature sources deviation of split frequencies below 0.01 and all branch and are listed in Table 1. lengths and substitution model parameters had poten- tial scale reduction factors less than 1.01 (Ronquist et DNA sequencing and editing. DNA amplifcations were al. 2012). The Maximum Likelihood analysis was done performed in a 25-μL reaction volume using GoTaq® in RAxML (Stamatakis et al. 2008) using the GTR+I+G Flexi DNA Polymerase. PCR products (5 μL) were visu- model with the above partitioning scheme with 1000 alized on a 1% agarose gel using ethidium bromide. Am- bootstrap pseudoreplicates on the CIPRES online portal plifed products (20 μL) were purifed using QIAquick (Miller et al. 2010).

115 Korneyev et al.: Phylogeny of Tephritis Genes ­­ - tory SIZK 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, CSCA 28S AATS, COI, 16S, Period, Deposi 110.962

110.29903 111.71139 111.63137 109.21239 109.36787

– – – – – – 40.612 33.1628 42.3805 78.4870 SIZK 28S AATS, COI, 16S, Period, 31.42421 35.32944 35.13545 31.95976 35.531111 47.206389 SIZK 28S AATS, COI, 16S, Period, 38.852222 46.998333 SIZK 28S AATS, COI, 16S, Period, 42.644563 74.51608042.503818 SIZK 78.403276 28S AATS, COI, 16S, Period, 42.644563 SIZK 74.516080 28S AATS, COI, 16S, Period, 50.511901 SIZK 30.261249 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, 42.644563 74.51608042.405358 SIZK 78.458351 28S AATS, COI, 16S, Period, 48.602686 SIZK 25.881431 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, 30.874722 51.523889 SIZK 28S AATS, COI, 16S, Period, S. Korneyev & S. Korneyev Korneyev V. S. Korneyev & S. Korneyev Korneyev V. S. Lampert, M. Savaris, A. L. Norrbom & N.E. Woodley & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. S. Lampert, & M. Savaris A.L. Norrbom S. Lampert, & M. Savaris A.L. Norrbom S. Korneyev & S. Korneyev Korneyev V. S. Lampert, & M. Savaris A.L. Norrbom S. Lampert, & M. Savaris A.L. Norrbom S. Korneyev & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. S.M. Blank & K. Kramp & S. Korneyev Korneyev V. 19 July 2017 20 July 2017 12 July 2017 12 July 2017 20 July 2017 15 July 2017 17 May 2014 17 May 24 May 2017 24 May 2017 26 May 13 June 2014 18 June 2017 20 June 2011 Smit J.T. 42.571917 2.110639 RMNH 28S AATS, COI, 16S, Period, Country: LocationCountry: Collection date Collector Lat. Long. : Leuk, along roadSwitzerland: 22 July 2004 Province, East Azerbaijan : Allen Norrbom nr Kaleybar Iran: Kurdistan, 5 km W Hossein Kurdistan, Iran: Abad USA: Cochise Co., Carr Canyon, USA: Cochise Co., Carr Canyon, AR Karakol Kyrgyzstan: ski resort Kashka-Suu Kyrgyzstan: 2017 3 August Irpin Ukraine: Orientals, Mont Pyrenees : Louis, La Llagonne USA: Coconino Co., Snowbowl, USA: Coconino Co., Snowbowl, AR parking area, USA: Summit and Wasatch Co; USA: Summit and Wasatch Uinta Mts, Kamas ESE 25 km; Hwy 150, UT ski resort Kashka-Suu Kyrgyzstan: 8 July 2017 Mary USA: Coconino Co., Lake Road (Rt. 3), AR Road, USA: Cochise Co., Paradise AR along road, wash Co., Rt. 191, ARUSA: Greenlee and Boyer-Ahmad Kohgiluyeh Iran: 17 July 2017 near Sisakht Province, Kyrgyzstan: Karakol gorge Karakol Kyrgyzstan: 27 July 2017 Ukraine: Ternopil region, near region, Ternopil Ukraine: Synkiv Kyrgyzstan: Kashka-Suu ski resort Kashka-Suu Kyrgyzstan: 17 July 2017 Host / reared or swept Host / reared (Anthemideae) Achillea millefolium / swept sp. (Anthemideae) / Tanacetum swept unknown, possibly Artemisia sp. unknown, (Anthemideae) / swept sp. (Anthemideae) / swept sp. (Anthemideae) / Pyrethrum swept (Anthemideae) vulgare Tanacetum / swept (Anthemideae) Anthemis arvensis / swept Corethrogyne possibly Corethrogyne unknown, / swept sp. (Astereae) sp., Erigeron Artemisia sp. (Anthemideae) / swept Chrysothamnus nauseosus / swept (Astereae) (Astereae) Baccharis sarothroides / swept / (Astereae) Baccharis salicifolia swept Achillea clypeolata (Anthemideae) / swept (Cichorieae) longirostre Tragopogon / swept Organism cf. dioscurea cf. Tephritis multiguttataTephritis Achillea sp. (Anthemideae) / swept tanaceti sp. nr. Tephritis Tephritis araneosa Tephritis spn. 4 Tephritis spn. 3 Tephritis tanaceti cf. Tephritis nigricauda cf. Tephritis stigmaticaTephritis Senecio spp. (Senecioneae) / swept cf. ovatipennis cf. Tephritis spn. 1Tephritis spn. 2 Tephritis / swept unknown labecula Tephritis baccharis Tephritis arizonaensis Tephritis sahandi Tephritis cf. conyzifoliae cf. Tephritis sibirica (Cichorieae) / reared gorge Karakol Kyrgyzstan: separata cf. Tephritis Picris rigida (Cichorieae) / swept 21 July 2017 Tephritis hurvitzi Tephritis Isolate SW040722 K14118 K14025 AR170719_01m1 K170720_02f1 K170803_06m1 U170618_04 RMNH.INS.545642 AR170712_04m1 US170524_04 K170727_02m1 K170708_14f1 AR170712_01m1 AR170720_01f1 AR170717_01f1 K14058 K170721_02m3 K16007 K170717_02m1 Sequence ID T_cf_dioscurea_ T_cf_dioscurea_ SW040722_CH T_multiguttata_ K14118_IR T_sp_nr_tanaceti_ K14025_IR T_araneosa_ AR170719_01m1_US T_spn4_K170720_ 02f1_KG T_spn3_K170803_ 06m1_KG T_cf_tanaceti_ U170618_04_UA T_cf_nigricauda_ 545642_FR T_stigmatica_ AR170712_04m1_US T_cf_ovatipennis_ US170524_04_US T_spn1_K170727_ 02m1_KG T_spn2_K170708_ 14f1_KG T_labecula_ AR170712_01m1_US T_baccharis_ AR170720_01f1_US T_arizonaensis_ AR170717_01f1_US T_sahandi_m1_ K14058_IR T_cf_conyzifoliae_ K170721_02m3_KG T_cf_divisa_ K16007_UA T_hurvitzi_ K170717_02m1_KG Table 3. Material. Table

116 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (1) 2020 Genes ­­ - tory SIZK 28S AATS, COI, 16S, Period, CSCA 28S AATS, COI, 16S, Period, CSCA 28S AATS, COI, 16S, Period, RMNH 28S AATS, COI, 16S, Period, Deposi 117.127 115.351

– – 39.341 38.818 42.3448 78.2326 SIZK 28S AATS, COI, 16S, Period, 42.29711 75.86995 SIZK 28S AATS, COI, 16S, Period, 42.598096 75.80945742.598096 SIZK 75.809457 28S AATS, COI, 16S, Period, 42.787072 SIZK 74.572389 28S AATS, COI, 16S, Period, 42.392630 SIZK 78.473630 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, 37.133333 44.86666742.787072 SIZK 74.572389 28S AATS, COI, 16S, Period, 42.392630 SIZK 78.473630 28S AATS, COI, 16S, Period, SIZK42.644563 28S AATS, COI, 16S, Period, 74.51608042.405358 SIZK 78.458351 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, 30.874722 51.523889 SIZK 28S AATS, COI, 16S, Period, 42.787072 74.57238942.787072 SIZK 74.572389 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, 38.852222 46.99833335.314167 SIZK 46.954722 28S AATS, COI, 16S, Period, 50.507139 SIZK 30.258750 28S AATS, COI, 16S, Period, SIZK 28S AATS, COI, 16S, Period, S. Korneyev & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. S. Korneyev & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. B.E. Altenhofer B.E. Altenhofer & S.M.Blank S. Korneyev & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. & S. Korneyev Korneyev V. S.M. Blank & K. Kramp 7 July 2017 7 July 2017 7 July 2017 7 July 2017 1 June 2017 14 July 2017 14 July 2017 23 July 2017 16 July 2017 26 May 2017 26 May 24 May 201224 May Smit J.T. 33.294667 35.759417 RMNH 28S AATS, COI, 16S, Period, 17 May 2014 17 May 20 May 2017 20 May 15 Sep. 2011 Smit J.T. 20 June 2017 15 June 2014 1 October 2016 Country: LocationCountry: Collection date Collector Lat. Long. Kyrgyzstan: Boom gorge Kyrgyzstan: Chon Aryk, near Kyrgyzstan: Bishkek Province, Azerbaijan West Iran: 10km west border, near Turkish Ziveh Chon Aryk, near Kyrgyzstan: Bishkek Afghanistan: Berganghak 24 June 2016 Skrylnik Yu. Kyrgyzstan: Jeti-Oguz Kyrgyzstan: and Boyer-Ahmad Kohgiluyeh Iran: near Sisakht Province, Kyrgyzstan: Orto-Tokoi Kyrgyzstan: Chon Aryk, near Kyrgyzstan: Bishkek Har (Mt.) Hermon Israel: Kyrgyzstan: Chon Aryk, near Kyrgyzstan: Bishkek Iran: East Azerbaijan Province, nr Province, East Azerbaijan Iran: Kaleybar USA: Lander Co; Desatoya Mts; USA: Lander Co; Desatoya SW 59 km; Hwy 722, NE Austin Iran: Kurdistan, Sanandaj, Abidar Kurdistan, Iran: Mt Irpin Ukraine: Netherlands: Ossendrecht, Stoppelbergen Pine Range; Co; White USA: Nye Hwy 6, NE Host / reared or swept Host / reared Sonchus paluster (Cichorieae) / reared acanthium (Cardueae) Onopordum / reared dolichotrichum Doronicum (Senecioneae) / reared , I. stenocalathia Inula grandis (Inuleae) / reared / swept unknown Lactuca tatarica (Cichorieae) / swept Helichrysum sp. (Gnaphalieae) / swept (Gnaphalieae) Phagnalon rupestre / swept arvensis (Calenduleae) Calendula arvensis / swept many diferent Asteraceae / swept Asteraceae diferent many Helichrysum sp. (Gnaphalieae) / swept Helichrysum arenarium (Gnaphalieae) / swept (Anthemideae) / swept possibly Arnica unknown, (Senecioneae) / swept diversifolia jonesi Organism Tephritis kovalevi Tephritis youngianaTephritis sp. (Cichorieae) / reared Youngia postica Tephritis Boom gorge Kyrgyzstan: cometaTephritis conuraTephritis / swept spp. (Cardueae) Cirsium arsenii Tephritis gorge Karakol Kyrgyzstan: / swept spp. (Cardueae) Cirsium kogardtauica Tephritis 31 July 2017 hyoscyamiTephritis maccus cf. Tephritis / swept spp. (Cardueae) Carduus bardanaeTephritis gorge Karakol Kyrgyzstan: tataricaTephritis / swept spp. (Cardueae) ski resort Kashka-Suu Kyrgyzstan: 27 July 2017 nozariiTephritis / reared (Cardueae) nivea Alfredia 17 July 2017 gorge Karakol Kyrgyzstan: / swept sp. (Cardueae) Cirsium 21 July 2017 Tephritis oedipus Tephritis discoidea Actinoptera tridensGoniurellia Capitites ramulosa (Inuleae) / swept Inula macrophylla Tephritis praecox Tephritis Trupanea cf. Trupanea Heringina arezoana Heringina guttata neesii Tephritis leavittensis cf. Tephritis Isolate K170714_05m1 K170714_02f1 K170707_31f2 K170731_03f1 K170707_21f1 K170727_01m1 K16001_f1 K170717_04f1 K170721_05f1 K170716_01m1 K170707_28 K170707_02 20120524 US170601_03 K14122 U161001 US170520_01f1 Sequence ID T_kovalevi_ T_kovalevi_ K170714_05m1_KG T_sp5_K170714_ 02f1_KG T_postica_K170707_ 31f2_KG T_cometa_K170731_03f1_ KG T_conura_K170723_01f1_KG K170723_01f1 T_arsenii_K14158__IRT_kogardtauica_ K14158 K170707_21f1_KG T_hyoscyami_ K170727_01m1_KG T_cf_maccus_K16001_f1_AF T_bardanae_K170717_04f1_ KG T_tatarica_K170721_05f1_ KG T_nozarii_K14057_IR K14057 T_oedipus_K170716_01m1_ KG Actinoptera_discoidea_ K170707_28_KG Goniurellia_tridens_ K170707_02_KG Capitites_ ramulosa_20120524_IL T_praecox_K14024_IR K14024 Trupanea_cf_jonesi_ US170601_03_US Heringina_arezoana_ K14122_IR Heringina_guttata_ U161001_UA T_neesii_545593_f1_NLT_leavittensis_ RMNH.INS.545593 US170520_01f1_US continued. 3 Table

117 Korneyev et al.: Phylogeny of Tephritis Genes ­­ - tory SIZK COI SIZK 28S AATS, COI, 16S, Period, SIZK COI SIZK COI RMNH COI RMNH COI RMNH COI RMNH COI RMNH COI RMNH COI RMNH COI RMNH COI Deposi 0.761

76.408

122.077

– 2.516914 2.061056

– – – – ­ ­ din T. Smit T. G. Ramel 38.750469 20.722714 RMNH COI D. Belgers & D. Belgers Smit J.T. G. Ramel P. van Hels van P. gen & J. 2008 2008 3 July 2014 Korneyev S. & V. 38.183 47.981 SIZK COI 16–22 June 30 Sep 2017 Allen Norrbom 39.018 20 July 2011 Korneyev S. & V. 47.933 24.116 RMNH COI 27 July 2013 Korneyev S. & V. 43.413 41.731 RMNH COI 26 May 201426 May Korneyev S. & V. 30.8747 51.5238 SIZK COI 28 May 200928 May Smit J.T. 42.872700 23 May 200923 May John Smit 42.191667 13 June 2011 24 June 2009 John Smit 43.364722 22.098056 RMNH COI 20 June 2011 Smit J.T. 42.572 2.111 RMNH COI 30 June 2012 John Smit 17 June 2009 Smit J.T. 43.299833 25 Feb – 2 Mar 25 Feb 27 August 201727 August S. Korneyev 38.514 13 August 201313 August Korneyev S. & V. 43.433 41.700 RMNH COI Country: LocationCountry: Collection date Collector Lat. Long. : Elodia Greece: The Netherlands, Rhenen, Blauwe Netherlands, Rhenen, Blauwe The Kamer : País Vasco, Alava, Vitoria- Alava, Vasco, Spain: País Gasteiz, Etxabarri-Urtupiña : Nis, Suva Planina, Serbia: Nis, Suva Jelasnicka, Klisura Afghanistan: Bande Amir Huesca, Spain: Aragón, S Embalse Sotonera Almudévar, 17 June 2016 Skrylnik Yu. Greece: Beabies Greece: Ukraine: Carpathians, HoverlaUkraine: 17 July 2011 Korneyev S. & V. USA: Putah Creek, CA USA: Putah Creek, Ukraine: Transcarpathian reg. reg. Transcarpathian Ukraine: Massif Carpathian Mts. Kuziy Orientals, Mont Pyrenees France: Louis, La Llagonne Netherlands: Colmlont, Wrakelberg Russia: North , Republic, Karachay–Cherkess southern part of town Teberda, Karachay- Caucasus, North Russia: Republic, Teberda, Cherkess Mt. Khatipara Park, USA: Maryland: Sandy Point sweeping area, Route 50 ramp Baccharis Iran: Qazvin, Alamut castleIran: Afghanistan: Bande Amir 9 June 2014 Guipúzcoa, San Vasco, Spain: País Korneyev S. & V. Sebastián, Igeldo 36.445 29 June 2016 Skrylnik Yu. 50.583 SIZK COI Host / reared or swept Host / reared Leontodon hispidus (Cichorieae) / swept Crepis biennis , C. blattarioides Crepis (Cichorieae) / swept unknown / swept unknown laciniata (Cichorieae) Scorzonera / swept S. oleraceus Sonchus asper , S. oleraceus (Cichorieae) / swept Doronicum austriacum Doronicum (Senecioneae) / reared strumarium (Heliantheae) / swept Achillea ptarmica (Anthemideae) / swept (Cichorieae) / swept (Cardueae) (Cardueae) umbrosa Cousinia sp. cf. / swept / swept unknown , Leontodon autumnalis , L. helveticus L. hispidus (Cichorieae) / swept Organism cf. cameo cf. Tephritis / swept Cousinia sp. (Cardueae) near Sisakht Pass Iran: Tephritis mutabilis Tephritis Tephritis crepidis Tephritis divisaTephritis Picris echioides (Cichorieae) / swept Tephritis recurrensTephritis sp. (Cichorieae) / swept Scorzonera Tephritis pallescens Tephritis pulchra Tephritis Tephritis formosa Tephritis Tephritis arnicae Tephritis Euaresta aequalis Euaresta dudichiTephritis speciosa (Inuleae) / reared Telekia matricariaeTephritis (Cichorieae) / swept foetida Crepis Tephritis angustipennis Tephritis vespertina Tephritis Tephritis validaTephritis Inula helenium (Inuleae) / swept Tephritis sauterinaTephritis / reared Aster alpinus (Astereae) subpuraTephritis / swept Baccharis sp. (Astereae) Tephritis alamutensis Tephritis robustaTephritis crinita Tephritis leontodontis / swept Cousinia sp. (Cardueae) Tephritis Vargehsaran Sarein, Iran: Isolate 30103300_ RMNH_545646 RMNH.INS.252199 K16005_m1_ Afgan RMNH.INS.544816 US170827_01 30103325_ RMNH_ INS_544815 RMNH.INS.555073 30103291_ RMNH_545637 30103355_ RMNH_544845 4007592937_ RMNH_556413 4007592956_ RMNH_556394 ML170719_01m1 K14108 K16003_f1 RMNH.INS.252292 Sequence ID T_cf_cameo_Iran_K14061 T_cf_cameo_Iran_K14061 K14061 T_mutabilis_0067 RMNH.INS.252029 T_crepidis_30103300_ RMNH_545646 T_divisa_0237 T_reccurens_252345_Serbia RMNH.INS.252345 T_pallescense_K16005_m1_ Afgan T_pulchra_252111_Spain RMNH.INS.252111 T_formosa_0071 T_formosa_0071 RMNH.INS.252033 T_arnicae_m1_ Ukraine_544816 Euaresta_aequalis_ US170827_01_US T_dudichi_30103325_ RMNH_INS_544815 T_matricariae_f1_ Netherlands_555073 T_angustipennis_30103291_ RMNH_545637 T_vespertina_30103355_ RMNH_544845 T_valida_4007592937_ RMNH_556413 T_sauterina_4007592956_ RMNH_556394 T_subpura_ML170719_01m1 T_alamutensis_Iran_K14108 T_alamutensis_Iran_K14108 T_robusta_Iran_K14192 T_crinita_Afgan_K16003_f1 K14192 T_leontodontis_0330Tepleo continued. 3 Table

118 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (1) 2020

Outgroup taxa. Species of the genera assigned to the “Higher Tephritinae” (sensu V. Korneyev 1999) (Euaresta, Actinoptera, Capitites, Goniurellia, Trupa­ nea), or Tephritis group of genera (sensu Merz 1999) Genes (Actinoptera, Capitites, Goniurellia, Trupanea) or the “groups 1 – 2” (sensu Han et al. 2006) (Euaresta, Go­ niurellia, Trupanea) have been included into analysis ­­ - as outgroups, along with Heringina, which was tested tory SIZK COI SIZK COI SIZK COI SIZK COI SIZK COI as a possible ingroup member of Tephritis. Deposi

Systematics of Asteraceae. The system used herein is 118.288 118.288 unk

110.75173 based on the tribal classifcation by F et al. (2009).

– – – The genus Doronicum, which is considered a genus incertae sedis basal to the tribe Senecioneae (Funk et

34.017 34.017 al. 2009) is provisionally considered to be a member 32.43562 of that tribe.

3. Results and discussion S. Korneyev, S. Korneyev, B. Brown, L. Gonsales & G.A. Kunn S. Korneyev, B. Brown, L. Gonsales & G.A. Kunn S. Lampert, & M. Savaris A.L. Norrbom

3.1. Molecular analysis 7 July 2017 Korneyev S. & V. 42.787072 74.572389 SIZK COI 31 July 2017 31 July 2017 26 July 2017 12 April 201712 April Korneyev V. 50.511901 30.261249 RMNH COI 11 April 201711 April J.D. Lopez 22 June 2009 Smit J.T. For the 44.851167 21.285306 concatenated RMNH COI dataset of the fve genes, phy- logenetic topology obtained with Bayesian inference is shown in Figure 1. The clade containing Tephritis, Capitites, Heringina, and T. praecox (the Tephritis aggregation lineage) was moderately supported in the Maximum Likelihood analyses (ML, bootstrap sup- port = 70), but very strongly supported as a mono- Country: LocationCountry: Collection date Collector Lat. Long. phyletic clade in the BI tree (BPP, Bayesian posterior probability = 1). In the ML analyses genus Capitites is Ukraine: Kyiv region, Irpin, region, Kyiv Ukraine: meadow Afghanistan: Bande Amir 29 June 2016 Skrylnik Yu. Kyrgyzstan: Chon Aryk, near Kyrgyzstan: Bishkek History USA: Los Angeles, Natural garden Museum of L. A. County, History USA: Los Angeles, Natural garden Museum of L. A. County, Houston USA: Texas: USA: Arizona: Pima Co., Mount USA: Arizona: Lemmon, Alder Campground, WP 325 Doliblato sands, Serbia: Belgrade, stop tweede placed within Tephritis, however, in the BI analyses it appears as the sister-group to the monophyletic group containing Tephritis s.str., Heringina, T. praecox, T. oe­dipus, and T. maccus, with strong support (BPP = 0.95). This corresponds well with the preliminary re- sults of the morphology-based phylogenetic analysis of S. Korneyev (2016) in which Capitites is the sister- Host / reared or swept Host / reared group to the monophyletic Tephritis s.str. + Heringina Hieracium lactuella , H. pilosella Hieracium (Cichorieae) / swept unknown / swept unknown (Cardueae) Cousinia tianschanica (Cardueae) / swept / (Astereae) Baccharis salicifolia swept Baccharis neglecta , B. halimifolia / swept (Astereae) , B. Baccharis glomerulifora / swept (Astereae) halimifolia Machaeranthera canescens Machaeranthera / swept (Astereae) dubius (Cichorieae) Tragopogon / swept + T. praecox clade. Within the Tephritis lineage, four major monophy- letic clades (I – IV in Fig. 1) are recognized. Clade I is represented by three morphologically distinct Tephri­ gladius admissa tis species plus Heringina, which in addition share no Organism obvious host-plant preferences, being associated with Tephritis ruralis Tephritis Tephritis cf. Tephritis Tephritis cf. Tephritis baccharis Tephritis palmeri Tephritis subpura Tephritis Tephritis signatipennis Tephritis heliophila Tephritis the Gnaphalieae (Heringina), Calenduleae (T. prae­ cox) or Cichorieae (T. oedipus and presumably T. mac­- cus). These species have mostly Mediterranean / distribution (except Heringina guttata, which Isolate has expanded its range on sandy dunes northwards to U170402_01_ Irpin_Ukr K16004_f1 K170707-29 US170831_07m1 AR170726_01f1 southern in the postglacial time). The species in Clade II do not seem to share any obvious morpho- logical synapomorphies and are diverse in their wing patterns; most of them infest host plants of the tribe Cardueae, except for T. kogardtauica and T. arsenii,

Sequence ID which infest members of the tribes Inuleae and Sene- cioneae. Clade III (BPP = 1) contains four morpho- T_ruralis_U170402_01_ Irpin_Ukr T_cf_gladius_K16004_f1_ Afgan2 T_cf_admissa_ K170707_29f1 T_baccharis_ US170831_07m1 T_palmeri_US170831_01f1 US170831_01f1 T_subpura_TX110426_01f1 TX110426_01f1 T_signatipennis_ AR170726_01f1 T_heliophila_0361Tephel RMNH.INS.252323 continued. 3 Table logically diverse species, but they share host plants

119 Korneyev et al.: Phylogeny of Tephritis PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA OR PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA Region Region Morphology, host Morphology, Morphology, host Morphology, Morphology, host Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology Morphology, host Morphology, Morphology Morphology Morphology, COI; Period; AATS; 28S AATS; COI; Period; Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; host Morphology, Morphology, host, COI Morphology, Morphology Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology host, COI; 16S; Period; Morphology, 28S AATS; host Morphology, Morphology Morphology, host, COI Morphology, Character Evidence Character Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host Morphology, Morphology, host Morphology, Morphology, host Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host Morphology, Morphology, host, COI Morphology, Morphology Morphology, host Morphology, Morphology, host Morphology, Morphology, host Morphology, host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host Morphology, Character Evidence Character Mohamadzade et al. 2015 Mohamadzade et al. 2015 Mohamadzade et al. 2015 Korneyev, Karimpour & Mohamadzade Korneyev, 2015 Mohamadzade et al. 2015 Merz 1994; Karimpour 2011; Korneyev 2016 Korneyev 2013 Source Korneyev 2016 Korneyev 2016 Source Tribe Tribe Inuleae Korneyev et al. 2016 Inuleae Inuleae Korneyev 2016; Merz 1994; 1992 host, COI Morphology, Cardueae & Kapoor 1988 Agarwal Cardueae 1974 Hardy Cardueae Korneyev & Evstigneev 2019 Cardueae 1993 Merz 1994; Korneyev & Kameneva Cardueae Merz 1994 Cardueae 1992 Merz 1994; Korneyev & Kameneva host, COI Morphology, Cardueae 1998 Sueyoshi Cardueae Merz 1994 Cardueae Kütük 2008 Cardueae Merz 1994 Cardueae Mohamadzade & Korneyev 2017 host, COI Morphology, Cardueae Cardueae Mohamadzade 2012 Cardueae 2013 Korneyev 2013; Shcherbakov Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Merz 1994 Cardueae Korneyev & Mohamadzade 2013 host Morphology, Cardueae Korneyev & 2019 Cardueae Korneyev & 2019 Cardueae White 1988; Merz 1994 Cichorieae Korneyev & Karyuk 2009 Cichorieae Mohamadzade et al. 2015 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Korneyev & 2019 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Hering 1937 Cichorieae Merz 1994 Cichorieae 1990 Korneyev & Kameneva Cichorieae Merz 1992b, 1994 Cichorieae Merz 1994 Cichorieae 2001 Merz 1994; Shcherbakov Cichorieae Merz 1994 Host Plant Host Plant Calenduleae Merz 1994; Smit 2006 Gnaphalieae Mohamadzade & Korneyev 2015 Gnaphalieae Mohamadzade & Korneyev 2015 Senecioneae Korneyev et al. 2015 Senecioneae Korneyev et al. 2015 Carduus onopordiodes Carduus Carduus edelbergii Carduus Cirsium obvallatum Cirsium , C. spinosissimum palustre Cirsium polyacanthum , C. incana maderensis Calendula arvensis Carduus acanthoides , C. crispus deforatus Carduus Lactuca tatarica Carduus nutans , C. thoermeri Carduus unknown Cirsium purpuratum Cirsium unknown unknown unknown swept from Lactuca scariola from swept Onopordum spp. Onopordum vulgare Cirsium Inula helenium unknown Helichrysum arenarium , C. vulgare , C. palustre arvense Cirsium sp. cf. umbrosa Cousinia sp. cf. , I. stenocalathia Inula grandis Doronicum dolichotrichum Doronicum unknown Helichrysum arenarium canum , C. libanoticum sorocephalum Cirsium unknown Doronicum austriacum , Arnica , Doronicum , D. hungaricum D. grandiforum speciosa , I. hirta Telekia Inula grandifora Host plant species Cirsium sp. Cirsium unknown , Ancathia igniaria nivea Alfredia Hypochaeris radicata Cousinia sp. Leontodon crispus , L. incanus tenuiforus Cousinia refracta Crepis albida Crepis Cousinia calathidia Picris hieracioides Cousinia sp. Leontodon hispidus Cousinia sp. Crepis foetida Crepis Hieracium lactuella , H. pilosella Hieracium Cousinia tianschanica Leontodon hispidus Youngia sp. Youngia Arctium minus Arctium , L. hispidus Leontodon autumnalis , L. helveticus unknown Jurinea baissunensis Leontodon hispidus Sonchus uliginosus Cousinia sp. Picris echioides Sonchus paluster Cousinia sp. biennis , C. blattarioides Crepis S. oleraceus Sonchus asper , S. oleraceus , A. lappa , A. minus nemorosum Arctium tomentosum , C. sibirica conyzifolia Crepis Host plant species : PA – Palearctic region; NEA – Nearctic region. – Palearctic region; NEA Abbreviations : PA Agarwal & Kapoor, 1988 & Kapoor, Agarwal atocoptera T. Hardy, 1974 Hardy, cardualis T. Korneyev & Evstigneev, 2019 & Evstigneev, Korneyev anthrax T. (Loew, 1844) conura (Loew, T. (Loew, 1844) praecox (Loew, T. T. hyoscyami (Linnaeus, 1758) T. T. oedipus Hendel, 1927 T. T. hendeliana Hering, 1944 T. Gentilini & V. Korneyev, 2006 † sophus Gentilini & V. Korneyev, T. Sueyoshi, 1998 longicauda Sueyoshi, T. Korneyev, Dirlbek, 2000 Korneyev, urelliosomima T. T. umbrosa Dirlbek & Dirlbek, 1968 umbrosa T. T. maccus Hering, 1937 T. (Loew, 1844) postica (Loew, T. 2008 Kütük, erdemlii T. (Loew, 1858) (Loew, valida T. Mohamadzade & S. Korneyev, 2015 Mohamadzade & S. Korneyev, gharalii T. 2015 H. arezoana Mohamadzade & Korneyev, 1840) cometa (Loew, T. Mohamadzade & Korneyev, 2017 alamutensis Mohamadzade & Korneyev, T. T. kogardtauica Hering, 1944 T. Korneyev, Khaganinia, Mohamadzade, Zarghani 2015 arsenii Korneyev, T. Mohamadzade & Korneyev, 2012 azari Mohamadzade & Korneyev, T. 1814) Heringina guttata (Fallen, acanthiophilopsis Hering, 1938 T. Korneyev, 2013 Korneyev, afrostriata T. T. arnicae (Linnaeus, 1758) T. 1939 dudichi Aczél, T. Species (for bold selected species sequences in this article) given are T. nozarii Mohamadzade, 2011 T. T. pallescens Hering, 1961 T. Portschinsky, 1892 tatarica Portschinsky, T. (Loew, 1844) (Loew, vespertina T. Korneyev, 2013 Korneyev, robusta T. (Loew, 1844) truncata (Loew, T. Korneyev, 2013 Korneyev, ochroptera T. (Loew, 1844) simplex (Loew, T. Korneyev, 2013 gladius Korneyev, T. T. separata Rondani, 1871 T. Korneyev, 2013 cameo Korneyev, T. T. mutabilis Merz, 1992 T. Portschinsky, 1892 Portschinsky, angulatofasciata T. (Loew, 1844) matricariae (Loew, T. (Loew, 1844) (Loew, ruralis T. T. admissa Hering, 1961 T. T. mariannae Merz, 1992 T. Korneyev & Korneyev 2019 & Korneyev youngiana Korneyev T. T. zernyi Hendel, 1927 zernyi T. (De Geer, 1776) leontodontis (De Geer, T. Séguy, 1930 theryi Séguy, T. Korneyev & Mohamadzade, 2013 tridentata Korneyev T. 1844) (Loew, fallax T. T. sonchina Hering, 1937 T. Korneyev & Korneyev, 2019 & Korneyev, Korneyev kyrghyzica T. divisa Rondani, 1871 T. Korneyev & Kameneva, 1990 & Kameneva, kovalevi Korneyev T. Korneyev & Korneyev, 2019 & Korneyev, ghissarica Korneyev T. Hendel, 1927 crepidis T. (Loew, 1844) (Loew, formosa T. (Schrank, 1803) bardanae (Schrank, T. conyzifoliae Merz, 1992 T. (Loew, 1846) (Loew, dilacerata T. Species (for bold selected species sequences in this article) given are

maccus Genus hyoscyami arnicae valida bardanae leontodontis formosa (b. g.) group Heringina group group group group (b. g.) group group Species Group Species Group I I I I II II III III Clade Clade species groups and linages. — 4. Tephritis Table

120 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (1) 2020 PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA Region Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host Morphology, Morphology, host Morphology, Morphology, host Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host Morphology, Morphology, host, COI Morphology, Morphology Morphology, host Morphology, Morphology, host Morphology, Morphology, host Morphology, host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host Morphology, Character Evidence Character Korneyev 2016 Korneyev 2016 Source Tribe Cardueae Mohamadzade 2012 Cardueae 2013 Korneyev 2013; Shcherbakov Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Korneyev 2013 Cardueae Merz 1994 Cardueae Korneyev & Mohamadzade 2013 host Morphology, Cardueae Korneyev & 2019 Cardueae Korneyev & 2019 Cardueae White 1988; Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Korneyev & 2019 Cichorieae Merz 1994 Cichorieae Merz 1994 Cichorieae Hering 1937 Cichorieae Merz 1994 Cichorieae 1990 Korneyev & Kameneva Cichorieae Merz 1992b, 1994 Cichorieae Merz 1994 Cichorieae 2001 Merz 1994; Shcherbakov Cichorieae Merz 1994 Host Plant Cirsium sp. Cirsium unknown , Ancathia igniaria nivea Alfredia Hypochaeris radicata Cousinia sp. Leontodon crispus , L. incanus tenuiforus Cousinia refracta Crepis albida Crepis Cousinia calathidia Picris hieracioides Cousinia sp. Leontodon hispidus Cousinia sp. Crepis foetida Crepis Hieracium lactuella , H. pilosella Hieracium Cousinia tianschanica Leontodon hispidus Youngia sp. Youngia Arctium minus Arctium , L. hispidus Leontodon autumnalis , L. helveticus unknown Jurinea baissunensis Leontodon hispidus Sonchus uliginosus Cousinia sp. Picris echioides Sonchus paluster Cousinia sp. biennis , C. blattarioides Crepis S. oleraceus Sonchus asper , S. oleraceus , A. lappa , A. minus nemorosum Arctium tomentosum , C. sibirica conyzifolia Crepis Sonchus arvensis Host plant species T. nozarii Mohamadzade, 2011 T. T. pallescens Hering, 1961 T. Portschinsky, 1892 tatarica Portschinsky, T. (Loew, 1844) (Loew, vespertina T. Korneyev, 2013 Korneyev, robusta T. (Loew, 1844) truncata (Loew, T. Korneyev, 2013 Korneyev, ochroptera T. (Loew, 1844) simplex (Loew, T. Korneyev, 2013 gladius Korneyev, T. T. separata Rondani, 1871 T. Korneyev, 2013 cameo Korneyev, T. T. mutabilis Merz, 1992 T. Portschinsky, 1892 Portschinsky, angulatofasciata T. (Loew, 1844) matricariae (Loew, T. (Loew, 1844) (Loew, ruralis T. T. admissa Hering, 1961 T. T. mariannae Merz, 1992 T. Korneyev & Korneyev 2019 & Korneyev youngiana Korneyev T. T. zernyi Hendel, 1927 zernyi T. (De Geer, 1776) leontodontis (De Geer, T. Séguy, 1930 theryi Séguy, T. Korneyev & Mohamadzade, 2013 tridentata Korneyev T. 1844) (Loew, fallax T. T. sonchina Hering, 1937 T. Korneyev & Korneyev, 2019 & Korneyev, Korneyev kyrghyzica T. divisa Rondani, 1871 T. Korneyev & Kameneva, 1990 & Kameneva, kovalevi Korneyev T. Korneyev & Korneyev, 2019 & Korneyev, ghissarica Korneyev T. Hendel, 1927 crepidis T. (Loew, 1844) (Loew, formosa T. (Schrank, 1803) bardanae (Schrank, T. conyzifoliae Merz, 1992 T. (Loew, 1846) (Loew, dilacerata T. Species (for bold selected species sequences in this article) given are

bardanae leontodontis formosa group (b. g.) group group Species Group II III III Clade continued. 4 Table

121 Korneyev et al.: Phylogeny of Tephritis PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA PA NEA NEA NEA NEA NEA NEA NEA NEA Region PA; NEA PA; Morphology, host Morphology, Morphology Host, COI; 16S; Period; AATS; 28S AATS; Host, COI; 16S; Period; Morphology Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Host, COI; 16S; Period; AATS; 28S AATS; Host, COI; 16S; Period; Morphology Morphology, host, COI Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI Morphology, Morphology Morphology, host, COI Morphology, Morphology, host Morphology, Morphology, host, COI Morphology, Morphology, host Morphology, Morphology, host Morphology, Morphology Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology Morphology, host, COI Morphology, Morphology, host, COI Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host Morphology, Morphology, COI Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host Morphology, Morphology Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; host, COI; 16S; Period; Morphology, 28S AATS; host, COI Morphology, Character Evidence Character Korneyev 2016 Korneyev 2016 Korneyev 2016 Jenkins & Turner 1989 Jenkins & Turner Korneyev 2016 Foote et al. 1993 Foote Korneyev 2016 Freidberg & Kugler 1989 Source Tribe Astereae 1989 Jenkins & Turner Astereae 1989 Jenkins & Turner Astereae 1989 Jenkins & Turner Astereae 1989 Jenkins & Turner Astereae 1989 Jenkins & Turner Astereae et al. 1993 Foote Astereae; Astereae; Cichorieae Merz 1994 Cichorieae Freidberg & Kütük 2002 Cichorieae Merz 1994 Cichorieae Freidberg & Kütük 2002 Cichorieae Freidberg & Kugler 1989 Cichorieae Merz 1994 Cichorieae Merz 1994 Host Plant Senecioneae Anthemideae Freidberg & Kugler 1989 Anthemideae Khaghaninia et al. 2011 Anthemideae Merz 1994 Anthemideae Merz 1994 Anthemideae Korneyev 2016 Anthemideae Merz 1992a Anthemideae Freidberg & Kugler 1989 Anthemideae Merz 1994 Anthemideae 1993; collections Korneyev & Kameneva host Morphology, Anthemideae Merz 1994 Otanthus maritimus unknown Achillea clypeolata unknown Anthemis arvensis Leucanthemum vulgare unknown Scorzonera hirsuta Scorzonera Achillea sp. sp. Scorzonera unknown Scorzonera laciniata Scorzonera Artemisia thuscula , B. halimifolia Baccharis glomerulifora Scorzonera kotschyi Scorzonera Pyrethrum santolinoides Pyrethrum unknown Tragopogon longirostre Tragopogon unknown Baccharis neglecta , B. halimifolia Tragopogon dubius Tragopogon Achillea millefolium Baccharis pilularis unknown Baccharis salicifolia spp., Senecio spp. Astereae Artemisia rutifolia Scorzonera hispanica Scorzonera unknown Baccharis sarothroides Ericameria nauseosa Achillea ptarmica Host plant species Loew, 1862 stictica Loew, T. (Loew, 1861) (Loew, spreta T. T. sahandi Mohamadzade et al., 2011 T. T. rydeni Hering, 1956 T. (Loew, 1856) nigricauda (Loew, T. T. neesii (Meigen, 1830) T. (Becker, 1908) nebulosa (Becker, T. T. scorzonerae Merz, 1993 scorzonerae T. (Becker, 1913) multiguttata (Becker, T. Loew, 1869 Loew, recurrens T. Korneyev & Dirlbek, 2000 mesopotamica Korneyev T. (Loew, 1844) (Loew, pulchra T. T. luteipes Merz, 1992 T. T. subpura (Johnson, 1909) subpura T. Freidberg & Kütük, 2002 & Kütük, merzi Freidberg T. Freidberg, 1981 jabeliae Freidberg, T. T. rufpennis Doane, 1899 T. Freidberg, 1981 hurvitzi Freidberg, T. T. hungarica Hering, 1937 T. T. palmeri Jenkins, 1989 T. T. heliophila Hendel, 1927 T. (Loew, 1856) dioscurea (Loew, T. T. californica Doane, 1899 californica T. T. crinita Hering, 1961 T. T. baccharis (Coquillett, 1894) T. stigmatica (Coquillett, 1899) T. 1869 Loew, brachyura T. T. carmen Hering, 1937 T. Freidberg, 1981 bimaculata Freidberg, T. Quisenberry, 1951 arizonaensis Quisenberry, T. 1959 labecula Foote, T. 1844) angustipennis (Loew, T. Species (for bold selected species sequences in this article) given are

labecula/ pulchra group baccharis group stigma­ angustipennis group tica Species Group IV IV IV IV Clade continued. 4 Table

122 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (1) 2020

in the tribe Cichorieae. Morphologically di- PA PA PA PA PA PA PA PA PA PA NEA NEA NEA NEA NEA verse members of Clade IV (BPP = 0.84) are Region associated with the tribes Anthemideae and Astereae, occurring in the Palaearctic and Nearctic Regions, except for T. stigmatica, associated with a few species of Astereae but primarily Senecioneae, and T. hurvitzi, which is the sister to the remainder of the clade, and represents a group of species strictly associated with host plants of the genera Tragopogon and Scorzonera (tribe Morphology, host, COI Morphology, Morphology, host Morphology, Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Morphology Morphology, COI; 16S; Period; AATS; AATS; COI; 16S; Period; Morphology, 28S Morphology, COI; 16S; Period; AATS; AATS; COI; 16S; Period; Morphology, 28S Morphology Morphology, COI; 16S; Period; AATS; AATS; COI; 16S; Period; Morphology, 28S Morphology, host Morphology, Morphology Morphology, host, COI; 16S; Period; host, COI; 16S; Period; Morphology, 28S AATS; Character Evidence Character Cichorieae). For the dataset based on COI, phyloge- netic topology obtained with Bayesian in- ference is shown in Figure 2, representing 64 species of the genus Tephritis, which is largely congruent with the phylogenetic tree based on the fve-genes dataset. Based on combined morphological char- acters, host plant preferences, and molecular Korneyev collected in 2017 Korneyev 2016 Source data of both datasets (fve genes and COI), Table 4 provides a species group classifca- tion for 106 described Tephritis species, in-

Tribe cluding the 35 species used in the fve-gene Astereae Merz 1994 Astereae Korneyev 2016 Astereae et al. 1993 Foote Astereae et al. 1993 Foote Host Plant Senecioneae et al. 1993 Foote Anthemideae Korneyev collected in 2017 Anthemideae Korneyev collected in 2017 Anthemideae Korneyev collected in 2017 Anthemideae et al. 1993 Foote Anthemideae et al. 1993 Foote Anthemideae Korneyev 2016 Anthemideae Merz 1994 phylogenetic analyses and 64 used in the COI analysis. The other 38 species are placed to the groups based on morphological simi- larity to some of the group members or both morphological and host similarity and with the species placed in the group based on mo- lecular analyses. Ten recognized groups of species are discussed under each of the four clades defned in the fve-gene analysis.

3.2. Discussion, clades and Aster alpinus Aster sp. Anthemis sp. sp. Pyrethrum Artemisia sp. unknown , aquafolia , Machaeranthera integrifolia M. canescens Arnica spp. Erigeron spp. Erigeron spp. Tanacetum Artemisia sp. Santolina chamaecyparissius unknown unknown Tanacetum vulgare; T. corymbosum T. vulgare; Tanacetum Host plant species species group recognition

3.2.1. Overview

In this study, we present phylogenetic analyses­ of the genus Tephritis inferred from two mi- tochondrial and three nuclear gene sequences from specimens collected in Europe, Middle East, and North America. Morphologically, the genus Tephritis is a homogenous, which impedes with carrying out a phylogenetic analysis based on morphological characters. For example, the male genitalia, which usu- ally serve as a source of a good number of phylogenetically meaningful characters, are almost identical. On the other hand, body and T. sauterina Merz, 1994 T. Richter, 1975 Richter, corolla T. T . sp.4 T. sp.3 T. sp.2 T. sp.1 Foote, 1960 signatipennis Foote, T. T. leavittensis Blanc, 1979 T. Foote, 1960 ovatipennis Foote, T. Foote, 1960 candidipennis Foote, T. T. araneosa (Coquillett, 1894) T. T. santolinae Hering, 1934 T. (Richter, 1995) (Richter, volkovitshi T. (Becker, 1908) (Becker, variata T. T. tanaceti Hering, 1956 T. Species (for bold selected species sequences in this article) given are wing coloration and patterns used for species identifcation are very variable and seem to angustipennis group be subject to frequent homoplasy.

Species Group The most widely-used morphological char­ acters that are used to group Tephritis species

IV are modifcations of the aculeus shape: leon­ Clade continued. 4 Table todontis with the deep apical notch (Merz

123 Korneyev et al.: Phylogeny of Tephritis

1992, 1993, 1995); pulchra with preapical steps (Frei- erately short (not transformed into a long trunk-like dberg & Kütük 2002); formosa with rounded blunt apex membranous tube) (SP) and sclerotized basal part of the (S. Korneyev & V. Korneyev 2019); maccus with acu­ glans uniformly smooth and having neither spines, nor leus moderately broad and sharply tapered subapically sclerotized appendages, nor other sclerotized sculpture to a short narrow apex (Mohamadzade et al. 2015). The (SP) (in contrast to Dectodesis Munro, Austrotephritis monophyly of each of the above species groups based on Hancock & Drew, Becker, Trupanea, characters not occurring elsewhere in the Tephritis group Benjamin, Hyalotephritis Freidberg, and of genera and thus considered synapomorphies is strong- Goniurellia, which usually have spinulose appendages ly supported by their host-plant relationships, belonging in the basal sinus of the glans). Such a defnition covers to the same genus or tribe of Asteraceae, in contrast to the species currently not included in Tephritis: Herin­ range of wing-pattern types, which varies widely (see gina guttata, H. arezoana, and Multireticula perspicil­ Freidberg & Kütük 2002 as an example). lata (Bezzi 1924), that differ only in a few characters, As has been previously shown for the Carpomyini fruit possibly autapomorphies, which do not contradict their fies, the morphological and molecular phylogenies are possible inclusion into Tephritis (broad, widely dark pat- often incongruent (Smith et al. 2006). Thus, the morpho- terned wing; occasionally setulose dorsal surface of vein logical characters are sometimes misleading when trying R4+5; partly shining or mainly yellow, black spotted ab- to identify monophyletic groups of species, which are ex- dominal tergites). tremely diverse in the body coloration and wing patterns. We consider them belonging in the Tephritis lineage On the other hand, the species groups within Tephritis are as a monophyletic taxon supported both by at least a sin- moderately supported by the association with certain host gle morphological synapomorphy (2 frontal setae vs. 3 plant taxa (genera, subtribes or tribes). It corresponds well frontal setae in most genera in the Tephritis group), in with the hypothesis that the early evolutionary events in addition to support via molecular-based phylogenetic such groups were the shifting to another, often distantly analysis. However, formally this inclusion is to be a part related but common and abundant host plant from an ex- of a wider forthcoming taxonomic revision. tinct host (compare with the Rhagoletis groups of species As has been previously noted (S. Korneyev 2013; well supported by the host plants and molecular phyloge- S. Korneyev & V. Korneyev 2019), some large complex- ny vs. morphology (Smith et al. 2006) and that ecological es of species with signifcant characters useful for identi- factors involving adaptations to and isolation of popula- fcation (e.g., the “apical fork” modifed into a dark apical tions on certain host plants have played as important a role crossband, mushroom-like spot or two apical spots sepa- in many cases as geographic isolation (Bush 1969, 1992) rated from remaining dark pattern) are clearly polyphyl- in the evolution of frugivorous tephritids. etic as shown by the current molecular-based analysis. The fve-gene molecular phylogenetic analysis shows The species groups with monophyly supported by the genus Tephritis (along with the species assigned to modifcations of the aculeus tip shape are the leontodon- the nominal genus Heringina as part of the in-group) tis, pulchra, formosa and maccus groups, as noted above. to be monophyletic (BPP = 0.95). It forms a monophy- About 1/3 of the described Tephritis species are not placed letic lineage with Trupanea, Goniurellia, and Capitites in any of the species groups discussed above (Table 4). (BPP = 0.99), which corresponds to the Tephritis group of Along with that, the phylogenetic analysis based on genera (sensu Merz 1999); the genus Capitites appears molecular data reveals several additional monophyletic here as a sister-group of the Tephritis (Fig. 1). This cor- species groups based on specialization for certain host roborates results obtained in the COI-based analysis with plant taxa. It concerns the angustipennis, arnicae and a wider set of taxa and partly by the morphological data valida species groups (Fig. 1), and certain lineages in the (S. Korneyev 2016 and unpublished data). angustipennis and hyoscyami groups strictly associated However, in some analyses, the position of Capitites with closely related plant taxa. differs alternatively being the sister-group of the Acan­ thiophilus – Trupanea – Tephritomyia lineage (Morgulis 3.2.2. Clade I (Fig. 1): Heringina and maccus 2017) or of Acanthiophilus – Trupanea – Tephritomyia – group Tephritis (Han, pers. comm.). Based on these results, the genus Tephritis can be defned as a monophyletic clade Each of the three lineages represented in Clade I (Fig. 1) within the Tephritis group of genera (Merz 1999) of the differ greatly from each other as well as from other species­ tribe Tephritini and a group of species with two pairs of Tephritis, in their wing patterns (Heringina, T. mac­ of frontal setae (synapomorphy (SA) of the genus), two cus), aculeus shape (Heringina, T. maccus, T. oedipus) or pairs of orbital setae (symplesiomorphy (SP)), and two phallus glans sclerotization (T. praecox). Monophyly of pairs of scutellar setae (SP), short spatulate labellum the clade is poorly supported and needs additional inves- (SP), wing apex with 3 large hyaline spots separated tigation. by two dark radiate bars (“apical fork”; SP; secondar- The subclade containing Heringina and T. praecox is ily modifed in several terminal lineages); male geni- fairly supported (BPP = 0.89) and includes two taxa mor- talia with phallus glans widely membranous in apical phologically different from each other and the remaining half and lacking the acrophallus (SA of the Tephritis species of Tephritis. The nominal genus Heringina, with group of genera), membranous part of the glans mod- two species, represents a lineage of rhizome-bud feeding

124 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (1) 2020

species (V. Korneyev et al. 2020, in prep.) associated with all the diversity of wing patterns). In molecular-based Helichrysum (tribe Gnaphalieae), whereas T. praecox is phylogenies, this lineage has the weakest support of the known to have fower head feeding maggots reared from four clades defned within Tephritis in the molecular Calendula (tribe Calenduleae), a tribe closely related phylogeny (BPP = 0.83). Most species within Clade II to Gnaphalieae. According to Rennwald et al. (2015), are associated with thistles (tribe Cardueae) showing no there are repeating references to Filago (Gnaphalieae) morphological synapomorphies. Clade II includes two as its additional host plant; these references can be traced well-supported monophyletic subclades within. down to J. Mik (1880), Hendel (1927), Mihályi (1960) The frst subclade includes Cirsium- and Carduus- unconfrmed and either based on misidentifcation of an associated species along with two species with non-Car- undescribed species or represent a real record from an dueae hosts. The Cardueae associated species: T. come­ alternative host plant. ta, T. conura, and T. hyoscyami share no morphological The subclade containing T. oedipus and T. maccus synapomorphies; each of these lineages represents sever- has moderately high support (BPP = 0.91). The mac- al morphs or host races currently considered conspecifc, cus group is clearly monophyletic and supported by a but showing very small differences in their mitochondrial unique morphological synapomorphy (aculeus moder- COI sequences: 1) T. cometa + T. acanthiophilopsis line- ately broad and sharply tapered subapically to a short age; 2) T. conura + unnamed host races (Seitz & Komma narrow apex: Fig. 3B) it was defned in Korneyev & 1984; Romstock & Arnold 1987; Diegisser 2005); 3) Dirlbek (2001). The group is represented in the fve gene T. hyoscyami + T. hendeliana. We consider these three phylogeny by T. maccus alone. It has a total of four ex- lineages corresponding to the hyoscyami species group tant species: T. azari Mohanadzade & Korneyev, 2012, sensu Merz (1995). Additional species closely related to T. gharalii Mohanadzade & Korneyev, 2015, T. maccus them, similar morphologically and also associated with Hering, 1937, and T. urelliosomima Korneyev & Dirl- Cirsium and Carduus, namely T. anthrax, T. atocoptera, bek, 2001; plus one extinct fossil species, T. sophus Gen- T. campana, T. cardualis, T. erdemlii, and T. umbrosa are tilini & Korneyev, 2006 (upper Miocene; the only fossil to be included in this group (S. Korneyev & Evstign- record of Tephritis (Gentilini et al. 2006)). Detailed in- eev 2019). T. longicauda Sueyoshi feeding in Cirsium formation about the group morphology and distribution fower heads in the Easetrn Palearctics show “intermedi- is given in Mohamadzade & S. Korneyev (2012) and ate” characters of T. cometa and T. conura (V. Korneyev Mohamadzade et al. (2015). No data on the host plants 1995; V. Korneyev & Ovchinnikova 2004) and may also of the species of the maccus group are known, except belong in this group. a sample of T. sp. cf. maccus swept in a stand of Lac­ Both the hyoscyami species group and the frst tuca scariola, its possible host plant. As the larvae of subclade appear to be paraphyletic, for the next two, T. oedipus feed in fower heads of Lactuca tatarica (tribe “non-thistle”, groups are nested within them. 1) Tephritis Cichorieae), these data could support a common plant kogardtauica, T. valida, and T. dudichi closely related ac- genus for the T. oedipus + T. maccus lineage and their cording to the COI tree (Fig. 2), as well as 2 undescribed sister-group relationship. new species, all feeding in fower heads of Inuleae, and Tephritis praecox, T. oedipus, T. maccus group spp. form the valida group. 2) Tephritis arsenii and T. arni­ and Heringina spp. have aberrant wing patterns, aculeus cae infesting Senecioneae plants form the arnicae group shape and / or sclerotization of phallus glans compared to (supported by the COI tree, Fig. 2). Species of both the rest of Tephritis species, and thus each can be consid- groups share the wide, subrectangular spots at apices of ered to be its sister-group, but neither the morphological veins R4+5 and M1+2, usually separated from the dark wing data, nor molecular phylogeny do not resolve polytomies pattern, with each other and T. hyoscyami + T. hendeli­ neither for separation them to genus-group taxa, nor for ana species that may also support their closer relation- lumping them (including synonymization of Heringina ships. Five-loci analysis reveals possible monophyly of and Tephritis). the T. kogardtauica + T. arsenii lineage, but its support is low (BPP = 0.65) and dubious. Moreover, in the COI 3.2.3. Clade II (Fig. 1): hyoscyami, arnicae, analysis, the arnicae group appears to be related much valida, and bardanae groups closer to T. ruralis, the species showing strong morpho- logical resemblance with T. arsenii and T. arnicae, but Clade II corresponds to the hyoscyami species group feeding in the Cychorieae. (sensu Merz 1995). The original defnition was based on Monophyly of the second subclade of Clade II two characters believed to be synapomorphies: oviscape (T. bardanae + T. tatarica + T. nozarii and their closest basally white setulose and “wing pattern with two nar- relatives) was revealed by both analyses to include spe- row rays along the apical portion of veins r4+5 and m1+2, cies very diverse morphologically but mostly associated giving a starshaped appearance”. He noticed that this with Cousinia, a vast paraphyletic genus with burdocks group may be further divided into two monophyletic (Arctium) nested within it. Monophyly of the bardanae subgroups, each with two species-groups (Merz 1995). group defned herein and corresponding to this lineage is The morphology-based monophyly of this group has not supported by the results of the COI-based phylogenetic been ever demonstrated (the frst character is present in analysis to include T. bardanae and T. alamutensis sis- most outgroups and the second is hardly applicable to ter taxa in the COI tree (with morphologically similar

125 Korneyev et al.: Phylogeny of Tephritis of Countries and States: AF – Afghanistan; CH – CH Afghanistan; – AF States: and Countries of Abbreviations posterior probability PP. Host plant associations are given as colored circles. — circles. colored as given are associations plant Host PP. probability posterior Abbreviations of Countries: AF – Afghanistan; CH – Confoederatio Helvetica (Switzerland); FR – France; IR – Iran; KG – Kyrgyzstan; NL – The recovered by Bayesian inference and Maximum Likelihood. Support values are recorded on each node as Bayesian posterior probability and maximum maximum and probability posterior Bayesian as node each on recorded are values Support Likelihood. Maximum and inference Bayesian by recovered Fig. 1. Phylogram based on data from two mitochondrial genes (COI, 16S) and three nuclear genes (ribosomal 28S, protein coding Period AATS) and Fig. 2. Phylogram based on data from one mitochondrial gene COI as recovered by Bayesian inference. Support values are recorded on each node as

likelihood bootstrap percentages PP/BP. Roman numerals indicate major clades discussed in likelihood the bootstrap text. percentages Host PP/BP. plant associations are given as colored circles. — – Ukraine; US . Netherlands; UA ↑ as ↓ Bayesian Confoederatio Helvetica (Switzerland); ES – Spain; FR – France; GR – Greece; IL – Israel; IR – Iran; IT – Italy; KG – Kyrgyzstan; NL – The Netherlands; Netherlands; The – NL Kyrgyzstan; – KG Italy; – IT Iran; – IR Israel; – IL Greece; – GR France; – FR Spain; – ES (Switzerland); Helvetica Confoederatio Texas. TX – – California; MD Maryland; Arizona; CA AZ – – Ukraine; US United States; RU – Russia; RS Serbia; UA

T. zernyi, T. ghissarica Korneyev & Korneyev, T. kyrghy­ and hardly contradicting their possible basal position zica Korneyev & Korneyev, T. ochracea, and T. afro­ (Fig. 2). Their host-plant associations are inknown; at striata). The bardanae group also includes T. tatarica most T. nozarii was reported to be swept from Cirsium and morphologically similar related species: T. admissa, (Mohamadzade 2012), which is not necessarily its host T. cameo, T. gladius, and T. robusta, which infest Cous­ plant. On the fve-gene tree T. nozarii, forms a monophy- inia and share a synapomorphy: a short apical crossband letic lineage with T. tatarica (BPP = 0.98). on the wing apex, which does not occur in the outgroups The last ungrouped species also belonging in the (S. Korneyev 2013). Clade II, is T. postica, which appears in an unresolved On the COI tree, the lineage corresponding to the lat- basal polytomy in the COI-based phylogram and a sis- ter assemblage also includes T. pallescens and T. nozarii, ter taxon to the hyoscyami species group but with a the two species with a very diffent, pale grid-like wing low support). It is associated with various species of the patterns; their topological position inside the lineage is genus Onopordum (Cardueae), which also supports its poorly supported (BPP of lineages inside < 0.7), vague, placement in the Clade II.

126 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (1) 2020

Judging from the host-plant preferences alone or also of veins R4+5 and M1+2, isolated from the remaining dark from the wing pattern, T. tridentata S. Korneyev et Mo- wing pattern, may also belong to this clade; as soon as no hamadzade Namin, 2013, an unplaced species associated DNA sequences are available, we place it here provision- with Jurinea (Cardueae) and having the spots at apices ally.

127 Korneyev et al.: Phylogeny of Tephritis

Fig. 3. Morphological synapomorphies of Tephritis species groups (modifcations of the aculeus tip shape). A: common aculeus shape for Tephritis (T. nigricauda); B: maccus group: aculeus sharply tapered subapically to a short narrow apex (T. maccus); C: leontodontis group: notched tip of the aculeus (T. separata); D: formosa group: aculeus subapically poorly sclerotized and apically narrowed and rounded (T. formosa); E: pulchra group: aculeus with pair of subapical steps (T. hurvitzi); F: Heringina: aculeus tip rounded with pair of steps on sides (H. guttata).

3.2.4. Clade III (Fig. 1): leontodontis and formosa Tephritis group of genera) and apically narrowed and groups rounded aculeus (Fig. 3D). T. dilacerata, T. formosa, T. ko­valevi, and T. sonchina have the oviscape entirely This clade has a high support in the fve-gene tree (BPP = covered by brown setulae, whereas T. youngiana Kor- 1), but the species included here share no synapomor- neyev & Korneyev, 2019 has a white-setulose oviscape phies. They all are associated with host plants of the tribe and is believed to be the sister-group to the other four Cichorieae (mainly Sonchus, Crepis, Picris or closely species, which also share association with fower heads related genera). The four species included in the fve gene of Sonchus, whereas T. youngiana is associated with analysis represent two species groups with high support Youngia (sometimes considered a synonym of Crepis). (BPP = 1) also in the COI analysis. Monophyly of T. formosa + T. kovalevi + T. youngiana merz (1995) defned the leontodontis group as hav- lineage is partly supported by the COI analysis (Fig. 2; ing aculeus apically not evenly rounded (with steps or in- BI = 0.77), with unresolved basal polytomy, which in- dentations, both states absent in the other Tephritis or in volves also morphologically different T. matricariae the outgroups). This multistate character cannot be con- (Loew), here placed to the leontodontis group. The for- sidered a single synapomorphy; instead, the aculeus tip mosa group provisionally includes a strictly endemic, shape provides several modifcations as synapomorphies little-known T. theryi Séguy from Morocco, which is be- of more restricted lineages considered below. lieved to be closely related to T. formosa. The leontodontis group, as defned here, includes T. crepidis, T. conyzifoliae, T. leontodontis, T. divisa, T. se- 3.2.5. Clade IV (Fig. 1): pulchra, baccharis and ­pa­rata, T. matricariae (Loew), T. mutabilis and based on angustipennis groups morphological data and host-plant preferences, also T. ma- ­­rian­nae, T. simplex (Loew), and T. truncata. These spe- This clade has moderate support (BPP = 0.84) and con- cies share at least one morphological synapomorphy not tains two major lineages, which correspond to the groups occurring elsewhere in the Tephritis group of genera, the considered below. deeply notched tip of the aculeus (Fig. 3C) (Merz 1995) The pulchra group (Merz 1993; Freidberg & Kütük and the group is strongly supported (BP = 1) in the COI 2002) corresponds to the T. hurvitzi lineage in this ana­ analysis (Fig. 2). In the fve-locus analysis, the leonto- lysis. It has been established based on two morpholo­ dontis group is represented by T. conyzifoliae and T. sep­ gical synapomorphies (oviscape with brown setulae arata, supporting its close relationship to the following only; aculeus with pair of subapical steps, both states group (Fig. 2). not occurring elsewhere in the Tephritis group of gen- Position of T. frauenfeldi Hendel, which has the wing era (Fig. 3E) and includes T. recurrens, T. hurvitzi, pattern very similar to T. leontodontis, and the aculeus T. heliophila, T. pulchra, T. scorzonerae, and T. crinita with shallow apical notch, but associated with Jurinea (a monophyletic clade supported by a neighbour-join- (Cardueae), remains unclear, as no DNA sequences are ing analysis based on COI (Smit et al. 2013), the COI available. analysis in this study (BI 0.94) (Fig. 2), and morpho­ The formosa group, as defned here, includes fve or logy: aculeus tip shape), and also T. carmen and T. mer­ six species with at least one synapomorphy: the subapi- zi. All the species except T. crinita (whose biology is cally poorly sclerotized (not occurring elsewhere in the unknown) induce communal fowerhead or stem galls

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in closely related Scorzonera and Tragopogon plants small size (see above), short aculeus and the cell r1 with 2 (tribe Cichorieae). hyaline spots. Among all the species of the genus, T. an­ The other subclade has moderately high support (BPP gustipennis has a circumboreal distribution from Europe = 0.87), containing a number of poorly supported line- through Siberia to Alaska, Canada, and south to Colorado ages, represented by species associated mainly with the in the USA (Foote et al. 1993). tribes Anthemideae and Astereae. In the fve-gene analysis, this group is represented by The baccharis group defned by Jenkins & Turner the three latter species, which form a monophyletic sub- (1989) is represented in this study by four morphologi- clade (BPP = 0.99), but the other species also associated cally diverse species (forming a well-supported subclade with Achillea and closely related Tanacetum, as T. sp. nr. (BPP = 0.98) in the Nearctic Region. This group includes tanaceti and T. sahandi, both are not pulled in this line- T. arizonaensis, T. baccharis, T. californica, T. palmeri, age, staying in an basal set of taxa with low support or T. rufpennis, and T. subpura (Jenkins & Turner 1989). “almost unresolved basal polytomy”. It is apparently a strictly monophyletic group represented Another part of this group also includes the species by the species are associated with hosts in the genus Bac­ either morphologically similar or poorly recogniable charis (tribe Astereae) and restricted in distribution to from T. nigricauda or associated with Leucanthemum ne Nearctic Region, but having no obvious synapomor- and related genera of the Anthemideae. In the fve-gene phies. The COI-based analysis (Fig. 2) shows that the lin- and COI-based analyses they are represented by T. sp. eage represented by T. arizonaensis and T. baccharis also nr. nigricauda and three additional undetermined species includes closely related T. palmeri and T. subpura; of mentioned by Smit et al. (2013). These species usually them, T. arizonaensis, T. californica, and T. palmeri have have a wide medial dark bar in cell cua, wide spots at an extremely short and wide “apical fork” in the wing the apices of veins R4+5 and M1+2 usually fused with each pattern, T. baccharis and T. rufpennis possess a further other and the remaining dark pattern, and a moderately modifed “apical dark”, but the wing pattern in T. sub­ short oviscape (at most as long as the 3 posteriormost ex- pura has a wing pattern with more typical apical fork. posed abdominal tergites). Additionally, T. bimaculata, Species in this group sometimes have additional white T. spreta, T. jabeliae, and T. mesopotamica may also be- frontal setae. The sister-lineage for the baccharis group long here. Despite their strong morphological similarity, is the Nearctic T. labecula, feeding in Ericameria (also species of this group show considerable genetic diversity, the tribe Astereae), but having a wing pattern strongly even falling into different lineages and giving the appear- resembling those of T. crinita, T. hurvitzi and T. merzi of ance of paraphyly of the group. the pulchra group. Furthermore, T. stigmatica, sister to The position of T. neesii, which also feeds in Leucan­ the rest of the species in the baccharis group+ labecula themum, appears to be basal to this and the following lin- lineage, has a similar wing pattern with a distinct stellate eage (Smit et al. 2013; S. Korneyev unpublished data). pattern connected to the pterostigma by a broad band, and A clade which includes T. araneosa, T. leavittensis, feeds in the fower head or stem galls of various Senecio and T. ovatipennis from the Nearctic Region and at least (tribe Senecioneae) (Novak et al. 1967; Goeden 1988). four undescribed new species from the Central Asian In the COI anaysis, this lineage unexpectedly contains mountains of the Palaearctic Region appears, on the Heringina with two species of predominantly Palaearctic contrary, to be strictly monophyletic, mostly Astereae- Middle East distribution and non-related hosts, which associated and nested among the Anthemideae-asso- looks extremely unlike and resulted from the method er- ciated species. Two of the undescribed species were ror. swept from Astereae (Aster, Erigeron, Chrysothamnus) The remaining species in Clade IV in the 5 gene at 2500 – 3500 m a.s.l. They mostly share blackish leg analysis form the angustipennis group (defned here), coloration, dark wing pattern, short oblique hyaline which comprises mainly smaller species (WL = 2.7 – 4.0 crossband distal to vein R1 apex, shining black tergite 6 mm) associated with plants of the tribes Anthemideae, and oviscape of female covered entirely by black setu- Astereae, and occasionally Senecioneae. Species in this lae. However, none of these characters occurs in all the group have similar grid-like wing patterns having two, species and thus can be considered a synapomorphy to three, or one large, often subrectangular hyaline spots in determine this group as a whole. We tentatively place cells r1, r2+3 and r4+5. Monophyly of this clade has very low other species into this group based on their morphologi- support on the fve gene tree (BPP = 0.56), but the mor- cal similarity, their host association, or on COI molecular phology and host plant preferences support this theory data, including the Nearctic T. candidipennis and T. sig­ of close relationship. In the COI-based analysis it shows natipennis, and possibly also the Palaearctic T. corolla paraphyly, with the pulchra group nested in it, and an and T. sauterina, the latter species infesting fower heads unresolved basal polytomy. of Aster alpinus (tribe Astereae). The species associated with various Achillea, Tan­ The poor resolution and high polytomy in this clade acetum and possibly Artemisia (tribe Anthemideae) here requires further studies and explanation. We can specu- considered angustipennis group with T. angustipen­ late if it may refect rather recent evolutionary events, nis, T. hungarica, T. rydeni, T. nebulosa, T. brachyura, such as a rapid radiation following Pleistocene glacia- T. dioscurea, T. tanaceti, and T. multiguttata, show no tion. This hypothesis is indirectly supported by the fact evident morphological synapomorphies, except possibly that Clade IV is the only part of the genus that has sub-

129 Korneyev et al.: Phylogeny of Tephritis

clades of either Holarctic or strictly Nearctic distribution, their help in NKMB () and support by the German Agency which could be formed by Trans-Beringean migration of of Academic Exchange (DAAD) in 2014. Our sincere thanks to ancestral populations. E. Altenhofer, S.M. Blank, D.A. Fachin, P.H. Kerr, E.J. Rodriguez, J.D. Lopez and K. Kramp for collecting material used in this work. We also thank two anonymous reviewers for their valuable com- 3.2.6. Tephritis incertae sedis ments. Mention of trade names or commercial products in this pub- lication is solely for the purpose of providing specifc information Most species of Tephritis in the analysis are here assigned and does not imply recommendation or endorsement. to ten species groups: angustipennis, arnicae, baccharis, bardanae, formosa, hyoscyami, leontodontis, maccus, pulchra, and valida. However, several remain unplaced: 6. References T. labecula, T. postica, T. stigmatica, T. oedipus, T. prae­ cox. Similarly, the species here assigned to Heringina are Agarwal M.L., Kapoor V.C. 1988. Four new species of fruit fies not forming a sister group to Tephritis and future stud- (Diptera: Tephritidae: Tephritini) together with redescription of ies have to consider whether Tephritis and Heringina are Trupanea inaequabilis Hering and their distribution in India. – synonyms. These studies should also include Austrote­ Journal of Entomological Research 12(2): 117 – 128. phritis, Capitites, Dectodesis, Heringina and address Aigbedion-Atalor P.O., Wilson D.D., Eziah V.Y., Day M.D., Pat- erson I.D. 2018. The distribution and abundance of the stem- the status of Tephritis praecox. About 70 species remain galling fy, connexa (Macquart) (Diptera: Tephri­ poorly known or unavailable for study and have neither tidae), a biological control agent of (L.) been analyzed nor placed into the any of species groups (Asteraceae), in Ghana. – African Entomology 26(2): 471 – 480. discussed here. Barr N.B., Cui L., McPheron B.A. 2005. Molecular systematics of nuclear gene period in genus Anastrepha (Tephritidae). – Annals of the Entomological Society of America 98(2): 173 – 180. Bush G.L. 1966. The , cytology and evolution of the ge- 4. Conclusions nus Rhagoletis in North America. – Bulletin of the Museum of Comparative Zoology 134: 431 – 562. Bush G.L. 1969. Mating behavior, host specifcity, and the eco- One hundred and six species of the genus Tephritis are logical signifcance of sibling species in frugivorous fies of the genus Rhagoletis (Diptera: Tephritidae). – American Naturalist placed in ten species groups (Table 4), based on morpho- 103: 669 – 672. logical, ecological or molecular features, whereas the re- Bush G.L. 1992. Host race formation and sympatric speciation in maining species (about 70) have not been placed into the Rhagoletis fruit fies (Diptera, Tephritidae). – Psyche 99: 335 – 357. any of species groups yet, as they remain poorly known Day M.D., Bofeng I., Nabo I. 2013. Successful biological control or unavailable for study. Future efforts will focus on in- of Chromolaena odorata (Asteraceae) by the gall fy Cecid­ ochares connexa (Diptera: Tephritidae). Papua New Guinea. Pp. creasing sampling density, and obtaining representative 400 – 408 in: Wu Y., Johnson T., Sing S., Raghu S., Wheeler specimens from geographically distant and biologically G., Pratt P., Warner K., Center T., Goolsby J., Reardon R. different populations and species. With these in hand, (eds), Proceedings of the XIII International Symposium on the molecular phylogenetic analysis in the context of mor- Biological Control of Weeds. – USDA Forest Service-FHTET, phology and host plant use should clarify hitherto unre- Morgantown, WV. 536 pp. Day M.D., Riding N., Senaratne K.A.D. 2016. The host specifc- solved issues of the taxonomy of the genus Tephritis and ity and climatic suitability of the gall fy related taxa such as Austrotephritis, Capitites, Dectode­ (Diptera: Tephritidae), a potential biological control agent for sis, Heringina and the status of Tephritis praecox. Chromolaena odorata (Asteraceae) in Australia. – Biocontrol Science and Technology 26(5): 691 – 706. Diegisser T. 2005. Artbildung via Wirtsrassen bei Tephritis conura (Diptera: Tephritidae). – Ph.D. thesis. Universität Mainz, Ger- 5. Acknowledgements many. Edgar R.C. 2004. MUSCLE: multiple sequence alignment with high accuracy and high throughput. – Nucleic Acids Research This work was fnancially supported by IIE, Fulbright Ukraine, Re- 32: 1792 – 1797. search and Development Program 2017 – 2018 (IIE Participant ID: Edwards P.B., Adair R.J., Holtkamp R.H., Wanjura W.J., Bruzz- PS00246781; Grantee ID: E0579598) fellowship for S. Korneyev ese A.S., Forrester R.I. 2009. Impact of the biological control and the Smith laboratory, Department of Entomology at Michigan agent (Tephritidae) on bitou bush (Chrysan­ State University. Some of the pictures were taken in Museum für themoides monilifera subsp. rotundata) in eastern Australia. – Naturkunde Berlin and fnancially supported by German Agency Bulletin of Entomological Research 99(1): 51 – 63. of Academic Exchange (DAAD (GR 6220 26202)) in 2014. This Foote R.H., Blanc F.L., Norrbom A.L. 1993. Handbook of the work could not have been possible without the generous help from Fruit (Diptera: Tephritidae) of America North of Mexico. – Dr Bernhard Merz and his signifcant impact in Tephritis identifca- Comstock Publishing Associates, Ithaca. xii + 571 pp. tion and taxonomy. We want to thank Samad Khaghaninia, Saeed Frauenfeld G.R. von 1857. Beitrage zur Naturgeschichte der Try- Mohamadzade Namin, Younes Karimpour, Babak Gharali, Doug peten nebst Beschreibung einiger neuer Arten. – Sitzungsber- Yanega, Emily A. Hartop, Lisa Gonzales, Giar-Ann Kung, Brian ichte der Akademie der Wissenschaften in Wien 22: 523 – 557. Brown, Martin Hauser, Alessandra Rung, Lucrecia Rodriguez, Freidberg A., Kugler J. 1989. Fauna Palaestina. Insecta IV. Dipte­ Morgan Jackson, Nicole E. Wonderlin, Courtney Larson, Rachel ra: Tephritidae. – Jerusalem: Humanities, Israel Academy of Sci- Osborn, Alexa R. Warwick, Marta Kolomayets, Inna Barysh, Anton ences. 212 pp. Senenko, Krystyna Kernytska, Elena P. Kameneva, Dmitry Minko Freidberg A., Kütük M. 2004. A new species of Tephritis from Tur- and Vladimir Lantsov for their valuable help. We thank Joachim key, with a key to the species of the Tephritis pulchra group. – Ziegler and Bernhard Schurian, Jenny Pohl and Sven Marotzke for Israel Journal of Zoology 48: 295 – 311.

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Authors’ contributions Electronic Supplement Files

All work in molecular lab was done by S. Korneyev under super- at http://www.senckenberg.de/arthropod-systematics vision and support of J. Smith and with assistance of D. Hulbert. DNA was extracted from specimens collected in 2004 – 2017. Most ASP_78-1_Korneyev_Electronic_Supplements.zip of the specimens were collected in Eastern Europe and Middle Asia DOI: 10.26049/ASP78-1-2020-05/1 (S. Korneyev and V. Korneyev), Western, Southern Europe and Is- rael (J. Smit). Material from North America is kindly provided by File 1: korneyev&al-tephrisphylogeny-asp2020-electronicsupple- A. Norrbom and S. Gaimari. Analysis was done by S. Korneyev ment1 ― Nexus fle with COI, Period, AATS, 16S and 28S se- under supervision of D. Hulbert, article was written by S. Korneyev quences for 5 gene tree shown on Fig. 1. and signifcantly improved by all coauthors. File 2: korneyev&al-tephrisphylogeny-asp2020-electronicsupple- ment2 ― Nexus fle with COI sequences for COI tree shown on Fig. 2.

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