Phylogenetic relationships of nonbiting midges in the subfamily Tanypodinae (Diptera: Chironomidae) inferred from morphology

FABIO LAURINDO DA SILVA; TORBJØRN EKREM

Article Alotanypus vittigera (Edwards) comb. nov.: adult redescription, immature description and a phylogenetic analysis of the genus (Diptera: Chironomidae: Tanypodinae) Abstract Anatopynia vittigera Edwards is transfered to Alotanypus. The male and female of A. vittigera comb. nov. are redescribed and immatures are described and illustrated. A cladistic analysis including one species of each Macropelopiini genus was conducted in order to assess the phylogenetic position of Alotanypus and to provide the first phylogenetic hypothesis for the genus. Adults and immatures were included in the analysis where discrete and continuos characters were considered. The cladistic analysis demonstrated that Alotanypus is a monophyletic genus, with Guassutanypus oliveirai as the sister group.

Systematic Entomology (2016), 41, 73–92 DOI: 10.1111/syen.12141 Phylogenetic relationships of nonbiting midges in the subfamily Tanypodinae (Diptera: Chironomidae) inferred from morphology F A B I O L A U R I N D O D A S I L V A 1 and T O R B J Ø R N E K R E M 2 1 Museum of Comparative Zoology, Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA, U.S.A. and 2 Department of Natural History, NTNU University Museum, Norwegian University of Science and Technology, Trondheim, Norway Abstract. The nonbiting midge subfamily Tanypodinae represents one of the most diverse lineages of Chironomidae. Despite the wide distribution and high diversity of tanypodine chironomids, the evolutionary history of the subfamily remains poorly understood. Here, we present the first phylogenetic analysis of the subfamily Tanypodinae based on morphological data. Cladistic analyses were conducted using 86 morphological characters from 115 species belonging to 54 tanypodine genera, including the eight currently recognised tribes: Anatopyniini, Clinotanypodini, Coelopyniini, Macropelopiini, Natarsiini, Pentaneurini, Procladiini and Tanypodini. We use characters from fourth-instar larvae, pupae and adults of both sexes. We examine the effects of implied weighting by reanalysing the data with varying values of concavity constant (k). Our analysis supports the monophyly of Tanypodinae with Podonominae as its sister group. All previously proposed tribes are recovered as monophyletic assemblages under a wide range of weighting factors. Under these conditions, the genus Fittkauimyia is the sister group of the remaining Macropelopiini and is erected as a new monobasic tribe, Fittkauimyiini trib.n. The tribe Pentaneurini is recovered as monophyletic with some internal relationships resolved. The genus Paramerina, recovered as sister of Reomyia + Zavrelimyia, is formally synonymised with Zavrelimyia syn.n., based on morphological similarity in all three life stages and treated as a subgenus of the latter. Finally, the recently suggested synonymies of Gressittius and Guassutanypus with Alotanypus and the establishment of the subgenera Conchapelopia (Helopelopia), Macropelopia (Bethbilbeckia), Monopelopia (Cantopelopia), Thienemannimyia (Hayesomyia) and Zavrelimyia (Reomyia and Schineriella) are investigated. Our results support all proposed changes, except for the subgenus-level status of Helopelopia and Cantopelopia. We suggest re-establishment of Helopelopia as a genus, but refrain from promoting genus-level status of Cantopelopia at present because the apparent sister-relationship between Monopelopia + Nilotanypus likely is due to wing vein reduction caused by miniaturisation. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid: zoobank.org:pub:DF012C17-AFB3-4904-83DC-30DD94D0B376. Correspondence: Fabio Laurindo da Silva, Museum of Comparative Zoology, rm. 406A, Harvard University, 26 Oxford Street, Cambridge, MA 02138, U.S.A. E-mail: laurindodasilva@fas.harvard.edu © 2015 The Royal Entomological Society 73 74 F. L. Silva and T. Ekrem Introduction Insects belonging to the family Chironomidae are true flies (order Diptera), and the most widely distributed free-living holometabolous insects (Ferrington, 2008). The immature stages of most species occur in freshwater, but numerous terrestrial or marine species are known (Sæther & Ekrem, 2003). The imagines are usually flying midges, and although copulation on the ground exists, most species reproduce through swarming in the air. The adult life stage of chironomids is short, and they have limited ability for aerial dispersal compared to many other freshwater insects. Presumably, the great species and habitat diversity in this family is a product of its antiquity, relatively low vagility and evolutionary plasticity (Ferrington et al., 2008), which makes the family not only a valuable source of indicator species for lentic and lotic aquatic ecosystems, but also a most interesting group for phylogenetic and biogeographical analyses. Although the distribution of the species in many genera is relatively well known, detailed analyses of distribution patterns and historical biogeography are rare in chironomids, especially from the Neotropical and Oriental regions. Chironomidae probably originated in the middle Triassic approximately 248–210 Ma (Cranston et al., 2010). It comprises at least 10 000 species in more than 400 genera (Armitage et al., 1995; Sæther et al., 2000) and roughly 6200 of these are known to science (P. Ashe, personal communication). Chironomidae is the most widespread of all aquatic insect families, occurring in all zoogeographical regions of the world (Silva et al., 2015). Phylogenetic studies on Chironomidae have a fairly long tradition, beginning with Goetghebuer (1914), who studied the relationships between groups in the family. However, it was not until Brundin’s monograph (1966) that species-level phylogenies of chironomid midges were produced (Ekrem, 2003). Since then, several groups have been investigated (e.g. Sæther, 1971, 1977, 1983, 1990, 2000; Brundin & Sæther, 1978; Adam & Sæther, 1999; Boothroyd & Cranston, 1999; Ekrem, 2003; Stur & Ekrem, 2006; Fu et al., 2010; Carew et al., 2011; Krosch et al., 2011; Krosch & Cranston, 2013). Cranston et al. (2012) reconstructed the phylogeny of Chironomidae based on DNA sequence data from multiple genes. However, within the subfamily Tanypodinae only the genera Alotanypus (Siri et al., 2011) and Labrundinia (Silva et al., 2015) have had phylogenetic hypotheses proposed at the species level. Tanypodinae is the third most speciose subfamily in the Chironomidae, with species distributed widely across most of the globe, occupying a diverse array of habitats including small streams and ponds to lakes and bays (Silva et al., 2011). The larvae of the majority of species are free-living and none are known to produce larval or pupal cases (Ashe et al., 1987). Generally regarded as predators, some species of Tanypodinae feed on diatoms and detritus (Oliver, 1971). Several species probably combine both types of feeding, with reliance on diatoms and detritus when prey items are scarce (Ashe et al., 1987). The generally predatory life style is reflected in the larval mouth apparatus, which differs from other Chironomidae in having a strong development of premental structures such as ligula and paraligulae (Cranston, 1995a). The subfamily comprises 54 genera within eight recognised tribes: Anatopyniini, Clinotanypodini, Coelopyniini, Macropelopiini, Natarsiini, Pentaneurini, Procladiini and Tanypodini. Tanypodinae was erected by Skuse (1889) primarily on the basis of adult males and their monophyly is well supported, with Podonominae as its sister group (Cranston et al., 2012). Postulated internal relationships at the generic level in Tanypodinae derive from Fittkau (1962), who formally erected the tribes Anatopyniini, Macropelopiini and Pentaneurini (Fig. 1). Roback & Moss (1977) established the tribes Natarsiini and Procladiini with an unusual phenetic method to chironomid systematics. Whereas the tribe Coelopyniini was erected based on the monobasic genus Coelopynia from Australia (Roback, 1982a), Siri & Donato (2015) analysed the Macropelopiini in a phylogenetic context and corroborated its monophyly. Phylogenetic analyses based on morphological characters in different chironomid groups, show that characters whose transformations are fully consistent with each other (i.e. objective unique synapomorphies) are very rare, particularly at the species level. Parallel and convergent development of character states is frequent and subsumed as homoplasy (Cranston, 1995b) and phylogenetic studies on Chironomidae based on morphological evidence usually include a high amount of homoplasy in the dataset. Characters that empirically appear to be more important to reconstruct genealogical relationships often do not have the influence on the result as they probably should have (Ekrem, 2003). Moreover, objective character weighting schemes often have not led to stable and trustworthy hypotheses (e.g. Ekrem, 2003). According to Sæther (1989), groups of recent radiation, such as Chironomidae, can be characterised by a high number of homoplasies in the postulated phylogeny, and lead to a Hennigian bush-type topology. In this scenario, the absence of stable character alternatives (synapomorphies) in the dataset is probable the main cause for resulting trees without topological resolution (Ekrem, 2003). Here, we examine the phylogenetic signal strength of morphological characters in defining major groups in Tanypodinae, despite the apparent high degree of morphological homoplasy in the subfamily. We present the first cladistic analysis of the subfamily using 115 species belonging to 54 genera and a diverse collection of morphological characters from immatures and imagines (86). Our goals are: (i) to examine the monophyly of Tanypodinae and its tribes, (ii) to identify characters that support these clades and (iii) to assess recent nomenclatural changes proposed by Cranston & Epler (2013) and Siri & Donato (2015), regarding genera in the tribes Macropelopiini and Pentaneurini. We also evaluate previous hypothesised relationships and classifications, and propose new ones. Finally, we provide diagnoses, notes on distribution and remarks on ecology and zoogeography for Tanypodinae. Material and methods Selection of taxa and coding of character states One-hundred-and-fifteen species representing 54 tanypodine genera were selected and examined for character state coding © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 Phylogenetic relationships in the subfamily Tanypodinae 75 Fig. 1. Fittkau’s proposed classification (1962) was the first to discuss the monophyly of major groupings within Tanypodinae. (File S1). These taxa represent the eight currently recognised tanypodine tribes (Table 1). The genera Aphroteniella (Aphroteniinae) and Podonomus (Podonominae) were selected as outgroups, as previous analyses suggest a close affinity between these subfamilies and Tanypodinae (Sæther, 2000; Cranston et al., 2012). The taxa Bethbilbeckia, Cantopelopia, Gressittius, Guassutanypus, Hayesomyia, Helopelopia, Reomyia and Schineriella were treated as separate genera to investigate the validity of the nomenclatural changes proposed by Cranston & Epler (2013) and Siri & Donato (2015). Morphological data were carefully considered and coded in a matrix (File S2) using Mesquite 2.75 (Maddison & Maddison, 2004). Wherever possible the fourth-instar larva, pupa and adults of both sexes were examined for each genera, but in some instances it was necessary to rely on published species descriptions as relevant material was unavailable for examination (File S1). Interspecific variability within the genera included in the analyses was coded as polymorphisms. Continuous characters (e.g. ratios) were divided into character states based on intervals of variation (File S2). Although we attempted to keep the number of autapomorphies low, some single character states are listed for certain species to indicate differences to other terminal taxa. We selected 86 morphological traits (40 from adults, 14 from pupae and 32 from larvae) (File S3). Most of the tanypodine specimens examined for coding of the matrix are deposited in the entomological collections the Zoologische © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 Staatssammlung München (ZSM), Florida Agricultural and Mechanical University (FAMU) and The Academy of Natural Sciences of Philadelphia (ANSP). The majority of the examined species are from the Holarctic Region, but species from other regions are included to provide a worldwide representation. Morphological terminology and abbreviations essentially follows that proposed by Sæther (1980). Phylogenetic analysis Phylogenetic analyses using parsimony were conducted on a dataset of 86 characters belonging to 62 terminals of Tanypodinae, as well as two outgroup genera. The matrix was analysed under implied weighting, implemented in tnt v1.1 (Willi Hennig Society Edition) (Goloboff et al., 2008). In this method, characters are weighted during tree searches and the weights applied to each character are summed to determine the fit, such that the cladogram with maximum total character fit is chosen as the most parsimonious cladogram (MPC) (Silva et al., 2015). Character fit is determined as a function of the values of the concavity constant (k), which controls how much a character is weighted against homoplasy (Legg et al., 2013); in general, as k increases, fit decreases (Goloboff, 1993). In this study, analyses with implied weighting were run with values of k ranging from 1 to 20. All characters were treated as unordered. Tree searches 76 F. L. Silva and T. Ekrem Table 1. Tribal and generic level classification of Tanypodinae. Anatopyniini Clinotanypodini Coelopyniini Macropelopiini Anatopynia Clinotanypus Coelotanypus Naelotanypus Coelopynia Alotanypusa Natarsiini Pentaneurini Natarsia Apsectrotanypus Bilyjomyia Brundiniella Chaudhuriomyia Derotanypus Fittkauimyia Macropelopiab Paggipelopia Psectrotanypus Radotanypus Wuelkerella Ablabesmyia Amazonimyia Amnihayesomyia Arctopelopia Australopelopia Chrysopelopia Coffmania Conchapelopia Denopelopia Guttipelopia Helopelopia Hudsonimyia Krenopelopia Labrundinia Larsia Procladiini Lobomyia Meropelopia Metapelopia Monopelopiac Nilotanypus Parapentaneura Pentaneura Pentaneurella Rheopelopia Telmatopelopia Telopelopia Thienemannimyiad Trissopelopia Xenopelopia Zavrelimyiae Tanypodini Djalmabatista Tanypus Laurotanypus Lepidopelopia Procladius Saetheromyia a Including Gressitius and Guassutanypus. Macropelopia (Bethbilbeckia). c Including Monopelopia (Cantopelopia). d Including Thienemannimyia (Hayesomyia). e Including Zavrelimyia (Paramerina, Reomyia and Schineriella). b Including were conducted using ‘Traditional Search’ options: 10 000 random addition sequences, TBR branch swapping and 100 trees saved per replicate. Under ‘Settings’, the General RAM was set to 50 Mb and the maximum number of trees to be held to 10 000. Clade support was assessed using symmetric resampling, as implemented in tnt. Both the absolute frequencies and the frequency differences (or GC values) were recorded (Goloboff et al., 2003), using the same ‘Traditional Search’ options given above with the default value of 33% change probability. We also conducted phylogenetic analyses using the software paup* 4.0b10 (Swofford, 1998) through an initial heuristic search with 50 random addition replicates on equally weighed characters keeping no more than 10 000 trees of suboptimal tree lengths, followed by a heuristic search with 500 random addition replicates on characters reweighted by their rescaled consistency index. Cladograms were rooted with Aphroteniella. FigTree 1.3.1 (Rambaut, 2009) was used to display the resulting trees. Results and discussion Monophyly of Tanypodinae in Chironomidae As expected, the subfamily Tanypodinae is confirmed as monophyletic (Fig. 2: GC = 95; absolute frequencies = 94). The group is recovered in all trees resulting from parsimony analyses of the morphological character matrix under implied weights with k values ranging from 1 to 20, and also in trees from paup* analyses of the unweighted and reweighted character matrices. Five morphological traits are recognised as synapomorphies for the Tanypodinae: (i) the female gonotergite IX is a fusion of tergite IX and gonocoxite IX, forming a strap-like structure; (ii) the female gonotergite IX has very few or no setae; (iii) the larval antenna is retractile; (iv) the larval ligula; and (v) paraligulae are well developed. Monophyly and relationships among and within Tanypodinae tribes Analyses using implied weights and values of k ranging from 12 to 20 each produced a single MPC. Each analysis recovered all previously proposed Tanypodinae tribes as monophyletic assemblages. The MPC produced by the k values ranges of 1–11 presented substantial divergence in the relationships among the taxa. Therefore, the MPCs found for k = 12–20 seem to be stable, with the topology of the single most parsimonious tree for the ingroup taxa being essentially unaffected by changes in k values. The MPC for k = 12 (fit = 12.74706, length 393, CI = 0.30, RI = 0.70) is shown in Fig. 2. In this tree, the Tanypodinae forms two well-supported groups: Pentaneurini + Natarsiini and non-Pentaneurini. Cranston et al. (2012), in a molecular phylogeny for Chironomidae, also recovered a well-supported monophyletic non-Pentaneurini group, which lead the authors to doubt the validity of the existing tribal substructure in the Tanypodinae. Parsimony analyses in paup* of the unweighted character matrix resulted in 312 856 cladograms of 345 steps with a strict consensus mostly showing large polytomies although Tanypodini, Clinotanypodini and Pentaneurini were monophyletic. An heuristic search on the matrix with characters reweighted according to their rescaled consistency index, however, returned 18 trees of 354 steps where all tribes were monophyletic and the relationship between them consistent with the result from the tnt analyses (Figure S1). In our study, the tribe Pentaneurini was recovered as sister to the tribe Natarsiini, being supported by two synapomorphies: (i) larva with reduced dorsomentum and (ii) body without © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 Phylogenetic relationships in the subfamily Tanypodinae 77 Fig. 2. Phylogenetic hypothesis for Tanypodinae, result from parsimony analysis of the morphological dataset under implied weights (k = 12); symmetrical resampling support values on branches: absolute frequencies/frequency differences (GC); negative values in square brackets can be interpreted as equivalent to zero support. © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 78 F. L. Silva and T. Ekrem lateral fringe of setae. Our analyses recovered Ablabesmyia with the Thienemannimyia group (Roback, 1971; Cranston & Epler, 2013; Silva et al., 2014) as its closest relative. Within Ablabesmyia four subgenera are currently recognised: Ablabesmyia, Asayia, Karelia and Sartaia (Roback, 1985; Oliveira & Fonseca-Gessner, 2008). With its homologous complex of dorsomedial lobes, Ablabesmyia is one of the most distinctive and well-defined genera within the tribe Pentaneurini. The tribe was recovered with some internal structuring, with the Thienemannimyia group including Amnihayesomyia, Arctopelopia, Australopelopia, Coffmania, Conchapelopia, Helopelopia, Lobomyia, Meropelopia, Metapelopia, Rheopelopia, Telopelopia, Thienemannimyia (including Hayesomyia) and Xenopelopia recovered as monophyletic. The genus Xenopelopia has been considered a specialised member of Thienemannimyia group (Roback, 1971) and is the sister group of the remaining taxa within this group in our tree. The group is characterised by the following features: (i) hypopygium with complex dorsomedial lobes (aedeagal lobes). These are very characteristic for each species and usually possess lateral and basal hairs of variable shape (Silva et al., 2014). However, they never form a complex of blades and filaments as in Ablabesmyia (Roback, 1971); (ii) abdominal segments of pupa with dense shagreen of longish, upright, mostly multi-branched or bifid spinules; and (iii) pseudoradula not linked to sclerotised zone basally. The Thienemannimyia group shows well-supported relationships with other Pentaneurini genera, such as Ablabesmyia, Hudsonimyia, Larsia, Pentaneurella and Trissopelopia. Treating the Thienemannimyia group as a separate tribe would render the remaining Pentaneurini paraphyletic. Thus, we suggest keeping a more informal name for this group of genera at present. Fittkau (1962) placed Natarsia, initially based on the evidence of the immature stages, in the tribe Pentaneurini. Later, Roback (1971) argued that the adult stages in most aspects showed affinities with the tribe Macropelopiini. The adult stage of Natarsia possesses some Macropelopiini characters, including an extended costa (Fig. 3E) and the presence of postnotal setae, and some Pentaneurini features, including a reduced number of teeth on the tibial spurs, and spatulate male claws. The pupa has a triangular anal lobe, characteristic of Pentaneurini, but is very distinctive in its abdominal chaetotaxy compared to other genera in Pentaneurini or Macropelopiini (Roback & Moss, 1977). The larva of Natarsia has characters, such as a dorsomentum with teeth reduced to a sclerotised complex, considered plesiomorphic for Pentaneurini (Fittkau, 1962). Based on the mixture of Macropelopiini and Pentaneurini features in Natarsia, Roback & Moss (1977) erected the monobasic tribe Natarsiini. In our results, Natarsiini was placed as sister group to the tribe Pentaneurini, occupying a basal position in relation to Pentaneurini genera as proposed by Fittkau (1962). Natarsiini is supported by three autapomorphies in the immatures: (i) pupal segment I–VII with lateral setae I and II grouped on a lateral tubercle; (ii) larval dorsomentum with teeth reduced to a sclerotised complex; and (iii) larva with lateral hairs in partial fringe. The tribe Macropelopiini was erected by Fittkau (1962) based upon characters from adult and immature stages. Actually, according to Spies (2005), the name Macropelopiini was originally coined by Zavřel (1929), with just the ending being different in the original spelling, which required a small correction to be in compliance with the International Code of Zoological Nomenclature (ICZN, 1999). Zavřel proposed ‘Macropelopiae’ for a family-group taxon, equivalent to a supertribe by current standards (Siri & Donato, 2015). Adult males belonging to the tribe Macropelopiini can be recognised by a costa produced for a distance equal to or longer than the length of RM (Murray & Fittkau, 1989) (see Fig. 3F, G). Although the pupae are distinguished mainly by the absence of thoracic comb (Siri & Donato, 2015), larvae of Macropelopiini are separable by a short second antennal segment. Roback (1971) argued that although Procladius shows many affinities to the members in Macropelopiini, it diverges enough to be placed in its own taxon. Consequently, Macropelopiini was subdivided into two subtribes: Macropelopina and Procladina. Based on a later numerical analysis, Roback & Moss (1977) established that Procladius is sufficiently distinct to merit consideration as a separate tribe apart from Macropelopiini (see below). In an analysis derived from morphological data, Siri & Donato (2015) recovered Macropelopiini as paraphyletic group with Fittkauimyia out of the tribe and sister group of Tanypus. Cranston et al. (2012) obtained comparable results derived from molecular data. Similarly, in our study, Macropelopiini was recovered with Fittkauimyia as a separate basal branch (see next paragraph). The monophyly of the group is supported by single synapormorphy: the outer fringe of the pupal anal lobe decreasing from base to apex and ending in small spines. The genus Fittkauimyia was erected by Karunakaran (1969) based on one species from Singapore. Roback (1971) when redescribing the genus (as Parapelopia) placed it in the tribe Macropelopiini. Cranston & Epler (2013) recognised that Fittkauimyia strongly diverges from all other Macropelopiini genera (and indeed all other Tanypodinae) in the structure of the dorsomentum, ligula and mandible. Consistent with our data, Fittkauimyia occupies the most basal position within the tribe Macropelopiini. The adults can be distinguished from all other tanypodines in having a semi-globose third segment of the maxillary palp, with strong setae on inner border. The pupae are recognised by the thoracic horn and the shape of the anal lobes, whereas the larva differs from other tanypodine genera by the characteristic dorsomentum. Based on the distinctiveness of Fittkauimyia through all developmental stages, a new tribe in Tanypodinae is proposed, Fittkauimyiini. The new tribe is supported by at least nine autapomorphies: (i) male maxillary palp segment 3 semi-globose; (ii) middle leg with spatulate and apically pectinate claws; (iii) thoracic horn perforate; (iv) pupa with abdominal segments II–VII fully fringed; (v) pupal anal lobe paddle-like with convergent internal margin; (vi) larval mandible with additional small dorsal and ventral teeth; (vii) larval ligula with point of inner lateral teeth strongly curved towards middle tooth; (viii) larval dorsomentum tripartite, concave and continuous; and (ix) larval dorsomentum with lateral complex of structures, comprising an inward- and an outward-directed tooth. The postulated new tribe is consistent with relationships derived from molecular data by Cranston © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 Phylogenetic relationships in the subfamily Tanypodinae 79 Fig. 3. Wings of adult male Tanypodinae, scale bar = 500 μm. (A) Thienemannimyia fusciceps (Edwards); (B) Ablabesmyia (Karelia) nilotica (Kieffer); (C) Pentaneurella katterjokki Fittkau & Murray; (D) Zavrelimyia melanura (Meigen); (E) Natarsia punctata (Fabricius); (F) Macropelopia fittkaui Ferrarese & Ceretti; (G) Psectrotanypus varius (Fabricius); (H) Anatopynia plumipes (Fries); (I) Coelotanypus sp.; (J) Procladius (Holotanypus) sp.; (K) Lepidopelopia annulator (Goetghebuer); (L) Tanypus (Tanypus) brevipalpis (Kieffer). et al. (2012) and from a phylogenetic analysis of the tribe Macropelopiini based on morphological data by Siri & Donato (2015). Although Fittkauimyia was recovered with Procladiini + Tanypodini as its closest relative, both of these studies recognised the genus as an independent clade separate from the tribe Macropelopiini. It has been almost two centuries since the first Anatopynia Fries, 1823 was described from Europe (Johannsen, 1905). Anatopynia was a wider concept historically (e.g. Edwards, 1931), and initially most members of the Macropelopiini were © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 placed in this genus. They were later transferred to other genera. Currently, there are no valid records of the presently defined genus from outside the Palaearctic Region (Cranston & Epler, 2013). The tribe Anatopyniini was erected by Fittkau (1962), who considered Anatopynia the most primitive or generalised taxon in the subfamily Tanypodinae. Adult males belonging to this tribe are readily recognised by the distinctive tibial spurs, which lack side teeth and the dense covering of setae on the head and thorax (Murray & Fittkau, 1989). The pupae can be distinguished by their triangular scale-like shagreen 80 F. L. Silva and T. Ekrem (Fittkau & Murray, 1986). Anatopyniini is the only tribe within Tanypodinae with five-segmented larval antennae (Cranston & Epler, 2013). According to our data, Anatopyniini is recovered with Fittkauimyiini and Macropelopiini as it closest relatives. The tribe is supported by eleven autapomorphies: (i) adult male antennal ratio (AR) higher than 5.00; (ii) adult male clypeus densely setiferous; (iii) adult male thorax with dense antepronotals and (iv) dense prealars; (v) adult male wing with macrotrichia present only in apical 1∕2 (Fig. 3H); (vi) lateral teeth of tibial spur absent; (vii) adult female with a distinct collar in spermathecal duct; (viii) pupa with shagreen of large scale-like spines; (ix) larval antenna five-segmented; (x) pecten hypopharyngis with multiple rows of teeth; and (xi) dorsomentum with 13–15 teeth. According to the accepted system of subdivisions within Tanypodinae (Fittkau, 1962; Roback, 1971; Sæther, 1977), the genera Clinotanypus, Coelotanypus and the later described Naelotanypus, known only as adult male, were placed in the same tribe, the so-called Coelotanypodini. However, based on ICZN (1999) Clinotanypodini Lipina, 1928 (originally proposed as a tribe ‘Clinotanypi’) currently is the valid name, whereas Coelotanypodini Fittkau, 1962 is a junior synonym (Spies, 2005). Larvae of Clinotanypus and Coelotanypus are morphologically similar in the labrum, antenna, maxilla, hypopharynx and mental complex, as well as in the modified mandibles (Cranston & Epler, 2013). The pupae share features such as branched D-setae and the lateral fringe on the outer border or the anal lobe (Fittkau & Murray, 1986). Adult males of Clinotanypodini can be distinguished by having tarsi with chordate fourth tarsomeres, an obvious comb on the fore tibia and a double comb on the hind tibia (Roback, 1982b). Roback (1971) divided Clinotanypus into two subgenera based on imaginal characters, the worldwide Clinotanypus and the Nearctic Aponteus, which the immature stages are unknown. In our study, Clinotanypodini is recovered with Coelopyniini as its sister group. The sister-relationship between Clinotanypodini and Coelopyniini may be related to the absence of wing macrotrichia, the cordiform fourth tarsomere of all tarsi and the hind tibia with apical comb with two parallel rows of bristles, in both taxa. Clinotanypodini is supported by five synapomorphies: (i) R1 and R4+5 widely separated; (ii) larval cephalic capsule apically narrow; (iii) larva with bladder-like labral sensory organs distinct at anterior margin of cephalic capsule; (iv) mandible with pecten mandibularis; and (v) ligula with 6–7 teeth. The adult stage of Coelopynia possesses a combination of Macropelopiini wing structure and Clinotanypodini leg characters, and was initially being placed within the latter tribe. Roback (1982a) noticed, however, that the tibial spurs with 3–4 lateral teeth more closely resemble those of some Pentaneurini and Tanypodini. Coelopynia has a unique pupal respiratory organ, with some superficial resemblance to those of Anatopynia and Procladius. The anal lobes show some similarity to those of Coelotanypus but the dorsal, lateral and ventral abdominal setae diverge from those found in the tribe Clinotanypodini (Roback, 1982a). The larval labrum lacks the flaps present in members of the tribes Clinotanypodini, Procladiini, Tanypodini and Macropelopiini as well as the numerous filaments seen in the labrum in Pentaneurini. The maxillary palp is elongate, resembling that of the Clinotanypodini, but the lacinia lacks the filaments present in the other mentioned tribes. Based on the singularity of the features seen in Coelopynia, Roback (1982a) proposed the monobasic tribe, Coelopyniini. In our study, the tribe is recovered with Clinotanypodini as its closest relative. The tribe is supported by four autapomorphies in the larva: (i) cephalic capsule with a distinct anteroventral gap; (ii) lacinia without fringes; (iii) labrum without flaps; and (iv) mandible exceptionally long and slender. As mentioned, Roback & Moss (1977) recognised that Procladius was more distinct from the other genera within Macropelopiini than either of the previous placements would indicate. Accordingly, they proposed the tribe Procladiini, including the genera Procladius and Psilotanypus (the latter currently a subgenus of Procladius). Later, Roback (1980) tentatively proposed the subtribes Djalmabatistina and Procladiina, which seems to have been overlooked by peers and not justified with the inclusion of Laurotanypus, Lepidopelopia and Saetheromyia in Procladiini (see below). Within Procladius, three subgenera are currently recognised: Holotanypus, Procladius and Psilotanypus (Roback, 1982c). The type species of Procladius, described from Australia, does not fall into either of the subgenera, which otherwise are separable in all stages (Cranston & Epler, 2013). In our study, the tribe Procladiini was recovered as monophyletic, including the genera Laurotanypus and Lepidopelopia, known only as adult males, which so far have had a doubtful position within this tribe (Sæther & Andersen, 2000; Ashe & O’Connor, 2009). The monophyly is supported by a single synapormorphy: the distance between MCu and FCu 1/2 as long as Cu1 (Fig. 3J). Laurotanypus was erected by Oliveira et al. (1992) for Laurotanypus travassosi based on one species from Brazil. This species resembles some species placed in Procladius (Psilotanypus) Kieffer. According to Spies et al. (2009), Laurotanypus should be compared with P. (Psilotanypus) stroudi (Roback, 1982d), in order to investigate possible synonymy. Moreover, P. (Psilotanypus) etatus (Roback, 1982d) possesses the adult scutal tubercle proposed as diagnostic for Laurotanypus, but its wing is darkened only around RM. Therefore, generic placement of these species remains tentative until their immature stages are discovered (Spies et al., 2009). Furthermore, Lepidopelopia together with Saetheromyia differs from other members of Procladiini in having a more cylindrical gonocoxite and a long tapering gonostylus without basal heel or setiferous inner lobe. These plesiomorphic features suggest that these genera constitute the sister group to the remaining genera in the tribe. Although in our analysis Procladius is the sister group to the remaining Procladiini, this may be attributed to the lack of character information from the unknown immatures of Laurotanypus, Lepidopelopia and partially from Saetheromyia. Procladiini was reconstructed with Tanypodini as its closest relative, refuting a sister-relationship between Macropelopiini and Procladiini previously proposed by Fittkau (1962) and Roback & Moss (1977). The proximity among members of Procladiini and Tanypodini was also indicated by Sæther & Andersen (2000). © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 Phylogenetic relationships in the subfamily Tanypodinae The tribe Tanypodini comprises the single genus Tanypus, which is morphologically very distinct from other tanypodine genera. The shape of the scutal tubercle and the pupal thoracic horn, the structure of the larval mentum, the shape of the mandible and maxilla, and the reduced pecten hypopharyngis separate Tanypus from all other genera. Tanypus possesses well-developed dorsomental plates that are joined medially, a feature seen in the Podonominae–Aphroteniinae line and most of the other Chironomidae. Based on this feature, Roback (1976) postulated that within Tanypodinae, the mentum of Tanypus represents the plesiomorphic condition, whereas in Pentaneurini, it represents the apomorphic one. Roback (1971) divided Tanypus into two subgenera based on imaginal characters, the worldwide Tanypus and the Nearctic Apelopia. According to our data, Tanypodini is recovered with Procladiini as it closest relative. The proximity among members of these tribes was also indicated by Cranston et al. (2012) and Siri & Donato (2015). The tribe is supported by four autapomorphies: (i) distance between MCu and FCu less than 1∕3 as long as Cu1 (Fig. 3L); (ii) pupal anal lobe rectangular-shaped; (iii) pseudoradula; and (iv) lateral vesicles absent. Monophyly of genera All terminal taxa included in the analyses are assumed, a priori, to be valid genera following the definitions given in two of the main monographs (Fittkau, 1962; Roback, 1971). We also assumed that each species chosen for character coding is congeneric with the type species of the genus in which it is placed. However, the taxonomic status of some of these genera was recently reviewed and therefore deserves to be discussed in light of our results. The taxonomic synonyms reviewed or proposed here that result in nomenclatural changes are listed as new combinations in the Appendix. The genus Helopelopia, treated originally as a subgenus of Conchapelopia, was given generic status based on immature characters by Fittkau & Roback (1983). However, Cranston & Epler (2013) suggested that Helopelopia should be treated as a subgenus of Conchapelopia, because a wider range of larval characters indicate that the generic diagnosis of Conchapelopia should be broadened. Helopelopia is recovered with Conchapelopia as its closest relative in our analyses, at least not refuting the subgeneric status of this taxon. According to B. Bilyj (personal communication), Helopelopia can be distinguished in the larval stage by the following combination of characters: the basal maxillary palp is longer than 65 μm, with two segments and an apical sensillum b; the mandible has a small but distinctly visible accessory tooth, and the pseudoradula is narrow with parallel sides reaching the base. In Conchapelopia the sides of the pseudoradula tapers to a wider base. In Helopelopia pupae, the thoracic horn is rather distinctly elongated with a narrow tubular atrium that bifurcates apically into two short, broad diverticula supporting a small, rounded to oval plastron. In Conchapelopia, the plastron is generally larger occupying about a 1∕3 of the lumen. It may be smaller as in C. pallens (=gonoides), but differs in having more, longer and narrow © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 81 apical diverticula (B. Bilyj, personal communication). The adult male of Helopelopia possesses a very distinctive hypopygium, bearing a modified gonostylus, which also warrants its separation from Conchapelopia. Moreover, the basomedial lobes between the gonocoxites are also different in having a separate ventral and dorsal filamentous lobes, whereas in Conchapelopia the two lobes are fused into one (B. Bilyj, personal communication). We believe that these distinct differences in all life stages are convincing arguments to accept Helopelopia and Conchapelopia as separate genera, at least until a species-level phylogeny is available for these taxa. Similarly, the genus Hayesomyia was initially described as a subgenus of Thienemannimyia (Roback, 1971). Based on diverging specimens from Ireland, Murray & Fittkau (1985) erected Hayesomyia as an independent genus, a status disputed by Cranston & Epler (2013), who argued that Hayesomyia should be treated as a subgenus in Thienemannimyia, due to the range in character variation present in the immature stages of this genus. Hayesomyia larvae are extremely difficult to distinguish from those of the Thienemannimyia group. Epler (2001) suggested that Hayesomyia may be separated from Thienemannimyia by its lower antennal ratio (AR < 5.0). However, this character falls within the average antennal ratio recorded for Thienemannimyia norena (AR 4.7, Roback, 1981). Hayesomyia pupae can be distinguished from Thienemannimyia by having an internally well-developed thoracic horn, holding a distinctive aeropyle. However, Roback (1981) described pupae of T. norena with a small, but distinctive aeropyle, suggesting that this character may not hold as diagnostic for Hayesomyia. The only character in the adult male that distinguishes Hayesomyia and Thienemannimyia is the presence of a scutal tubercle in Hayesomyia. Nonetheless, Kobayashi (2003) recognised difficulty in observing this structure, as he was able to see the scutal tubercle in only four out of twenty-two examined specimens of H. tripunctata. Thus, this trait appears unreliable in distinguishing the two genera. In our study, Hayesomyia is sister to Thienemannimyia, thus not refuting the classification proposed by Cranston & Epler (2013). We recovered Paramerina, Reomyia, Schineriella and Zavrelimyia as a monophyletic group with Schineriella as sister to Paramerina + Reomyia + Zavrelimyia in the tnt analyses and without internal structure in the paup* analyses. Roback (1986) described the genus Reomyia for western North American Zavrelimyia wartinbei (Roback, 1984). The pupa of Reomyia was included in the Holarctic key to Tanypodinae pupae as Tanypodinae genus III (Fittkau & Murray, 1986). According to Epler (2001), who informally described the larva, Reomyia is very similar to Zavrelimyia and may not be separable from that genus in the larval stage. With regard to the pupa, Reomyia can be distinguished by a subtle difference in the D setae, which are thin and elongate. In Zavrelimyia these setae are shorter and rounded (Epler, 2001). In the adult male, Reomyia specimens differs from Zavrelimyia only in the presence of a scutal tubercle. However, at least one species of Zavrelimyia has a scutal tubercle (B. Bilyj, personal communication). Epler (2001) suggested the possibility of reuniting Reomyia and Zavrelimyia. This status was adopted by Cranston & Epler (2013). 82 F. L. Silva and T. Ekrem Likewise, Schineriella was erected for Tanypus schineri by Murray & Fittkau (1988) with descriptions and generic diagnosis for the adult male and pupa. Although technically undescribed, the larva of Schineriella is known from the Netherlands and resembles those of Paramerina and Zavrelimyia. Schineriella differs from Paramerina essentially in the presence of an undivided maxillary palp. However, P. togavicea, from Japan, has a single segmented maxillary palp (see below), thus bridging the gap between the two genera. The granulose area of muscle attachment at the base of the ligula is narrow and nearly oval in Schineriella, different from the usually broader band in Zavrelimyia that reaches the full basal width of the ligula. However, some species of Zavrelimyia possess a narrower granulose area, erasing this character as diagnostic for Schineriella. Schineriella pupae may be distinguished from Zavrelimyia by having a wide thoracic horn against the narrower and elongate thoracic horn possessed by the latter. Nonetheless, a few morphotypes of Zavrelimyia have a larger thoracic horn, suggesting also that this character is unsuitable for separation at the generic level. In the adult male, there may be a subtle difference in the wing venation as Schineriella is considered not to possess R3 , whereas Zavrelimyia has R3 fading out distally. However, at least one species of Zavrelimyia has this vein reduced or absent (Fittkau, 1962). Cranston & Epler (2013) suggested that Schineriella should be considered a subgenus of Zavrelimyia. Paramerina was erected for Palearctic Tanypus cingulatus by Fittkau (1962). The placement of Paramerina as a subgenus of Zavrelimyia was first proposed by Hamilton et al. (1969), but was overlooked by peers and both genera were kept separate until Cranston & Epler (2013) discussed the possibility of uniting both genera. Paramerina differs from Zavrelimyia only in the segmented basal maxillary palp of the larva. However, Niitsuma et al. (2011) reared a larva of a new Paramerina species from Japan with an undivided basal palp segment, indicating that this diagnostic character does not hold. Coffman & Ferrington (1996) and Ferrington et al. (2008) also indicated that both maxillary palp forms were present in Paramerina. Unfortunately, the single segmented form has not been confirmed associated with adults through rearing (only pupae). According to B. Bilyj (personal communication), another distinguishing trait in the larvae is the shape of the apical claws on the posterior parapods. In Zavrelimyia one claw is bifid and in Paramerina all claws have previously been regarded as simple (e.g. Fittkau & Roback, 1983) even if Roback (1972) already had described bifid claws in P. smithae. Epler (2001) described the larva of Z. bifasciata without the typical bifid claw, and more recently Niitsuma et al. (2011) found that the larva of P. togavicea with the single segmented palp also had a bifid apical claw, thus looked like a typical Zavrelimyia. Moreover, some species in both genera have similar arrangements, in relation to the ventral pore, of the ventral cephalic setae, which usually are distinct at the generic level (Cranston & Epler, 2013). In the pupa, only two characters have been used for separation of the two genera: the presence or absence of spinules on the inner margin of the anal lobe and the relative length of the male genital sacs versus the anal lobes. In Zavrelimyia the inner margins of anal lobes are smooth and the male genital sacs are shorter than the anal lobes. Again, P. smithae possess a few spinules on the inner margin of the anal lobes and the male genital sacs end near the tips of the anal lobes, thus providing a gradual transition in this character (B. Bilyj, personal communication). In the adult male, the structures used for separating the two genera are: the presence or absence of large setae on TIX and the shape of the tibial spur on the foreleg. In Paramerina, the TIX is considered not to have large setae, except for P. hanseni described by Roback (1971: 274), which possesses three setae on TIX. The shape of the fore tibial spur in P. smithae appears intermediate between the typical elongate shape and the lyrate shape present in Zavrelimyia (B. Bilyj, personal communication). Based on the results from our phylogenetic analyses and the above morphological comparison, we believe that the concept of Zavrelimyia should be broadened and include Paramerina, Reomyia and Schineriella as subgenera. Cantopelopia was described by Roback (1971) for Cantopelopia gesta with descriptions and a generic diagnosis for the adult male. Epler & Janetzky (1999) referred to an ‘apparently undescribed, species of Monopelopia’, which was later recognised as the larva of C. gesta (Epler, 2001). Because the larvae of C. gesta are very similar to those of some Monopelopia, particularly concerning the coloration of claws on posterior parapods, and Cantopelopia had been assumed more closely related to Paramerina, this caused an interesting situation in understanding the relationship between these genera. Jacobsen (2008) distinguished the pupa of Cantopelopia based on the absence of a subterminal spine in the thoracic horn. However, except for Monopelopia caraguata, nearly all species of Monopelopia have a thoracic horn not holding a subterminal spine. The differences between adult males of Cantopelopia and Monopelopia are the two tibial spurs on middle and hind legs of Cantopelopia (Monopelopia has only a single tibial spur) and the apically wide gonostylus of Cantopelopia compared to the apically attenuate gonostylus of all described Monopelopia species (Epler, 2001). Based on the aforementioned considerations, Cranston & Epler (2013) treated Cantopelopia as a subgenus of Monopelopia. In our study, Cantopelopia was recovered with Nilotanypus + Monopelopia as its closest relative. However, the sister-relationship between Nilotanypus and Monopelopia may be attributed to the absence of R2+3 in both genera, a character possibly related to small size. A phylogenetic analysis with this feature deactivated recovered Cantopelopia as sister to Monopelopia, agreeing with the nomenclatural action proposed by Cranston & Epler (2013). Fittkau & Murray (1988) described Bethbilbeckia for southeastern Nearctic Bethbilbeckia floridensis with descriptions and generic diagnosis for the adult male, pupa and larva. However, based on the resemblance with Macropelopia in all life stages, Cranston & Epler (2013) treated Bethbilbeckia as a subgenus of the latter. Bethbilbeckia larvae resemble those of Macropelopia. According to Fittkau & Murray (1988), the larva of this genus could be distinguished from Macropelopia by the five-segmented antenna and the ring organ in bottom 1∕3 of maxillary palp. However, Bethbilbeckia shares the location of the ring organ with Macropelopia and actually has four antennal segments. According to Watson (2010), there is an extended © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 Phylogenetic relationships in the subfamily Tanypodinae membranous area between the second and third antennal segments, which apparently has been mistaken for a third segment by Fittkau & Murray (1988). Thus, based on larval morphology, there is no reason for keeping these genera separated. The only trait in the pupae that has been used to separate the two genera is the D1 setae, which arise from prominent tubercles in Bethbilbeckia, larger than those usually present in Macropelopia. Adult males of Bethbilbeckia can be distinguished from those of Macropelopia only by subtle differences that can be attributed to the smaller body size of Bethbilbeckia compared to Macropelopia. For instance, the uniserial arrangement of the temporal (inner and outer vertical and postorbital) setae in Bethbilbeckia may be related to its smaller size. Watson (2010), in a review of Bethbilbeckia, argued that the lack of several characters considered diagnostic for Macropelopia, such as the fore tibial comb of the adult male, the ventral swelling of the male gonostylus and the apically expanded granulation of the larval pseudoradula (Fittkau & Roback, 1983; Murray & Fittkau, 1989), should be enough to retain both genera. In our analyses, Bethbilbeckia was recovered with Macropelopia as its closest relative, thus not contesting Cranston & Epler’s classification (2013). Nevertheless, the most appropriate classification of these genera/subgenera remains uncertain and need to be substantiated by further studies using molecular data. Guassutanypus, described by Roque & Trivinho-Strixino (2003) for southeastern Brazil Guassutanypus oliveirai, was synonymised with Alotanypus by Cranston & Epler (2013). According to Roque & Trivinho-Strixino (2003), larvae of Guassutanypus can be distinguished from Alotanypus by the higher number of dorsomental teeth and the ring organ situated between proximal and middle 1∕2 of basal segment of maxillary palp. However, considering the morphological diversity in Alotanypus, these traits may not be reliable to separate the latter from Guassutanypus. Moreover, both genera have similar arrangements of the ventral cephalic setae, particularly in relation to the ventral pore, which are commonly different at the genus level (Roque & Trivinho-Strixino, 2003; Cranston & Epler, 2013). Guassutanypus pupae may differ from those of Alotanypus by the unique thoracic horn, which has a characteristic convolute atrium. However, Gressittius, recently considered as a junior synonym of Alotanypus (see below), has similar thoracic horn, and also A. kuroberobustus from Japan (Niitsuma, 2005). Thus, this character cannot be regarded as diagnostic for Guassutanypus. Adult males of Guassutanypus may be distinguished from Alotanypus by the absence of preepisternal setae (Roque & Trivinho-Strixino, 2003). However, these setae are also absent in A. dalyupensis and A. kuroberobustus (Siri & Donato, 2015). Similarly, Gressittius described by Sublette & Wirth (1980) based on specimens from New Zealand was also placed in Alotanypus as a result of phylogenetic analyses of the tribe Macropelopiini (Siri & Donato, 2015). The genera can be discriminated in the immature stages by differences in their larval antennal ratio (AR 4.1) and pupal thoracic horn (with a convolute atrium). The wing pattern was used to separate Gressittius from the remaining Tanypodinae in the adult males. According to Siri & Donato (2015), the characters used to define Gressittius overlap with those of Alotanypus and justify a synonymy © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 83 of the two genera. In our analyses, Alotanypus, Gressittius and Guassutanypus were recovered as a monophyletic group, supporting Gressittius and Guassutanypus as junior synonyms of Alotanypus. Chaudhuriomyia, described by Paul & Mazumdar (2015) for eastern Himalaya Chaudhuriomyia binduensis, is very similar to Apsectrotanypus and may not be readily separable from that genus. According to Paul & Mazumdar (2015), larvae of Chaudhuriomyia can be distinguished from Apsectrotanypus by the larger pore size in the S7 cephalic seta, the absence of an intersegmental band between antennal segment 2 and 3, the unequal mental teeth, the slightly inwardly bent inner tooth of the ligula and the long procercus. Except for the absence of an intersegmental band in the antenna, we believe that the remaining characters are too subtle and not reliable to retain these taxa as separate genera. Chaudhuriomyia pupae seem to be the stage with more consistent distinguishing traits. The D1 seta on tergites II–VII arise from a large sclerotised tubercle and the presence of a small aeropyle permit the distinction of the genus from Apsectrotanypus. For the adult males, there are no reliable distinguishing features between Chaudhuriomyia and Apsectrotanypus. Paul & Mazumdar (2015) indicate that Chaudhuriomyia may be distinguished by having spatulate claws and tergite IX with a few setae against the pointed claws and the hairy tergite IX possessed by species in Apsectrotanypus. However, we question if these traits are enough to justify separation at the genus level. In the results from our tnt analyses using implied weights, Chaudhuriomyia and Apsectrotanypus do not group as closest relatives. However, results from the paup* analyses on the reweighted character matrix group these two genera as sisters, and combined they are related to Bilyjomyia + Bethbillbeckia + Macropelopia (Figure S1). The proximity among these genera was also indicated by Paul & Mazumdar (2015). Although, we refrain from synonymising Chaudhuriomyia with Apsectrotanypus at present, we consider its genus-level status doubtful. The most suitable classification of this taxon remains uncertain and need to be corroborated with additional studies using molecular and morphological data at the species level. Taxonomy Fittkauimyiini trib.n. http://zoobank.org/urn:lsid:zoobank.org:act:14F36F90-4AEE4CC3-A740-181BA2281288 Type genus. Fittkauimyia Karunakaran 1969: 75, here designated. Composition. Fittkauimyia carranquensis Dantas & Hamada, 2013; F. crypta Serrano & Nolte, 1996; F. disparipes Karunakaran, 1969; F. mayumiae Dantas & Hamada, 2013; F. nipponica Ueno, Takamura & Nakagawa, 2005; F. olivacea Niitsuma, 2004; F. petersi (Freeman, 1955); F. serta (Roback, 1971). Diagnosis. Adult: Medium-sized, wing length about 2.0 mm. Head with verticals irregularly uniserial, postorbitals uniserial. 84 F. L. Silva and T. Ekrem Eyes not iridescent. Palp segment 3 semi-globose, with strong setae on inner border. Antenna with apical flagellomere offset, 4× as long as broad. Antennal ratio (AR) 1.4–2.3. Thorax with antepronotum reduced, without V-shaped emargination; with 1–2 lateral antepronotals. Acrostichals few; dorsocentrals 10–12, uniserial; anepisternals and dorsal postnota1 present and distinct; scutal tubercle present. Wing membrane with dense or scattered macrotrichia, unmarked. Costa strongly produced; R2+3 present and distinct; RM slightly beyond MCu; FCu before MCu. Legs with tibial spurs elongate, cone-shaped. Claws on fore and hind legs normal, sharp; claws on middle leg spatulate and apically pectinate. Pulvilli present. Male hypopygium tergite IX without setae. Male gonocoxite elongate, subcylindrical and at least 2× as long as broad, with dorsal lobe-like setiferous swellings. Male gonostylus simple, slightly tapered at apex. Female genitalia with three globular seminal capsules; spermathecal ducts separate for their complete length. Pupa: Small–medium sized. Cephalothorax with thoracic horn large, flattened, gradually expanding to the apical 2∕3, narrowing towards apex; outer border convex, inner border straight; external membrane with dispersed, pointed spines. Horn sac filling the entire lumen, perforate. Plastron plate oval, 1.5× as wide as high. Thoracic comb and basal lobe absent. Abdominal tergite I with indistinct or absent scar. Shagreen spinules short, solitary and pointed distally. Segment I with 3 D and 2 L setae, all simple; D1 on segments III–VII robust and pointed, arising from sclerotized tubercles; D2 and D3 on segments III–IV as long as the segment and hooked apically. Segment I without lateral fringe. Segments II–VII fringed; S VIII with five LS setae. Anal lobe about 2× as long as broad; outer border convex, fringed beyond the distal macrosetae; inner border fringed in the apical 1∕ ; anal lobe points converging and overlapping distally. Anal 2 macrosetae without adhesive sheaths. Larva: Medium-sized. Head oval; cephalic index about 0.75. Antennal ratio about 5.4. Mandible moderately curved, gradually narrowed towards apex; apical tooth indistinctly delimited; mola nonprotruding, with separate sets of numerous accessory teeth. Maxilla with basal segment apically narrowed, with ring organ at base of distal 1∕ . Dorsomentum with continuous, concave-arched, toothed 3 plate weakly subdivided into median and lateral sections; median section with protruding, apically notched middle tooth and three rounded teeth on each side; lateral section with outwardly-curved tooth as large as third lateral in median section, and larger, inwardly-curved tooth with pointed apex. Pseudoradula of uniform width in apical part, partly with coarse granulation. Ligula with five teeth, about 2× as long as apical width, basal 1∕3 with strongly narrowed sclerotized part; row of teeth weakly concave; middle tooth shorter and more slender than others; point of inner tooth strongly inclined towards middle tooth. Paraligula slender, with a longer inner point and 2–3 short outer points. Pecten hypopharyngis with teeth of inner 1∕2 subequal in length. Body with dense lateral fringe of swim-setae; anal tubules conical, half as long as posterior parapod. Procercus with eight setae. All claws of posterior parapod light, many large claws with fine spinules on inner side. Diagnoses, distribution and ecological notes Although used in many published papers, most tribes of subfamily Tanypodinae have never been formally diagnosed. Exceptions are Coelopyniini and Macropelopiini described by Roback (1982a) and Siri & Donato (2015), respectively. Here we give diagnostic characters for all Tanypodinae tribes, including the newly erected Fittkauimyiini. The diagnostic features presented here are not necessarily synapomorphies, but characters or combination of characters that are characteristic. Distribution and ecological notes are also provided. Anatopyniini Anatopyniini is monobasic with adult males readily recognised by having an antennal ratio (AR) higher than 5.00. The dense covering of setae on head and thorax, and a wing with macrotrichia present only in apical 1∕2 (Fig. 3H) is characteristic to the tribe. The distinctive tibial spur without lateral teeth is a further feature that allows easy recognition of members of Anatopyniini. Pupae can be distinguished by the unique thoracic horn, the triangular scale-like shagreen and the shape of the anal lobe that is unlike any other tribe in the Tanypodinae. Anatopynia is the only genus of Tanypodinae with five-segmented larval antennae. The larvae are large and resemble those of Macropelopiini. These diverge particularly in having multiple rows of teeth on pecten hypopharyngis, a large number of dorsomental teeth and mandible with two accessory teeth next to each other (Cranston & Epler, 2013). Larvae of Anatopyniini live in the littoral zones of small lakes and ponds. Anatopynia plumipes is the only species known with certainty. It is recorded in the western Palaearctic from Central to Northern Europe. Clinotanypodini Adult males of Clinotanypodini are readily recognised by the presence of anepisternals, tarsi with chordate fourth tarsomeres, hind tibia with apical comb of bristles in two parallel rows. The wing is without macrotrichia, but the membrane is densely covered with microtrichia; the vein R2 is not connected to R3 , ending in the wing membrane (Fig. 3I). Tarsal claws are pointed. Pupae of this tribe can be distinguished by the abdominal segments having forked dorsal and Sa setae, paddle-like anal lobe with straight internal margin, without apical spines. An anal lobe with lateral fringe of setae on the outer margins is a further feature that allows easily recognition of pupae in this tribe. Larvae of Clinotanypodini are characterised by dorsomental teeth arranged in longitudinal rows on the M appendage, ligula with 6–7 teeth, and well-developed lateral fringe of setae on the larval body. The group has worldwide distribution with Naelotanypus so far being recorded only in Colombia and Surinam. Larvae occur in or on bottom sediments in marshes, ponds, lakes, artificial impoundments and in the slower portions of streams and rivers. They prefer soft sediments and can be found in waters that have been organically enriched. In Asia, larvae of Clinotanypus may be abundant in rice fields, ditches and pools in drying rivers (Cranston & Epler, 2013). © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 Phylogenetic relationships in the subfamily Tanypodinae Coelopyniini Coelopynia is the only genus of the tribe Coelopyniini. Adult males of this genus can be distinguished by the absent scutal tubercle, the wing with sessile Cu1 and complete R2 + 3 , and by having elongate tibial spurs with 3–4 lateral teeth. Tarsi with chordate fourth tarsomeres and lacking tibial comb on the foreleg is characteristic for the tribe. The pupae can be recognised by the typical thoracic horn without a thoracic comb and an abdominal scar. An anal lobe with a lateral fringe of setae on inner and outer margins is distinctive for pupae in Coelopyniini. The larvae are characterised by the wedge-shaped cephalic capsule with a distinct anteroventral gap, a labrum without flaps, nonfringed lacinia, five-toothed ligula, pectinate paraligulae and an exceptionally long and slender mandible. Larvae of Coelopyniini have so far only been occasionally recorded in Australia, usually in deep lakes, billabongs and in both lentic and lotic waters. Fittkauimyiini Adult males of Fittkauimyiini can be recognised by the unique semi-globose third segment of the maxillary palp and the distinctive shape of the claws on the middle leg. Pupae are readily recognised by the perforate thoracic horn, the fully fringed abdominal segments II–VII and the paddle-like anal lobe with convergent internal margin. The larvae of Fittkauimyiini differ strongly from all other tanypodines in the arrangement of the dorsomental teeth in a concave arc and the shape of the mandible, ligula and dorsomentum. There are no other tanypodine larvae where the mandible has additional small dorsal and ventral teeth, or the ligula has curved inner lateral teeth. The tripartite dorsomentum with lateral complex of structures, comprising an inwardly and an outwardly directed tooth is distinctive for the tribe. Larvae of Fittkauimyiini live in rivers and in the littoral region of lakes, generally in tropical and subtropical regions throughout the world (Cranston & Epler, 2013). The group has also been collected from decayed wood and leaves accumulated on the bottom of a small stream in the Amazon rainforest (Dantas & Hamada, 2013). The larvae are ‘sit-and-wait’ predators, with a diet that includes oligochaetes (Serrano & Nolte, 1996). 85 tribe. The larvae of Macropelopiini may be characterised by an elongate mandible, a short second antennal segment (except for Alotanypus antarcticus), dorsomental teeth in well-developed transverse plates, and a body with a developed lateral fringe of setae. The tribe Macropelopiini presents worldwide distribution, with larvae commonly being found in depositional substrates of springs, brooks, bugs, cool seeps, headwater and slow-flowing rivers. Larvae of Alotanypus seem to tolerate a wide range of conditions, including very acidic waters (Cranston & Epler, 2013). Natarsiini The placement of Natarsia, with its combination of Macropelopiini and Pentaneurini characters, has been challenging to many chironomid taxonomists. Adult males belonging to this tribe can be recognised by noniridescent eyes, uniserial postoculars, fore tibia with apical comb of bristles, spatulate claws and bare tergite IX. The pupae of Natarsiini resemble those of Krenopelopia in the thoracic horn. However, the paired L setae with L1 much longer than L2 , the reduced LS setae on segment VIII and the very short anal macrosetae in Natarsiini clearly separate pupae from those of Krenopelopia (Fittkau & Murray, 1986). Additionally, lateral setae I and II group on a tubercle on segments I–VII and is distinctive for the tribe. The larvae of Natarsiini are easily recognised by having the ring organ in apical 1∕3 of the basal segment of the maxillary palp, a dorsomentum with teeth reduced to a sclerotised complex, a thick and short antenna (about 1∕3 length of head and twice the length of the mandible), a large molar tooth of the mandible, a pseudoradula with fine granules not in longitudinal rows and with partial fringe of lateral hair on the body. Natarsiini are mainly Holarctic in distribution, but has relatively recently been recorded in the Oriental Region in China (Cheng & Wang, 2006). The tribe was found in more than 70% of 18 lakes studied in the Yucatan Peninsula, Mexico (Vinogradova & Riss, 2007). Larvae inhabit small streams, springs, marshes and the littoral zone of montane or northern lakes. Most of them exhibit hygropetric behaviour in small standing waters including Sphagnum bogs (Vallenduuk & Moller-Pillot, 2007). Natarsia larvae were also reported from ‘wet soils’ in the Czech Republic by Frouz & Matěna (2000). They may tolerate organic and toxic discharges, particularly sewage (Hudson et al., 1990). Macropelopiini Pentaneurini Adult males of tribe Macropelopiini can be distinguished by the presence of anepisternals, a costa produced beyond R4+5 by a distance at least as long as RM (Fig. 3F, G), and a tergite IX with distal margin straight to weakly concave. Pupae belonging to the tribe can be recognised by absence of thoracic comb, a large dorsal setae on the abdominal segments, arising from exceptionally prominent tubercles and a paddle-like anal lobe with divergent internal margin. An anal lobe with a decreasing outer fringe from base to apex ending in small spines is an additional trait that allows recognition pupae in this © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 This is the largest tribe in Tanypodinae, comprising 30 genera. Adult males of Pentaneurini stand apart from other tanypodines by the absence of anepisternals and postnotals, a costa ending at R4+5 , or at most produced for a distance shorter than twice the length of RM (Fig. 3A–D) (except for Australopelopia). The pupae are easily distinguished by the presence of an elongate scar on tergite I, and a triangular anal lobe without a fringe of lateral setae. An anal lobe with apical chitinised spines is a further character that permits easy recognition of members in 86 F. L. Silva and T. Ekrem this tribe. Larvae of Pentaneurini are recognised by an elongate cephalic capsule, a dorsomentum without well-developed teeth and a body without lateral fringe of setae. The group has worldwide distribution, being found in a wide range of environments, such as hot springs, herbaceous marshes, Sphagnum bogs, ditches, ponds, the littoral zone of lakes and the slower moving portions of streams and rivers. Larvae of Ablabesmyia janta and A. tucuxi are known to be symbiotic on Mollusca (Unionidae: Quadrula) (Roback, 1982e) and Porifera (Metaniidae: Drulia) (Fusari et al., 2012). Moreover, Monopelopia larvae have been reported to occur in bromeliad phytotelmata (Epler & Janetzky, 1999). Hudsonimyia larvae are hygropetric, living in shallow-water streams flowing slowly over granite outcrops covered with algae, moss and detritus (Roback, 1979; Caldwell & Soponis, 1982). Procladiini Adult males of Procladiini can be easily recognised by the much longer, at least 1∕2 as long as Cu1 , petiole or stalk on wing vein Cu (Fig. 3J, K). The pupae belonging to Procladiini are very distinct on the genus level, being recognised at the tribal level only by the quadrate to semicircular anal lobe. The larvae of Procladiini are distinguished by the rotund head capsule, well-developed dorsomental tooth plates, mandible with large basal tooth, ligula darkened over the distal 1∕2, pectinate paraligulae and body with well-developed lateral fringe of setae. The tribe has worldwide distribution with Procladius prevalent in the Holarctic Region. Djalmabatista occurs in the New World and in the Afrotropical and Australasian regions. The genus has not been reported in the western Palaearctic, but pupal exuviae have been found in a river in the transition zone between the Palaearctic and southwest China (Cranston & Epler, 2013). Saetheromyia has only been recorded from Japan, whereas Laurotanypus has a Neotropical distribution. Lepidopelopia is endemic to the Afrotropical Region. Larvae occur in the bottom sediments of bogs, ponds, lakes, streams and rivers. A few species also colonize the profundal zone of large, deep lakes. Procladius can be found in heavily polluted conditions, where larvae are subject to numerous deformities (Warwick, 1990). Tanypodini Tanypus is the only genus in the Tanypodini. Adult males of this genus are readily recognised by having an ovoid scutal tubercle. A petiole or stalk between MCu and FCu that is < 1∕3 as long as Cu1 is only found in Tanypus (Fig. 3L). The pupae of Tanypus present a high degree of variation characteristic of an apomorphic group (Fittkau & Murray, 1986). They can be characterised by the bulbous thoracic horn and the reduced, rectangular anal lobe. The larvae of Tanypodini stand apart from other tanypodines by the singular structure of the mentum, the absence of a pseudoradula and lateral vesicles, the shape of the mandible (the median tooth seems small compared to the lateral teeth) and maxilla, and the reduced pecten hypopharyngis. Tanypus is species-rich and global in distribution; larvae live in or on soft sediments in standing and slowly flowing waters, especially in temperate to warm regions, where they can tolerate high nutrient content and salinities (Cranston & Epler, 2013). Representatives of some species of Tanypus have been reported from hot springs at temperatures up to 44.5∘ C (Thienemann, 1954). The larvae of this group prey on and suck out the body fluids on the soft parts of other chironomid larvae (the head capsule is not engulfed as in many other Tanypodinae), although they may also ingest plant material and algae (Roback, 1976; Epler, 2001). Dietary studies indicate high detritus content (Silva et al., 2008), even though predation undoubtedly occurs. Remarks on zoogeography Chironomidae are a cosmopolitan family of nonbiting midges occurring in all zoogeographical regions of the world including Antarctica (Ashe & O’Connor, 2009). It is the most widely distributed group of insects, having adapted to nearly every type of aquatic or semiaquatic environment. The range of conditions under which chironomids occur is more extensive than of any other family of aquatic insects (Ferrington et al., 2008). Representatives of Tanypodinae have been recorded from nearly all zoogeographical regions, with the majority of its genera occurring in more than on region. The wide distributional amplitude displayed by species of Tanypodinae is related to the very extensive set of morphological, physiological and behavioural adaptions found among the members of this subfamily. Currently, Tanypodinae has 54 recognisable genera, 8 of which are further divided into subgenera: Ablabesmyia, Clinotanypus, Macropelopia, Monopelopia, Procladius, Tanypus, Thienemannimyia and Zavrelimyia. Only 13 genera, Ablabesmyia, Apsectrotanypus, Clinotanypus, Conchapelopia, Fittkauimyia, Larsia, Macropelopia, Monopelopia, Nilotanypus, Procladius, Tanypus, Thienemannimyia and Zavrelimyia are known from all zoogeographical regions. The Nearctic and Palearctic regions, which together constitute the Holarctic Region, have high faunal similarity. Forty-two genera are found in the Holarctic, with 29 genera common to both regions. Genera occurring in the Nearctic not yet recorded from the Palaearctic are: Coelotanypus, Denopelopia, Helopelopia, Hudsonimyia, Pentaneura and Radotanypus. Genera currently represented in the Palaearctic, but not yet registered from the Nearctic are: Amnihayesomyia, Anatopynia, Coffmania, Lobomyia, Pentaneurella, Saetheromyia and Telmatopelopia. In contrast, the Neotropical and Oriental regions, which theoretically should have rich generic faunas (Ashe et al., 1987), are among the least known, which may reflect the lack of work done on the tanypodine fauna in these regions. Most Neotropical and Oriental species are only known from one or two countries with many only known from the original type-locality. The Neotropical Region has seven exclusive genera: Amazonimyia, Laurotanypus, Metapelopia, Naelotanypus, Paggipelopia, Parapentaneura and Wuelkerella, whereas Chaudhuriomyia is currently the only endemic genus to the Oriental Region. Although the numbers of described tanypodine genera recorded from the Neotropical and Oriental regions are relatively low, © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 Phylogenetic relationships in the subfamily Tanypodinae some efforts have recently been made to describe and document new genera from these areas. Amazonimyia, Chaudhuriomyia, Metapelopia, Paggipelopia and Wuelkerella were all described within the last three years (Suárez & Sublette, 2012; Silva et al., 2014; Paul & Mazumdar, 2015; Silva & Wiedenbrug, 2015; Siri & Donato, 2015). Twenty-two genera are recorded from the Oriental Region (Ashe & O’Connor, 2009). However, the discovery of genera already known from other regions is to be expected in both the Neotropical and Oriental regions (Ashe et al., 1987). The tanypodines, as well as other Chironomidae, of the Australasian Region is comparatively poorly known – both as adults and immatures (Roback, 1982a). Nineteen genera are present in the region, with Australopelopia and Coelopynia being the only ones endemic to the area. Likewise, the tanypodine fauna of the Afrotropical Region probably is poorly known, with also 19 genera recorded. Despite the magnitude of the African continent, only two tanypodine genera are exclusive to the region: Chrysopelopia and Lepidopelopia, both known only as adult males. In summary, the Tanypodinae generic diversity progresses from high latitudes to low latitudes (i.e. from temperate regions towards the tropics) and there is also an increase in species diversity and numbers of specimens. Similar pattern is also exhibited by the subfamily Chironominae. In contrast, Diamesinae, Orthocladiinae, Podonominae and Prodiamesinae show the opposite pattern, with richer diversity in the temperate regions (Ashe et al., 1987). This model may reflect the adaptations of the species to prevailing environmental conditions, such as oxygen concentration, water temperature, climate and altitude. The subfamily Telmatogetoninae is species-poor and predominantly marine and does not fit either pattern, probably because a distinct array of other factors may interfere in its distribution. Ecological knowledge of the remaining subfamilies, Aphroteniinae, Buchonomyiinae, Chilenomyiinae and Usambaromyiinae, is still very fragmented and their low generic and species richness prevent any ecological or zoogeographical generalisations. Conclusion This is the first study to include all Tanypodinae genera in a phylogenetic analysis using morphological data. The results find substantial congruence with previous ideas concerning relationships within the subfamily. The monophyly of Tanypodinae is well supported by five morphological synapomorphies, with Podonominae as its sister group. All previously proposed tribes are recovered as monophyletic under a wide range of applied weighting regimes. Under these conditions, the genus Fittkauimyia occupies a basal position relatively to genera within Macropelopiini and is established as a new monobasic tribe, Fittkauimyiini, supported by at least nine synapomorphies. The tribe Pentaneurini is recovered with some internal structuring such as a monophyletic Thienemannimyia group. The genus Paramerina is recovered as sister of Reomyia + Zavrelimyia with Schineriella as the closest relative of these three genera combined. Thus, Paramerina is here regarded as subgenus of Zavrelimyia. Laurotanypus and Lepidopelopia, which so far had © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 87 a doubtful placement, are recovered as members of the tribe Procladiini. Recent nomenclatural changes suggesting the synonymisation of Gressittius and Guassutanypus with Alotanypus and the subgeneric status of Bethbilbeckia (in Macropelopia), Hayesomyia (in Thienemannimyia), and Reomyia and Schineriella (in Zavrelimyia) are corroborated. Helopelopia, a subgenus of Conchapelopia, is re-established as a distinct genus in the Thienemannimyia group. This study provides an initial outline for understanding the evolutionary history of Tanypodinae. Although, only morphological data were used to reconstruct the phylogeny, the results provide a reasonably robust hypothesis to be further tested with molecular data. As we have shown, there is phylogenetic signal in morphological data at the generic level in Tanypodinae and future studies should combine evidence from different sources in order to dilute the bias of homoplasy and maximising the information content in phylogenetic analyses. Supporting Information Additional Supporting Information may be found in the online version of this article under the DOI reference: 10.1111/syen.12141 Figure S1. Strict consensus tree from the resulting 18 cladograms based on parsimony searches on a reweighted character matrix in paup*. Characters were reweighted according to their rescaled consistency index. File S1. Morphological exemplar taxa. File S2. Morphological character list. File S3. Morphological character matrix. Acknowledgements We would like to thank Martin Spies and Marion Kotrba, who enabled us to study the valuable material preserved in the Zoologische Staatssammlung München (ZSM). Thanks to Sofia Wiedenbrug for her assistance on pupal character coding, to Paula F. M. Rodrigues for useful comments and valuable suggestions on the phylogenetic analyses and to Caroline S. N. Oliveira and Susana T. Strixino for providing us with important material. Thanks also to Augusto Siri for sharing the manuscript of their work on the phylogeny of Macropelopiini before publication and to Bohdan Bilyj for insightful comments on the first version of our manuscript. F. L. Silva was supported by a Postdoctoral Fellowship from the Coordination for the Improvement of Higher Education Personnel (CAPES). References Adam, J.I. & Sæther, O.A. (1999) Revision of the genus Nilothauma Kieffer, 1921 (Diptera: Chironomidae). Entomologica Scandinavica Supplement, 56, 1–107. 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Accepted 8 July 2015 First published online 30 August 2015 APPENDIX New generic placements Based on the discussion and conclusions in the present manuscript, the following nomenclatural actions for all species names in Bethbilbeckia, Conchapelopia (Helopelopia), Cantopelopia, Gressittius, Guassutanypus, Hayesomyia, Paramerina, Reomyia and Schineriella are proposed (previous placement in parenthesis): Alotanypus antarcticus comb.n. (Hudson, 1892: 43). (Gressittius) Alotanypus oliveirai comb.n. (Roque & TrivinhoStrixino, 2003: 161). (Guassutanypus) Alotanypus umbrosus comb.n. (Freeman, 1959: 406). (Gressittius) Helopelopia cornuticauda comb.n. (Walley, 1925: 277). (Conchapelopia (Helopelopia)) Helopelopia pilicaudata comb.n. (Walley, 1925: 277). (Conchapelopia (Helopelopia)) Macropelopia (Bethbilbeckia) floridensis comb.n. (Fittkau & Murray, 1988: 254). (Bethbilbeckia) Monopelopia (Cantopelopia) gesta comb.n. (Roback, 1971: 270). (Cantopelopia) Monopelopia (Cantopelopia) meilloni comb.n. (Freeman, 1955: 31). (Cantopelopia) Monopelopia (Cantopelopia) robacki comb.n. (Lehmann, 1979: 14). (Cantopelopia) Thienemannimyia (Hayesomyia) aquila comb.n. (Cheng & Wang, 2006: 36). (Hayesomyia) Thienemannimyia (Hayesomyia) cinctuma comb.n. (Cheng & Wang, 2006: 38). (Hayesomyia) Thienemannimyia (Hayesomyia) fengkainica comb.n. (Cheng & Wang, 2006: 41). (Hayesomyia) Thienemannimyia (Hayesomyia) galbina comb.n. (Cheng & Wang, 2006: 43). (Hayesomyia) © 2015 The Royal Entomological Society, Systematic Entomology, 41, 73–92 91 Thienemannimyia (Hayesomyia) rotunda comb.n. (Cheng & Wang, 2006: 46). (Hayesomyia) Thienemannimyia (Hayesomyia) senata comb.n. (Walley, 1925: 276). (Hayesomyia) Thienemannimyia (Hayesomyia) triangula comb.n. (Cheng & Wang, 2006: 48). (Hayesomyia) Thienemannimyia (Hayesomyia) trina comb.n. (Cheng & Wang, 2006: 50). (Hayesomyia) Thienemannimyia (Hayesomyia tripunctata comb.n. (Goetghebuer, 1922: 59). (Hayesomyia) Thienemannimyia (Hayesomyia) zayunica comb.n. (Cheng & Wang, 2006: 55). (Hayesomyia) Zavrelimyia (Paramerina) ababae comb.n. (Harrison, 1991: 64). (Paramerina) Zavrelimyia (Paramerina) ampliseta comb.n. (Hazra, Saha, Mazumdar & Chaudhuri: 332). (Paramerina) Zavrelimyia (Paramerina) anomala comb.n. (Beck & Beck, 1966: 344). (Paramerina) Zavrelimyia (Paramerina) aucta comb.n. (Johannsen, 1931: 498). (Paramerina) Zavrelimyia (Paramerina) cingulata comb.n. (Walker, 1856: 172). (Paramerina) Zavrelimyia (Paramerina) clara comb.n. (Hazra, Saha, Mazumdar & Chaudhuri: 335). (Paramerina) Zavrelimyia (Paramerina) divisa comb.n. (Walker, 1856: 192). (Paramerina) Zavrelimyia (Paramerina) dolosa comb.n. (Johannsen, 1931: 500). (Paramerina) Zavrelimyia (Paramerina) edwardsi comb.n. (Freeman, 1955: 28). (Paramerina) Zavrelimyia (Paramerina) fasciata comb.n. (Sublette & Sasa, 1994: 6). (Paramerina) Zavrelimyia (Paramerina) fragilis comb.n. (Walley, 1925: 205). (Paramerina) Zavrelimyia (Paramerina) fittkaui comb.n. (Lehmann, 1981: 13). (Paramerina) Zavrelimyia (Paramerina) hanseni comb.n. (Roback, 1971: 274). (Paramerina) Zavrelimyia (Paramerina) ignobilis comb.n. (Johannsen, 1932: 496). (Paramerina) Zavrelimyia (Paramerina) inficia comb.n. (Chaudhuri & Debnath, 1985: 167). (Paramerina) Zavrelimyia (Paramerina) levidensis comb.n. (Skuse, 1889: 281). (Paramerina) Zavrelimyia (Paramerina) longepis comb.n. (Freeman, 1955: 32). (Paramerina) Zavrelimyia (Paramerina) mauretanica comb.n. (Fittkau, 1962: 333). (Paramerina) Zavrelimyia (Paramerina) meilloni comb.n. (Freeman, 1955: 31). (Paramerina) Zavrelimyia (Paramerina) minima comb.n. (Kieffer, 1911: 365). (Paramerina) Zavrelimyia (Paramerina) nigromarmorata comb.n. (Goetghebuer, 1935: 361). (Paramerina) Zavrelimyia (Paramerina) okigenga comb.n. (Sasa, 1990: 141). (Paramerina) 92 F. L. Silva and T. Ekrem Zavrelimyia (Paramerina) okimaculata comb.n. (Sasa, 1990: 142). (Paramerina) Zavrelimyia (Paramerina) parva comb.n. (Freeman, 1961: 617). (Paramerina) Zavrelimyia (Paramerina) quininficia comb.n. (Chaudhuri & Debnath, 1985: 169). (Paramerina) Zavrelimyia (Paramerina) septemguttata comb.n. (Freeman, 1955: 28). (Paramerina) Zavrelimyia (Paramerina) smithae comb.n. (Sublette, 1964: 100). (Paramerina) Zavrelimyia (Paramerina) taylori comb.n. (Roback, 1982c: 102). (Paramerina) Zavrelimyia (Paramerina) togavicea comb.n. (Sasa & Okazawa, 1992: 213). (Paramerina) Zavrelimyia (Paramerina) testa comb.n. (Roback, 1971: 272). (Paramerina) Zavrelimyia (Paramerina) vaillanti comb.n. (Fittkau, 1962: 335). (Paramerina) Zavrelimyia (Paramerina) valida comb.n. (Paul, Hazra & Mazumdar, 2013: 499).(Paramerina) Zavrelimyia (Paramerina) yunouresia comb.n. (Sasa, 1989: 152). (Paramerina) Zavrelimyia (Reomyia) wartinbei comb.n. (Roback, 1986: 283). (Reomyia) Zavrelimyia (Schineriella) schineri comb.n. (Murray & Fittkau, 1988: 247). 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