Four new neotropical species oF TrigonomeTopus Macquart , 1835 ( Diptera , lauxaniiDae )

The genus Trigonometopus Macquart (1835) currently includes 13 species, found in most biogeographic regions, being absent from the Afrotropical and Australian Regions. The genus included six known Neotropical species and four new species are herein added to the genus: T. assisensis sp. nov., T. boraceiensis sp. nov., T. lourdesae sp. nov., and T. mariae sp. nov., the first two from the State of São Paulo, the other two respectively from the states of Mato Grosso do Sul and Santa Catarina. A key for the exclusively Neotropical species of the genus is provided. Key-Words: Description; New taxa; Neotropical region; Systematics. 1830 (from a French locality not stated by Meigen, maybe Bordeaux according to Papp, 1984); T. vittatus Loew, 1869 (from Georgia, USA); T. canus Meijere, 1916 (from Indonesia, Java, considered a questionable species by Papp, 2007); T. fuscipennis Hendel, 1912 (from Samoa); T. ishidae Matsumura, 1918 (from Japan; S.D. Gaimari, person. comm.); and T. semibrunneus Malloch, 1929 (from Niue, American Samoa). Knab (1914a) reviewed the species from the New World, described two new species and presented a key. His characterization of the genus, howeveralong head with a triangular profile, its horizontal frons, face strongly receding, and antenna inserted at frontal apex-is broad and vague, as pointed by Stuckenberg (1971). Malloch (1923) described one new species from Cuba and presented a key for the identification of the species he had seen-the key most used for the genus in the region so far. INTRODUCTION The genus Trigonometopus was described by Macquart (1835) to include Tetanocera frontalis Meigen, 1830, at that time known from France. The genus now includes 13 species found in the Palearctic, Nearctic, Oriental e Neotropical regions-the genus is noticeably absent in the Afrotropical and Australian regions. There are six species known from the Neotropical region: T. albifrons Knab, 1914a; T. albocostatus Hendel, 1912; T. angustipennis Knab, 1914a; T. immaculipennis Malloch, 1923; T. punctipennis Coquillett, 1898, and T. rotundicornis Williston, 1896. T. punctipennis was originally described from the Nearctic region, but was registered for Mexico and Costa Rica (Gaimari & Silva, 2010b). The valid non-Neotropical species, according to Pape & Thompson (2013), are: T. eborifacies Shatalkin, 1997 (from Russia); T. frontalis Meigen, http://dx.doi.org/10.1590/0031-1049.2015.55.0## Volume 55(##):A‐L, 2015 1. Universidade Federal do Paraná – UFPR. Programa de Pós-Graduação em Entomologia, Setor de Ciências Biológicas, Departamento de Zoologia. Caixa Postal 19.020; CEP 81531-980, Curitiba, PR, Brasil. E-mail: angelbyo@outlook.com 2. Universidade Estadual Paulista – UNESP. Faculdade de Ciências Agrárias e Veterinárias de Jaboticabal, Departamento de Morfologia e Fisiologia Animal. Via de Acesso Professor Paulo Donato Castellane, s/n, Vila Industrial, CEP 14884-900, Jaboticabal, SP, Brasil. E-mail: vcsilva@fcav.unesp.br The taxon as a family group based on Trigonometopus has been studied mainly by Knab (1914b), Malloch (1929), Wheeler (1956), Stuckenberg (1971), Shatalkin (1997, 2000), Sasakawa (1998, 2002, 2005) and Papp (2007), besides many others who have just described new species. A suprageneric taxon has been proposed to gather Trigonometopus and some other genera, ranked as subfamily by Becker (1905), but considered as tribe by Papp (2007). It would include, according to Papp (2007), the following Old World genera: Diplochasma Knab; Kerteszomyia Malloch; Luzonomyza Malloch; Maquilingia Malloch; Neotrigonometopus Malloch; Protrigonometopus Hendel; Shatalkinella Papp; Trigonometopsis Malloch; Trigonometopus Macquart; and Tetroxyrhina Hendel. Papp (2007) reviewed the Trigonometopini providing a better characterisation of its Old World genera. The tribe can be characterized by a number of features: an elongated head, in most genera longer than high; a broad, setose fronto-orbital plate (an obvious synapomorphy of the clade); the thorax more or less elongated; abdomen never strongly flattened; tergite 7 fused to the epandrium, at least medially (another synapomorphy of the tribe); male genitalia with no surstylus nor surstylar lobe on epandrium; phallus wholly membranous; and three spermathecae. Shatalkin (1997) considered Tetroxyrhina a subgenus of Trigonometopus, but pointed many differences between the two taxa. He indicated as Palaearctic species of Trigonometopus s. str. just T. frontalis (Meigen, 1830) and T. eborifacies Shatalkin, 1997, refering as diagnostic features for this taxon the first flagellomere pointed apically, fore fronto-orbital setae situated on level of anterior margin of eye, two sternopleural setae, yellow halteres, epandrium forming a ring structure as a result of the ventral expansion of its lateral sectors. Papp (2007) elevated Tetroxyrhina to the generic level, and it seems to be the most speciose genus of the tribe. Gaimari & Silva (2010b) presented a key to the Neotropical Lauxaniidae genera that includes Trigonometopus. Diagnostic features for the genus in the Neotropical region would be the head sharply pointed anteriorly; fronto-facial angle less than 90°; eye at least slightly longer than high; postsutural intra-alar seta absent; wing hyaline or marked; crossvein dm-cu beyond middle of wing; cell dm with rounded apex. This paper is a contribution to the knowledge of the Neotropical Lauxaniidae, a large acalyptrate family still poorly known in the Neotropics, with the description and illustration of four new species of Trigonometopus from Brazil. The paper discusses interesting genitalic features of males and females that may be particularly useful in a worldwide revision of the genus. MATERIALS AND METHODS This study utilized material collected by the authors and specimens borrowed from the “Coleção Padre Jesus Santiago Moure”, Universidade Federal do Paraná, Curitiba, Brazil (DZUP), and from the Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP). Specimens of one of the new species were collected by the first author in Assis in the State of São Paulo. The type specimens were deposited at the collections of DZUP and MZUSP. The study also includes material collected in the “SISBIOTA Diptera CNPq/FAPESP” project in the State of Mato Grosso do Sul, housed at the MZUSP. For the study of the morphology of male and female terminalia, the post-abdomen was removed from a dry specimen and placed in a cold 10% KOH solution for 24-48 hours to soften the tissues, and then rinsed in distilled water and dehydrated in an increasing ethanol series (30%, 50%, 70% and 95%). It was then bleached with lactophenol and stored in a vial with glycerin, which was fixed to the insect pin. Illustrations were made using a Leica DM 2500 microscope and Leica MZ 12,5 stereomicroscope, both equipped with a camera lucida. The morphological terminology follows Gaimari & Silva (2010b) and Cumming & Wood (2009).


INTRODUCTION
The genus Trigonometopus was described by Macquart (1835) to include Tetanocera frontalis Meigen, 1830, at that time known from France.The genus now includes 13 species found in the Palearctic, Nearctic, Oriental e Neotropical regions-the genus is noticeably absent in the Afrotropical and Australian regions.There are six species known from the Neotropical region: T. albifrons Knab, 1914a; T. albocostatus Hendel, 1912; T. angustipennis Knab, 1914a;T. immaculipennis Malloch, 1923;T. punctipennis Coquillett, 1898, andT. rotundicornis Williston, 1896.T. punctipennis was originally described from the Nearctic region, but was registered for Mexico and Costa Rica (Gaimari & Silva, 2010b).The valid non-Neotropical species, according to Pape & Thompson (2013), are: T. eborifacies Shatalkin, 1997 (from Russia); T. frontalis Meigen, The taxon as a family group based on Trigonometopus has been studied mainly by Knab (1914b), Malloch (1929), Wheeler (1956), Stuckenberg (1971), Shatalkin (1997Shatalkin ( , 2000)), Sasakawa (1998Sasakawa ( , 2002Sasakawa ( , 2005) ) and Papp (2007), besides many others who have just described new species.A suprageneric taxon has been proposed to gather Trigonometopus and some other genera, ranked as subfamily by Becker (1905), but considered as tribe by Papp (2007).It would include, according to Papp (2007), the following Old World genera: Diplochasma Knab; Kerteszomyia Malloch; Luzonomyza Malloch; Maquilingia Malloch; Neotrigonometopus Malloch; Protrigonometopus Hendel; Shatalkinella Papp;Trigonometopsis Malloch;Trigonometopus Macquart;and Tetroxyrhina Hendel. Papp (2007) reviewed the Trigonometopini providing a better characterisation of its Old World genera.The tribe can be characterized by a number of features: an elongated head, in most genera longer than high; a broad, setose fronto-orbital plate (an obvious synapomorphy of the clade); the thorax more or less elongated; abdomen never strongly flattened; tergite 7 fused to the epandrium, at least medially (another synapomorphy of the tribe); male genitalia with no surstylus nor surstylar lobe on epandrium; phallus wholly membranous; and three spermathecae.Shatalkin (1997) considered Tetroxyrhina a subgenus of Trigonometopus, but pointed many differences between the two taxa.He indicated as Palaearctic species of Trigonometopus s. str.just T. frontalis (Meigen, 1830) and T. eborifacies Shatalkin, 1997, refering as diagnostic features for this taxon the first flagellomere pointed apically, fore fronto-orbital setae situated on level of anterior margin of eye, two sternopleural setae, yellow halteres, epandrium forming a ring structure as a result of the ventral expansion of its lateral sectors.Papp (2007) elevated Tetroxyrhina to the generic level, and it seems to be the most speciose genus of the tribe.Gaimari & Silva (2010b) presented a key to the Neotropical Lauxaniidae genera that includes Trigonometopus.Diagnostic features for the genus in the Neotropical region would be the head sharply pointed anteriorly; fronto-facial angle less than 90°; eye at least slightly longer than high; postsutural intra-alar seta absent; wing hyaline or marked; crossvein dm-cu beyond middle of wing; cell dm with rounded apex.
This paper is a contribution to the knowledge of the Neotropical Lauxaniidae, a large acalyptrate family still poorly known in the Neotropics, with the description and illustration of four new species of Trigonometopus from Brazil.The paper discusses interesting genitalic features of males and females that may be particularly useful in a worldwide revision of the genus.

MATERIALS AND METHODS
This study utilized material collected by the authors and specimens borrowed from the "Coleção Padre Jesus Santiago Moure", Universidade Federal do Paraná, Curitiba, Brazil (DZUP), and from the Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP).
Specimens of one of the new species were collected by the first author in Assis in the State of São Paulo.The type specimens were deposited at the collections of DZUP and MZUSP.The study also includes material collected in the "SISBIOTA Diptera CNPq/FAPESP" project in the State of Mato Grosso do Sul, housed at the MZUSP.
For the study of the morphology of male and female terminalia, the post-abdomen was removed from a dry specimen and placed in a cold 10% KOH solution for 24-48 hours to soften the tissues, and then rinsed in distilled water and dehydrated in an increasing ethanol series (30%, 50%, 70% and 95%).It was then bleached with lactophenol and stored in a vial with glycerin, which was fixed to the insect pin.Illustrations were made using a Leica DM 2500 microscope and Leica MZ 12,5 stereomicroscope, both equipped with a camera lucida.
Diagnosis: head triangular in profile (Fig. 1A); antenna inserted at frontal apex; face strongly protruding; eye longer than high; fronto-facial angle slightly higher than 45°; fronto-orbital setae situated on level of anterior margin of eye; proclinate setulae on the anterior part of frons; anepimeron bare; postsutural intra-alar seta absent; ctenidium absent; wing sapromyziform; wing hyaline or marked; crossvein dm-cu beyond middle of wing; base of R 4+5 bare; cell dm with rounded apex.
Etymology: The specific epithet assisensis was used as a homage to the type-locality, the city of Assis, where a large number of the type specimens were collected.To be treated as a noun in apposition.

Etymology:
The specific epithet boraceiensis is a homage to the type-locality, "Estação Biológica de Boracéia", where the holotype was collected.To be treated as a noun in apposition.
Trigonometopus mariae sp.nov.(Fig. 4) Chaetotaxy: fore coxa with 3 apical setae; fore femur with one posterodorsal, one posteroventral and one dorsal rows of setae; fore tibia with 2 apical setae: 1 anterodorsal and 1 ventral; mid coxa with 2 basal setae and 1 apical setae; mid femur with 1 anterior row of setae, 1 apical posterodorsal seta, 1 apical posterior seta; mid tibia with 2 apical setae: 1 anterodorsal and 1 anteroventral; hind coxa with 2 apical setae: 1 anterior and 1 lateral; hind femur thickened, with 1 strong anterodorsal seta; hind tibia with 1 apical anterodorsal spine, 1 ventral spine.Wing (Fig. 4D): Hyaline, infuscated at costal margin, spotted at both end of crossvein dm-cu, two spots along R 4+5 near its apex; costa sapromyziform; crossvein r-m located in the midpoint of discal medial cell; crossvein dm-cu located before midpoint of cell r 4+5 ; longitudinal veins parallel; vein A 1 short, not reaching wing margin.Comments: Abdomens dissected and stored in vials with glycerin and pinned with the specimens.This species is similar to T. punctipennis, differing by the frons being longer (1,5 times longer than wide in T. punctipennis), brown (brown with yellow central stripe in T. punctipennis), first flagellomere oval (pointed apically in T. punctipennis), scutum brownish yellow, darker at lateral margins (brownish yellow with central gray dorsocentral vittae through entire length and dark brown laterally in T. punctipennis), wing infuscated at costal margin (infuscated at cell bm and base of basal radial cell and r-m spotted in T. punctipennis).

DISCUSSION
This genus has considerably wide distribution in the world, but it is still poorly known in the Neotropical region.This paper almost doubles the number of previously known species for the Neotropics, suggesting that there is the need for collecting effort in more diverse habitats in different parts of the region.The actual number of Neotropical species of Trigonometopus is likely much higher.
Among the new species, only T. assisensis sp.nov.has four spermathecae, a feature considered by Gaimari & Silva (2010a) to be an apomorphy for the Eurychoromiinae and as well a feature sporadically found in other lauxaniid genera.It has been also observed in the Neotropical lauxaniine genera Cephalella Malloch (Silva, 1999) and Neogriphoneura Malloch (Mello & Silva, 2008), and in the Australian species Australinina geniseta (Malloch), Homoneura (Homoneura) angustigena Kim and H. (H.) eurymelon Kim (Kim, 1994).This feature is still insufficiently known in many species and genera of the family and a more extensive comparative analysis would better show the evolution of the number of spermathecae in the Lauxaniidae.Shatalkin (1997) and Papp (2007) considered that the following features define Trigonometopus: first flagellomere pointed apically, anterior fronto-orbital seta situated on level of anterior margin of eye and structure of the epandrium forming a ring as a result of the ventral expansion of its lateral sectors.These features are quite variable, however, among the Neotropical species or are even absent in some species (or the information is not available).The apex of the flagellomere is pointed only in T. assisensis and T. punctipennis, although not forming a small beak, as in T. frontalis.The position of the anterior fronto-orbital setae is more variable among the Neotropical species, but five of the species share the state in T. frontalis, while there is no information about T. albocostatus for this character.The shape of the epandrium is not known in six of the Neotropical species, but in five of the species the abdomen certainly does not form a ring.
Other characters always mentioned for the genus are the number and position of dorsocentral setae and the number of katepisternal setae: the dorsocentral setae are variable, but in the Nearctic species T. vittatus (no available information for T. albocostatus) and in all Neotropical species there is only one katepisternal seta, while T. punctipennis shares with T. frontalis a pair of katepisternal setae.This shows that part of the features formerly taken as diagnostic for the genus actually apply to only part of Trigonometopus, not being shared by part or all of the Neotropical diversity of the genus.The set of Neotropical species diverge from the rest of the genus to a certain degree.A detailed study of the relationships within Trigonometopus is necessary to put the Neotropical species into proper context within the genus.