Cordiluroides Albuquerque (Diptera: Muscidae): new species and key to the Neotropical genus

Here we describe three new species of Cordiluroides Albuquerque, one striking new species from Brazil, Minas Gerais, C. albitarsata sp. nov., and two from Peru, Cuzco, C. inca sp. nov. and C. wayqechensis sp. nov. Additionally, photographs from the types of C. listrata Albuquerque, 1954, C. megalopyga Albquerque, 1954 and C. insularis (Williston, 1896) are included. With the new species, Cordiluroides is now known from 11 species exclusively distributed in the Neotropical region with records from Bolivia, Brazil, Colombia, Costa Rica, Mexico, Saint Vincent, and Peru. We give an updated identification key to species from the Neotropical region. Finally, we describe for the first time the external morphology and genitalia of the Cordiluroides species using Scanning Electron Microscope (SEM) images. A R T I C L E I N F O Article history: Received 31 March 2020 Accepted 22 September 2020 Available online 30 October 2020 Associate Editor: Sarah Oliveira


Introduction
Cordiluroides Albuquerque, 1954, is an exclusively Neotropical genus of Muscidae with records from Bolivia, Brazil, Colombia, Costa Rica, Mexico, Saint Vincent, and Peru (Löwenberg-Neto and de Carvalho, 2013). The genus was proposed by Albuquerque (1954) for three species: C. listrata, C. megalopyga, and C. neotropica, the later one synonymized with C. listrata by Couri and Pamplona (1992). In this study also was proposed a new combination of Cordylura geniculata Macquart, 1851 to Cordiluroides. Later, Pont (1972) proposed two new combinations for the genus: Cordiluroides bicolor (Bigot, 1885) and Cordiluroides insularis (Williston, 1896), originally described in Dialyta Meigen, 1826 andCoenosia Meigen, 1826 respectively. After, Couri and Gonçalves (2002) added a new species C. paulistensis to the genus. Finally, Couri et al. (2006) recorded the genus for the first time from Costa Rica and revised the status of Cordiluroides bistriata and C. vittifera, removing them from the synonym of C. insularis, so that the genus has now eight valid species.
Species of the genus can be easy distinguished by the high insertion of antenna, above middle of the eye, the presence of only one pair of postsutural intra-alar setae, lower calypter transverse, and abdomen elongated (Albuquerque, 1954). There are three identification keys to the genus: Couri and Pamplona (1992) and Couri and de Carvalho (2002) for the Neotropical species, and Couri et al. (2006) for the Costa Rica species. There is no information about the biology and immature stages.
Here we describe one striking new species from Minas Gerais, Brazil and two new species collected from Peru. We also provide photos from the type material of C. insularis Williston, 1896, C. listrata Albuquerque, 1954, and C. megalopyga Albuquerque, 1954. An updated key to the Neotropical species of Cordiluroides also furnished. Herein, we describe for the first time the external morphology and genitalia of the Cordiluroides species using Scanning Electron Microscope (SEM) images.

Material
This study was mostly based on Cordiluroides specimens from Wayqecha Cloud Forest Biological Station (Cuzco, Peru), collected by Malaise trap in June 2012, which are deposited in the National Museum of Natural History (USNM) and the Padre Jesus Santiago Moure entomological collection, Universidade Federal do Paraná, Curitiba, Brazil (DZUP). The material from Parque Nacional do Itatiaia (Rio de Janeiro and Minas Gerais states, Brazil) was collected using Malaise traps placed on "Brejo da Lapa" area, located in the highest part of the park, as part of the project "Conhecer para proteger: Medindo a riqueza de insetos do Parque Nacional do Itatiaia -BIOTA-FAPERJ" coordinated by Marcela Monné (Museu Nacional, UFRJ). This material was identified based on the available Neotropical species keys and prepared at the Diptera laboratory of Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ/UFRJ).

Dissections and terminology
Pinned dry specimens were examined under the stereomicroscope and the terminalia were examined after being removed from the abdomen, cleared with cold potassium hydroxide (KOH) 10% for 24 hours, then transferred to acetic acid, after being dehydrated in 70% alcohol and then placed in glycerin. The terminalia were dissected, analyzed and illustrated under the optical microscope with the help of a camera lucida. After examination, the terminalia were placed in glass vials that were fixed to the original pinned specimen.

Images
The entire specimens or extracted male genitalia were used in Scanning Electron Microscope (SEM). For the photographs, the entire specimens or male genitalia were dehydrated in a graded series of alcohol, critical-point dried and sputter-coated with gold. Photographs were taken using a JEOL JSM 6360-LV at Centro de Microscopia Eletrônica, Universidade Federal do Paraná, Curitiba, Paraná, Brazil.
Photographs of the adult habitus of new species from Peru and Brazil were stacked using an auto-montage setup acquired at Laboratório de Sistemática e Bioecologia de Coleoptera (UFPR), and a Leica M205 C, version v 4.8.0 at Museu Nacional, Universidade Federal do Rio de Janeiro (UFRJ), respectively. Photographs of the type material were stacked using an auto-montage setup acquired at Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro and Natural History Museum, London.

Taxonomy
Key to species (Modified from de Carvalho, 2002 andCouri et al., 2006) 1 Apical scutellar seta very reduced or absent ( Fig. 1e and  7 (6) Abdomen of male very elongated, tergites 1+2 and 3 translucent yellow and with a median dorsal triangular brown cloud; tergites 4 and 5 brown, sometimes tergite 4 also translucent yellow at base; abdomen of female not elongated, brown, with lateral yellow areas on tergites 1+2 and 3 and grey pruinose on tergites 4 and 5; wings   Diagnosis. The presence of white setae (beard) in the gena and postgena in male (Fig. 4c) and female, fore tarsus with tarsomeres 2-5 flattened and white in male (Fig. 4a) easily distinguish Cordiluroides albitarsata sp. nov. from the other Cordiluroides. Ground color brown with grey pollinosity. Head (Fig. 4a-c and f): Frons brown with a discrete golden pruinosity in center in male and more homogeneous dark brown in female; fronto-orbital plate, face and gena silver pruinose; gena and postgena in male and only postgena in female with white setae (beard); antennae and arista brown, apex of pedicel lighter in some specimens; palpus dark brown. Thorax (Fig. 4d-e): Scutum with 5 brown vittae, a little larger in female. Upper and lower calypters white; halter yellow. Wing: Infuscated (Fig. 4d). Legs: All coxae yellow with white pollinosity, fore femur yellow, mid and hind femora yellow with apical fifth brown, all tibia yellow, all tarsi dark brown, except fore tarsus in male with tarsomere 1 dark brown, except on tip, tarsomere 2 white, with a brown area at base, tarsomeres 3-5 white (Fig. 4a); pulvilli light brown, claws black, ligther at base. Abdomen: With a brown middle stripe and lateral brown clouds in all tergites.
Head. Interocular space about one-third of head-width at the level of anterior ocellus. Four pairs of frontal setae, the two lower ones short and the two superior ones long and backwards directed; inner vertical seta long; ocellar setae very short; antennae inserted a little above middle of eyes; pedicel reaching oral margin; arista short pubescent. Gena narrow. Vibrissa strong and long. Proboscis with developed teeth.
Legs. Fore femur with a row of long and sparse setae on pd and pv surfaces; fore tibia with one long median p setae, one preapical d and two apicals, one p and one pv. Mid femur with two ad setae on middle third, 3-4 long and sparse v setae, posterior surface with two preapical setae; mid tibia with one long median ad setae, one long median pd, one long preapical d and one apical ad; hind femur with a sparse ad row with about six setae, av surface with four sparse setae, preapical setae on d, ad and pd surfaces; hind tibia with one long median ad seta, one long d on apical-third, one av on apical-third, one preapical ad and one apical v strong.
Head. Interocular space about one-third of head-width at level of anterior ocellus. Eyes sparsely ciliated (Fig. 10c). Four pairs of frontal setae with different lengths; one pair of long reclinate orbital setae; ocellar setae long (Fig. 10b), similar in length to the reclinate orbital. Antennae inserted a little above middle of eyes; flagellomere about 3 times as long as pedicel; arista short pubescent (Fig. 10c-d). Gena narrow. Vibrissa strong and long (Fig. 10b). Proboscis with developed teeth.
Legs. Fore femur with a row of sparse setae on pd and pv. Fore tibia with one long median p seta; one d, p preapical seta; one pv apical seta, these three shorts. All pulvilli and claws developed. Mid femur with 4 long sparse pv setae; p with two preapical setae. Mid tibia with a submedian a to av seta; one d, p, v and pv preapical seta, the v is larger than the others. Hind femur with 4 long apical ad setae; one apical d, pd seta and pv with 3 long, thin and sparse setae. Hind tibia with one long median pd, ad seta; one long submedian d seta; one ad, av preapical seta.
Length. Male: body: 2.8mm mm; wing: 3.0mm. Head. Interocular space about one-third of head-width at level of anterior ocellus. Eyes sparsely ciliated. Four pairs of frontal setae of different lengths; one pair of long reclinate orbital setae; ocellar setae long, similar in length to the reclinate orbital. Antennae inserted a little above middle of eyes; flagellomere about 4 times as long as pedicel; arista short pubescent. Cheek narrow. Vibrissa strong and long (Fig. 2b). Proboscis with developed teeth.
Legs. Fore femur with 4-8 long pd setae; 2 long basal a setae; row of 11 long v setae and 4 short basal pv setae. Fore tibia with one long median p seta; one d, p preapical seta; one pv apical seta, these three shorts. All pulvilli and claws well developed. Mid femur with 2 long basal a setae; 3 long basal av setae; 4 long sparse pv setae; p with two preapical setae. Mid tibia with a submedian a to av seta; one d, p, v and pv preapical seta, the v is longer than the other setae. Hind femur with 4 long apical ad setae; one apical d, pd seta and pd, pv with 5 long, and sparse setae, in av the three basal are less developed. Hind tibia with one long median pd, ad seta; one long submedian d seta; one ad, av preapical seta.

Discussion
With the three new species described in this study, Cordiluroides comprises 11 species distributed around the Neotropical region, Bolivia, Brazil, Colombia, Costa Rica, Mexico, Saint Vincent, and Peru (Supplementary material- Table S1). Despite the remarkable morphology of the genus, the monophyly of the Cordiluroides is unclear. In the phylogenetic analysis hypothesis to Coenosiini proposed by Couri and Pont (2000), Cordiluroides was placed in a monophyletic clade together with Neodexiopsis Malloch, 1934, based only one synapomorphy: the hind femur with three preapical dorsal setae. However, there is no study available in literature discussing the phylogenetic relationship between species of Cordiluroides.
Describing new species is an important step towards the taxonomic and phylogenetic understanding of a group, furthermore the description of the Cordiluroides albitarsata, C. inca, and C. wayqechensis, contributes with new morphological characteristics for the genus. Cordiluroides albitarsata presents white setae (beard) in the gena and postgena (Fig. 4c) and fore tarsus with tarsomeres 2-5 flattened and white in male (Fig. 4a). While Cordiluroides inca, and C. wayqechensis present the apical scutellar seta reduced, these characteristics enlarging the diagnostic characteristics to species of Cordiluroides. In addition, Cordiluroides albitarsata increasing the knowledge about genus distribution with the first Cordiluroides specie's record to the Brazilian state of Minas Gerais.