Revista Brasileira de Entomologia Revista Brasileira de Entomologia
Rev Bras Entomol 2016;60:217-26 - Vol. 60 Núm.3 DOI: 10.1016/j.rbe.2016.05.004
Systematics, Morphology and Biogeography
Review of the New World genus Cholomyia (Diptera, Tachinidae), with a new species from Costa Rica
Marcelo Domingos de Santis, , Silvio Shigueo Nihei
Universidade de São Paulo, Instituto de Biociências, Departamento de Zoologia, São Paulo, SP, Brazil
Recebido 01 Março 2016, Aceitaram 23 Maio 2016
Abstract

The tachinid genus Cholomyia presents Neotropical and Nearctic distribution with three species: C. acromion (Wiedemann, 1824), C. filipes (Walker, 1857), and C. inaequipes Bigot, 1884. In the present paper, all species are reviewed and redescribed, and a new species from Costa Rica is described, C. zumbadoisp. nov. An identification key based on males is provided. For the first time, the male terminalia of all species, and the female terminalia and first instar larva of C. inaequipes are described and illustrated. Finally, based on the detailed morphological study we discuss the systematic placement of Cholomyia into Myiophasiini-Tachininae. A list of host–parasite records is synthesized.

Keywords
Larva, Myiophasiini, Redescription, Tachininae, Taxonomy
Introduction

The genus Cholomyia was erected by Bigot (1884) for his new species, C. inaequipes, based on five males from Mexico. Later, two other species were transferred to the genus: C. acromion (Wiedemann, 1824) and C. filipes (Walker, 1857) (Guimarães, 1971). All species are distributed exclusively in the Neotropical region, except for C. inaequipes, which also occurs in the Nearctic region. The genus was previously placed in the Urodexiini-Dexiinae (Townsend, 1936; Guimarães, 1971) and in the latest Nearctic Catalogue (O’Hara and Wood, 2004) was transferred to the Myiophasiini-Tachininae.

C. filipes (Walker, 1857) is known only from its original description and then listed in the Neotropical catalogue (Guimarães, 1971). C. acromion (Wiedemann, 1824) and C. inaequipesBigot, 1884 are better known, for they parasitize the curculionid Conotrachelus Dejean, 1835, an important pest that attacks grown plums, apples and peaches (O’Brien and Couturier, 1995). Data on immature stages of C. inaequipes, parasitizing the plum-curculio Conotrachelus nenuphar (Herbst, 1797) were published elsewhere (Quaintance and Jenne, 1912). On the other hand, the male and female terminalia have never been described.

In the present paper, the genus Cholomyia is reviewed. The valid species are redescribed, and a new species is described from Costa Rica, C. zumbaboisp. nov. An identification key is provided based on male specimens. Descriptions and illustrations of the male terminalia of all species and the female terminalia and first instar larva of C. inaequipes are presented for the first time. Finally, based on the detailed morphological study, we discuss the systematic placement of Cholomyia into the tribe Myiophasiini. A list of host–parasite is synthesized from literature.

Material and methods

The examined material is deposited in the following institutions: The Natural History Museum, London, United Kingdom (BMNH); Coleção Entomológica do Instituto Oswaldo Cruz, Rio de Janeiro, Rio de Janeiro, Brazil (CEIOC); Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica (INBio); Muséum National d’Histoire Naturelle, Paris, France (MNHN); Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ); Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (MZSP); Universidade Estadual de Feira de Santana, Feira de Santana, Brazil (UEFS) and Colección Museo de Historia natural, Universidad de la Amazonia, Florencia, Colombia (UAM-E). Other repositories cited in the text: Museo di Zoologia dell’Universita di Torino, Torino, Italy (MZUT); National Museum of Natural History, Washington, United States (USNM) and Zoological Museum, Copenhagen, Denmark (ZMUC). The labels of the type material are represented with quotes (”) to indicate the same label, slash (/) to line break, and semicolon to indicate a new label.

The following abbreviations are used for the position of the setae/setulae on legs: A – anterior; AD – anterodorsal; AV – anteroventral; D – dorsal; P – posterior; PD – posterodorsal; PV – posteroventral; and V – ventral.

Male and female terminalia and first instar larva were clarified in 10% solution of KOH at room temperature for 24 about hours, then neutralized with acetic acid (50%), and washed with distilled water and a series of ethanol solutions of increasing concentrations. After examination on microscope temporary slides, the terminalia were stored in plastic microvial with glycerin and attached to the respective specimen. A permanent microscope slide with the first-instar larvae was mounted for examination and measurements, and both slide and adult pinned specimen were properly labelled.

Morphological terminology follows Cumming and Wood (2009), but for antennal morphology follows Stuckenberg (1999), and for male terminalia Tschorsnig (1985). Female terminalia and larva morphology follow Cantrell (1988), but the term cephaloskeleton of Courtney et al. (2000) is used.

Species were identified by morphological characters using the original description and redescriptions, for C. inaequipes and C. acromion, and by previously identified specimens by the museum of origin, for C. filipes. Host records mentioned in this study were gathered form the literature. New data from specimens labels were not found. Geographical data were gathered based on specimens’ labels and literature records. Those records without original geographic coordinates were georeferenced using GeoNames Search (http://www.geonames.org/export/geonames-search.html) and GeoHack (http://toolserver.org/∼geohack/). The distribution map was produced using the software Diva-Gis 7.5.

ResultsGenus CholomyiaBigot, 1884

(Fig. 1)

Fig. 1.
(0.37MB).

Distributional map of Cholomyia species.

CholomyiaBigot, 1884: 42 (also 1884: xxxvii). Type species: C. inaequipesBigot, 1884, by monotypy.

AcromiodexiaTownsend, 1931a: 335. Type species: Musca acromionWiedemann, 1824, by original designation.

References. Wulp, 1891: 246 (revision); Townsend, 1892: 275 (key to North American Tachinidae); Williston, 1908: 356 (key to North American Diptera); Coquillett, 1910: 522 (catalogue); Townsend, 1927: 222 (key to South American “muscoids”); Aldrich, 1929: 13 (notes and catalogue); Curran, 1934 (key to British Guyanan Tachinidae); Townsend, 1936: 43, 45 (key to Urodexiini genera); Townsend, 1939: 107 (redescription of Cholomyia); 97 (redescription of Acromiodexia); Guimarães, 1971: 105 (catalogue); Wood, 1987 (key to Neartic Tachinidae genera); O’Hara and Wood, 2004: 275 (catalogue); Wood and Zumbado, 2010: 1387 (key to Central American Tachinidae genera); O’Hara, 2014: 18 (list).

Description. Eyes very close to each other in males, with frons tapering towards the vertex, not approximate in females. Fronto-orbital plate well developed in profile, twice the parafacial width, at midlength. Parafacial setulose with slight pale pruinosity. Eye bare. Ocellar setae proclinate and well differentiated from the adjacent setae; postocellar setae proclinate about ¼ the length of ocellar setae; inner vertical setae straight and barely differentiated from postocular setae; outer vertical setae proclinate and long. Gena height 0.4mm in average. Genal dilation with pale pruinosity and covered with pale setulae. Facial ridge bare. Antenna usually brownish yellow, almost reaching lower facial margin and inserted below middle of eyes. Arista densely plumose with long inner dorsal trichia and with same length as postpedicel. Strong and convergent vibrissa; 1–2 developed subvibrissal setae. Palpus filiform in male, more rounded in female. Scutum with yellow or white pruinosity on presutural portion. Prosternum bare. Notopleuron with anterior seta longer than posterior seta (about twice longer). Acrostichal setae 3+1 (postsutural seta weak). Dorsocentral setae 2+3. Intra-alar setae 1+1. Proepimeral setae 2. Proepisternal setae 2. Anatertige setulose. Wing hyaline. Costal spine absent. Vein M1 ending at wing margin close to tip. Rs node setulose ventrally. Upper calypter whitish; lower calypter yellowish. Male with mid leg extremely elongate, pulvilli and tarsal claws elongated. Fore femur with row of PD and PV setae. Hind coxa bare on posterodorsal margin. Abdomen cylindrical in male (oval in females) and bicolored: syntergite I+II and tergite III usually occasionally yellow, tergite IV with light brown posterior margin (thin band) (entirely black in females), and tergite V black with two lateroventral silver pruinose areas. Syntergite I+II with lateral marginal setae. Tergites IV and V with 2 posterior marginal pair of setae and 1 median marginal pair of setae. Male terminalia with cerci enlarged, slightly convex and setulose at base. Surstylus broad, and usually slightly rounded at base. Base of pregonite sub-triangular, postgonite rod-shaped with strong setae, beginning at base of basiphallus. Distiphallus short with numerous spinules on apical membranous portion and basiphallus elongated. Ejaculatory apodeme fan-shaped.

Diagnosis. Cholomyia differs from myiophasiine Gnadochaeta mainly by the plumose arista (pubescent in Gnadochaeta), parafacial pale setulose (setulae longer and black in Gnadochaeta), notopleuron with anterior seta about twice longer than posterior seta (equal size in Gnadochaeta), mid legs elongate in males (not elongate in Gnadochaeta) and abdomen and legs almost entirely yellow (both entirely black in Gnadochaeta).

Key to species of Cholomyia (males)

  • 1.

    Mid leg approximately twice the length of the other legs (Figs. 23, 38); two katepisternal setae; postpronotum with two basal setae; abdominal syntergite I+II entirely yellow (Figs. 22, 23, 37, 38) ………………… 2

    Mid leg approximately 1.5× the length of the other legs (Figs. 3, 15); one katepisternal seta; postpronotum with 1 anterior seta and 2–3 basal setae; abdominal syntergite I+II with a light brown transverse band on posterior margin (Figs. 2, 3, 14, 15) ……………………… 3

  • 2.

    Anepisternum with 2 upward setae on anterodorsal corner. Surstylus with rounded enlargement at apex in posterior view (Fig. 29). Ejaculatory apodeme narrow at base and wide at apex (about twice the basal width), and 1.6× longer than cerci (Fig. 26) (USA, Mexico, Guatemala, Guyana, Venezuela, Colombia, Peru, Brazil) ……………… C. inaequipesBigot, 1884

    Anepisternum with 4–5 set of upward directed setae on anterodorsal corner. Surstylus with no apical enlargement in posterior view (Fig. 44). Ejaculatory apodeme narrow at base and very wide at apex (about 4.5× the basal width), 1.3× longer than cerci (Fig. 41) (Costa Rica) ………… C. zumbadoisp. nov.

  • 3.

    Thorax light brown (Figs. 3, 4), prescutum with thin white pruinosity (Fig. 2); anepisternum with 1 upward setula on anterodorsal corner; abdominal tergite III with 1 median marginal pair of setae, usually strong (Peru, Guyana, Brazil) … C. acromion (Wiedemann, 1824)

    Thorax dark brown (Figs. 14, 15), prescutum with dense white pruinosity (Fig. 14); abdominal tergite III without median marginal seta (Costa Rica, Guyana, Colombia, Brazil) … C. filipes (Walker, 1857)

C. acromion (Wiedemann, 1824)

(Figs. 2–13)

Figs. 2–9.
(0.39MB).

Cholomyia acromion (Wiedemann), ♂: 2, dorsal habitus; 3, lateral habitus; 4, head, lateral view; 5, head, frontal view. 6–9, ♂ terminalia: 6, ejaculatory apodeme, lateral view; 7, aedeagus and hypandrium, lateral view; 8, epandrium, surstylus and cerci, lateral view; 9, epandrium, surstylus and cerci, posterior view. (Legends: basiph, basiphallus; cer, cerci; distph, distiphallus; ejac apod, ejaculatory apodeme; ep, epandrium; hypd, hypandrium; phapod, phallapodeme; pregt, pregonite; posgt, postgonite; sur, surstylus). Scale bar from figures 2 to 5: 1mm; figures 6 to 9: 0.1mm.

Figs. 10–13.
(0.37MB).

Cholomyia acromion (Wiedemann), ♀: 10, dorsal habitus; 11, lateral habitus; 12, head, lateral view; 13, head, frontal view. Scale bar: 1mm.

M. acromionWiedemann, 1824: 47. Holotype ♀ (ZMUC; not examined). Type locality: “South America”. Wiedemann, 1830: 412 (redescription).

Acromiodexia acromion; Townsend, 1936: 45 (comments).

C. acromion; Emden, 1950: 203 (host record); Parker et al., 1953: 12 (host record); Parker, 1953: 54, 66 (host record; figure of puparium, posterior spiracles and cephaloskeleton of first instar); Guimarães, 1971: 105 (catalogue); Guimarães, 1977: 30 (host–parasite catalogue); O’Brien and Couturier, 1995: 234 (host record).

Diagnosis. C. acromion differs from C. inaequipes and C. zumbadoi, and it resembles more with C. filipes, by having the postpronotal lobe with 1 anterior and 2–3 basal setae, one katepisternal seta, and abdominal syntergite I+II with a narrow light brown transverse band on distal margin. Moreover, C. acromion differs from C. filipes by having the thorax light brown, prescutum with thin white pruinosity, and abdominal syntergite I+II with median marginal pair of setae.

Redescription

Male (Figs. 2–5)

Body length: 6.7mm in average (6.3–7.2mm; n=4). Wing length: 6.5mm in average (5.9–6.9mm; n=4).

  • Colouration. Genal groove yellow. Antenna yellow. Proboscis and palpus yellow. Thorax light brown, with white pruinosity on prescutum. Fore and mid legs brown, hind leg yellowish with posterior half black, and all legs with tarsus, claw and pulvillus light brown. Abdomen translucent yellow, but with a light brown thin band on posterior margin of syntergite I+II, a larger band on tergite III (about ¼), and tergite IV with black posterior margin, about distal ⅔ of width.

  • Head. (Figs. 4, 5) Frontal vitta tapering towards the apex. Gena, when seen in profile, about 0.3–0.4 times as height as eye. Gena and parafacial with white pruinosity.

  • Thorax. (Figs. 2, 3) Postpronotal lobe with 1 anterior and 2–3 basal setae (nearly aligned). Supra-alar setae 1+1. Postalar setae 2. Scutellum with 1 basal (weak), one subapical (weak) and one apical (weak or absent) pairs of setae. Anepisternum with 4–5 strong setae and without upward setulae on anterodorsal corner. Katepisternal setae 1 (posterior).

  • Wing. R4+5 setulose dorsally from Rs node until half of distance to r-m crossvein, and R1 setulose dorsally until distal 1/3. Rs node with 2–4 setulae ventrally.

  • Legs. (Fig. 3) Fore femur with row of PD and PV setae; fore tibia with 1 submedian and 2 preapical setae on PD surface. Mid leg elongate with 1.5× of length of other legs, 13.5mm in average (12.2–14.5mm; n=3); mid femur with 2 AV supramedian setae, 1 D supramedian seta; and mid tibia with 2 AD setae at apical third and 1 P seta on median third. Hind femur with a row of PD and AD setae, 1 D seta on basal third, 1 PV seta on median third and 2 preapical AD setae; and hind tibia with 1 PD seta on median third, 1 median AV seta, 1 median AD seta and 1 preapical seta on PV and on PD surfaces.

  • Abdomen. (Figs. 2, 3) Syntergite I+II with 1 to 2 pairs of lateral marginal setae. Tergite III with 1 lateral marginal and 1 median marginal pairs of setae. Tergites IV and V with 2 lateral marginal and 1 median marginal pairs of setae.

  • Male terminalia. (Figs. 6–9) Cerci tapered and well separated at apex in posterior view, curved and with tip narrowed internally in profile. Surstylus narrowing near the apex in posterior view (Fig. 9), concave and somewhat curved posteriorly in profile (Fig. 8). Base of pregonite sub-triangular, postgonite rod-shaped with strong setae, beginning at base of distiphallus (Fig. 7). Ejaculatory apodeme widening slightly at apex, and conspicuously supersized, about 1.3× length of cerci (Fig. 6).

Female (Figs. 10–13)

Body length: 5.1mm in average (4.8–5.6mm; n=3); wing length: 4.6mm in average (4.5–4.8mm; n=3). (Figs. 12–13).

  • Differs from male by the following: Fronto-orbital plate with reclinate orbital setae 2, and proclinate orbital seta 1. Ocellar setae not developed and straight, inner vertical setae well developed and reclinate; outer vertical setae developed, about ½ length of inner vertical seta. Thorax dark brown (sometimes light brown), with white pruinosity on scutum and scutellum. Postpronotal setae 2. Supra-alar setae 2. Katepisternal setae 1–2 (anterodorsal and posterodorsal). Mid leg dark brown and not elongate as in males, 6–7mm (Fig. 11). Pulvillus not elongated (Fig. 10). Abdomen oval with anterior thin band of white pruinosity on tergites II–V; syntergite I+II dark yellow, tergite III black, with anterior margin dark yellow, tergites IV and V entirely black, the latter with conspicuous pointed tip.

  • Examined material. GUYANA: Mazaruni: Cuyuni-Mazaruni, 1 ♀, 16.viii.1937, Richards & Smart col. (BMNH); “British Guiana”: [Pres. by Imp. Int. Ent. B. M. 1927], 1 ♀, F. A. Squire col. (BMNH). BRAZIL: Minas Gerais: Cambuquira, 1 ♂, 2.1941, Lopes & Gomes col. (CEIOC); Rio de Janeiro: Grajaú, 1 ♀, undated, H. Souza Lopes col. (MNRJ); idem, 1 ♂, 1.i.1939, [H. Souza] Lopes col. (MZSP); idem, 1 ♂, 22.ix.1940, [H. Souza] Lopes & Oliveira col. (BNMH); idem, 1 ♂, 1.v.1947, [H. Souza] Lopes col. (MZSP); Rio de Janeiro, 1 ♂, fevereiro.1939, [Serviço Febre Amarela, M. E. S. Bras] (MZSP).

  • Distribution. Peru, Guyana, Brazil (Minas Gerais, Rio de Janeiro, São Paulo). (Fig. 1).

C. filipes (Walker, 1857)

(Figs. 14–21)

Figs. 14–21.
(0.35MB).

Cholomyia filipes (Walker), ♂: 14, dorsal habitus; 15, lateral habitus; 16 head, lateral view; 17 head, frontal view. 18–21, ♂ terminalia: 18, ejaculatory apodeme, lateral view; 19, aedeagus and hypandrium, lateral view; 20, epandrium, surstylus and cerci, lateral view; 21, epandrium, surstylus and cerci, posterior view. Scale bar from figures 14 to 17: 1mm; figures 18 to 21: 0.1mm.

Dexia filipesWalker, 1857: 202. Holotype ♂ (BMNH; not examined). Type locality: “Brazil”.

C. filipes; Guimarães, 1971: 105 (catalogue).

Diagnosis. C. filipes differs from C. inaequipes and C. zumbadoi, and it resembles more with C. acromion, by having the postpronotal lobe with 1 anterior and 2–3 basal setae, one katepisternal seta, and abdominal syntergite I+II with a narrow light brown transverse band on distal margin. Moreover, C. filipes differs from C. acromion by having the thorax dark brown, prescutum with dense white pruinosity, and abdominal syntergite I+II without median marginal setae.

Redescription

Male (Figs. 14–17)

Body length: 6.6mm in average (5.0–7.3mm; n=8). Wing length: 6.2mm in average (5.2–6.8mm; n=8).

  • Colouration. Genal groove dark yellow. Antenna light yellow. Proboscis and palpus yellow. Thorax dark brown, with dense white pruinosity on prescutum. Fore leg with yellow femur black, mid and hind femur yellow (except for posterior margin blackish on distal ¼), and all legs tibia and tarsus black. Abdomen light yellow, but light brown thin band on posterior margin on syntergite I+II and tergite III (about ¼), and tergite IV with black posterior margin, about ⅔ of width.

  • Head. (Figs. 16, 17) Frontal vitta tapering towards the apex. Gena, when seen in profile, about 0.2–0.3 times as height as eye. Gena and parafacial with pale pruinosity.

  • Thorax. (Figs. 14, 15) Postpronotal lobe with 1 anterior and 2–3 basal setae (nearly aligned). Supra-alar setae 1+2. Postalar setae 1 or 2. Intrapostalar seta present (weak). Scutellum with 1 basal, 1 subapical (or absent), 1 discal (weak) and 1 apical (decussate) pair of setae. Anepisternum with 5–6 strong setae and with 1 upward setula on anterodorsal corner. Katepisternal seta 1 (posterior).

  • Wing. R4+5 setulose dorsally from Rs node until half of distance to r-m crossvein. R1 setulose dorsally (scarce) until distal ⅓. Rs node with 2 setae ventrally.

  • Legs. (Fig. 15) Fore femur with row of PD and PV setae; fore tibia with 1 submedian and 2 preapical setae on PD surface. Mid leg elongated, with 1.5× of length of other legs 12.3mm in average (9.2–15.8mm; n=5); mid femur with 1 submedian and 1 preapical on PD surfaces and mid tibia with 2 supramedian AD setae at apical third, 1 PV setae on median third. Hind femur with row of PD and AD setae, 1 PD seta on basal third, 1 PV seta on median third and 2 preapicals seta on AD; hind tibia with 1 PD seta on median third, 1 median AD seta, 2 preapical PV surface.

  • Abdomen. (Figs. 14, 15) Syntergite I+II with 1 pair of lateral marginal setae. Tergite III with 1 pair of median marginal. Tergites IV and V with 2 lateral marginal and 1 median marginal pairs of setae.

  • Male terminalia. (Figs. 18–21) Cerci tapered at apex, touching each other in posterior view (Fig. 21), with narrowed and subtly curved edge internally in profile (Fig. 20). Surstylus narrowing near the apex in posterior view (Fig. 21), concave and somewhat curved posteriorly in profile (Fig. 20). Base of pregonite sub-triangular, and postgonite rod-shaped, with strong setulae, beginning at base of distiphallus (Fig. 19). Ejaculatory apodeme with wide apex and conspicuously supersized, with 1.6× length of cerci (Fig. 18).

Female. Unknown.

  • Examined material. COSTA RICA: 1 ♂, 1921, P. Serre col. (MNHN). Guyana: “Brit. Guiana”: 2 ♂, 1908, K. S. Wise col. (BMNH). COLOMBIA: Florencia: 1 ♂, 10.ix.2010, Yardany Ramos-Pastrana col. (UAM-E). BRAZIL: Rondônia: Monte Negro, Setor Chacareiro, S10°16′21.4″ W63°20′45.2″, 1 ♂, 10.ii.2012, Malaise, Lamas, Nihei & eq. col. [SISBIOTA CNPq/FAPESP] (MZSP); Amazonas: Manaus, 1 ♂, vi.1955, Elias & Roppa col. (MNRJ); Pará: Oriximiná, 1 ♂, 12–13.i.1968, [Exp. Perm. Amaz.] (MZSP); Mato Grosso: Chap.[Chapada] dos Guimarães, Vale da Benção, S15°25′10.7″ W055°47.22′9″, 1 ♂, 17.i.2012, Coleta Manual, Lamas, Nihei & eq. col. [SISBIOTA CNPq/FAPESP] (MZSP); idem, 1 ♂, 19.i.2012, Coleta Manual, Lamas, Nihei & eq. col. [SISBIOTA CNPq/FAPESP] (MZSP); Goiás: Anápolis, 1 ♂, VII.1955, [Serviço Febre Amarela, M. E. S. Bras] (possible colector) (MZSP); Mato Grosso do Sul: Serra da Bodoquema, Fazenda Califórnia, Rio da Gruta, S20°42′28″ W055°51′07″, 1 ♂, 09.xii.2011, Nihei, Gudin, Sato, Moll col. [SISBIOTA CNPq/FAPESP] (MZSP).

  • Distribution. Costa Rica, Guyana, Colombia (Florencia), Brazil (Rondônia, Amazonas, Pará, Mato Grosso, Goiás, Mato Grosso do Sul). (Fig. 1)

  • Remarks. José H. Guimarães during his studies at USNM in 1967 made some personal notes based on records from J.M. Aldrich performed in the BMNH in 1929, which he reports D. filipes as synonym of C. inaequipes. However, Guimarães (1971) in the Neotropical catalogue still kept C. filipes as a valid species. In our review, we recognized both C. filipes and C. inaequipes as separate and valid species. In fact, C. filipes shares more similarities with C. acromion than with C. inaequipes.

C. inaequipesBigot, 1884

(Figs. 22–36)

Figs. 22–29.
(0.48MB).

Cholomyia inaequipes Bigot, ♂: 22, dorsal habitus; 23, lateral habitus; 24 head, lateral view; 25 head, frontal view. 26–29, ♂ terminalia: 26, ejaculatory apodeme, lateral view; 27, aedeagus and hypandrium, lateral view; 28, epandrium, surstylus and cerci, lateral view; 29, epandrium, surstylus and cerci, posterior view. Scale bar from figures 22 to 25: 1mm; figures 26 to 29: 0.1mm.

Figs. 30–36.
(0.45MB).

Cholomyia inaequipes Bigot, ♀: 30, dorsal habitus; 31, lateral habitus; 32 head, lateral view; 33 head, frontal view. 34, ♀ terminalia; 35, larvae, and 36, cephaloskeleton. Scale bar from figures 30 to 33: 1mm; figures 34 to 35: 0.1mm; figure 36: 0.05mm.

Musca longipesFabricius, 1805: 398 (preoccupied Scopoli, 1763). Holotype ♂ (ZMUC; not examined). Type locality: “Brazil”.

C. inaequipesBigot, 1884: 42 (also 1884: xxxvii). Lectotype ♂ (BMNH; not examined). Type locality: “Mexico”.

Thelairodes basalisGiglio-Tos, 1893:3. Holotype ♀ (MZUT; not examined). Type locality: “Mexico”.

C. nigricepsWilliston, 1908: 353, Fig. 146 (figure of head and legs). Holotype ♂ (whereabouts unknown). Type locality: “unknown”.

Metadexia flavipes Johnson in Coquillet: 1905:78. Nomen nudum.

References. Wulp, 1891: 247 (redescription of C. inaequipes); Brauer, 1897: 365 (inaequipes as synonym of longipes); Johnson in Coquillett, 1905:78 (C. flavipes as junior synonym of T. basalis); Pierce, 1908: 381 (host record); Johnson, 1912: 102 (notes of C. longipes); Quaintance and Jenne, 1912: 150 (notes, hosts and figure of puparium); Pierce et al., 1912: 78 (host record); Brooks, 1922: 7, 11, 13, 16 (host record); Mutchler and Weiss, 1925: 10, 13 (host records); Aldrich, 1929: 13 (redescription and comments); Snapp, 1930: 77, 79 (host record); Townsend, 1931b: 93 (C. inaequipes as synonym of C. longipes); Townsend, 1936: 45 (comments about C. inaequipes); Curran, 1934: 502 (comments on female of C. inaequipes); Fattig, 1949: 12 (host record); Armstrong, 1958: 17 (host record); Patton, 1958: 36 (distributional records); Gibson, 1964: 525 (host record); Guimarães, 1971: 105 (catalogue); Arnaud, 1978: 15, 144, 611 (host–parasite catalogue); Maier, 1980: 61 (host record); O’Hara and Wood, 2004: 275 (catalogue); Jenkins et al., 2006: 438 (host record); Wood and Zumbado, 2010: 1401 (comments).

Diagnosis. C. inaequipes differs from C. acromion and C. filipes, and it resembles more with C. zumbadoi, by having the mid leg approximately twice the length of the other legs, two katepisternal setae, postpronotum with two basal setae and abdominal syntergite I+II entirely yellow. Moreover, C. inaequipes differs from C. zumbadoi by having the anepisternum with 2 upward setae on anterodorsal corner, surstylus with rounded enlargement at apex in posterior view, ejaculatory apodeme narrow at base and wide at apex (about twice the basal width), about 1.6× longer than cerci and pregonites without narrowed basal portion.

Redescription

Male (Figs. 22–25)

Body length: 8mm in average (7.1–8.8mm; n=30). Wing length: 8.2mm in average (7.1–9mm; n=30).

  • Colouration. Genal groove dark yellow. Proboscis and palpus pale yellow. Antenna yellow. Thorax dark brown, with pruinosity usually yellow on prescutum. Fore and hind legs with femur yellow (except mid distal region black), and all legs with tibia and tarsus black. Abdomen yellow, but light brown thin band on posterior margin of tergite III (about ¼) and tergite IV with black posterior margin, about distal ⅔ of width.

  • Head. (Figs. 24, 25) Frontal vitta narrow. Gena, when seen in profile, about 0.4–0.5 times as height as eye. Gena and parafacial with white pruinosity. Parafacial with thin pale pruinosity.

  • Thorax. (Figs. 22, 23) Postpronotal lobe with 2 basal setae. Supra-alar setae 1+1. Postalar setae 1 or 2. Scutellum 1 one basal, 1 discal (weak or absent), 1 subapical and 1 apical (usually decussate) pairs of setae. Anepisternum with 5–6 strong setae and with 2 upward setulae on anterodorsal corner. Katepisternum 2 (posterior and anterior).

  • Wing. R4+5 setulose dorsally from Rs node until half of distance to r-m crossvein, and R1 setulose dorsally until distal ⅓ (rarely beyond). Rs node with 3–4 setae ventrally.

  • Legs. (Fig. 23) Fore femur with row of PD and PV setae, fore tibia with 2 PD setae at apical third and 1 PD on median third, and 1 D and 1 PV preapical setae. Mid leg very elongate twice longer than other legs, 18.8mm in average (14.9–20.5mm; n=30); mid femur with 4 AD setae supramedian and 1 PV seta preapical, mid tibia with 2 supramedian AD setae and preapicals 1 D, 1V and 1 P. Hind femur with a row of AV and AD setae, 1 submedian D seta, and 1 preapical P seta; tibia with 1 supramedian AD seta, 1 median AD seta, 1 median AV seta, 1 submedian D seta and 1 DV preapical seta.

  • Abdomen. (Figs. 22, 23) Syntergite I+II with 2 pairs of lateral marginal setae. Tergite III (rarely) with 1 pair of lateral marginal seta. Tergites IV and V with 2 pairs of lateral marginal setae and 1 pair of median marginal setae.

  • Male terminalia. (Figs. 26–29) Cerci tapered at apex, touching each other or not, in posterior view (Fig. 29), with a subsequent narrowing and internally curved in profile (Fig. 28). Surstylus with a subsequent narrowing near the apex in posterior view (Fig. 29), and somewhat curved posteriorly in profile (Fig. 28). Base of pregonite sub-triangular, postgonite rod-shaped with strong setae, beginning at base of basiphallus (Fig. 27). Ejaculatory apodeme wide at apex, and conspicuously supersized, about 1.6× length of cerci (Fig. 26).

Female (Figs. 30–33)

Body length: 8mm in average (8–8.1mm; n=2); wing length: 6.6mm in average (6.5–6.8mm; n=2).

  • Differs from male by the following: (Figs. 32, 33) Fronto-orbital plate with reclinate orbital setae 2, and proclinate orbital setae 1. Ocellar setae not developed and straight, inner vertical setae well developed and reclinate; outer vertical setae developed about ½ length of inner vertical seta. (Figs. 30, 31) Prescutum with thin white pruinosity, denser on posterior half. Mid leg dark brown and not elongate as in males, 9.2mm (n=1). Pulvillus not elongated. Abdomen oval, not translucent; yellow but abdominal tergite III (with 1 developed marginal median seta) only posterior half black, tergites IV and V entirely black, the latter with conspicuous pointed tip.

  • Female terminalia. (Fig. 34) Tergite 6 dorsally with 2 pairs of spiracles, 6th spiracle on membrane ventrally and 7th spiracle near ventral end of tergite, with several setae. Tergite 7, with a few setae dorsally and laterally. Tergite 8 as narrow strip, with one pair of setae laterally. Sternite 5 subrectangular with some scarce setae. Sternite 6 well developed as a complete ventral shield, with several small setae. Sternite 7 with setae on posterior margin and without small setae. Sternite 8 with one long and one small seta on posterior margin. Sternite 10 without lingulae, but with lateral projections dorsally and several scarce setae. Cerci well developed, sub-circular, with several setae apically. Syntergite 9+10 absent.

First instar larva (Fig. 35)

Body length: 1.029mm in average (1.008–1.062mm). [70–80 larvae were measured from two ♀]

  • Colourless except for the pigmented cephaloskeleton and ventral cuticular spines on each segment posterior margin (without cuticular scales). Along all the segments, complete bands of minute spines (encircle all the segment). Antenna not visible. Posterior spiracle as a simple round opening in the 11th segment, with short internal tubule visible through the cuticle. The 12th segment is prolonged into a conical tail. Cephaloskeleton (Fig. 36) with fine mouth hook, slightly ventrally curved and sharply pointed, with well developed accessory sclerite. Long intermediate region, with salivary gland thin and long (almost as long as intermediate region), with posterior portion enlarged. Dorsal cornu larger and somewhat longer than ventral cornu, and both less pigmented than anterior parts of skeleton.

  • Examined material. UNITED STATES OF AMERICA: Washington: Dayton: 1 ♂, no date, C.W. Johnson col. (MZSP). COLOMBIA: Florencia: 2 ♂, 21.x.2011, Yardany Ramos-Pastrana col. (UAM-E). PERU: Huarochirí: 1500m, 1 ♂, 1.5.1970, C. Kofytkowski col. (MZSP). BRAZIL: Rondônia: Vilhena: 1 ♀, xi.1973, Alvarenga & Roppa col. (MNRJ); Amazonas: Manaus, 1 ♂, 4.iv.1946, Elias & Roppa col. (MNRJ); R. Ita: 1 ♂, vi.1950, J. C. M. Carvalho col. (MNRJ); Pará: Canidé: Gurupi, 1 ♂, 27.ii.1966, Malkin col. (MZSP); Concórdia do Pará: Santana, Floresta dos Macacos, 1 ♂, ix.1969, [Exp. Ferm. Amaz.] (MZSP); Bahia: Cachoeira, Faz Vila Real: 1 ♂, 23.vii.2007, E. Alvin & et al. col. (USFS); Mato Grosso: Fazenda Primavera, Rio Paraná: 1 ♂, x.1954, Rabello col. (MZSP); Tapirapé [Serra do Urubu Branco–possibly]: 1 ♂, i.1940, Carvalho col. (MNRJ); Goiás: Anápolis, 1 ♂, iv.1.1936 (no collector data), Goiânia, 1 ♂, viii.1943, Freitas e Nobre col. (MZSP); Mato Grosso do Sul: Corumbá, 4 ♂, iii.1949, M. P. Barreto col. (MZSP); Minas Gerais: Passa Quatro, 3 ♂, 1903, E. R. Wagner (MNHN), Lassance, 1 ♂, 20 a 31.i.1939, Martins, Lopes e Mangabeira col. (MZSP); Rio de Janeiro: Angra dos Reis (Japuhyba): 1 ♂, xii.1940, Oliveira & Venfel (CEIOC); Barra da Tijuca: 1 ♂, 19.6.1994, Rocha Barros Santos Machado col. (MNRJ); Itatiaia: 5 ♂, iv.1945, M. P. Barretos, col., idem, 24.i.1934, J. F. Zikán. col.; Nova Friburgo, Mury: 1 ♂, xii.1976, 3 ♂, 1/15.iv.1964, 3 ♂, 1/15.iv.1964, 3 ♂, 22–24.xii.1964, 1 ♂, 1–31.Jan.1965, 16 ♂, xii.1965, 1 ♂, 1–2.Nov.1970, 1 ♂, iii.1972, 5 ♂, xii.1974, 1 ♂, xii. 1975, 1 ♂, xii.1976, Gred & Guimarães, cols. (MZSP); Petrópolis: 6 ♂, xi.1929 (MZSP), idem, 1 ♂, 4.69, H.S. Lopes col. (MNRJ); São Paulo: Barueri: 1 ♂, 23.ii.1955, 1 ♂, 23.iv.1957, 1 ♂, 16.ii.1965,1 ♂, 16.i.1966, K. Lenko col. (MZSP); Cássia dos Coqueiros, Cajuru: 3 ♂, x.1954, M.P. Barreto col. (MZSP), idem, 1 ♂, xi.1954, M. P. Barreto col. (MZSP); Mogi das Cruzes: 1 ♂, nov.1939, M. Carrera col. (MZSP); Guarujá: 1 ♂, 25/30.i.1941, M. Carrera col. (MZSP), idem, 1 ♂, i.1942, M. Carrera col. (MZSP); Salesópolis: Estação Biológica de Boracéia, 850m: 1 ♂, 30.i.1968, Rabello col., idem, 9 ♀, 21.v–28.vi.2008 (malaise), Nihei, Fiqueiredo & Andrade col. (MZSP); São Paulo: 1 ♂, 9.iv.1933, M. T. Leite col. (MZSP); Santa Catarina: Nova Teutônia [27°11′S 52°23′W]: 2 ♂, 14.5.1936, Fritz Plaumann col. (MZSP), idem, 1 ♂, 10.10.1937, Fritz Plaumann col. (MZSP).

  • Distribution. United States of America (Arkansas, Arizona, California, Connecticut, Florida, Gainesville, Indiana, Kansas, Kentucky, Louisiana, Massachusetts, Maryland, Missouri, New Jersey, New York, Ohio, Pennsylvania, Texas, Virginia, Washington, West Virginia and Wisconsin), Mexico (Vera Cruz), Guatemala, Guyana, Venezuela, Colombia (Florencia), Peru, Brazil (Pará, Mato Grosso, Goiás, Mato Grosso do Sul, Minas Gerais, Rio de Janeiro, São Paulo, Santa Catarina). (Fig. 1)

  • Remarks. This species presents the widest geographical distribution, ranging from northern USA (Wisconsin) to southern Brazil (Santa Catarina) (Fig. 1), and has the most extensive list of host records.

  • M. longipes Fabricius is synonym of C. inaequipes Bigot. Wulp (1891: 247), commented: “(…) it is possible that [C. inaequipes] is M. longipes Fabri. (…)”. Brauer (1897: 365) stated the possible synonym between both species, and Aldrich (1929: 13) formally proposed such synonymy: “(…) inaequipes Bigot, new, is equals M. longipes Fabricius preoccupied.” Townsend (1931: 93) also agreed with this synonymy: “Musca longipes Fab. preoc. (…) is Cholomyia inaequipes Big. (…)”, and reproduced this in his Manual of Myology (Townsend, 1939). However, some other authors placed C. longipes as synonym of C. acromion (e.g., Parker, 1953; Guimarães, 1971). Aldrich (1929) and Townsend (1931) examined and compared the types, while Parker (1953) and Guimarães (1971) did not mention about types examination.

C. zumbadoisp. nov.

(Figs. 37–44)

Figs. 37–44.
(0.29MB).

Cholomyia zumbadoi sp. nov., ♂: 37, dorsal habitus; 38, lateral habitus; 39 head, lateral view; 40 head, frontal view. 41–44, ♂ terminalia: 41, ejaculatory apodeme, lateral view; 42, aedeagus and hypandrium, lateral view; 43, epandrium, surstylus and cerci, lateral view; 44, epandrium, surstylus and cerci, posterior view. Scale bar from figures 37 to 40: 1mm; figures 41 to 44: 0.1mm.

Diagnosis. C. zumbadoi differs from C. acromion and C. filipes, and it resembles more with C. inaequipes, by having the mid leg approximately twice the length of the other legs, two katepisternal setae, postpronotum with two basal setae and abdominal syntergite I+II entirely yellow. Moreover, C. zumbadoi differs from C. inaequipes by having the anepisternum with 4–5 upward setae on anterodorsal corner, surstylus without concavity and curved externally in profile view, ejaculatory apodeme narrow at base and very wide at apex (about twice the basal width), about 1.3× longer than cerci and pregonites with basal portion narrowed.

Description

Male (Figs. 37–40)

Body length: 8.3mm in average (8.0–8.8mm; n=4). Wing length: 8.4mm in average (8.0–8.5mm; n=4).

  • Colouration. Genal groove dark yellow. Antenna yellow. Proboscis and palpus pale yellow. Prescutum usually with yellow pruinosity. Thorax dark brown. Fore leg with yellow femur, mid and hind femur yellow (except for blackish posterior margin about distal ¼), and all legs with tibia and tarsus black. Abdomen yellow, but light brown thin band on posterior margins on tergite III (about ¼) and tergite IV with black posterior margin, about distal ⅔ of width.

  • Head. (Figs. 39, 40) Frontal vitta narrow. Gena, when seen in profile, about 0.4–0.5 times as height as eye. Gena and parafacial with thin pale pruinosity.

  • Thorax. (Figs. 37, 38) Postpronotal lobe with 2 basal setae. Supra-alar setae 1+2. Postalar setae 1 to 2. Scutellum with one basal, one discal (weak or absent), one subapical and one apical (generally decussate) pairs of setae. Anepisternum setae with 5–6 of strong setae and with 4–5 developed upward setae on anterodorsal corner. Katepisternal setae 2 (anterior and posterior).

  • Wing. R4+5 setulose dorsally from Rs node until half of distance to r-m crossvein and R1 fully setulose dorsally (scarce in the median region). Rs node with 5 setulae ventrally.

  • Legs. (Fig. 38) Fore femur with row of PD and PV setae, fore tibia with 2 PD and 1 PV setae on median third, and 1 D and 1 PV preapical setae. Mid leg very elongate, twice longer than other legs, 20.5mm in average (20.3–20.8mm; n=4), mid femur with 4 supramedian AD setae, 1 preapical PV seta; mid tibia with 2 supramedian AD setae, and 1 D, 1V and 1 P preapical setae. Hind femur with a row of AV (usually) and AD setae, 1 submedian D seta, and 1 preapical P seta; hind tibia with 1 supramedian AD seta, 1 median AD seta, 1 median AV seta, 1 submedian D seta and 1 DV preapical seta.

  • Abdomen. (Figs. 37, 38) Syntergite I+II with 2 pairs of lateral marginal setae. Tergite III (rarely) with 1 lateral marginal pair of seta. Tergites IV and V with 2 pairs of lateral marginal setae and 1 pair of median marginal setae.

  • Male terminalia. (Figs. 41–44) Cerci tapered at apex, touching each other or not, in posterior view (Fig. 44), narrowed tip without curvature in profile (Fig. 43). Surstylus without narrowing near the apex, in posterior view (Fig. 44), slightly curved posteriorly, with absence of concavity, in profile (Fig. 43). Base of pregonite sub-triangular with basal portion narrowed, postgonite rod-shaped with strong setae, beginning at the base of distiphallus (Fig. 42). Ejaculatory apodeme very wide at apex, with conspicuously supersized, about 1.3× length of cerci (Fig. 41).

Female. Unknown.

  • Type material examined. Holotype ♂ (INBio) labelled as “COSTA RICA, Prov. Alajuela, PN/Volcán Tenório, Albergue Heliconias,/send. Mirador 1000m. 16. Jul. al 04/.ago.2000, J. D. Guitiérrez, Manual. L N_298575_423400 #62544″; “INB0003304285/INBIOCRI COSTA RICA”; “Cholomyia zumbadoi/Santis, M. det./Out – 2013”; “Holótipo” [red label with black borders].

  • Paratypes. ♂ (INBio). “COSTA RICA, Prov. Alajuela, PN/Volcán Tenório, Alberue Heliconias,/send. Mirador 1000m. 16. Jul. al 04/.ago.2000, J. D. Guitiérrez, Manual. L N_298575_423400 #62544″; “INB0003304285/INBIOCRI COSTA RICA”; “Parátipo” [green label with black boards]. ♂ (INBio). “COSTA RICA, Prov. Alajuela, Upala,/Bijagua, Falda, N.E./V. Tenório, 700–1200m, 28.may.2001, M. Alfro/Libre L_N_295900_426650 #63212″; “INB0003333473/INBIOCRI COSTA RICA”; “Parátipo” [green label with black boards]. ♂ (INBio). “COSTA RICA, Prov. Alajuela, Upala,/Bijagua, Sector Alto Los Brenes. 700/m. 26 ABR – 11 MAY 2001. J. D./Gutiérrez. Red com Aguameil./L_N_300600_422800 #62187”; “INB0003333473/INBIOCRI COSTA RICA”; “Parátipo” [green label with black boards].

  • Type locality. Costa Rica, Alajuela, P.N. Volcán Tenório.

  • Distribution. Costa Rica (Alajuela) (Fig. 1)

  • Etymology. This species is named in honour of Dr. Manuel A. Zumbado, curator of Diptera at the Instituto Nacional de Biodiversidad (INBio).

Host–parasite records

On the systematic placement of Cholomyia

In a detailed comparative study of the male terminalia of Tachinidae, Tschorsnig (1985: 99) characterized the terminalia of members of the tribe Myiophasiini, where he pointed out: (1) a large epandrium (in Tschorsnig, 1985: Fig. 46 can be noticed that the apical region is short and not tapered in lateral view); (2) cerci separated at apex (by his Fig. 46, the proximal region is not fully fused); (3) the surstylus is narrow and curved internally; (4) pregonite with a plate-like form with setulae and in front of basiphallus (Tschorsnig, 1985: Fig. 133); and (5) the ejaculatory apodeme is large and fan-shaped. Tschorsnig's observations were based on two species of Angiorhina, which is currently a junior synonym of Gnadochaeta (sensu O’Hara and Wood, 1998).

The male terminalia of Cholomyia species mostly agree with the tribal characterization provided by Tschorsnig (1985): (1) the epandrium is large with its elongated apical region and tapered in posterior view; (2) cerci are separated at the apex but with fully fused proximal region; (3) the surstylus is narrow, but slightly curved internally (although in C. zumbadoisp. nov. it is rectilinear); (4) the pregonite has a sub-triangular form, and is in front of basiphallus, but it lacks setulae, and is expanded on the distal region; and finally, (5) the ejaculatory apodeme is large and also has a fan shape. Therefore, based on your detailed analysis of Cholomyia species, including male terminalia, the systematic placement of Cholomyia, transferred from the Urodexiini (sensu Guimarães, 1971) to the Myiophasiini (sensu O’Hara and Wood, 2004) is supported and confirmed herein.

Besides the external morphology and male terminalia, characters from immature stage also support the placement of Cholomyia into the Myiophasiini. By comparing the 1st instar larva and cephaloskeleton of C. inaequipes with those of Gnadochaeta globosa (Townsend, 1892), illustrated and described by Townsend (1942) and Bissel (1945), we could observe some resembling features shared by both species. The 1st instar larva of G. globosa has the ventral cuticular spines on each segment of the posterior margin, the posterior spiracles located at the 11th segment, lacks a spine-like microtrichia in all segments, and the last segment is prolonged into a conical tail, exactly as C. inaequipes (Fig. 35). The cephaloskeleton of G. globosa has a down-curved and sharply pointed mouth hook, also present in C. inaequipes (Fig. 36) and in C. acromion (Parker, 1953: Fig. 60). The well-developed accessory sclerite and the salivary gland with the same posterior enlargement strongly resemble those observed in C. inaequipes (Fig. 36).

In the first cladistic analysis of the family performed by Cerretti et al. (2014), the tribe Myiophasiini was only represented by some species of Gnadochaeta Macquart (G. puncticeps Zetterstedt, 1859 and some spp.), and was recovered as sister group of three genera of Palpostomatini, forming a monophyletic group, “clade B” (Gnadochaeta+Palpostomatini) of Cerretti et al. (2014). The only synapomorphy that supports this group is the presence of a membranous medial surface of male pregonite (105:1). Despite regarding the presence of this membrane as doubtful (we believe this could it be a weaker sclerotization), we considered this character state as present in Cholomyia. Although the taxonomic sampling was extremely biased by Palearctic taxa and the Myiophasiini was underrepresented by a single genus in Cerretti et al. (2014), the genus Cholomyia seems to be placed in the clade B. However, not conclusively in Myiophasiini because the autapomorphy of Gnadochaeta, the presence of a posteromedial connection between pregonites (108:1), that could be a synapomorphy of this tribe, is not present in Cholomyia (even though the two homoplasies for the Palpostomatini are not present in the genus Cholomyia).

Conflicts of interest

The authors declare no conflicts of interest.

Acknowledgements

We would like to thank the curators Carlos Lamas (MZSP), Christophe Daugeron (MNHN), Freddy Bravo (UEFS), Jane Costa (CEIOC), Manuel Zumbado (INBio), Márcia Couri (MNRJ), Nigel Wyatt (BMNH) and Yardany Ramos-Pastrana (UAM-E), for the loan of material. Thanks to Rodrigo Dios and two anonymous reviewers for suggestions on an earlier version of this manuscript. Financial support from CNPq (Proc. No. 101723/2013-0) and FAPESP (Proc. No. 2013/05131-6). This work was partially funded by the SISBIOTA-DIPTERA research project (CNPq Proc. No. 563256/2010-9, FAPESP Proc. No. 2010/52314-0).

References
Aldrich, 1929
J.M. Aldrich
Further studies of types of American Muscoid flies in the collection of the Vienna Natural History Museum
Proc. U. S. Natl. Mus., 74 (1929), pp. 1-34
Armstrong, 1958
T. Armstrong
Life-history and ecology of the plum curculio, Conotrachelus nenuphar (Hbst.) (Coleoptera: Curculionidae), in the Niagara Peninsula, Ontario
Can. Entomol., 90 (1958), pp. 8-17
Arnaud, 1978
P.H. Arnaud Jr.
A Host–Parasite Catalog of North American Tachinidae (Diptera)
United States Department of Agriculture. Miscellaneous Publication, (1978)
Bigot, 1884
J.M.F. Bigot
[Description d’un nouveau genre et d’une nouvelle espèce de Diptères de la famille des Dexidae]. Bulletin des séances et bulletin bibliographique de la Société entomologique de France 4, 42. [Note: Also published in 1884, Bulletin des séances et bulletin bibliographique de la Société entomologique de France (ser. 6) 4: xxv–xxvii]
(1884)
Bissel, 1945
T.L. Bissel
Myiophasia globosa (tns.), tachinid parasite of the cowpea curculio
Ann. Entomol. Soc. Am., 38 (1945), pp. 417-440
Brauer, 1897
F. Brauer
Beiträ ge zur Kenntniss der Muscaria schizometopa und Beschreibung von zwei Hypoderma-Arten
Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Classe, 106 (1897), pp. 329-382
Brooks, 1922
F.E. Brooks
Curculios that attack the young fruits and shoots of Walnut and Hickory
U.S. Dep. Agric. Bull., 1066 (1922), pp. 1-16
Cantrell, 1988
B.K. Cantrell
The comparative morphology of the male and female postabdomen of the Australian Tachinidae (Diptera) with descriptions of some first-instar larvae and pupae
Invertebr. Taxon., 2 (1988), pp. 81-221
Cerretti et al., 2014
P. Cerretti,J.E. O’Hara,D.M. Wood,H. Shima,D.J. Inclan,J.O. Stireman
Signal through the noise? Phylogeny of the Tachinidae (Diptera) as inferred from morphological evidence
Syst. Entomol., 39 (2014), pp. 335-353
Coquillett, 1910
D.W. Coquillett
The type species of the North American genera of Diptera
Proc. U. S. Natl. Mus., 37 (1910), pp. 499-647
Courtney et al., 2000
G.W. Courtney,B.J. Sinclair,R. Meier
Morphology and terminology of Diptera larvae
pp. 85-161
Cumming and Wood, 2009
J.M. Cumming,D.M. Wood
Morphology and terminology
pp. 9-50
Curran, 1934
C.H. Curran
The Diptera of Kartabo, Bartica district, British Guiana, with descriptions of new species from other British Guiana localities
Bull. Am. Mus. Nat. Hist., 6 (1934), pp. 287-532
Emden, 1950
F.I. Van Emden
Dipterous parasites of Coleoptera
Entomol. Mon. Mag., lxxxvi (1950), pp. 182-206
Fabricius, 1805
C.F. Fabricius
Systema antliatorum secundum ordines, genera, species
Brunsvigae, (1805)
Fattig, 1949
P.W. Fattig
The Larvaevoridae (Tachinidae) or parasitic flies of Georgia
Emory Univ. Mus. Bull., 8 (1949), pp. 1-40
Gibson, 1964
L.P. Gibson
Biology and life history of acorn-infesting weevils of the genus Conotrachelus (Coleoptera: Curculionidae)
Ann. Entomol. Soc. Am., 57 (1964), pp. 521-526
Giglio-Tos, 1893
E. Giglio-Tos
Diagnosi di nuovi generi e di nuove specie di Ditteri. VIII
Bollettino dei Musei di Zoologia ed Anatomia Comparata della Reale Università di Torino, 8 (1893), pp. 1-11
Guimarães, 1971
J.H. Guimarães
Family Tachinidae
pp. 1-333
Guimarães, 1977
J.H. Guimarães
Host–parasite and parasite–host catalogue of South American Tachinidae (Diptera)
Arq. Zool., 28 (1977), pp. 1-131
Johnson, 1912
C.W. Johnson
Dipterological notes
Psyche, 19 (1912), pp. 102-103
Jenkins et al., 2006
D.A. Jenkins,R.F. Mizell III,D. Shapiro-Ilan,T. Cottrell,D. Horton
Invertebrate predators and parasitoids of plum curculio, Conotrachelus nenuphar (Coleoptera: Curculionidae) in Georgia and Florida
Fla. Entomol., 89 (2006), pp. 435-440
Maier, 1980
C.T. Maier
Quince curculio Conotrachelus crataegi (Coleoptera, Curculionidae) developing in apple a new host in southern New England, USA
Proc. Entomol. Soc. Wash., 82 (1980), pp. 59-62
Mutchler and Weiss, 1925
A.J. Mutchler,H.B. Weiss
New Jersey Department Agriculture, (1925)pp. 1-22
O’Brien and Couturier, 1995
C.W. O’Brien,G. Couturier
Two new agricultural pest species of Conotrachelus (Coleoptera: Curculionidae: Molytinae) in South America
Ann. Soc. Entomol. Fr., 3 (1995), pp. 227-235
O’Hara, 2014
J.E. O’Hara
World genera of the Tachinidae (Diptera) and their regional occurrence. Version 8.0. PDF document
(2014)
O’Hara and Wood, 1998
J.E. O’Hara,D.M. Wood
Tachinidae (Diptera): nomenclatural review and changes, primarily for America north of Mexico
Can. Entomol., 130 (1998), pp. 751-774
O’Hara and Wood, 2004
J.E. O’Hara,D.M. Wood
Catalogue of the Tachinidae (Diptera) of America north of Mexico
Mem. Entomol., 18 (2004), pp. 1-410
Patton, 1958
C.N. Patton
A catalogue of the Larvaevoridae of Florida
Fla. Entomol., 41 (1958), pp. 29-39
Parker, 1953
H.L. Parker
Bollettino del Laboratorio di Entomologia Agraria Filippo Silvestri, (1953)pp. 45-73
Parker et al., 1953
H.L. Parker,P.A. Berry,A.S. Guido
Host–parasites list of insects reared in the South American parasites laboratory during period 1940–1946
Revista de la Associación (Federación) Rural del Uruguay (Ingenieros Agrónomos), 92 (1953), pp. 1-101
Pierce, 1908
W.D. Pierce
A list of parasites known to attack American Rhynchophora
J. Econ. Entomol., 1 (1908), pp. 380-396
Pierce et al., 1912
W.D. Pierce,R.A. Cushman,C.E. Hood
The insect enemies of the cotton boll weevil
U.S. Department Agriculture, Bureau of Entomology Bulletin, n. ser. 100, (1912)pp. 1-99
Quaintance and Jenne, 1912
A.L. Quaintance,E.L. Jenne
The plum curculio. Bulletin of the United States Department of Agriculture
Bureau Entomol., 103 (1912), pp. 1-250
Scopoli, 1763
G.A. Scopoli
Entomologia Carniolica exhibens insecta Carnioliae indigena et distributa in ordines, genera, species, varietates: methodo Linnaeana
(1763)
Snapp, 1930
I.O. Snapp
Life History and Habits of the Plum Curculio in the Georgia Peach Belt. Technical Bulletin
United States Department of Agriculture 188, (1930)pp. 1-90
Stuckenberg, 1999
B.R. Stuckenberg
Antennal evolution in the Brachycera (Diptera), with a reassessment of terminology relating to the flagellum
Studia Dipterol., 6 (1999), pp. 33-48
Townsend, 1892
C.H.T. Townsend
The North American Genera of Calyptrate Muscidæ. Paper III
Trans. Am. Entomol. Soc., 19 (1892), pp. 273-278
Townsend, 1927
C.H.T. Townsend
Synopse dos generos muscoideos da região húmida tropical da America, com generos e especies novas
Rev. Mus. Paul., 15 (1927), pp. 203-385
Townsend, 1931a
C.H.T. Townsend
New genera and species of American oesitromuscoid flies
Rev. Entomol., 1 (1931), pp. 313-354
Townsend, 1931b
C.H.T. Townsend
Notes in American oestromuscoid types
Rev. Entomol., 1 (1931), pp. 65-104
Townsend, 1936
C.H.T. Townsend
Manual of Myology. Part IV
Charles Townsend & Filhos, (1936)
Townsend, 1939
C.H.T. Townsend
Manual of Myology. Part IX
Charles Townsend & Filhos, (1939)
Townsend, 1942
C.H.T. Townsend
Manual of Myology. Part XII
Charles Townsend & Filhos, (1942)
Tschorsnig, 1985
H.P. Tschorsnig
Taxonomie forstlich wichtiger Parasiten: Untersuchungen zur Struktur des mänlichen Postabdomens der Raupenfliegen (Diptera, Tachinidae)
Stuttgarten Beiträge zur Naturkunde, 383 (1985), pp. 1-137
Walker, 1857
F. Walker
Characteres of undescribed Diptera in collection of W. W. Saunders
Trans. R. Entomol. Soc. Lond., (1857), pp. 190-235
Wiedemann, 1824
C.R.W. Wiedemann
Munus rectoris in Academia Christiana Albertina aditurus analecta entomologica ex Museo Regio Havniensi
Kiliae [= Kiel], (1824)
Wiedemann, 1830
C.R.W. Wiedemann
Aussereuropäische zweiflüge Insecten, Hamm
(1830)
Williston, 1908
S.W. Williston
Manual of North American Diptera
3rd edition, J.T. Hathaway, (1908)
Wood and Zumbado, 2010
D.M. Wood,M.A. Zumbado
Tachinidae (Tachinid flies, parasitic flies)
pp. 1343-1417
Wulp, 1891
F.M. Van der Wulp
Family Muscidae (cont.)
Biologia Centrali-Americana,
Corresponding author. (Marcelo Domingos de Santis mrclsantis@gmail.com)
Copyright © 2016. Sociedade Brasileira de Entomologia
Rev Bras Entomol 2016;60:217-26 - Vol. 60 Núm.3 DOI: 10.1016/j.rbe.2016.05.004