Research Article |
Corresponding author: Diego J. Inclán ( diegojavier.inclanluna@studenti.unipd.it ) Academic editor: Pierfilippo Cerretti
© 2014 Diego J. Inclán, John O. Stireman.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Inclan DJ, Stireman III JO (2014) A new species and synonymy of the Neotropical Eucelatoria Townsend and redescription of Myiodoriops Townsend. ZooKeys 464: 63-97. https://doi.org/10.3897/zookeys.464.8155
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The New World tropics represents the most diverse region for tachinid parasitoids (Diptera: Tachinidae), but it also contains the most narrowly defined, and possibly the most confusing, tachinid genera of any biogeographic region. This over-splitting of genera and taxonomic confusion has limited progress toward our understanding the family in this region and much work is needed to revise, redefine, and make sense of the profusion of finely split taxa. In a recent analysis of the Neotropical genus Erythromelana Townsend, two species previously assigned to this genus, Euptilodegeeria obumbrata (Wulp) and Myiodoriops marginalis Townsend were reinstated as monotypic genera. In the present study, we demonstrate that Euptilodegeeria obumbrata (Wulp), previously assigned to three different genera, represents in fact a species of the large New World genus Eucelatoria Townsend, in which females possess a sharp piercer for oviposition. We also show that the species Eucelatoria carinata (Townsend) belongs to the same species group as Eucelatoria obumbrata, which we here define and characterize as the E. obumbrata species group. Additionally, we describe Eucelatoria flava sp. n. as a new species within the E. obumbrata species group. Finally, we redescribe the genus Myiodoriops Townsend and the single species M. marginalis Townsend.
Exoristinae , Blondeliini , Euptilodegeeria , Erythromelana , Machairomasicera , Hypostena , Tachinidae , Diptera , Parasitoid
The New World tropics represents one of the most biodiverse regions of the world, but its flora and fauna remains poorly known. This is particularly true for flies in the family Tachinidae, where the Neotropical fauna represents more than 35% of the total described species (
An example of the taxonomic instability of Neotropical tachinid genera is witnessed in the species Euptilodegeeria obumbrata (Wulp). This species was first classified in the former tachinid genus Hypostena by
Similar to the situation described above, although somewhat less confusing, is the situation of the other species recently excluded from Erythromelana by
In our recent revision of the Neotropical Erythromelana (
This revision was based on 28 adult specimens from four collections. Additional Nearctic and Neotropical taxa in the genus Eucelatoria from the NMNH, CNC and JOS collections were examined for comparison. Additional specimens of Blondelia Robineau-Desvoidy, Celatoria Coquillett, Myiopharus Brauer & Bergenstamm, Opsomeigenia Townsend, Euthelyconychia Townsend, Lixophaga Townsend and Vibrissina Rondani in the JOS collection were also examined for comparison. Acronyms used in the text for the collections and museums from which specimens were borrowed appear below, with their names and respective curators.
BMNH Natural History Museum, Department of Entomology, London, UK; N.P. Wyatt.
CNC Canadian National Collection of Insects, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada; J.E. O’Hara.
INBio National Biodiversity Institute of Costa Rica, Department of Entomology, Santo Domingo de Heredia, Costa Rica; M. Zumbado.
NMNH National Museum of Natural History, Department of Entomology, Smithsonian Institution, Washington, USA; N.E. Woodley.
JOS Private collection of John O. Stireman III, housed at Wright State University, Dayton, Ohio, USA.
Adult specimens were examined with a Nikon SMZ1000 stereoscopic microscope equipped with an ocular micrometer and a digital Nikon Coolpix 8800 camera (Nikon, Tokyo, Japan). To create images with a greater depth of field, 15‒30 photos of each specimen/structure at different focal points were taken. Final photos were compiled into a single image using the image stacking software CombineZM (
Descriptions and redescriptions of species follow terminology and abbreviations used in the Manual of Central American Diptera (
Male terminalia of tachinids provide some of the best characters for taxonomic studies at the species level. Dissections were performed according to the procedure described by
Morphological traits of 17 Eucelatoria specimens (14 males and 3 females), and 11 Myiodoriops specimens (5 males and 6 females) were measured. Additionally, male terminalia from 6 Eucelatoria and 2 Myiodoriops specimens were dissected. In species descriptions, the number of specimens for which particular characters were measured is given by “N”. When possible, means “x” are reported for continuous characters.
Data from each type specimen and other specimens examined are cited exactly as they appear on the label, with each line separated by a diagonal slash (/) and information for each individual label enclosed within quotation marks. Additional information not appearing on the label is enclosed within brackets. Finally, the depository is cited in parentheses.
Maps were created using SimpleMappr (
Eucelatoria Townsend, 1909: 249. Type species: Tachina (Masicera) armigera Coquillett, 1889, by original designation.
Euptilodegeeria Townsend, 1931: 465. Type species: Hypostena obumbrata Wulp, 1890, by original designation. Syn. n.
See
In the recognition of the genus Eucelatoria provided by
Eucelatoria obumbrata (Wulp, 1890), comb. n.
Eucelatoria carinata (Townsend, 1919).
Eucelatoria flava Inclán & Stireman sp. n.
Diagnosis
The Eucelatoria obumbrata species group can be distinguished from other species of Eucelatoria and other blondeliines (see discussion section below) using a combination of character states: (1) presence of sexual patches on the ventral portions of abdominal tergites 4 and 5 of males, (2) wing vein R4+5 setose from its base nearly to crossvein r-m in both sexes, and (3) a piercing ovipositor formed by abdominal sternite 7 in the female. Additional distinguishing traits include: mid-dorsal depression reaching only half way to hind margin of syntergite 1+2 and short spine-like setae on the ventral edge of the tergite 4 in females. This group can be easily separated from Erythromelana, in which E. obumbrata was formerly included (
Geographic distribution and seasonal occurrence
Species in the E. obumbrata species group are widely distributed in the Neotropical Region, from southern Mexico to Ecuador (Fig.
Discussion
Eucelatoria is a diverse new world tachinid genus, with Central and South America harboring most of the species. The genus belongs to a core clade of Blondeliini, along with Blondelia, Celatoria, Vibrissina and several other genera, that share the derived traits of females with a midventrally keeled abdomen, often with short stout bristles, and sternite seven modified into a hook-like piercer. Boundaries between genera within this group are less clear (
Each of the species treated here, E. obumbrata, E. carinata and E. flava sp. n., possesses at least some of the key traits of the Blondelia-group clade, including the keeled, spined abdomen with sternite 7 modified as a piercer in females, and well developed, anteriorly curved postgonites in males (
In the last revision of Eucelatoria,
We found three additional specimens from Costa Rica that belong to this species group, but each one is sufficiently morphologically distinct that it appears to be an undescribed species close to E. obumbrata. Each of the three specimens exhibits slight but distinct differences in the external morphology and male terminalia, but it remains unclear if these differences represent extensive intra-species variation or distinct species. Therefore, we leave these specimens undescribed until additional material is available to describe them as new or determine whether they are allied with a described species.
1 | Abdomen mostly or wholly black, with at most yellow laterally on tergites 1+2 to 4, males with median discal setae present on tergites 3 and/or 4 | 2 |
– | Abdomen wholly yellow, median discal setae absent on tergites 3 and 4 | E. flava sp. n. |
2 | Eyes densely haired, abdomen mostly black, with yellow only laterally on tergites 1+2 to 4, males with median discal setae present on tergite3 and/or tergite4 | E. obumbrata (Wulp) |
– | Eyes sparsely haired, abdomen wholly black (only known from a single female) | E. carinata (Townsend) |
Hypostena obumbrata Wulp, 1890: 143.
Euptilodegeeria obumbrata (Wulp):
Erythromelana obumbrata (Wulp):
Lectotype male, by designation of
10 specimens examined. 2 males labeled: “Co-type”, “♂”, “Omilteme,/ Guerrero,/ 8000 ft. [feet]/ July H. H. Smith.”, “Central America/ Pres. By F.D. Godman,/ O. Salvin/ 1903-172.”, “B.C.A. Dipt. II./ Hypostena obumbrata v.d.W”, “PARALECTOTYPE/ Of Hypostena obumbrata Wulp./ Designated 1980/ D.M. Wood”, “ Cotype/ 23967 U.S.N.M.”, “USNM 2049536”, “Eucelatoria/ obumbrata (Wulp)/ det. D.J. Inclán/ & J.O. Stireman”, “DI81NM”, “DI82NM” (NMNH); 2 males, as above except without the last label, “DI79NM”, “DI78NM” [ 1 specimen with terminalia dissected] (NMNH); 1 male, as above except without the “Cotype/…” labeled and having one extra label “Euptilodegeeria obumbrata/ Det. CHTT”, “DI80NM” (NMNH); 1 male, same as above except without the last two labels and the paralectotype label was attached in 1979, “ DI105BM” (BNHM); 1 male, same as above except without the last two labels, the location label “Xucumanatlan [miss spelled Xocomanatlan]/ Guerrero/ 7000 ft./ July. H.H. Smith” and the paralectotype label was attached on 1979, “DI106BM” (BNHM); 1 male and 2 females, “Omilteme,/ Guerrero,/ 8000 ft. [feet]/ July H. H. Smith.”, “Central America/ Pres. By F.D. Godman,/ O. Salvin/ 1903-172.”, “Eucelatoria/ obumbrata (Wulp)/ det. D.J. Inclán/ & J.O. Stireman”, “DI109BM” [male with terminalia dissected], “DI108BM”, “DI107BM” (BNHM).
This species can be distinguished from E. flava sp. n. by the primarily black coloration of the abdomen, with yellow coloration being restricted to the sides of tergites 1+2, 3, and 4. This contrasts with the entirely yellow abdomen of E. flava. Eucelatoria obumbrata usually bears median discals on tergite 3 and/or tergite 4, but these are absent in E. flava. The terminalia are similar between these species, but differ in several subtle respects including: the basal section of sternite 5 is distinctly shorter and broader basally in E. obumbrata; the surstylus, in lateral view, is equal to the cercus in length or slightly longer, whereas in E. flava it is markedly longer. In posterior view, the lateral margins of the cerci are narrowed linearly until the apical cleft, whereas in E. flava they are abruptly constricted below the upper lobes; the pregonite of E. obumbrata is relatively rectilinear, whereas that of E. flava triangular in shape, with a relatively broad at base, and strong narrowing toward apex. Females differ from E. carinata in having yellow coloration laterally on tergites 1+2, 3, and 4 (all black in E. carinata), densely haired eyes, more sparsely bristled palpi, and silvery parafrontals (bronzy in E. carinata).
Redescribed from 11 males (including the lectotype and 4 paralectotypes), and 2 females, unless otherwise noted as “N”.
Length: males, 6.2–7.1 mm (x = 6.8 mm); females, 6.1–7.0 mm (x = 6.5 mm).
Head (Fig.
Thorax (Fig.
Legs entirely black. Tarsal claws longer than 5th tarsomere in male and shorter than 5th tarsomere in female. Mid tibia with 1 anterodorsal seta, 2 posterodorsal setae, and 1 ventral seta. Hind tibia with anterodorsal setae uneven in length and not closely spaced; 2 well-developed posterodorsal setae, rarely with 1 additional shorter seta; 2 well-developed anteroventral setae. Upper and lower calypteres brownish-yellowish. Wing varied from light to dark fumose on cells sc, r1, r2+3, and sometimes on r4+5. Females with nearly hyaline wings. Wing vein R4+5 dorsally setose from its base nearly to crossvein r-m, and R1 bare, rarely only with 1 or 2 setae. Vein M smoothly curved at bend and ending at wing margin, separately from vein R4+5.
Abdomen (Figs
Male terminalia (N = 2, Figs
Female terminalia (Fig.
Specimens of E. obumbrata have been collected in southwestern Mexico (Fig.
Holotype male, labeled: “ ECUADOR, Napo [Province]/ 7 km. s. [South] Baeza/ 20-25.II.79/ G. &M. Wood 2000m”, “HOLOTYPE/ Eucelatoria/ flava/ Inclán & Stireman [red label]”, “DI244CA [specimen ID]” (CNC).
Paratype, 1 male: “DI12CA” (CNC). As above, except the identification type label reads “PARATYPE/ Eucelatoria/ flava/ Inclán & Stireman [yellow label]”.
From the Latin flava, meaning yellow, in reference to the yellow abdomen that distinguishes this species from its close related species, E. obumbrata.
This species is morphologically very similar to E. carinata and E. obumbrata, but can be easily separated by the abdominal coloration. Eucelatoria flava sp. n. has a yellow abdomen, which contrasts with the abdomen of E. carinata that is entirely black and E. obumbrata that is primarily black, with yellow coloration confined to the sides of styntergite 1+2, and tergites 3 and 4. Additionally, median discal setae are lacking on tergites 3 and 4 in males of this species where they are present on tergites 3 and/or 4 in males of E. obumbrata. The eyes of this species are sparsely and short-haired, contrasting with the densely and long-haired eyes of E. obumbrata and from the sparsely, but long-haired eyes of E. carinata.
Described from 2 males, unless otherwise noted as “N”.
Length: 6.6–6.7 mm.
As described for E. obumbrata except for:
Head (Fig.
Thorax (Fig.
Wing varied from light to dark fumose on cells c, sc, r1, r2+3, and r4+5. Wing vein R4+5 dorsally setose from its base until nearly the crossvein r-m, and R1 bare.
Abdomen (Figs
Male terminalia (N = 1, Figs
The only two known specimens of E. flava sp. n. were collected in highland cloud forest at about 2000 m in altitude on the eastern slope of the Andes of Ecuador (Fig.
Machairomasicera carinata Townsend, 1919: 578.
Eucelatoria carinata (Townsend):
Holotype female, labeled: “Manchi Ecuador/7000 ft/22-XI” [no year, but given as 1910 in description], “CHT Townsend/ Collector”, “Below/ Manchi Ec/Nov 22”, “Type No. 22247/U.S.N.M.”, “Machairomasicera/carinata/♀ Det CHTT 1”, “Eucelatoria/ carinata (Townsend)/ det. D.J. Inclán/ & J.O. Stireman” (NMNH).
This species can be distinguished from E. flava sp. n. and E. obumbrata by the entirely black coloration of the abdomen, which contrasts with the entirely yellow abdomen of E. flava, and the yellow and black abdomen of E. obumbrata. It also differs from females of E. obumbrata in having sparsely haired eyes, more densely bristled palpi, strongly infuscated wing veins, and a bronze tinted parafacial (dull silver in known females of E. obumbrata).
Length: 6.7 mm.
As described for E. obumbrata except for:
Head (Fig.
Thorax (Fig.
Wing moderately fumose on anterior half around veins C, Sc, R1 and R4+5, light infuscation also present along veins M, CuA1, and dm-cu. Wing vein R4+5 dorsally setose from its base until nearly the crossvein r-m, and R1 bare.
Abdomen (Fig.
The only known specimen of E. carinata was collected in Ecuador. The specimen was collected in the Andes Mountains at about 7000 ft (2100 m). The locality of the specimen reads “Below Manchi”, but it is unclear what this name refers to.
Myiodoriops Townsend, 1935: 227. Type species: Myiodoriops marginalis Townsend, 1935: 227, by original designation.
Myiodoriops marginalis Townsend, 1935.
Myiodoriops can be separated from other blondeliine genera (see discussion section below) using a combination of external characters and traits of the male terminalia including: 2 katepisternal bristles, 2 postpronotal setae (or, if a small inner seta is present, all three arranged in a line or broad arc), sparsely haired eyes, facial ridge with hairs on lower 1/3 or less, vein M ending in R4+5 vein just before wing margin or in wing margin very close to R4+5, lack of proclinate orbital setae in males, the mid-dorsal depression extending nearly to the hind margin of tg1+2, absence of a piercing structure in females, and short, spine-like setae on the anteriorly on the apex of the surstyli.
Myiodoriops is superficially similar to the E. obumbrata species group and to the genus Erythromelana in size, shape, and general appearance, which may explain the former grouping of these taxa into a single genus. However, it can be separated from these taxa using external morphological traits. It differs from the genus Eucelatoria generally in lacking the apomorphic piercing structure and associated short spines on ventral margins of abdominal tergites in females and absence of median discal setae on abdominal tergites 3 and 4, and it specifically lacks the apomorphic traits of the E. obumbrata group of R4+5 bristled nearly to crossvein r-m and sex patches in the male. Myiodoriops can be separated from Erythromelana by having the vibrissa inserted slightly above the lower facial margin (subtended by one or more setae), vein M ending in R4+5 vein or in wing margin very close to R4+5, and the mid-dorsal depression extending nearly to the hind margin of tg1+2. Additionally, Myiodoriops has only 2 katepisternal setae, which differs from Eucelatoria and from most species of Erythromelana which have 3 (see
The presence of short spines on the tip of the surstylus is reminiscent of Myiopharus (see
Redescribed from 5 males (including the type M. marginalis) and 6 females.
Length: males, 5.1–5.8 mm (x = 5.42 mm); females, 3.9–5.1 mm (x = 4.54 mm).
Head: Parafacial covered with dull silver pruinescence. Fronto-orbital fig and vertex black in ground color, covered with silver pruinescence appearing grayish from certain angles, usually with faint sparsely golden pruinescence dorsally. Frontal vitta usually entirely black, sometimes fading to dark-brown toward antenna. Pedicel black and first flagellomere black, covered with fine microtrichia and appearing grayish. Arista long, with minute setae, black with brown on basal 1/3 or less, thickened on basal 1/4 or less. Fronto-orbital fig with 5–7 medioclinate frontal setae in male, 4–7 in female; 3 reclinate inner orbital setae in males, 2 in females; female with 2 proclinate outer orbital setae, male without outer orbitals. Vertex with one reclinate inner and usually one lateroclinate outer vertical seta, the latter often barely or undifferentiated from the row of postocular setae in both sexes. Inner orbital and vertical setae usually about twice the length of frontal setae. Ocellar setae well-developed, proclinate. Parafacial bare and narrow with the narrowest point about equal to the widest portion of the palpus in males; in females narrower, about the basal width of the palpus. Facial ridge with hairs on basal 1/3 or less, and lower margin of face descending slightly below the level of vibrissa. Subvibrissal ridge short, usually with 1 to 3 setae; postgena narrow, with a distinct but small genal dilation. Posteroventral part of the head with the majority of white-yellowish fine setae and posterodorsal part of the head with one row of black setae behind the postocular row. Palpus brownish to black in color, distinctly swollen apically, more markedly in females.
Thorax: Shiny black in ground color; presutural scutum with evident white pruinescence, postsutural scutum with much sparser pruinescence revealing underlying black ground color. In dorsal view, only the presutural scutum appears grayish; whereas in lateral view the postsutural scutum appears grayish as well. Faint white pruinose stripes on presutural scutum leaving 4 black vittae; the inner 2 vittae longer and thinner, almost 1/2 the width of each of the outer 2 vittae. Prosternum with several hair-like setae. Postpronotum with 2 or 3 setae, when 3, the inner most is reduced in size and together they form a broadly obtuse angle, ca. 130–150°. Proepisternum bare. Katepisternum with 2 setae. The first postsutural supra-alar seta smaller than the notopleural setae. Scutellum with 3 pairs of setae, without apical setae or with one small hair-like pair.
Legs entirely black. Tarsal claws longer than 5th tarsomere in male and shorter than 5th tarsomere in female. Mid tibia with 2 posterodorsal setae, and 1 ventral seta. Hind tibia with anterodorsal setae uneven in length and not closely spaced; 2 well-developed posterodorsal setae, rarely with 1 additional shorter seta; 2 anteroventral setae. Upper and lower calypters translucent yellow-brownish. Wing length nearly equal to body length. Wing usually hyaline, rarely light fumose on the anterior edge. Wing vein R4+5 dorsally setose only at its base, and R1 bare. Vein M smoothly curved at bend and ending in vein R4+5 near the wing margin or separately in the margin closely approximated to vein R4+5.
Abdomen: Mostly black with yellow laterally on tg1+2 to tg4 on males, fully black in females. Transverse bands of sparse white pruinosity usually on the anterior 1/4 of tg1+2 to tg5. Mid-dorsal depression of tg1+2 extending to marginal setae and nearly to hind margin. One pair of median marginal setae on tg1+2 and tg3; a row of median marginals on tg3 to tg5; 1 pair of lateral marginal setae on tg1+2 and tg5; discal setae absent in both sexes. Sternites completely overlapped by tergites.
Male terminalia: Sternite 5 with median cleft smoothly V-shaped; apical lobes narrowed to broad points at their apices. The anterior margin of st5 clearly concave. The basal section of st5 distinctly shorter than the length of the apical lobes. Hypandrial arms separated. Pregonite curved anteriorly and tapered to a narrow rounded tip. Postgonite distinctively curved anteriorly, with narrow, almost pointed apex. Epiphallus small, hidden between the pregonites. Surstylus, in lateral view, broad, anteriorly curved and narrowed toward the apex, considerably longer than cercus. Surstylus with several short spine-like setae on the anterior side of its apex. Cercus, in lateral view, broad, slightly concave along anterior margin and narrowed only on the posterior margin of the apex. In posterior view, the cerci with long rectilinear upper lobes, nearly as long as the medial section + apical cleft combined. Apices of cerci, in posterior view, with excavated inner margins. Lateral margins of cerci without a constriction towards the apical section; apical cleft well defined. Distiphallus divided at base into long and a broader sclerotized portion with a toothed margin anteriorly.
See the distribution of Myiodoriops marginalis below, except for four undescribed specimens (see discussion below) that were collected in Brazil, Peru and Argentina (Fig.
The phylogenetic affinities of M. marginalis are unclear. As indicated in the diagnosis, there is little reason to believe that the species belongs with its former congeners in the genus Erythromelana or Eucelatoria, nor does it appear to be closely related to these taxa (see also
The genus description is based primarily on the specimens available for the known species M. marginalis. However, we found four specimens from Peru, Brazil and Argentina that belong to this genus, but they appear represent one or more undescribed species near M. marginalis. We have included these specimens in the genus description to cover all the generic variability, but we did not describe these specimens given the limited material and their poor condition. Additionally, of these four specimens, three are females and each is from a different locality. These four specimens exhibit slight differences in external morphology (e.g., parafacial width and abdominal coloration), but it is unclear if these differences represent intraspecific variation, male-female dimorphism, or actual differences between species. Therefore, we leave these specimens undescribed until additional material is available that can be used to help establish their identity.
Myiodoriops marginalis Townsend, 1935: 227;
Erythromelana marginalis (Townsend):
Holotype male labeled: “HOLO-/TYPE”, “Type [red label]”, “Pariká/ Ruhununí/ B. Guiana/ Jan. 1934 [hand written]”, “Mycos/ 4401 [hand written]”, “Press. By/ J.G. Myers/ B.M. 1940-24” “Myiodoriops/ marginalis TT [hand written]/ DetCHTT ♂”, “Myiodoriops/ marginalis Townsend/ det. D.J. Inclán/ & J.O. Stireman” (BNHM).
Seven specimens examined. 1 male labeled: “St. Augustine,/ Trinidad, BWI./ 1. 24. 60”, “Myiodoriops/ marginalis [hand written]”, “DI240CA” (CNC). 1 male labeled: “St. Augustine,/ Trinidad, BWI./ JAN 8 1960”, “F. D. Bennett/ Collector”, “X P. (77)/ near/ Myiodoriops [hand written]”, “Myiodoriops/ marginalis Townsend/ det. D.J. Inclán/ & J.O. Stireman”, “DI241CA” (CNC). 1 male labeled: “PIARCO/ Trinidad, BWI./ OCT. 29. 1953.”, “Collector/ F. J. Simmonds”, “77 [hand written]”, “Myiodoriops/ n. sp. ♂ [hand written]”, “Myiodoriops/ marginalis Townsend/ det. D.J. Inclán/ & J.O. Stireman”, “DI243CA” (CNC). 1 female labeled, same as previous except by “OCT. 29. 1953”, without sp. ID, “DI74CA” CNC. 1 male labeled: “W. ARIMA/ TRINIDAD/ 26-8-1964”, “Myiodoriops/ marginalis Townsend/ det. D.J. Inclán/ & J.O. Stireman”, “DI73CA” (CNC). 1 female labeled: “St. Augustine,/ Trinidad, BWI./ II. 17. 60”, “Myiodoriops/ marginalis Townsend/ det. D.J. Inclán/ & J.O. Stireman”, “DI39CA” (CNC). 1 female labeled: same as previous except by “II. 28. 60”, “DI236CA” (CNC).
See diagnostic section for the genus Myiodoriops.
Redescribed from 5 males (including the type M. marginalis) and 3 females.
Length: males, 5.1–5.8 mm (x = 5.42 mm); females, 3.9–4.53 mm (x = 4.21 mm).
As described for the genus except:
Head (Fig.
Specimens of M. marginalis have been collected only from Guyana in northern South America, and from the southern Caribbean islands of Trinidad and Tobago (Fig.
Identification of the Eucelatoria obumbrata species group, Myiodoriops and Erythromelana using Wood and Zumbado (2011).
All three genera should readily key to couplet 114 (along with nearly all blondeliines) in
114 | Vein R4+5 setose on dorsal surface halfway or more from its base at junction of R2+3 and R4+5 to crossvein r-m (Figs 158, 160, 161) | 115 |
– | Vein R4+5 dorsally with few setae at base only, not extending halfway to crossvein r-m | 129 |
115 | Eye with conspicuous ommatrichia, each longer than combined diameter of four or more eye facets (as in Fig. 20) | 116 |
– | Eye apparently bare | 120 |
116 | Facial ridge bristled on lower half or more, with row of erect bristles along most of length (Figs 21–24) | 117 |
– | Facial ridge bare except for few small recumbent bristles above vibrissa [specimens of some species of the E. obumbrata species group have fine setae nearly to one-half the height of the facial ridge, but these are short and hair-like above the lower third] | 118 |
... | ||
118 | Lateral scutellar bristles parallel to one another and shorter than subapical bristles (as in Fig. 130); ventral surfaces of abdominal tergites 4 and 5 of male each with patch of appressed black hair (sex patch, Fig. 165) | 118a |
– | Lateral scutellar bristles divergent and about as long as subapical bristles (Fig. 127); ventral surfaces of abdominal tergites 4 and 5 of male with or without patches of appressed hair | 119 |
118a | Male with a pair of proclinate orbital bristles; female abdomen and ovipositor unmodified; Two katepisternal bristles | Leptostylum Macquart |
– | Male without pair of proclinate orbital bristles; female with short stout bristles on the ventral margins of tergites, sternite 7 modified into sharp, hook-like piercer, usually concealed between ventral edges of tergites; usually three katepisternal bristles | Eucelatoria Townsend, in part |
... | ||
120 | Ventral katepisternal bristle as large as, or larger than, anterodorsal katepisternal bristle (rarely only slightly thinner) and situated close to upper margin of midcoxa, within no more than twice its diameter from coxal margin (Fig. 118); vein A1 ending at wing margin (Fig. 160), although apex of vein may be thin and easily overlooked without transmitted light or light reflected from upper surface | 121 |
– | Ventral katepisternal bristle absent or distinctly smaller than anterodorsal katepisternal bristle and usually situated closer to anterodorsal bristle than to midcoxa (intermediate or closer to coxa in a few Actia and Ceromya), but not as close to coxa as twice its diameter (Fig. 117); vein A1 ending in membrane before reaching margin of wing (Fig. 161) | 122 |
... | ||
122 | Vein R4+5 setulose dorsally from base to well beyond crossvein r-m (Fig. 161) | 123 |
– | Vein R4+5 without setulae beyond crossvein r-m | 124 |
... | ||
124 | Scutellum lacking both lateral and discal bristles (as in Fig. 132); basal portion of proboscis when extended longer than prementum (Fig. 82), and membrane between lower genal margin and clypeus thickened, forming convex paraclypeal sclerite (as in Fig. 80) (not visible if proboscis is retracted into base of head); labella extending forward | Ginglymia Townsend |
– | Scutellum with lateral and discal bristles; basal portion of proboscis shorter than prementum, and membrane between lower genal margin and clypeus without sclerite; labella either padlike or extending posteriorly | 125 |
125 | Facial ridge with row of erect bristles on basal half or more | 126 |
– | Facial ridge bare except for few small setae above vibrissa [specimens of some species of the E. obumbrata species group have fine setae nearly to one-half the height of the facial ridge, but these are short and hair-like above the lower third] | 127 |
... | ||
127 | Veins R4+5 and M ending separately on either side of wing apex relatively far apart (Fig. 158) | Chaetostigmoptera Townsend, in part |
– | Veins R4+5 and M both ending before wing apex (as in Fig. 148) | 128 |
128 | Both lateral and subapical scutellar bristles long, stout, divergent (as in Fig. 131); vibrissa subtended by one or more subvibrissal bristles below it (as in Figs 20–22); three postsutural supra-alar bristles present, middle one largest | Italispidea Townsend |
– | Lateral scutellar bristles either lacking or short and thin; subapical bristles divergent or convergent; vibrissa with or without one or more subvibrissal bristles below it; two or three postsutural supra-alar bristles present | 128a |
128a | Lateral scutellar bristles either lacking or short, thin, convergent; subapical bristles also convergent, crossed medially; vibrissa arising from anteroventral corner of head without subvibrissal bristles below it (as in Fig. 25); postsutural supra-alar bristles reduced to two: the true first bristle absent; the apparent first, therefore, the larger of the two (Fig. 99). Males without obvious sex patches on abdominal tergites 4 and 5; female without short stout bristles on the ventral margins of tergites and without sternite 7 modified into a piercer | Ischyrophaga Townsend |
– | Lateral scutellar bristles present, short, and parallel or divergent; subapical bristles divergent; vibrissa subtended by one or more subvibrissal bristles below it; usually 3 postsutural supra-alar bristles; males with sex patches on the ventral surfaces of abdominal tergites 4 and 5; female with short stout bristles on the ventral margins of tergites, sternite 7 modified into sharp, hook-like piercer, usually concealed between ventral edges of tergites | Eucelatoria Townsend, in part |
129 | Eye with conspicuous ommatrichia, each longer than combined diameter of four or more eye facets (as in Fig. 20) | 130 |
– | Eye apparently bare | 134 |
130 | Parafacial with row of stout erect bristles along entire length (Fig. 37); base of vein R4+5 with single large bristle (as in Figs 156, 159) | Eulasiona Townsend |
– | Parafacial lacking row of erect bristles; base of vein R4+5 with more than one small bristle | 131 |
131 | Vibrissa arising at level of lower margin of head (as in Fig. 25); usually with two postpronotal bristles (as in Fig. 93), rarely with three; middorsal depression on abdominal syntergite 1+2 not extending back to hind margin of syntergite | Erythromelana Townsend, in part |
– | Vibrissa arising above level of lower margin of head, with at least one subvibrissal bristle (Fig. 20); three or more postpronotal bristles present; middorsal depression on abdominal syntergite 1+2 extending back to hind margin of syntergite (as in Figs 186, 188) | 132 |
... | ||
134 | Facial ridge setose on lower half or more, with row of erect bristles or hairs or both along most of length | 135 |
– | Facial ridge bare except for few small recumbent bristles above vibrissa | 150 |
... | ||
150 | Median discal bristles present on tergites 3 and 4 | 151 |
– | Median discal bristles absent from tergites 3 and 4 | 160 |
... | ||
160 | Eye exceptionally large, covering almost all of side of head; distance between eye and lower margin of head less than twice width of palpus (as in Fig. 14); ocellar triangle not raised to form tubercle; ocellar bristles arising beside or in front of anterior ocellus, their bases about as far apart as posterior ocelli | Sphaerina Wulp |
– | Eye smaller, distance between eye and lower margin of head greater than twice width of palpus; ocellar triangle raised; ocellar bristles arising behind anterior ocellus, their bases closer together than posterior ocelli | 161 |
161 | Vibrissa arising from anteroventral corner of head (Fig. 25), with at most one subvibrissal bristle below it; parafacial very narrow; lateral scutellar bristle short or lacking (Fig. 132); postsutural supra-alar bristles usually reduced to two, true first bristle absent (as in Fig. 99) | 162 |
– | Vibrissa arising above anteroventral corner of head (Fig. 20), subtended by one or more subvibrissal bristles; parafacial narrow or broad; lateral scutellar bristle well developed (as in Figs 130, 131); postsutural supra-alar bristles three or more, middle one largest (as in Figs 100–104) | 163 |
162 | Arista plumose (Fig. 25); genal dilation extending forward to about vibrissal angle, anterior genal seta thus arising close to base of vibrissa; midtibia at most with small anterodorsal seta scarcely longer than width of tibia; lateral scutellar bristles lacking | Phyllophilopsis Townsend, in part |
– | Arista bare; genal dilation distinctly separated from vibrissal angle by gap of membrane, so that single subvibrissal seta distinctly separated from genal setae; midtibia with well-developed anterodorsal seta; lateral scutellar bristles present | Erythromelana Townsend, in part |
163 | Lateral scutellar bristles at least four-fifths as long and as straight as subapical scutellar bristles, strongly divergent (as in Fig. 131); parafacial extremely narrow; with two reclinate orbital bristles, markedly different from each other in size (as in Fig. 19) | Italispidea Townsend, in part |
– | Lateral scutellar bristles about two-thirds (or less) as long as subapical scutellar bristle (as in Fig. 130); parafacial broader; reclinate orbital bristles more numerous or more uniform in size | 164 |
164 | Ocellar setae minute, shorter than length of ocellar triangle; frontal and reclinate orbital bristles forming single even row, increasing in size toward vertex usually regularly (as in Figs 65, 66), or with abrupt increase in some species; body pale ochreous brown | Ophirion Townsend |
– | Ocellar setae present, longer than ocellar triangle; frontal and reclinate orbital bristles, if arising in single row, usually varying in size, with largest frontal bristles in middle of row (as in Figs 63, 64); body color usually brown or black, except on sides of abdomen | 165 |
165 | Veins M and R4+5 each ending separately on either side of wing apex (Fig. 158) | Chaetostigmoptera Townsend, in part |
– | M and R4+5 both ending anterior to wing apex (as in Fig. 156) | 165a |
165a | Male with two pairs of proclinate orbital setae (as in females); usually 2 reclinate orbital setae; three postpronotal bristles arranged in a triangle or strong arc; 2 or 3 katepisternal bristles | Myiopharus Brauer & Bergenstamm, in part |
– | Male without proclinate orbital setae; usually 3 reclinate orbital setae; 2 apparent postpronotal bristles, innermost bristle reduced or absent, when present, the three are arranged in a broad arc forming an angle of > 120°; 2 katepisternal bristles | Myiodoriops Townsend |
We would like to thank to Jim O’Hara, Monty Wood (Invertebrate Biodiversity, Agriculture and Agri-Food Canada, CNC), Norm Woodley (Systematic Entomology Laboratory, NMNH), Nigel Wyatt (Natural History Museum, Department of Entomology, BMNH) and Manuel Zumbado (National Biodiversity Institute of Costa Rica, INBio) who provided specimen loans. We also thank Matt Duncan for helping with some of the images and Z. “Kai” Burington for useful discussions concerning Eucelatoria. Thanks to Pierfilippo Cerretti, Jim O’Hara and one anonymous reviewer for carefully reviewing and editing this manuscript. This work was supported by a PhD Fellowship from the CARIPARO Foundation to DJ Inclán and a U.S. NSF grant (DEB 1020571) to JO Stireman.