Corresponding author: Wayne N. Mathis (
Academic editor: Rudolf Meier
The genus
Flies of the genus
The phylogenetic position of
The nomenclatural history of the genus and its included species is uncomplicated, as would be expected for a little-known genus that comprises few species. When
Most of the species considered herein are newly described, and in their individual treatments, we have included all specimens that were available and that could be reliably identified. Specimens included in type series are more restricted, particularly those for which identifications are based primarily on characters of the male terminalia. Other specimens are listed with the appropriate species under the section “Other specimens examined.” Type specimens of all nominate taxa were studied. Label data accompanying each holotype are quoted verbatim with a slash mark to separate data of one label from another. Clarifying or interpretive comments are included within brackets.
Descriptions of species are composite if specimens other than the holotype were available. For the most part, information given in the generic description is not repeated in the species descriptions. The descriptive terminology follows that published in the recent
We have adopted the nomenclature of
Because specimens are small, usually less than 3.50 mm in length, study and illustration of structures of the male terminalia and female reproductive organs required use of a compound microscope. Observation of detailed structures of the male terminalia was sometimes needed to confirm a species’ identification.
Dissections of male terminalia were performed following
Distribution maps were made using ESRI ArcView7 GIS 3.2. Longitude and latitude coordinates were obtained for the locality where each specimen was collected and entered into a Microsoft Excel© spreadsheet. If unavailable directly from specimen labels, longitude and latitude were estimated using gazetteers and maps, usually available electronically, to determine the geographical coordinates.
Three head ratios are used commonly in the descriptions and are defined here for the convenience of the user (ratios are averages of three specimens).
1. Head ratio: head height (distance from peristomal margin to vertex)/head width. The measurements were taken from the head from an anterior view.
2. Facial ratio: interantennal width (shortest distance between antennal bases)/facial height (distance in the middle of the face between the ptilinal suture and the ventralmost margin of the face). The measurements were taken from the head from an anterior view.
3. Frontal ratio: frontal width (straight line distance between compound eyes at the level of the anterior ocellus)/frontal height (distance between the ptilinal suture (anterior margin of the frons) and the vertex. The measurements were taken from the head with an orientation to make the frons flat in the field of view.
The author’s contributions generally adhered to the following division of labor: most of the descriptive work was done by the first two authors, and the species names are attributed to them; the third author contributed information on the female reproductive system and observations on mating behavior. All three participated in field work together in Ecuador, where many of the specimens were collected and where observations and preliminary dissections were made.
Although this study was based primarily on specimens in the National Museum of Natural History, Smithsonian Institution (USNM), numerous others were borrowed, particularly primary type specimens of the species described previously. To our colleagues and their institutions listed below who lent specimens, we express our sincere thanks. Without their cooperation this study could not have been completed.
American Museum of Natural History, New York, New York (David A. Grimaldi)
Academy of Natural Sciences of Philadelphia, Pennsylvania (Jon K. Gelhaus, and Jason D. Weintraub)
California Academy of Sciences, San Francisco, California (Paul H. Arnaud, Jr.)
Canadian National Collection, Ottawa, Canada (Owen Lonsdale, J. Frank McAlpine)
University of Guelph, Guelph, Ontario, Canada (Stephen A. Marshall)
Insituto Nacional de Biodiversidad, Santo Domingo, Costa Rica (Manuel Zumbado)
Instituto Nacional de Pesquisas da Amazônia, Manaus, Amazonas, Brazil (José Albertino Rafael, Rosaly Ale-Rocha)
Zoologische Staatssammlung München, München, Germany (Marion Kotrba)
To give perspective to this revision, we are providing diagnoses for the family
The concept of
1 | Fronto-orbital seta 1; ocellar setae present. Costa short, extended to vein R4+5; vein CuA2 weak or lacking, lacking a cell |
|
– | Fronto-orbital setae 2; ocellar setae absent. Costa long, extended to vein M; vein CuA2 usually well developed, usually with a distinct cell |
|
The subfamily
Although
1 | Frons with 1 pair of interfrontal setae; eyes bare. Katepisternum with 2 subequal setae. Hindfemur bearing anterodorsal, preapical seta | |
– | Frons lacking interfrontal setae; eyes microsetulose, sometimes sparsely. Katepisternum bearing 1 prominent seta. Hindfemur lacking anterodorsal, preapical seta | 2 |
2 | Fronto-orbital setae reclinate or occasionally mesoclinate, lacking a proclinate seta; medial vertical seta present with proclinate orientation; face in profile angulate, dorsal surface flattened. Supra-alar seta lacking; lateral scutellar setae 1 pair, apical. Crossvein bm-cu lacking, cells bm and dm confluent; vein CuA2 weak or lacking; cell |
|
– | Fronto-orbital setae comprising 1 proclinate and 1 reclinate setae; medial vertical seta absent; face in profile shallowly and vertically arched, lacking a flattened, dorsal area. Supra-alar seta present, well developed; lateral scutellar setae variable but usually 2 pairs (only an apical pair in |
Known only from the New World tropics.
Specimens of
Our field work and sampling, regardless of the collecting technique, also indicates that two to four species frequently occur sympatrically at the same microhabitat. We observed that one species at these sites usually predominates in numbers of individuals. How the various species partition the habitat and what their population structure is are basic questions that remain unanswered.
Several species exhibit considerable sexual dimorphism, especially in the width and coloration of the face. Males in these species tend to have wider faces (hypercephaly), i.e., larger facial ratios, and frequently there is a distinctive colorational pattern. The facial pattern usually also involves microtomentum or its absence in additional to color. These details are included in descriptions of appropriate species.
Within the subfamily
1. Midtibia with an apical, anteroventral spine-like seta.
2. Arista bipectinate (
3. Face bearing a dorsoclinate pair of setae, these usually inserted above other facial setae.
1. Frons bearing a pair of interfrontal setae that are usually slightly reclinate to dorsoclinate. The interfrontal setae, as described, are unique to
2. Forefemur with 1-3 posteroventral setae on apical half.
3. Scutellum bearing a single pair of marginal setae, these apical (also in some species of
4. Reclinate fronto-orbital seta inserted behind proclinate fronto-orbital seta.
5. Each tibia with a dorsoapical seta.
6. Hindfemur with a subapical dorsal seta.
7. Anepisternum with 1–2 setae along posterior margin (relatively common in other taxa of Asteioinea).
8. Several characters of the male terminalia.
1 | Large facial setae arranged in a single transverse row of about 8 setae; forefemur bearing 1 large, posteroventral seta at apical 1/3; antenna mostly yellowish orange, pedicel and basal flagellomere with some blackish coloration dorsally | 2 |
– | Large facial setae arranged in 2-3 irregular, transverse rows; forefemur bearing 1-3 subequal, posteroventral setae at apical 1/3; antenna mostly black (base of basal flagellomere pale in males of |
4 |
2 | Face above transverse carina moderately microtomentose, appearing dull to at most subshiny; tarsomeres of foreleg mostly yellowish, apical tarsomere dark colored; forebasitarsus not compressed | 11 |
– | Face above transverse carina sparsely microtomentose to bare, appearing shiny; tarsomeres of foreleg dark colored, mostly blackish, forebasitarsus slightly compressed | 3 |
3 | Face with receded, ventral portion short, height less than width of antennal pedicel. Forecoxa mostly whitish yellow; mid- and hindfemora with basal 1/3–1/2 yellowish, contrasted with blackish brown apical portion | 12 |
– | Face with receded, ventral portion long, height about equal to width of antennal pedicel. Forecoxa black; mid- and hindfemora mostly to entirely blackish brown, at most with basal 1/6 becoming pale | 10 |
4 | Interfrontal seta long, at least 2/3 length of lateral vertical seta. Basal flagellomere variable, frequently about twice as long as high | 5 |
– | Interfrontal seta short, about l/2 length of lateral vertical seta. Basal flagellomere short, about as long as high (the |
16 |
5 | Ventral projection of pedicel long, length of projection subequal to length of pedicel without considering projection; face of female mostly black, at least dorsally, projected forward, bulbous, evenly arched horizontally, very shallowly arched vertically anterior to rounded, dorsal projection; wing generally strongly infumate (the |
6 |
– | Ventral projection of pedicel short, length of projection conspicuously less than length of pedicel without considering projection; face of female generally angulate, dorsal 2/3 more or less evenly slopped anteroventrally to a transversely arched ridge, thereafter nearly vertical to oral margin; wing faintly infumate to completely hyaline | 11 |
6 | Antenna of male black (basal flagellomere yellowish basally); face of male with medial portion black, black coloration extended to ventral margin of face, microtomentum sparse or lacking; antennal groove lacking subshiny area extended obliquely medioventrally nearly to oral margin | 7 |
– | At least basal flagellomere, apex of ventral extension of pedicel, and ventral margin of face of male pale colored, mediodorsal area of face densely microtomentose, silvery white; antennal groove with a subshiny, black area extended obliquely medioventrally nearly to oral margin | 8 |
7 | Midfacies of male with a conical prominence, shaped like an inverted V with a rounded vertex; face of female with extreme lateral margins yellowish; forefemur of male with basal 2/3 yellow, apical 1/3 black, lacking a preapical ring | 4 |
– | Midfacies of male with a shallowly arched prominence; forefemur of male mostly black, with at most basal 1/4 pale | 7 |
8 | Antenna of male yellow | 9 |
– | Scape and pedicel of male mostly brown to brownish black, only basal flagellomere partially to mostly yellowish | 9 |
9 | Basal flagellomere of male distinctly two toned, with basal half yellow, apical half black | 5 |
– | Basal flagellomere of male yellow, not two toned | 10 |
10 | Middle and hind femora mostly yellow, at most with apical 1/3 to 1/4 black | 6 |
– | Middle and hind femora mostly black, at most with basal 1/4 to 1/3 yellowish | 8 |
11 | Forecoxa black | 12 |
– | Forecoxa whitish yellow to yellow | 14 |
12 | Dorsal, transverse row of facial setae consisting of 1 pair of well-developed, dorsoclinate setae; basal flagellomere of male pale, yellowish on basal 1/2. Wing darkly infumate | 18 |
– | Dorsal, transverse row of facial setae with some ventroclinate setae in addition to large dorsoclinate setae; antenna of male entirely black. Wing either mostly hyaline or very faintly infumate | 13 |
13 | Face bicolored, ventral, receded portion yellow, dorsal portion black; distance between antennal bases of males nearly twice length of antenna (Mexico) | 16 |
- | Face black, ventral, receded portion with some silver microtomentum that becomes denser toward oral margin; distance between antennal bases of males about equal to length of antenna (Brazil) | 17 |
14 | Clypeus yellow (Dominica) | 15 |
– | Clypeus black | 15 |
15 | Parafacial and ventral portion of face adjacent to parafacial yellow (Bolivia (La Paz: Guanay), Brazil (São Paulo: Araçatuba, Córrego Azul)) | 14 |
- | Parafacial and ventral portion of face adjacent to parafacial black (Brazil. Rio de Janeiro: Floresta da Tijuca) | 13 |
16 | Surstyli fused medially | 2 |
– | Surstyli separate medially | 17 |
17 | Surstylus in lateral view straight, parallel sided ( |
1 |
– | Surstylus in lateral view swollen basally, thereafter parallel sided ( |
3 |
Moderately small flies, body length 2.70 mm (holotype).
Illustrations of
Distribution of
The holotype male is labeled “W of Bogota COLOMBIA/M R Wheeler collector/USNM ENT 00118284 [plastic bar code label]/
Colombia. Bogota: Bogota (west;
(
The specific epithet,
This species, represented by two males only, is known only from Colombia and is the only species of
Moderately small to medium-sized flies, body length 2.35–3.10 mm.
Illustrations of
Illustrations of
Distribution of
The holotype male is labeled “Cachi C[osta] R[ica][,] 9 III 1910 [9 Mar 1910][,] P P Calvert/Valley of Rio Naranjo/HoloTYPE 6069 [pink]/475 ♂/TYPE No. 6064 [number written across right end of label] Planinasus AMBIGUUS E.T.Cresson,Jr. [pink; species name and number handwritten].” The holotype is double mounted (minuten pin in rectangular card block), is in moderately poor condition (wings glued together), and is deposited in the ANSP (6069).
Costa Rica. Cartago: Cachí, Valley of Rio Naranjo (
(
This widespread species was previously known only from Costa Rica and Peru (
Small to moderately small flies, body length 1.80–2.55 mm.
Illustrations of
Distribution of
The holotype male is labeled “
(
The species epithet,
This species, like
Moderately small to medium-sized flies, body length 2.50–3.70 mm.
Same as male except as follows:
Illustrations of
Internal female reproductive organs of
Internal female reproductive organs of Planinasus kotrbae sp. n., photos, left lateral view.
Distribution of
The holotype male is labeled “ECUADOR. Prt. Or[e]ll[a]na: RioTiputini [Biodiversity Station] (
Ecuador. Orellana: Rio Tiputini Biodiversity Station (
(
The specific epithet,
Marion Kotrba and W. N. Mathis observed the mating behavior of
Having access to a rearing cage facilitated observations in the laboratory as well as temporarily holding specimens that were to be dissected. Keeping females to be dissected in the cage ensured their freshness, which makes dissecting easier, and the preparations that resulted were excellent. The tissue is pliable and extraneous material, such as tracheoles, can easily be removed.
This species is a member of the
Antennal bases of males more approximate, distance between slightly more than antennal length; face of males with a subshiny, black area extended obliquely medioventrally from ventral margin of antennal groove nearly to oral margin
Moderately small to medium-sized flies, body length 2.15–3.05 mm.
Illustrations of
Distribution of
Same as male except as follows:
The holotype male is labeled “Brasil (MA[RANHÃO]), Mirador Parque Est. Mirador Base da Geraldina/Armadilha Suspensa 23.ix.2006 [23 Sep 2006], F. Limeira de Oliveira, col./USNM ENT 00118277 [plastic bar code label]/
Brazil. Maranhão: Parque Estadual Mirador, Base da Geraldina (
(
The specific epithet,
This species is very similar to
Moderately small flies, body length 2.35–2.55 mm.
Illustrations of
Moderately small to medium-sized flies, body length 2.10–3.10 mm.
Same as male except as follows:
Illustrations of
Distribution of
The holotype male is labeled “
Trinidad and Tobago. Tobago. St. John: Parlatuvier (
(
The specific epithet,
This species is known from northern South America, including Trinidad and Tobago. It keys out near
Moderately small to medium-sized flies, body length 2.35–3.10 mm.
Same as male except as follows:
Illustrations of
Distribution of
The holotype male is labeled “Caripito [
Venezuela. Monagas: Caripito (
(
Adults of this species were fairly common on vegetation in the understory of gallery forests along the Rio Manu. The flies occurred on prominent, frequently sun-lit, broad leaves, especially in stream bottoms, where males displayed and courted females and drove off other males.
This species is very similar to
Moderately small to medium-sized flies, body length 2.10–3.05 mm.
Illustrations of
Distribution of
Same as male except as follows:
The holotype male is labeled “
Ecuador. Orellana: Rio Tiputini Biodiversity Station (
(
The specific epithet,
Moderately small flies, body length 2.10–2.75 mm.
Illustrations of
Distribution of
As in male except as follows: Head generally narrower, head ratio 0.65–0.67; frontal ratio 0.55–0.56; facial ratio 0.35–0.45.
The holotype male is labeled “PERU.Madre de Dios: Manu, Rio Manu, 250 m[,] Pakitza,
Peru. Madre de Dios: Río Manu, Pakitza (
(
The specific epithet,
Structures of the male terminalia of this species and those of
Small to moderately small flies, body length 1.60–2.80 mm.
Illustrations of
Illustrations of
Distribution of
As in male except as follows: Head generally narrower, head ratio 0.60–0.62; frontal ratio 0.51–0.54; facial ratio 0.26–0.29.
The holotype male is labeled “
Dominican Republic. La Vega: near Jarabacoa, Salto Guasara (
(
The species epithet,
Variation in wing coloration is particularly evident in specimens of this species and may be related to the age of specimens, older specimens having darker wings, but also to underlying genetic variation. The shape of the surstylus and hypandrium readily distinguish this species from congeners.
Moderately small flies, body length 2.10–2.80 mm.
Illustrations of
Distribution of
As in male except as follows: Head generally narrower, head ratio 0.60–0.62; frontal ratio 0.61–0.64; facial ratio 0.27–0.31.
The holotype male is labeled “
Guyana. Conservation of Ecological Interactions and Biotic Associations (CEIBA; ca. 40 km S Georgetown;
(
The specific epithet,
See “Remarks” under
Moderately small to medium-sized flies, body length 2.60–3.15 mm.
Illustrations of
As in male except as follows: Head generally slightly narrower, head ratio 0.65–68; frontal ratio 0.44–48; facial ratio 0.34–37. Face mostly uniformly bluish black, silvery white microtomentum more evident.
The holotypes male is labeled “Floresta da Tijuca- BRASIL/30 XII 1991 A. BAPTISTA R. BAPTISTA em Marantaceae [date and name of plant family handwritten]/USNM ENT 00118276 [plastic bar code label]/
Brazil. Rio de Janeiro: Rio de Janeiro, Floresta da Tijuca (
(
The specific epithet,
This species is most similar to
Moderately small to medium-sized flies, body length 2.20–3.00 mm (holotype 2.38 mm).
Illustrations of
Distribution of
As in male except as follows: Head generally slightly narrower, head ratio 0.67–69; frontal ratio 0.41–44; facial ratio 0.41–44.
The holotype male is labeled “BOLIVIA: Santa Cruz Dept. Ichilo Prov. Hotel Flora Y Fauna[,] 4–6 km SSE Buena Vista[,]
Bolivia. Santa Cruz: Ichilo, Buena Vista (4–6 km SSE; Hotel Flora y Fauna;
(
The specific epithet,
See “Remarks” under
Moderately small to medium-sized flies, body length 2.55–3.25 mm.
Illustrations of
As in male except as follows: Head generally slightly narrower, head ratio 0.70–75; frontal ratio 0.63–0.67; facial ratio 0.31–0.36.
The holotypes male is labeled “DOMINICA, W.I[.] trail,1mi.n.junc. rds to Rosalie & Castle Bruce, 1300' Apr. 23, 1966[,] R.J. Gagne/Bredin-Archbold-Smithsonian Bio.Surv.Dominica/USNM ENT 00118283 [plastic bar code label]
West Indies. Dominica. St. David: 1.6 km N of junction of roads to Rosalie and Castle Bruce (
(
The specific epithet,
Externally this species is very similar to
Moderately small to medium-sized flies, body length 2.45–3.15 mm.
Illustrations of
Illustrations of
Distribution of
As in male except as follows: Head generally narrower, head ratio 0.63–0.65; frontal ratio 0.43–0.45; facial ratio 0.49–0.53.
The holotype male is labeled “
Mexico. Chiapas: Cacahoatán (7 km N;
(
The species epithet,
This species is known thus far only from southern Mexico. We suspect that it will be found to be more widespread once there is better sampling from other Central American countries. It is the only species of the
Moderately small to medium-sized flies, body length 2.50–3.50 mm.
Illustrations of
Distribution of
As in male except as follows: Head generally narrower, head ratio 0.86–89; frontal ratio 0.54–0.57; facial ratio 0.33–0.36.
The holotype male is labeled “
Brazil. São Paulo: Mogi das Cruzes, Serra do Itapeti (
(
The species epithet,
In the key to species, this species is close to
Medium-sized flies, body length 3.00–3.15 mm.
Illustrations of
As in male except as follows: Head generally narrower, head ratio 0.70; frontal ratio 0.56; facial ratio 0.35.
The holotype male is labeled “PERU.Madre de Dios: Manu, Erika (near Salvacion),550m,5–6 Sept1988,AFreidberg/
Peru. Madre de Dios: Río Manu, Erika (near Salvación;
(
The specific epithet,
The male abdomen of this species is atypical of other congeners in having apparent fusion of some sternites (sternites 4+5) and a well-developed sternite 6 that is broadly produced medioposteriorly to form a shallowly bifurcate projection. We have observed this condition only in this species, and our observation is limited to the single holotype male.
Moderately small flies, body length 2.20 mm; wing length 1.71 mm.
The holotype female is preserved in Dominican amber (Dominican Republic. specific provenance unknown) and is deposited in the AMNH (type number DR-8-208).
Erin Kolski prepared the distribution maps, and the inked illustrations were carefully rendered by Elaine R. S. Hodges. For reviewing a draft of this paper we thank Amnon Freidberg and an anonymous reviewer. We are also grateful to David Challinor (deceased), former Assistant Secretary for Research, Smithsonian Institution, and Stanwyn G. Shetler, former Deputy Director of the National Museum of Natural History, for financial support to conduct field work and study primary types through grants from the Research Opportunity Fund.
Field work on the West Indies was funded largely by grants from the Biodiversity Program (Biological Surveys and Inventories, BSI), National Museum of Natural History, Smithsonian Institution ((Lynne R. Parenti and George R. Zug, former chairs). We are also grateful to the Smithsonian Institution’s Biological Diversity of the Guianas Program (publication series number 192; Vickie A. Funk, Director; Carol Kelloff, Coordinator) for supporting field work in Guyana.
Some of the research and field work was generously funded by a grant from the Partnerships for Enhancing Expertise in Taxonomy program (PEET), National Science Foundation (PEET 952-1773). Some of the field work in Brazil was funded by the FAPESP (Fundação de Amparo à Pesquisa do Estado de São Paulo), which we also acknowledge and thank. Final aspects of this research, especially the preparation of the manuscript for publication, were supported by the Sabrosky Fund, Department of Entomology, Smithsonian Institution.
Recent field work in Brazil (December 2009-June 2010) that resulted in the vast majority of specimens studied in this paper was supported by a grant from CNPq (Visiting Researcher/Process number 401609/2009-0), which we gratefully acknowledge and thank. We thank Dianne Mathis for helping with all views of the production of this paper, especially the field work in Brazil.