Research Article |
Corresponding author: James B. Whitfield ( jwhitfie@life.illinois.edu ) Academic editor: Jovana M. Jasso-Martínez
© 2025 James B. Whitfield, Jose L. Fernandez-Triana, Caroline Boudreault.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Whitfield JB, Fernandez-Triana JL, Boudreault C (2025) Two new Neotropical genera of Miracinae (Hymenoptera, Braconidae) with unusual metasomal morphology. Journal of Hymenoptera Research 98: 509-524. https://doi.org/10.3897/jhr.98.150254
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Two morphologically unusual new neotropical genera of Miracinae, Paramomirax Whitfield and Fernandez-Triana, and Fusimirax Whitfield and Fernandez-Triana, are here proposed and described. The two new genera differ strikingly from previously known Miracinae in having completely different patterns of desclerotization of the anterior metasomal tergites. Paramomirax, so far known only from paramo vegetation at high elevation in Colombia, is represented by P. peckorum Whitfield, new species. Fusimirax, so far known only from lower elevation forests in the Dominican Republic, is described from three species, F. masneri Whitfield & Fernandez-Triana, new species, F. robusta Whitfield, new species, and F. gracilis, Whitfield, new species. None of the four species has any recorded hosts yet, although all previous Miracinae with known hosts attack leaf-mining and bark-mining moth larvae.
Colombia, Dominican Republic, Fusimirax, leaf-mining Lepidoptera, paramo, Paramomirax
The Miracinae are, based on described species, a small subfamily of Braconidae classified within the “microgastroid” lineage, which is now well established as a monophyletic lineage based on both morphological and molecular, including genome-scale, data (
The currently recognized genera (Centistidea Rohwer, Mirax Haliday and Rugosimirax Ranjith & van Achterberg), despite their proposed diagnostic differences, all possess the current set of shared derived characters for Miracinae familiar to braconid taxonomists: antennal flagellomeres fixed at 12; maxillary palpal segments 5 and 6 fused so that palpi appear 4-segmented (ancestral segments 1 and 2 also fused as with many other braconids); wings with a distinctive reduced venation (Fig.
A Centistidea sp., female, CNC, dorsal view of metasoma B (different) Centistidea sp., female, CNC, dorsal habitus C Rugosimirax expectata Ranjith and van Achterberg, dorsal view of mesosoma and anterior portion of metasoma (modified slightly from
Here we report the discovery of two new Neotropical groups of Miracinae which both fall clearly outside the usual metasomal tergite morphology recognized as characteristic for Miracinae, and describe them as new genera, along with distinct species found within them. Nevertheless, these groups share all the other traits typical for Miracinae, so we assign them to Miracinae unambiguously. Clearly these groups ultimately require further analysis of their phylogenetic placement using molecular data, once suitable material of them is available. We describe them here to highlight them since their unusual morphology suggests a somewhat broader definition of the subfamily, as featured below.
As far as is known, all Miracinae are koinobiont endoparasitoids of leaf-mining and bark-mining Lepidoptera (
All specimens included in this study were found in the pinned collections of the Canadian National Collection of Insects, Arachnids and Nematodes in Ottawa (
Morphological terminology generally follows Whitfield (1997), adapted to be consistent with the Hymenoptera Anatomy Ontology (http://portal.hymao.org/projects/32/public/ontology/) terms.
The two genera below differ in metasomal morphology somewhat from the previously accepted diagnosis for Miracinae in the degree and pattern of desclerotization of anterior tergites and laterotergites, and in the shapes and sculpturing of the remaining sclerotized plates. They do not resemble each other in these differences, as will become clear below, and also exhibit several other unique combinations of features not found elsewhere in Miracinae. In particular, the delineation of Miracinae based on the y-shaped pattern of anterior metasomal sclerites as found in Centistidea, Mirax and Rugosimirax can no longer be sustained, although the separation of anterior metasomal spiracles onto the (mostly desclerotized) laterotergites, along with fixation upon 12 flagellomeres in the antenna and 4 maxillary palpal segments, as well as the characteristic reduced wing venation, still hold for all known Miracinae as defined here.
1 | Anterior metasomal tergites extensively desclerotized, leaving Y- or T-shaped sclerotized bridges (“skeleton”) composed of elongate T1 and portions of T2+3 (Fig. |
2 |
– | Anterior metasomal tergites less extensively desclerotized; T1, T2 and T3, if discretely defined, sculptured AND sclerotized bridges not Y- or T-shaped (Figs |
4 |
2 | Propodeum anteriorly with longitudinal carinae on either side submedially but without medial carina (Fig. |
Rugosimirax Ranjith & van Achterberg |
– | Propodeum either with longitudinal medial carina, at least anteriorly (Figs |
3 |
3 | Propodeum with longitudinal medial carina (Fig. |
Centistidea Rohwer (as recognized by authors who treat it as separate from Mirax) |
– | Propodeum smooth medially, without longitudinal medial carina | Mirax Haliday |
4 | Metasomal T1–3 smooth and essentially unsculptured, but not desclerotized except laterotergites of T1 (Figs |
Paramomirax , gen. nov. |
– | Metasomal tergites 2 and 3 fused into a sculptured syntergite resembling some tergites Microgastrinae (T1 still as in other Miracinae) (Figs |
Fusimirax , gen. nov. |
As with the following new genus, Paramomirax shares with other Miracinae the antenna with flagellomeres fixed at 12, the proximal 1–3 of which have no placodes, the more distal ones possessing one rank of placodes, ancestral maxillary palpal segments 5 and 6 fused to reduce the number of segments to four, the characteristic reductions in wing venation (Figs
Paramomirax peckorum Whitfield, paratype female CNC5342831 A hypopygium and ovipositor mechanism, ventral view B lateral habitus C head, dorsal view D head, anterior view E fore wing F mesosoma and metasoma, dorsal view G metasomal and ovipositor mechanism, dorsal view H head, mesosoma and anterior metasomal tegite 1, dorsal view.
The form of the anterior metasomal segments (desclerotization patterns, Y-shaped skeleton of remaining sclerotized tergites (Fig.
The genus name makes reference to the high altitude paramo habitat of the type species, Paramomirax peckorum sp. nov., described below.
Holotype. • Female, Colombia: Quindio, 21 km E. Calarca, 6-III-1974, 10,000’, S. & J. Peck, CNC5342830. Deposited in Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa (
Paratype. • 1 female, same data as holotype, CNC5342831. Deposited in
Size. Body length (excluding ovipositor) 2.3 mm; fore wing length 2.7 mm.
Coloration. Head and mesosoma black except lighter brownish lower genae; antenna black at extreme base (especially scape), pedicel distally and flagellum brown; mandibles and palpi pale brown to golden; legs proximally mostly dark brown, lightening distally on femora and most of tibiae, darker brown tarsi; pigmented parts of wing venation dark brown; metasoma fairly evenly deep brown dorsally, slightly lighter ventrally; ovipositor sheaths deep brown.
Morphology. Face indistinctly punctate, smooth medially; antennae slightly shorter than body, flagellomeres 8–11 only 1.2–1.3 × longer than wide; pronotum laterally with sharp but weakly impressed upper and lower grooves; mesoscutum shiny, very indistinctly punctate, with notauli very weakly indicated on extreme anterior face; scutellum convex, weakly punctate, slightly longer than anteriorly wide, subtriangular but with curved scrobe separating it from mesoscutum; mesopleuron mostly convex and shiny, without any clear central grooves; metanotum mostly sunken relative to shiny raised medial boss; propodeum weakly sculptured, with percurrent longitudinal medial carina, crossed by angular transverse carina at just past midlength; anterior tergite 1 (T1) narrow anteriorly, expanding into elongate oval posterior 0.75; laterotergal region of T1 pigmented light brownish but sclerotized and strongly striate; T2 and T3 shiny, smooth and evenly sclerotized; typical miracine Y-shaped structure visible only as slightly differently colored areas hidden beneath cuticle; laterotergal regions of T2 and T3 evenly sclerotized as on dorsal parts.
T4 evenly sclerotized and shiny, similar to T2+3; hypopygium evenly sclerotized, darker distally, somewhat acuminate but with blunt tip; ovipositor nearly straight, sheaths as long as hind tibiae and setose over entire distal 0.75.
Male. Not known.
Variation. Only two very similar females from the same locality are known.
Only one locality so far, in paramo vegetation over 3,000 m.
Not known, but host presumed to be a small leaf-mining caterpillar as with other Miracinae.
This species has by far the fewest desclerotized regions of the anterior metasoma, and the longest ovipositor, of any miracine yet discovered. It is also one of the largest (roughly 10–20% larger than average).
The specific epithet is meant to honor the collectors, Stewart and Jarmila Kukalova-Peck.
As with the above new genus, Fusimirax shares with other Miracinae the antenna with flagellomeres fixed at 12, maxillary palpal ancestral segments 5 and 6 fused to reduce the number of segments to four, the characteristic reductions in wing venation (Figs
Fusimirax masneri Whitfield and Fernandez-Triana, holotype female A lateral habitus B head and prothorax, anteroventral view C fore wing D head and anterior of mesonotum, dorsal view E antenna F metasoma and ovipositor mechanism, lateral view G propodeum and metasoma, dorsal view H head, mesosoma and anterior of metasoma, dorsal view.
Fusimirax gracilis Whitfield, paratype male CNC5342838 A lateral habitus B head and prothorax, anteroventral view C wings D head and anterior of mesonotum, dorsal view E mesosoma and metasoma, dorsal view F head, mesosoma, dorsal view .
The form of the anterior metasomal segments (desclerotization patterns, Y-shaped skeleton of remaining sclerotized tergites (Fig.
The genus name makes reference to the fusion of T2 and T3 into an obvious syntergite.
1 | Mesoscutum light orange-brown, contrasting with darker mesosomal regions (Fig. |
Fusimirax masneri Whitfield & Fernandez-Triana, sp. nov. |
– | Mesoscutum dark brown to black, concolorous with other mesosomal regions (Figs |
2 |
2 | Syntergite of T2+3 less than 2 × as long as width at posterior end, surface roughly rugose with weak overlay of longitudinal sculpture (Fig. |
F. robusta Whitfield, sp. nov. |
– | Syntergite of T2+3 slightly greater than 2 × as long as width at posterior end, surface coarsely longitudinally carinate (Fig. |
F. gracilis Whitfield, sp. nov. |
Holotype. • Female, Dominican Republic: Barahona Sra. Bahoruco, Loma Remigio, 800 m, cloud forest III-1991, L. Masner, CNC5342832 (
Paratypes. None.
Size. Body length 2.0 mm; fore wing length 2.2 mm.
Coloration. Head dorsally bark brown, shading to much lighter ventrally; especially clypeus; mouthparts and palpi pale yellowish; scape and pedicel yellowish, remainder of antenna brown; mesosoma anteriorly (pronotum and propleuron, dorsal portion of mesopleuron, tegulae and mesoscutum) light yellow-brown, remainder darker brown; legs all pale yellowish; pigmented portions of wing venation pale brownish, including pterostigma; metasoma anteriorly with whitish laterotergites and lateral edges of T2/T3 syntergite; T1 and most of syntergite dark brown; T4 and succeeding terga paler brown; ovipositor sheaths darker brown, especially distally.
Morphology. Face very weakly and indistinctly punctate throughout; raised medially; antennae slightly longer than body, slender, with even distal flagellomeres at least twice as long as broad; propleuron with dorsal groove weakly crenulate, ventral groove broad, arcuate and smooth; mesoscutum weakly sculptured, with clear notauli over anterior 0.4 of its length (Fig.
Male. Not known.
Variation. Only one female specimen available.
So far, only one locality at 800 m in the Dominican Republic. The label says “cloud forest”, but this seems probably not quite accurate based on the locality and elevation.
Not known, but host presumed to be a small leaf-mining caterpillar as with other Miracinae.
The strikingly colored mesoscutum and metasoma combination is unique.
The specific epithet honors the great hymenopterist Lubomir Masner, the collector who sampled in the Dominican Republic over multiple years.
Holotype. • Female, Dominican Republic: Barahona Sra. Bahoruco, Loma Remigio, 800 m, cloud forest III-1991, L. Masner, CNC5342833 (
Paratypes. None.
Size. Body length 1.9 mm; fore wing length 2.1 mm.
Coloration. Head, mesosoma (except lighter brown pronotum and propleuron) and metasomal syntergite dark brown to blackish, posterior segments of metasoma lighter brown; scape light yellow-brown; remainder of antenna dark brown; mandible light brown, palpi pale yellowish; legs entirely light yellow-brown; tegulae and pigmented parts of wing venation including pterostigma light brownish; laterotergites of T1 and T2 almost white, contrasting with dark syntergite; ovipositor sheaths deep brown.
Morphology. Face smooth, with raised medial ridge; antennae slightly longer than body, with apical flagellomeres roughly twice as long as broad; propleuron with dorsal and ventral grooves represented by very fine and poorly impressed lines; mesoscutum with satiny reflections and distinctly punctate over most of surface except posterolateral corners; notauli indistinct, even anteriorly; scutellum polished and raised centrally, weakly convex, clearly longer than anteriorly broad; mesopleuron weakly roughened, convex, with slight weak depression centrally but no obvious grooves; metanotum mostly sunken, with some abortive cross-ridges but raised medial boss not extending to anterior edge; propodeum finely sculptured, with percurrent longitudinal medial carina splitting at midlength to produce a small elongate areola-like structure, met laterally by angled transverse carinae; T1 narrow anteriorly, widening to elongate-oval raised broader posterior end with some sculpturing; laterotergal region of T1 whitish, with central area of striation; T2 and T3 fused into elongate sculptured syntergite; T2 portion bell-shaped (wider posteriorly) with lateral flanges at grooved T2/T3 junction; T3 roughly parallel-sided; laterotergal regions of T2 and T3 whitish, with some striations; T4 less strongly sclerotized and smooth; hypopygium evenly sclerotized and subtriangular with sharp tip; ovipositor nearly straight; sheaths roughly half as long as hind tibiae with setae over distal half.
Male. Unknown.
Variation. Only one female known.
Same locality as F. masneri (see remarks there).
Not known, but host presumed to be a small leaf-mining caterpillar as with other Miracinae.
The shape of the syntergite is diagnostic, and the propodeal carinae are unique as well.
The epithet “robusta” refers to the anterior metasomal syntergite shape, not to the overall body shape, which is perhaps only slightly more robust than the other species.
Holotype. • Female, Dominican Republic: Rancho Arriba, 26-III-1978, 400 m, L. Masner, CNC5342834 (
Paratypes. • 3 males, Dominican Republic: Baharona, 4 km N. Paraiso, 150 m, 22-III-91, L. Masner, CNC5342835, CNC5342836, CNC5342837; • 2 males, Dominican Republic: Colonia, Cord. Central, 1000 m, 25-III-1978, L. Masner (LM), CNC5342838, CNC5342839. All deposited in
Size. Body length 1.6 mm; fore wing length 1.7 mm.
Coloration. Head, mesosoma (except lighter brown propleuron) and metasomal syntergite dark brown, T1 light yellowish and posterior segments of metasoma lighter brown; mandibles and palpi pale yellowish; scape pedicel and proximal two flagellomeres pale yellow-brown, remaining flagellomeres shading gradually to very slightly darker brown at distal end of antenna; legs entirely pale yellow-brown; tegulae and pigmented portions of wing venation (including pterostigma) very pale yellow-brown; laterotergites of T1 and T2 whitish; entire venter of metasoma very pale except slightly darker anterior half of hypopygium; ovipositor sheaths pale yellow-brown.
Morphology. Face rather smooth, convex to a weak ridge medially; antennae slightly shorter than body, slender, with apical flagellomeres about 2 × as long as broad; propleuron with smooth dorsal and ventral grooves joined together posteriorly by another smooth groove; mesoscutum with satiny reflections and sparsely but distinctly punctate except smoother posterior margin; notauli poorly defined but extending over anterior half; scutellum smooth, nearly flat and longer than broad; mesopleuron very faintly sculptured, with poorly-defined groove/depression centrally; metanotum mostly deeply sunken, with abortive cross-carinae and medial boss raised only over posterior half; propodeum weakly sculptured, with percurrent longitudinal medial carina crossed by distinct angled transverse carinae just past midlength; T1 very narrow and elongate, abruptly broadening at posterior end to meet T2; laterotergal region of T1 strongly whitish, with central striation; T2 and T3 fused to form longitudinally sculptured elongate syntergite which broadens gradually to T2/T3 flared and weakly grooved junction, parallel-sided over T3; laterotergal regions of T2 and T3 whitish, with some regions of striation; T4 evenly sclerotized but weakly pigmented and translucent; hypopygium evenly sclerotized to fairly sharp tip; ovipositor nearly straight; sheaths less than half as long as hind tibiae with setae over distal half.
Male (Fig.
Variation. As described above for males.
So far known only from Dominican Republic, but extending from 150 m to 1000 m elevation, thus overlapping elevationally with the other species of this genus.
Not known, but host presumed to be a small leaf-mining caterpillar as with other Miracinae.
Based on available material, this may be the most common Fusimirax, but we have so few overall that it would be difficult to extrapolate much.
The specific epithet “gracilis” refers to the small and delicate body of these wasps, especially pronounced in this species but true to some extent for most Miracinae.
JBW would like to thank the CanaColl Foundation for funding his visit to the