Research Article |
Corresponding author: Gabriel A. R. Melo ( garmelo@ufpr.br ) Academic editor: Michael Ohl
© 2023 Diego N. Barbosa, Gabriel A. R. Melo.
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:
Barbosa DN, Melo GAR (2023) Revision of Bethylinae from Dominican amber, with description of a new genus (Hymenoptera, Bethylidae). Journal of Hymenoptera Research 96: 167-180. https://doi.org/10.3897/jhr.96.100862
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Fossil species of bethyline wasps from Dominican amber are revised. Based on a cladistic analysis of female morphological data we recognize a new genus and species, †Crassibethylus dominicanus gen. et sp. nov., which came out as sister group of the clade Odontepyris + Prosierola. We also transfer to Bethylidae a fossil taxon originally described in Sclerogibbidae, placing †Pterosclerogibba Olmi as a new junior synonym under Goniozus Förster, in the new combination †Goniozus antiquus (Olmi). Additionally, the third species of Bethylinae known from Dominican amber, †Goniozus respectus Sorg, is diagnosed and illustrated with color images of the holotype. An identification key to the species of Bethylinae from Dominican amber is provided.
Aculeata, aptery, Chrysidoidea, Lepidoptera, Miocene
Bethylinae have their diversity represented by 540 species distributed into 11 genera, and recorded around all zoogeographical regions (
The females of Bethylinae are idiobiont ectoparasitoids mostly of lepidopterous mature larvae. The majority of host records are based on species of Goniozus, and according to
In Bethylinae, there is a reduced sexual dimorphism, thus being easy to establish sex associations in most groups. The genera belonging to Bethylinae are recognized based on the following combination of characteristics: antenna with 10 or 11 flagellomeres; mandible thick, short, with four sharp apical teeth; clypeus extending posterad into frons, with clypeal carina strongly outcurved in profile; dorsal pronotal area with posterior margin outcurved medially; legs with tarsal claws bifid and strongly angled; forewing with first abscissa of vein Rs+M tubular, minimally as a stub; sting with posterior area of second valvifer with anterior slender projection (
The phylogenetic relationships within Bethylinae have received the attention from previous studies conducted by
In the present work we revise the fauna of Bethylinae known from Dominican amber. Our starting point was an intriguing fossil inclusion that at first sight suggested a female sclerogibbid wasp due to its distinctly swollen profemora. Further study revealed that instead it represented a Bethylinae, showing many features in common with Odontepyris and Prosierola, and which is described here as new genus. During our investigations we also concluded that †Pterosclerogibba antiqua, a Dominican amber fossil described by
The amber piece containing the fossil inclusion that prompted this study is deposited in the Departament of Zoology, of the Universidade Federal do Paraná, Curitiba, Brazil (
Morphological terminology follows mainly
We used the key proposed by
In order to establish the phylogenetic position of the bethyline taxon under investigation, we used
The tree searches followed all parameters used by
The specimen images were obtained by a LEICA DFC295 digital camera, and the stacking was carried using the software Zerene Stacker. The figures were prepared in GIMP (2.10.18). The images of the type specimen of †Goniozus respectus Sorg were provided by the depositary institution.
The implied weighting analysis, under K=13.3, resulted in six most parsimonious trees, whose strict consensus is shown in Fig.
Although we did not code the fossil bethyline under investigation as possessing a gibbous mesopleuron, it has a considerably swollen mesopleuron, therefore reinforcing its similarity to the clade Prosierola + Odontepyris. Also, it shares with Prosierola the posterior margin of the pronotum slightly produced backward medially (8:1). Considering its unique position and close relationship with Prosierola and Odontepyris, we propose a new fossil genus to accommodate this Dominican amber bethyline.
Order Hymenoptera Linnaeus, 1758
Superfamily Chrysidoidea Latreille, 1802
Family Bethylidae Haliday, 1839
†Crassibethylus dominicanus sp. nov.
Maxillary palp with four articles; prosternum shorter than area of procoxa; pro- and metafemora strongly swollen, profemur lenticular-shaped, about 0.6× as wide as long; posterior margin of pronotum slightly produced backward medially; mesoscutellar groove present as a narrow sulcus; forewing vein 2Cu present; insertion of metacoxae displaced anteriorly between mesocoxal insertion; ventral carina of metasomal petiole straight.
Female. Head. Head as long as wide; antenna with 11 flagellomeres; clypeus with median clypeal carina continuous with frontal carina, lateral lobe not evident; anterior margin of frons projected anteriorly. Eye large, occupying most of lateral portion of head. Ocelli located close to vertex crest; anterior angle of ocellar triangle obtuse. Vertex straight. Mesosoma. Pronotum with dorsal pronotal area longer than mesoscutum, its posterior margin slightly produced backward medially. Mesoscutum without notaulus; parapsidal signum absent. Mesoscutellar groove as a narrow sulcus, lacking enlarged lateral pits. Metapectal-propodeal complex coriaceous; metapostnotum indistinct, apparently without metapostnotal median carina and metapostnotal-propodeal suture; posterior transverse carina present. Mesopleuron somewhat swollen in dorsal view, coriaceous, not sculptured; mesopleural pit shallow. Prosternum relatively large, somewhat rhombic-shape. Wings. Forewing with costal cell conspicuous; second radial cell closed, short, almost triangular; length of vein Rs2 as long as vein M2; length of vein Rs2+M2 as long as vein 1Rs2; stigma well developed, bula present. Legs. Pro- and metafemora strongly swollen, profemur lenticular-shaped, about 0.6× as wide as long. Insertion of metacoxae displaced anteriorly between mesocoxal insertion. Tarsal claws strongly curved, with a large basal expansion. Metasoma. Sting with 3vv narrow and long.
The genus name is an allusion to the strongly swollen profemora, from Latin crassus, thick, fat, stout, and Bethylus, the type genus of Bethylinae.
Holotype
female, amber inclusion from the Dominican Republic (
Holotype. Female. Body length 3.2 mm; forewing 1.95 mm long. Color. Wings clear hyaline; antenna, mandible, palpi, tegula, legs, and wing venation light castaneous; metasoma castaneous; head and mesosoma dark castaneous. Head. Head as long as wide in dorsal view, triangular-shaped in both frontal and lateral views. Mandible narrow and evenly wide along its extension. Clypeus with median lobe rounded, well projected; median carina well developed, continuous with frontal carina; lateral lobe inconspicuous. Antenna with flagellar pubescence sparse, with few outstanding erect setae; scape 3.0× longer than pedicel; pedicel as long as first flagellomere; flagellomeres subequal in length. Eye not protruding, elliptical, large, and glabrous; area between anterior margin of eye and demarcation of median clypeal lobe clearly concave. Frons coriaceous and finely punctate. Posterior ocelli close to vertex crest; anterior angle of ocellar triangle obtuse; anterior ocellus aligned with upper tangent of eyes. Vertex crest straight, sparse thick setae at corner. Hypostomal carina straight. Mesosoma. Pronotum wider than long, coriaceous and finely punctate; anterior corner angled. Metapectal-propodeal complex with lateral marginal carina. Wings. Forewing with R2 cell longer than 1Cu2 cell; only one longitudinal flexion line present, not forming rectangular area. Metasoma. About 2.3× longer than wide, polished, whole surface glabrous. Genitalia with 3vv narrow and longer than wide.
Male. unknown.
The specific epithet dominicanus is an allusion to the Dominican Republic, country from the where the amber piece originated.
Goniozus Förster, 1856: 95. Type species: Bethylus claripennis Förster, 1851.
Parasierola Cameron, 1883: 197. Type species: Parasierola testaceicornis Cameron, 1883.
Progoniozus Kieffer, 1905: 105. Type species: Perisemus floridanus Ashmead, 1905.
Perisierola Kieffer, 1914: 533. Type species: Parasierola gallicola Kieffer, 1905.
†Pterosclerogibba Olmi, 2005: 186. Type species: †Pterosclerogibba antiqua Olmi, 2005, syn. nov.
Goniozus Förster is the second most speciose genus in Bethylinae, being composed by 174 described species, three of them representing fossil taxa, two from Baltic amber and one from Dominican amber (
The fossil fauna of Dominican amber is interpreted here as containing two species of Goniozus. In addition to †G. respectus, described by
†Pterosclerogibba antiqua Olmi, 2005: 186. Female holotype. Dominican Republic: amber from La Toca mine (GPCO, not examined).
Antenna, tibia, and tarsi light castaneous; head and mesosoma castaneous; metasoma dark-castaneous to nearly black. Head longer than wide, not punctured; clypeal carina strong, but not extending over frons; antenna relatively short, apical portion moniliform; malar space as long as eye length; eye small, without pilosity; ocellar triangle with a right anterior angle; vertex concave, apparently without thick setae. Mesosoma without conspicuous setae; posterior margin of dorsal pronotal area concave posteriorly; mesoscutum shorter than dorsal pronotal area, not punctured, notaulus absent; mesopleuron not gibbous; mesoscutellum as long as mesoscutum, not punctured; metapectal-propodeal complex as long as mesonotum. Forewing with clearly developed pterostigma, venation apparently reduced. Proleg with a swollen femur, almost 2.0× longer than wide, metafemora not distinctly swollen. Metasoma without visible setae; sting curved downward.
Within Bethylinae, Olmi’s species can be accommodated in Goniozus based on its general morphology and especially for lacking diagnostic features found in other genera of Bethylinae. Considering that Goniozus, as currently defined (see
†Goniozus respectus Sorg, 1988: 86. Female holotype. Dominican Republic: amber from unknown mine (SGN, examined through photographs).
Antenna, clypeus and legs light castaneous; head and mesosoma dark-castaneous nearly black. Head as long as wide, sparsely punctured; antenna with flagellomeres with large sensilla, distinctly moniliform; malar space shorter than eye length; eye large, without pilosity; ocellar triangle with anterior angle obtuse; vertex straight, with few scattered setae. Mesosoma setae as long as head setae; posterior margin of dorsal pronotal area concave; mesoscutum shorter than dorsal pronotal area, punctured, notaulus absent; mesoscutellum as long as mesoscutum, punctured; metanotum with metascutellum visible; metapectal-propodeal complex longer than mesonotum, lateral marginal carina of the metapectal-propodeal complex present. Forewing with clearly developed pterostigma, 1M2 cell, Rs2 vein not reach margin. Legs with profemur almost 3.0× longer than wide, metafemora not distinctly swollen. Metasoma with setae slightly longer than those on mesosoma.
The species was described by
1 | Profemur and metafemur strongly swollen, profemur about 0.6× as wide as long; mesopleuron swollen; insertion of metacoxae displaced between mesocoxae | † Crassibethylus dominicanus gen. et sp. nov. |
– | Profemur and metafemur not as swollen, profemur much less than 0.6× as wide as long; mesopleuron not swollen; metacoxae inserted posteriorly to mesocoxal insertion | 2 (Goniozus) |
2 | Head longer than wide, with malar space as long as eye length; eye small; ocellar triangle with anterior angle right; vertex concave; notauli absent; metapectal-propodeal complex as long as mesonotum; profemur almost 2.0× longer than wide | † G. antiquus (Olmi) |
– | Head as long as wide, with malar space shorter than eye length; eye large; ocellar triangle with anterior angle obtuse; vertex straight; mesoscutum with notauli; metapectal-propodeal complex longer than mesonotum; profemur almost 3.0× longer than wide | † G. respectus Sorg |
The discovery of an enigmatic bethylid wasp in Dominican amber has provided us with an opportunity to reassess the taxa currently known from this fossil deposit. The inclusion of the new fossil in the dataset assembled by
Our phylogenetic results also shed light on the age and biogeography of the Odontepyris + Prosierola clade. While the species of Odontepyris are distributed in the Old World, being presently recorded from the Afrotropical, Australian, Oriental and Palearctic regions, Prosierola is restricted to the New World, including the Greater Antilles (
Although recognition of a new taxon at genus level in Bethylinae does not solve the paraphyletic condition of Goniozus, we think that simply subsuming Odontepyris and Prosierola (and for that matter †Crassibethylus gen. nov. as well) within Goniozus is not a reasonable solution to the problem. These taxa have distinct morphologies and occupy distinct regions of the globe. Future work on Bethylinae should focus on establishing the limits of the species groups within Goniozus, so that a new genus-level classification could be proposed on more solid ground. There are already some genus-group names available, currently synonymized under Goniozus, that could be reinstated for some of these lineages.
It is also important to call attention to additional contributions of our work to the understanding of the evolution within Sclerogibbidae. The removal of †Pterosclerogibba from Sclerogibbidae to Bethylidae, placed here as a synonym of Goniozus, makes the fauna of Sclerogibbidae known from Dominican amber restricted to †Probethylus poinari, a taxon described from an apterous female and exhibiting a condition similar to that found in the extant species (
We would like to thank Magno S. Ramos for providing the original files of his dataset and for discussing some characters of Bethylinae; Brunno B. Rosa for his help in photographing the fossil inclusion described here; and Mónica M. Solórzano Kraemer for providing photographs of the type specimen of †Goniozus respectus. Financial support has been received through a research scholarship from CNPq (grant 313588/2021-0).
Character matrix
Data type: Morphogical (Excel spreadsheet)
Explanation note: Morphological character matrix.
Phylogenetic tree
Data type: (PDF file)