Research Article |
Corresponding author: Gabriel A. R. Melo ( garmelo@ufpr.br ) Academic editor: Michael Ohl
© 2020 André L. Martins, 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:
Martins AL, Melo GAR (2020) Revision of the fossil species of Thaumatodryinus Perkins from Dominican amber, with a new combination and description of a new species (Hymenoptera, Dryinidae). Journal of Hymenoptera Research 79: 77-88. https://doi.org/10.3897/jhr.79.57686
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The fossil species of Thaumatodryinus from Dominican amber are studied, and the first revision is presented with a key to the known taxa. We recognize three species, T. miocenicus Olmi, 1995, T. priscus (Olmi, 1998), comb. nov. and T. fuscescens sp. nov. The current classification of the genus and relationships between fossil and living species are discussed. Comments on the host records for Thaumatodryinus are presented.
Aculeata, Chrysidoidea, Miocene, Thaumatodryininae, wasps
In recent years, a significant number of species of Dryinidae have been described from different fossiliferous deposits (
Thaumatodryininae is a small subfamily of Dryinidae, known to attack nymphs of the auchenorrhynchous Flatidae (
In this study, we contribute for the knowledge of Dryinidae by reviewing the fossil species of Thaumatodryinus from Dominican amber and propose a new species for the genus. Furthermore, the status of Dryinus priscus Olmi, 1998 is reinterpreted based on our comparative morphological study.
The amber pieces studied here are deposited in the Universidade Federal do Paraná, Curitiba, Brazi (
Morphological terminology follows
In the description and diagnosis the following abbreviations were used: POL, refers to the minimum distance between the inner edges of the lateral ocelli; OL, refers to the minimum distance between the inner edges of the lateral ocellus and the median ocellus; OOL, refers to the minimum distance from the outer edge of the lateral ocellus to the eye inner margin; OPL, refers to the minimum distance from the posterior edge of a lateral ocellus to the occipital carina; and TL, refers to the minimum distance from the posterior edge of eye to the occipital carina. The measurements provided for ocellar ratio, antennomeres and for the fore leg articles represent relative values.
Study of the inclusions was carried out in a Leica M125 stereomicroscope. Color images of Thaumatodryinus fuscescens sp. nov. were obtained by a LEICA DFC295 digital camera, and those of T. miocenicus Olmi and Dryinus priscus Olmi by a Nikon DS-Ri2 using NIS-Elements D–Z-series 11*. Image stacking was carried using the software Zerene Stacker. The figure plates were prepared using Adobe Photoshop (version Cs6).
Subfamily Thaumatodryininae Olmi, 1984
Thaumatodryinus koebelei Perkins, 1905, by monotypy.
Among living and fossil dryinids, the species of Thaumatodryinus can be recognized by the following combination of characters: mandible with four teeth in both sexes, teeth progressively larger from anterior to posterior ends along cutting edge; outer surface of mandible with a distinct basal sulcus, extending from the anterior to the posterior condyle; mid portion of anterior margin of clypeus straight or convex; occipital carina complete in fossil taxa and in males of most living species, females of many living taxa with incomplete carina; maxillary and labial palpomeres in proportion 6:3; antenna filiform in both sexes; flagellomeres 3–7 of female with single set of rhinaria on its mid length, flagellomere 8 with two sets of rhinaria along its length; rhinaria on flagellomeres 5–8 each with four long setae, two at each side; pronotum saddle-shaped and crossed by strong transverse impression at central portion, pronotal lobe reaching tegula; anterior margin of mesoscutellum with a foveate groove; fore wing with 2r-rs shorter than 3Rs&4Rs; female with tarsal claw simple and with basal expansion, male with bifid claw, without basal expansion; apex of 3rd tarsomere of fore leg with variable number of thick bristles in female; chela with rudimentary claw, enlarged claw with one row of lamellae and two subapical teeth laterally on outer surface; female hind coxa with a distinct basal projection ventrally; dorsal and posterior surfaces of propodeum convex; tibial spur formula for both sexes 1/1/2.
1 | Eyes hemispherical, about 1.1× longer than wide; maximum head width about 1.9× distance between inner margins of tegulae; antenna with 1st flagellomere as long as 2nd (Fig. |
Thaumatodryinus fuscescens sp. nov. |
– | Eyes slightly longer, about 1.3–1.4× width; maximum head width about 1.7× distance between inner margins of tegulae; proportion between 1st and 2nd flagellomeres variable; POL shorter than 2.5× OL; OOL about 5× OPL; lower margin of clypeus convex; pronotum surface mostly granulate; sculpture of mesoscutum variable; notauli not extending along entire length of mesoscutum; anterior margin of mesoscutellum with a groove | 2 |
2 | Antenna with 1st flagellomere shorter than 2nd (Fig. |
Thaumatodryinus miocenicus Olmi |
– | Antenna with 1st flagellomere almost as long as 2nd (Fig. |
Thaumatodryinus priscus (Olmi) comb. nov. |
Thaumatodryinus fuscescens sp. nov. is characterized by the body predominantly testaceous, except head and part of pronotum and remainder of mesosoma black; eyes hemispherical, about 1.1× longer than wide; maximum head width about 1.9× distance between inner margins of tegulae; frontal line present; mid portion of clypeus with anterior margin straight; antenna with 1st flagellomere as long as 2nd; POL 3.2× OL; OOL about 6× OPL; occipital carina complete; notauli percurrent, extending from anterior to posterior margins of mesoscutum.
Female holotype (Fig.
†Thaumatodryinus fuscescens sp. nov., female holotype. A habitus, dorsal view B right antenna, dorsal view C head and mesosoma, antero-ventral view D head and mesosoma, dorsal view E apical portion of flagellum, with details of flagellomeres 5–8 F–G Apical portion of fore leg, with details of the chela. Scale bars: 2 mm (A), 0.5 mm (B), 0.5 mm (C–D), 0.3 mm (E–G).
Thaumatodryinus fuscescens sp. nov. is more similar to the extant Neotropical fauna than to the other two species found in the Dominican amber. It has in common with some of the living species a convex vertex and slightly elevated ocellar triangle, straight lower clypeal margin, complete occipital carina, percurrent notauli and anterior margin of mesoscutellum with a narrow row of small foveae. Among the fossil species studied here, T. fuscescens sp. nov. is more similar to T. miocenicus and differs from T. priscus by the head shape with eyes protuberant on the lateral side, frontal line present and ratio of OL short than TL. The new species differ from T. miocenicus and T. priscus by the granulated integument of the head; clypeus with lower margin straight; 1st flagellomere as long as 2nd; hemispherical-shaped eyes; and percurrent notauli. The chela of T. fuscescens is closed and for this reason it is not possible to determine the number of lamellae in the enlarged claw and in the 5th tarsomere.
The species is named in reference to its overall dark coloration, from the Latin fuscus, dark, dusky, and -escens, beginning of, becoming.
Holotype female, in amber from the Dominican Republic (piece
Thaumatodryinus miocenicus
Olmi, 1995: 263. Female holotype. Dominican Republic: amber from unknown mine, probably El Valle mine (
Female holotype, 5.6 mm (see fig. 1–2 in
While in the original description
Female holotype, in amber from the Dominican Republic: amber from an unknown mine (probably El Valle mine) DR-99-146 HT (
Dryinus priscus
Olmi, 1998: 77. Female holotype. Dominican Republic: amber from unknown mine, probably El Valle mine (
Female holotype, 4.0 mm (see fig. 55 in Olmi 1998). Thaumatodryinus priscus is easily recognized by the body testaceous, except by the dark petiole and mesoscutellum apparently black or brown. Antenna with dense and short setae and rhinaria present in flagellomeres 3–8. Head slightly convex, apparently granulated; eye somewhat bulging; frontal line absent; mid portion of clypeus with anterior margin convex. Pronotum granulated; mesoscutum granulated, notauli incomplete, reaching approximately 0.7 of mesoscutum length; mesoscutellum shorter than mesoscutum, with sculpture not clearly evident; anterior margin of mesoscutellum with broad, shallow groove; metanotum shorter than mesoscutellum, with indistinct sculpture; propodeum reticulate rugose and distinctly long, about as long as high in lateral view. Ocellar ratio: POL= 5; OL= 2.5; OOL= 10; OPL= 2; TL= 3. Antennomeres in following proportions: 9:5:20:19:23:27:20:15:10:12. Eye elongated, about 1.4× longer than wide; maximum head width about 1.7× distance between inner margins of tegulae; pronotum shorter than head (15:26); mesoscutum as long as pronotum (16:15). Fore wing with one dark transverse band and vein 2r-rs shorter than 3Rs&4Rs (11:21). Fore leg segments in following proportions: 19 (coxa): 20 (trochanter): 45 (femur): 30 (tibia): 11 (1th tarsomere): 4 (2nd tarsomere): 6 (3th tarsomere): 21 (4th tarsomere): 31 (5th tarsomere); 5th tarsomere slightly longer than enlarged claw (31:27) and with two rows of numerous lamellae (exact number hard to observe); enlarged claw apparently with one subapical tooth and with one row of numerous lamellae (exact number hard to observe). Propodeum distinctly long, about as long as high in lateral view, dorsal surface as long as posterior surface.
This species was originally described in the genus Dryinus by Olmi (1998) and redescribed as such by
†Thaumatodryinus priscus (Olmi, 1998), female holotype. A habitus, lateral view B habitus, ventral view C head and mesosoma, lateral view D head and mesosoma, dorsal view E head and mesosoma, ventral view F apical portion of fore leg, with details of the chela. Scales: 1 mm (A), 1 mm (B), 0.5 mm (C), 0.5 mm (D–F).
Female holotype, in amber from Dominican Republic: amber from an unknown mine (probably El Valle mine) DR-14-341 (
All fossil species of Thaumatodryininae are currently known only from Dominican amber. Species described originally in Thaumatodryinus from Baltic amber by Brues (1923, 1933) have been all transferred to Harpactosphecion, a distantly related fossil genus belonging to the Dryininae (
We document here the presence of three species of Thaumatodryinus in Dominican amber. Two of them, T. fuscescens sp. nov. and T. miocenicus, resemble the extant species in their protuberant and hemispherical eyes, and a more compact propodeum. Thaumatodryinus priscus is more divergent, possessing more elongated eyes and a longer propodeum. In general, it has a more slim, elongated body and this might have contributed to its interpretation as a Dryinus by Olmi (1998). Also, the amber piece containing the holotype has been further polished after the original description, allowing for a better view of the inclusion, as can be seen in the photographs provided here.
As far as it is known, species of Thaumatodryinus attack only the fulgoroid family Flatidae and host records are known for four species of Thaumatodryinus from the Australian, Nearctic, Neotropical and Oriental regions (
We would like to thank James Carpenter and David Grimaldi for granting access to the collections under their care and for their kind help during ALM’s visit to the American Museum of Natural History (