Research Article |
Corresponding author: Ralph Peters ( r.peters@zfmk.de ) Academic editor: Michael S. Engel
© 2015 Marcel Bläser, Lars Krogmann, Ralph Peters.
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:
Bläser M, Krogmann L, Peters RS (2015) Two new fossil genera and species of Cerocephalinae (Hymenoptera, Chalcidoidea, Pteromalidae), including the first record from the Eocene. ZooKeys 545: 89-100. https://doi.org/10.3897/zookeys.545.6470
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Cerocephalinae (Chalcidoidea, Pteromalidae) is a small group of parasitoid wasps characterized by a number of derived diagnostic features. Their hosts are endophytic beetles. So far, 43 species of Cerocephalinae have been described, including one fossil species from the Miocene. In this study, we add two new genera and species from Baltic and Dominican amber to the fossil record. Tenuicornus dominicus gen. et sp. n. is the second genus described from Dominican amber, and Pteropilosa lailarabanorum gen. et sp. n., described from Baltic amber, represents the oldest record of the subfamily, pushing the minimum age of Cerocephalinae back to the Eocene. Diagnostic characters of both species are discussed in comparison with other Cerocephalinae. An updated key to extant and fossil Cerocephalinae is presented.
Tenuicornus , T. dominicus , Pteropilosa , P. lailarabanorum , Miocene, Baltic amber, Dominican amber
Cerocephalinae is one of the smaller, yet most distinctive subfamilies of Pteromalidae (Hymenoptera, Chalcidoidea). Cerocephaline wasps are comparatively easy to recognize by a prominence in the intertorular area that can be a longitudinal carina or a tooth-like structure of varying size, by typical wing vein length ratios, and by two spurs on the hind tibia. Including the fossil record, only 43 species of Cerocephalinae in 14 genera have been described (
Recent studies (
Cerocephalinae stand out as particularly underrepresented in the fossil record, with only one representative from Dominican amber, Dominocephala vibrissae Krogmann, 2013, known so far. In this study, two additional fossils representing two new genera and species are described. One of the new genera was found in Dominican amber (same as Dominocephala vibrissae), which is estimated to be 20–15 million years old (
Terminology follows
Pteropilosa lailarabanorum gen. et sp. n. in dorsolateral view, showing general habitus and diagnostic characters. A General habitus B Fore wing with distinct pilosity on disc, with comparatively short marginal vein, and without a tuft of erect setae on the parastigma (pst)C Head and mesosoma, showing surface sculpture on pro-, and mesonotum, and distinct frenal line (fl)D Head and antennae, showing tooth-shaped intertorular prominence (ip) positioned slightly below insertion of antennae; antennae inserted well above ventral margin of eyes, segments 3–6 of the 6-segmented funicle distinctly transverse.Scale: 500 µm (A); 200 µm (B–D).
Intertorular area with prominence that can be a carina or tooth-shaped (also referred to as inner antennal process (iap)). Fore wing disc bare or with setae reduced to setal bases (pilose only in Pteropilosa gen. n.), with marginal fringe, fore wing always with thickened juncture between marginal vein and submarginal vein (parastigma), marginal vein elongate, postmarginal and stigmal vein shortened, postmarginal vein usually shorter than stigmal vein. Mesonotum with complete notauli. Hind tibia with two spurs.
This key is modified after
1 | Antenna with five funicular segments in females and six funicular segments in males | 2 |
– | Antenna with six funicular segments in females (Fig. |
4 |
2 | Antenna shortened, with all funicular segments transverse; fore wing without a tuft of erect setae on parastigma (apterous species occur) | Choetospilisca Hedqvist, 1969 [USA, Brazil, India] |
– | Antenna of normal size, with all funicular segments longer than or about as long as wide; fore wing with or without a tuft of erect setae on parastigma | 3 |
3 | Fore wing with a tuft of erect setae on parastigma | Theocolax Westwood, 1832 (in part) [Cosmopolitan] |
– | Fore wing without a tuft of erect setae on parastigma (as in Fig. |
Acerocephala Gahan, 1946 (in part) [USA, Samoa] |
4 | Fore wing disc distinctly pilose (Fig. |
Pteropilosa gen. n. [fossil, Eocene Baltic amber] |
– | Fore wing disc bare, setae (if present) reduced to setal bases | 5 |
5 | Head, pronotum and mesoscutum sculptured; fore wing without a tuft of erect setae on parastigma | 6 |
– | Head dorsally and mesoscutum entirely smooth and polished; pronotum usually polished but sometimes partly sculptured; fore wing with or without a tuft of erect setae on parastigma | 10 |
6 | Mandibles enlarged and conspicuous, at least 1/3 as long as head capsule | 7 |
– | Mandibles of normal shape and dimensions, less than 1/3 as long as head capsule | 9 |
7 | Mandibles without a basal process | Muesebeckisia Hedqvist, 1969 [Brazil] |
– | Mandibles with a basal process | 8 |
8 | Mandibles with two distinct teeth | Gnathophorisca Hedqvist, 1969 [Brazil] |
– | Mandibles with three distinct teeth | Gahanisca Hedqvist, 1969 [Brazil] |
9 | Antenna inserted at level of ventral margin of eye | Neosciatheras Masi, 1917 [Seychelles] |
– | Antenna inserted much higher than ventral margin of eye | Sciatherellus Masi, 1917 [Seychelles] |
10 | Mandible elongate and conspicuous, 1/3 to 3/4 as long as head capsule | 11 |
– | Mandible of normal size, far less than 1/3 as long as head capsule | 13 |
11 | Head with row of conspicuously elongate setae extending dorsally from the lower facial process; fore wing without a tuft of erect setae on parastigma | Dominocephala Krogmann, 2013 [fossil, Miocene Dominican amber] |
– | Head without a row of conspicuously elongate setae; fore wing with or without a tuft of erect setae on parastigma | 12 |
12 | Mandible two-dentate; fore wing with a tuft of erect setae on parastigma | Paralaesthia Cameron, 1884 [Panama] |
– | Mandible four-dentate; fore wing without a tuft of erect setae on parastigma | Acerocephala Gahan, 1946 (in part) [USA, Samoa] |
13 | Antenna inserted distinctly below level of ventral margin of eye; head in dorsal view parallel-sided | Theocolax Westwood, 1832 (in part) [Cosmopolitan] |
– | Antenna inserted at or slightly below level of ventral margin of eye; head in dorsal view not parallel-sided | 14 |
14 | Fore wing with a tuft of erect setae on parastigma | 15 |
– | Fore wing without a tuft of erect setae on parastigma | 16 |
15 | Antenna with all funicular segments transverse, i.e., shorter than wide; propodeum with median carina | Paracerocephala Hedqvist, 1969 [Zaire] |
– | Antenna with all funicular segments longer than wide or at least as long as wide; propodeum without median carina | Cerocephala Westwood, 1832 [Cosmopolitan] |
16 | Antenna with all funicular segments longer than wide, cylindrical, almost parallel-sided | Laesthiola Bouĉek, 1993 [USA] |
– | Funicular segments different; most funicular segments shorter than wide or as long as wide | 17 |
17 | Face deeply impressed, with long setae lateral to the impression of face; intertorular prominence nail-like and positioned distinctly above toruli (Fig. |
Tenuicornus gen. n. [fossil, Miocene Dominican amber] |
– | Face convex, without long setae lateral to the impression of face; intertorular prominence variable in shape but never nail-like and positioned at level of or below toruli | Neocalosoter Girault & Dodd, 1915 [Southern Hemisphere, Neotropics, Nearctic, Philippines] |
Tenuicornus dominicus gen. et sp. n., showing general habitus and diagnostic characters. A General habitus in frontolateral view, showing rather long exerted part of the ovipositor B Head and mesosoma in dorsal view, mesosoma with polished mesonotum, foveolate notauli, and absent frenal line C Head in frontolateral view, with deep facial impression and enlarged setae at each side of lateral margin of the impression D Head and antennae in frontolateral view, at slightly different angle than C, with nail-shaped inner antennal process (iap) visible and highlighted with a red line; antennae inserted at ventral margin of eyes, with long scape, and 6-segmented, clavate funicle E Fore wing without a tuft of erect setae at parastigma (pst) and without pilosity or setal bases on wing disc. Scale: 500 µm (A); 200 µm (B–D).
Pteropilosa lailarabanorum sp. n.
Female holotype, preserved in Eocene Baltic amber (56-34 Ma). Holotype deposited in the amber collection of the State Museum of Natural History, SMNS collection number BB-2815.
Funicle of female 6-segmented with funicular segments 3-6 distinctly transverse, i.e., shorter than wide (Fig.
The generic name Pteropilosa is composed of two parts. The first being Ptero-, which is derived from the old-Greek “pteryx”, meaning “wing”; the last letters -pilosa are derived from the Latin “capillosus”, meaning “hairy”. It can be roughly translated as “hairy wings”, referring to the most striking unique feature of this new genus. The generic name is feminine in gender.
See genus.
Female: total body length 1.62 mm; length of head 0.32 mm, of mesosoma 0.64 mm, of metasoma 0.86 mm. Body without metallic luster. Head: height 0.35 mm. Shape of head round (globose) and head smooth, without sculpture; no depression on face; mandibles small (not measurable). Eyes rather big and slightly egg-shaped, 0.22 mm high and 0.19 mm wide. Intertorular prominence tooth-shaped and positioned between toruli, but slightly below level of toruli. Antennae: inserted well above ventral margin of eyes (in upper third of eyes); scape length 96 µm, pedicel length 48 µm, pedicel rather short and stout; funicle 6-segmented. First funicular segment wider than long, F2 longer than wide. F3-F6 wider than long, distally broadened club-like (F1: length: 33 µm × width: 37 µm; F2: 40 µm × 38 µm; F3: 47 µm × 56 µm; F4: 48 µm × 69 µm; F5: 48 µm × 71 µm; F6: 54 µm × 77 µm); all funicle segments with sensilla. Clava egg-shaped, 0.14 mm long and 81 µm wide. Wings: fore wing disc pilose, without fuscous band. Elongate admarginal setae absent. Fore wing length 1.19 mm and width 0.33 mm; submarginal vein 0.42 mm long, marginal vein 0.33 mm and postmarginal vein 76 µm long, stigmal vein curved, 76 µm long; stigma slightly thickened and uncus visible. Hind wing length 0.98 mm and width 0.23 mm, only two hamuli visible. Legs: length of hind tibia 0.24 mm, hind femur 0.35 mm, hind trochanter 46 µm, hind coxa 0.15 mm. The other legs cannot be measured due to the position of the specimen within the amber. Mesosoma: pronotum, mesoscutum and mesoscutellum completely reticulated. Frenal line clearly visible and foveolate; frenum 0.55 times as long as mesoscutellum. Length of pronotum 0.25 mm, of mesoscutum 0.13 mm, of mesoscutellum 50 µm, of frenum 29 µm. Axillae large, medially connected. Metanotum with distinct foveae; lateral panel of metanotum wide and smooth. Metascutellum thin; not reaching anterior margin of metanotum. Propodeum coarsely reticulated. Metasoma: petiole (Mt1) small, transverse, hardly recognizable. Metasomal tergites Mt2 to Mt9 smooth, Mt2: 0.23 mm, Mt3: 42 µm, Mt4: 0.15 mm, Mt5: 0.13 mm, Mt6: 67 µm, Mt7 0.10 mm, Mt8/9: 70 µm. Ovipositor 0.12 mm (top view) exerted beyond end of gaster.
Following the latest available identification key of
Named after Laila and Raban Ohlhoff, the grandchildren of the private donor.
Tenuicornus dominicus sp. n.
Female holotype, preserved in Lower Miocene Dominican amber (20-15 Ma). Holotype deposited in the amber collection of the Senckenberg Forschungsinstitut und Naturmuseum Frankfurt am Main, Germany, collection number SMF Be 2395.
Funicle of female 6-segmented and clavate with F1–F2 longer than wide and F3–F6 wider than long and distally broadening; scape elongate (1/3 of total antennal length) (Fig.
The first letters of the generic name Tenui- are derived from the Latin word “tenuis”, meaning “thin” or “sharp”. The last letters -cornus of the generic name are derived from the Latin word “cornus”, meaning “horn”. The generic name is male in gender and refers to the thin inner antennal process.
See genus.
Female: total body length 2.07 mm; length of mesosoma 0.92 mm, of metasoma 1.02 mm. Body without metallic luster. Head: height 0.39 mm, width 0.38 mm. Face deeply impressed. Shape of head cuboid; mandibles not visible, hidden in facial depression. Margin of facial depression with rough surface and with a thickened seta on each side (length of setae 85 µm). Inner antennal process (iap) positioned distinctly above level of toruli; nail-like, 0.11 mm long, with basis wider than apex, orientated anteroventrally. Eyes large and egg-shaped, 0.25 mm high and 0.2 mm wide. Antennae: inserted at ventral margin of eyes; scape long and slightly curved, length 0.25 mm; pedicel length 75 µm, rather short and stout; funicle 6-segmented with first two funicular segments longer than wide (F1: length: 71 µm × width: 52 µm; F2: 68 µm × 60 µm) and F3-F6 wider than long, distinctly broadening distally, club-like (F3: 59 µm × 72 µm; F4: 60 µm × 74 µm; F5: 58 µm × 80 µm; F6: 52 µm × 86 µm). Clava egg-shaped: 97 µm long and 77 µm wide. Wings: fore wing long and slender (length 1.53 mm and width 0.55 mm); submarginal vein 0.57 mm long, marginal vein 0.38 mm long, postmarginal vein 0.11 mm long, stigmal vein 99 µm long; stigma slightly thickened and uncus visible. Wing disc bare, setal bases absent. Elongate admarginal setae present. Hind wing length 1.27 mm and width 0.27 mm, three hamuli present. Mesosoma: pronotum and anterior part of mesoscutum with slight traces of strigulate surface sculpture; mesonotum otherwise polished. Notauli foveolate. Length of pronotum 0.37 mm, length of mesoscutum 0.22 mm, length of mesoscutellum 0.22 mm; no frenal line. Axillae medially connected. Mesopleuron (in lateral view) height 0.29 mm, width 0.36 mm. Prepectus enlarged and triangular in shape (0.14 mm × 0.17 mm). Length of metanotum 39 µm; length of propodeum 72 µm. Propodeum without a median carina or plicae. Legs: coxae strong and stout. Femora and tibiae long and slender (Fe1: 0.45 mm; Fe2: 0.32 mm; Fe3: 0.41 mm; Ti1: 0.39 mm; Ti2: 0.49 mm; Ti3: 0.42 mm). Metasoma: petiole (Mt1) short and transverse, hardly visible. Metasomal tergites Mt2 to Mt9 smooth, Mt2: 0.28 mm, Mt3: 0.17 mm, Mt4: 0.15 mm, Mt5: 0.16 mm, Mt6: 0.11 mm, Mt7 60 µm, Mt8/9: 71 µm. Ovipositor 0.23 mm (top view) exerted beyond end of gaster.
The two genera that are most similar to Tenuicornus are Neocalosoter and Cerocephala. Tenuicornus runs to Neocalosoter in the key of
The name dominicus is derived from the amber deposit in which the fossil was found.
Pteropilosa lailarabanorum features a variety of characters that are unique within Cerocephalinae. The most striking of these characters is the wing pilosity (Fig.
Since both fossils were found without syninclusions (in contrast to Dominocephala), it is impossible to speculate about the host species of the newly described genera. However, the short and stout ovipositors and the overall similarity of body size and appearance with extant taxa of Cerocephalinae suggest that both genera are, like all known extant representatives, parasitoids of endophytic beetles.
Although all three known fossils are expected to be parasitoids of endophytic beetles, there are already hints of differential host specialization. The ovipositor of Tenuicornus is further exserted beyond the end of the gaster than in the other two fossils, which might serve as an indicator that this genus has a different host biology than D. vibrissae and P. lailarabanorum.
All three known cerocephaline fossils are females. This is most likely connected to the association with wood-boring beetles, which represent the vast majority of known cerocephaline hosts. Females visit trees for oviposition and have a much higher chance of being trapped in resin than males. Members of the family Ptinidae, a common host group of Cerocephalinae (
Tenuicornus has a very conspicuous single thickened and enlarged seta on each side of the facial impression (Fig.
The description of Pteropilosa pushes the minimum age of Cerocephalinae from the Miocene to the Eocene and hints to an earlier host shift to endophytic beetle parasitoids than previously thought (
We are grateful to Rainer Ohlhoff (Eggenstein-Leopoldshafen) for the donation of the female holotype of Pteropilosa lailarabanorum, and to Mónica M. Solórzano Kraemer (Senckenberg Forschungsinstitut und Naturmuseum) for the loan of material. Karin Wolf-Schwenninger (SMNS) kindly prepared and grinded the amber specimens and Günter Bechly granted access to the amber collection at SMNS. We thank John Heraty and an anonymous reviewer for their critical comments that helped improving the manuscript. Also, we thank Heather Humphrey (Universität Bonn) for linguistic help.