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. 2014 May 2;9(5):e94056.
doi: 10.1371/journal.pone.0094056. eCollection 2014.

Folding wings like a cockroach: a review of transverse wing folding ensign wasps (Hymenoptera: Evaniidae: Afrevania and Trissevania)

István Mikó  1 , Robert S Copeland  2 , James P Balhoff  3 , Matthew J Yoder  4 , Andrew R Deans  1
Affiliations

Folding wings like a cockroach: a review of transverse wing folding ensign wasps (Hymenoptera: Evaniidae: Afrevania and Trissevania)

István Mikó et al. PLoS One. .

Abstract

We revise two relatively rare ensign wasp genera, whose species are restricted to Sub-Saharan Africa: Afrevania and Trissevania. Afrevania longipetiolata sp. nov., Trissevania heatherae sp. nov., T. hugoi sp. nov., T. mrimaensis sp. nov. and T. slideri sp. nov. are described, males and females of T. anemotis and Afrevania leroyi are redescribed, and an identification key for Trissevaniini is provided. We argue that Trissevania mrimaensis sp. nov. and T. heatherae sp. nov. populations are vulnerable, given their limited distributions and threats from mining activities in Kenya. We hypothesize that these taxa together comprise a monophyletic lineage, Trissevaniini, tr. nov., the members of which share the ability to fold their fore wings along two intersecting fold lines. Although wing folding of this type has been described for the hind wing of some insects four-plane wing folding of the fore wing has never been documented. The wing folding mechanism and the pattern of wing folds of Trissevaniini is shared only with some cockroach species (Blattodea). It is an interesting coincidence that all evaniids are predators of cockroach eggs. The major wing fold lines of Trissevaniini likely are not homologous to any known longitudinal anatomical structures on the wings of other Evaniidae. Members of the new tribe share the presence of a coupling mechanism between the fore wing and the mesosoma that is composed of a setal patch on the mesosoma and the retinaculum of the fore wing. While the setal patch is an evolutionary novelty, the retinaculum, which originally evolved to facilitate fore and hind wing coupling in Hymenoptera, exemplifies morphological exaptation. We also refine and clarify the Semantic Phenotype approach used in previous taxonomic revisions and explore the consequences of merging new with existing data. The way that semantic statements are formulated can evolve in parallel, alongside improvements to the ontologies themselves.

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Conflict of interest statement

Competing Interests: The authors have declared that no competing interests exist.

Figures

Figure 1
Figure 1. CLSM micrographs of the right wing base of Evaniidae (anterior to the top).
A: Evania albofascialis. B: Afrevania longipetiolata sp. nov.
Figure 2
Figure 2. Fore wing of Afrevania longipetiolata sp. nov.
A: CLSM micrograph of the fully unfolded fore wing, anterior to the left. B: Brightfield image of the fully unfolded fore wing, anterior to the left. C: Brightfield image of the folded fore wing, anterior to the top.
Figure 3
Figure 3. SEM micrographs of Afrevania longipetiolata sp. nov.
A: Anteroventral region of the face, anterior view, dorsal to the top. B: mandible, posterior view, distal to the left. C: Mesosoma, lateral view, anterior to the left.
Figure 4
Figure 4. Brigthfield images of Afrevania longipetiolata, anterior to the left.
A: Head and mesosoma, lateral view. B: Dorsal region of the metapectal-propodeal complex, lateral view. C: Habitus, posterodorsal view. D: Posterolateral region of the metapectal-propodeal complex, posterodorsal view.
Figure 5
Figure 5. Brightfield images of the fore wing of Evaniidae, anterior to the left.
A: Evania albofascialis. B: Trissevania anemotis. C: Rothevania valdiviana. D: Brachygaster minutus.
Figure 6
Figure 6. Brightfield images of the head of Trissevaniini, anterior view, dorsal to the top.
A: Trissevania anemotis Kieffer, 1913. B: Trissevania slideri sp. nov. C: Afrevania longipetiolata sp. nov. D: Afrevania leroyi Benoit, 1953.
Figure 7
Figure 7. Brightfield images of the head and the mesosoma of Trissevania species, dorsal view, anterior to the left.
A: Trissevania anemotis Kieffer 1913. B: Trissevania slideri sp. nov.
Figure 8
Figure 8. Brightfield images of Trissevania mrimaensis sp. nov., lateral view, anterior to the left.
A: Head and mesosoma. B: dorsolateral region of the mesothorax.
Figure 9
Figure 9. Brightfield images showing the proximal part of the male antenna of Trissevania species, distal to the left.
A: Trissevania slideri sp. nov. B: Trissevania hugoi sp. nov.
Figure 10
Figure 10. Brightfield images of Afrevania longipetiolata, anterior to the left.
A: Dorsolateral view. B: Dorsal view.
Figure 11
Figure 11. Brightfield images of Trissevania mrimaensis sp. nov., anterior to the left.
A: Head and mesosoma, dorsal view. B: Habitus, dorsolateral view.
Figure 12
Figure 12. Brightfield images of the head and the mesosoma of Afrevania species, dorsal view, anterior to the left.
A: Afrevania longipetiolata sp. nov. B: Afrevania leroyi Benoit 1953.
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Grants and funding

This research was funded by the U. S. National Science Foundation (grants DBI-0850223, DEB-0842289) and by the Phenotype Research Coordination Network (NSF DEB-0956049). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.