Multi-gene phylogeny and divergence estimations for Evaniidae (Hymenoptera) – our latest open access publication


Sharanowski BJ, Peixoto L, Dal Molin A, Deans AR. 2019. Multi-gene phylogeny and divergence estimations for Evaniidae (Hymenoptera) PeerJ 7:e6689 DOI: 10.7717/peerj.6689

Abstract.—Ensign wasps (Hymenoptera: Evaniidae) develop as predators of cockroach eggs (Blattodea), have a wide distribution and exhibit numerous interesting biological phenomena. The taxonomy of this lineage has been the subject of several recent, intensive efforts, but the lineage lacked a robust phylogeny. In this paper we present a new phylogeny, based on increased taxonomic sampling and data from six molecular markers (mitochondrial 16S and COI, and nuclear markers 28S, RPS23, CAD, and AM2), the latter used for the first time in phylogenetic reconstruction. Our intent is to provide a robust phylogeny that will stabilize and facilitate revision of the higher-level classification. We also show the continued utility of molecular motifs, especially the presence of an intron in the RPS23 fragments of certain taxa, to diagnose evaniid clades and assist with taxonomic classification. Furthermore, we estimate divergence times among evaniid lineages for the first time, using multiple fossil calibrations. Evaniidae radiated primarily in the Early Cretaceous (134.1–141.1 Mya), with and most extant genera diverging near the K-T boundary. The estimated phylogeny reveals a more robust topology than previous efforts, with the recovery of more monophyletic taxa and better higher-level resolution. The results facilitate a change in ensign wasp taxonomy, with Parevania, and Papatuka, syn. nov. becoming junior synonyms of Zeuxevania, and Acanthinevania, syn. nov. being designated as junior synonym of Szepligetella. We transfer 30 species to Zeuxevania, either reestablishing past combinations or as new combinations. We also transfer 20 species from Acanthinevania to Szepligetella as new combinations.

A photographic catalog of Ceraphronoidea types at the Muséum national d’Histoire naturelle, Paris – our latest open access publication

Screen Shot 2019-02-28 at 1.41.14 PM

Trietsch C, Mikó I, Deans AR (2019). A photographic catalog of Ceraphronoidea types at the Muséum national d’Histoire naturelle, Paris (MNHN), with comments on unpublished notes from Paul Dessart. European Journal of Taxonomy 0(502). DOI: 0.5852/ejt.2019.502

Abstract.—The majority of Ceraphronoidea (Insecta: Hymenoptera) species were described in the late 1800s and early 1900s, with most of these early descriptions relying on text alone. Few type specimens have been illustrated and even fewer have been photographed, posing a challenge to taxonomists working on the group today. Here, we attempt to remove the barriers obstructing Ceraphronoidea research by creating a photographic catalog of the type specimens present at the Muséum national d’Histoire naturelle (MNHN) in Paris, France. We discuss the history of the ceraphronoid specimens present in the collection and provide comments on unpublished species notes from former Ceraphronoidea taxonomist Paul Dessart. We synonymize Ceraphron myrmecophilus Kieffer, 1913 syn. nov. with Aphanogmus abdominalis (Thomson, 1858) (Hymenoptera: Ceraphronidae) based on the male genitalia morphology, body shape and presence of foveae on the median length of the mesoscutellum. We also report the discovery of the missing male holotype of Ceraphron testaceus (Risbec, 1953) (Hymenoptera: Ceraphronidae) and several potential types of Aphangomus aphidi (Risbec, 1955) (Hymenoptera: Ceraphronidae).

Revision of Trassedia (Hymenoptera: Ceraphronidae), an evolutionary relict with an unusual distribution – our latest publication

Screen Shot 2018-12-11 at 11.24.33 AM

Mikó I, Trietsch C, van de Kamp T, Masner L, Ulmer JM, Yoder MJ, Zuber M, Sandall EL, Baumbach T, Deans AR (2018) Revision of Trassedia (Hymenoptera: Ceraphronidae), an evolutionary relict with an unusual distribution. Insect Systematics and Diversity 2(6): 4 DOI: 10.1093/isd/ixy015

Abstract.—Ceraphronoidea is composed of two, seemingly well-defined families, Ceraphronidae and Megaspilidae. The position of Trassedia Cancemi 1996 within the superfamily is unclear, as this genus shares characteristics of both families. For instance, Trassedia possess both the pterostigma form characteristic of Megaspilidae, and the Waterston’s evaporatorium, a structure unique to Ceraphronidae. Trassedia was known only from a single specimen of T. luapi Cancemi 1996 from Madagascar. We describe nine new species: Trassedia australiensis Mikó and Masner sp. nov. (Australia), Trassedia yanegai Mikó and Trietsch sp. nov. (Thailand), Trassedia brasiliensis Masner and Mikó sp. nov. (Brazil), Trassedia nigra Masner and Mikó sp. nov. (Brazil), Trassedia nigrorufus Mikó and Masner sp. nov. (Panama), Trassedia guianensis Mikó and Masner sp. nov. (French Guiana), Trassedia angustioculus Mikó and Masner sp. nov. (French Guiana), and Trassedia pilosus Masner and Mikó sp. nov. (Costa Rica), and Trassedia gauldi Mikó and Masner sp. nov. (Costa Rica and Brazil). To illuminate the morphological concepts presented here, we provide SR-µCT and confocal laser scanning microscopy (CLSM)-based 3D reconstructions. The Waterston’s organ is sexually dimorphic in Trassedia; it is unpaired in males and paired in females. We describe modifications of the metasoma apex that align with the enlarged hind tarsi, a leg phenotype peculiar to Trassedia and the unique subdivision of the first valvifer. We report the presence of the occipital depression in Trassedia and describe how this structure is involved in a secondary articulation between the head and the mesosoma. We discuss the possible function and phylogenetic relevance of the pterostigma in Ceraphronoidea. Based on its Southern Hemisphere distribution we hypothesize that Trassedia’s presence predates the break-up of Gondwana.

The Insect Collectors’ Code – our latest publication

A vehicle outfitted for fieldwork, primarily to collect Odonata, from a 1959 sampling trip by George H. and Alice F. Beatty. Photo probably by the Beattys

Check out our latest publication, in American Entomologist:

Trietsch C, Deans AR (2018) The Insect Collectors’ Code. American Entomologist 64(3): 156–158 DOI: 10.1093/ae/tmy035

Can’t access the article? See a preprint here: (123KB PDF)

A new megaspilid wasp from Eocene Baltic amber – our latest open access publication


Mikó I, van de Kamp T, Trietsch C, Ulmer JM, Zuber M, Baumbach T, Deans AR. (2018A new megaspilid wasp from Eocene Baltic amber (Hymenoptera: Ceraphronoidea), with notes on two non-ceraphronoid families: Radiophronidae and StigmaphronidaePeerJ 6:e5174

Abstract.—Ceraphronoids are some of the most commonly collected hymenopterans, yet they remain rare in the fossil record. Conostigmus talamasi Mikó and Trietsch, sp. nov. from Baltic amber represents an intermediate form between the type genus, Megaspilus, and one of the most species-rich megaspilid genera, Conostigmus. We describe the new species using 3D data collected with synchrotron-based micro-CT equipment. This non-invasive technique allows for quick data collection in unusually high resolution, revealing morphological traits that are otherwise obscured by the amber. In describing this new species, we revise the diagnostic characters for Ceraphronoidea and discuss possible reasons why minute wasps with a pterostigma are often misidentified as ceraphronoids. Based on the lack of ceraphronoid characteristics, we remove Dendrocerus dubitatus Brues, 1937, Stigmaphronidae, and Radiophronidae from Ceraphronoidea and consider them as incertae sedis. We also provide some guidance for their future classification.

Ceraphron krogmanni (Hymenoptera: Ceraphronidae), a new species from Lower Saxony – our latest open access publication

the male genitalia of this wasp species are large and unusual

Ulmer JM, Miko I, Deans AR (2018) Ceraphron krogmanni (Hymenoptera: Ceraphronidae), a new species from Lower Saxony with unusual male genitalia. Biodiversity Data Journal 6: e24173. DOI: 10.3897/BDJ.6.e24173


Male genitalia phenotypes of Ceraphron (Jurine, 1807) are informative for species delimitation, but due to their minute size, these characters have not been used extensively. Recent developments in visualisation techniques, e.g. confocal laser scanning microscopy and high resolution bright field imaging, allow for more thorough examination of these minute anatomical structures and the development of a robust, male genitalia-based taxonomic system. We also establish a character set, a template, that will facilitate future revisions of these wasps.

New information

Ceraphron krogmanni sp. nov. is described with outsized male genitalia and multiple diagnostic traits that are unique amongst Ceraphron species.

Unique extrication structure in a new megaspilid – our latest open access publication

wasp glued to a point made from cardstock

Trietsch C, Mikó I, Notton D, Deans A (2018) Unique extrication structure in a new megaspilid, Dendrocerus scutellaris Trietsch & Mikó (Hymenoptera: Megaspilidae). Biodiversity Data Journal 6: e22676.

Abstract.—A new species, Dendrocerus scutellaris Trietsch & Mikó (HymenopteraMegaspilidae), is described here from male and female specimens captured in Costa Rica. This species is the only known ceraphronoid wasp with a straight mandibular surface and raised dorsal projections on the scutellum, called the mesoscutellar comb. It is hypothesised that the function of the mesoscutellar comb is to aid the emergence of the adult from the host, especially since the mandibles lack a pointed surface to tear open the pupal case. The authors also provide phenotypic data in a semantic form to facilitate data integration and accessibility across taxa and provide an updated phenotype bank of morphological characters for megaspilid taxonomic treatments. In updating this phenotype bank, the authors continue to make taxonomic data accessible to future systematic efforts focusing on Ceraphronoidea.

Translucent cuticle and setiferous patches in Megaspilidae – our latest open access publication


Trietsch C, Mikó I, Ulmer JM, Deans AR (2017) Translucent cuticle and setiferous patches in Megaspilidae (Hymenoptera, Ceraphronoidea). Journal of Hymenoptera Research 60: 135-156. DOI: 10.3897/jhr.60.13692

Abstract.—All Ceraphronoidea have metasomal patches of translucent cuticle and setae that have never been investigated before, despite their potential behavioral and phylogenetic relevance. To understand the internal and external morphology of these structures, specimens were examined using a broad array of histology-based methods, including transmission electron microscopy (TEM), scanning electron microscopy (SEM), confocal laser scanning microscopy (CLSM) and serial block-face scanning electron microscopy (SBFSEM). For the first time, the setiferous patches are shown to be associated with exocrine glands in Ceraphronoidea. The proposed glandular function is the secretion of pheromones, with the setae above the pore openings serving as a surface for evaporation. The translucent cuticle is morphologically distinct from the setiferous patches; structures resembling lamellar bodies were found underneath the translucent cuticle, and may be associated with photoreceptors or endocrine glands. The locations of translucent cuticle on the metasoma are unique to different families and genera within Ceraphronoidea, and could be useful for inferring phylogenetic relationships. The character distribution suggests that the genera Trassedia and Masner are more closely related to Ceraphronidae than Megaspilidae. We found similar structures containing translucent cuticle in Orussidae and Ichneumonoidea, indicating that these structures are potentially a rich character system for future phylogenetic analysis in Hymenoptera.

Malpighian tubules in Trichoplusia ni – our latest open access publication

CLSM volume rendered micrograph showing the Malpighian tubules and the urinary bladder of Trichoplusia ni. Image by Loren Rivera-Vega (source)

Rivera-Vega L, Mikó I (2017) Know your insect: Malpighian tubules in Trichoplusia ni (Lepidoptera: Noctuidae). Research Ideas and Outcomes 3: e11827. DOI: 10.3897/rio.3.e11827

Abstract.—Malpighian tubules are mainly known to be involved in excretion. However, recent studies have begun to look into other potential roles including detoxification, immunity, host establishment, etc. In this case study, we observed the Malpighian tubules of the cabbage looper (Trichoplusia ni) using confocal laser scanning microscopy. We also discuss other functions that Malpighian tubules are known for (i.e. silk-like and gall-inducing secretions) as well as the similarities between Malpighian tubules and salivary glands in endoparasitic Hymenoptera.

Reflections on teaching insect biodiversity

Last semester I taught my Insect Biodiversity and Evolution course in a slightly new way, and now that it’s over I have a chance to revisit the experience and read my student evaluations. Overall I have to say that I am quite happy! I do see lots of room for improvements, of course, but first a little context …

Three ongoing situations drove me to revamp the course: (1) I get a lot of requests from colleagues to share my teaching materials, and I often found myself uncomfortable with their state (not always clear, sometimes with images of questionable provenance); (2) so many TAs have worked on the materials (often improving, sometimes meddling) that they lost some cohesion; (3) the collection exercise was never quite right, requiring so many specimens/taxa that the resulting product was often not usable for research or teaching (poor preps, sloppy labels, … rushed work).

My goal was to rebuild the course, almost from the ground up, and avail the new materials in such a way that they could be iteratively improved, commented on, and used by anyone. See the results at our GitHub repo. I hoped to release the new materials under a Creative Commons Attribution license (CC BY), to maximize peoples’ ability to refine the materials. I think it’ll be a long time before that can really happen, as many of the images I used are licensed in a way that doesn’t allow commercial use or derivatives.

grad students sitting on stone steps inside a dorm
The ENT 432 crew, at Raven’s Roost at Powdermill Nature Reserve — lots of smiles! September 2016. Photo (CC BY 2.0) by Hillary Morin

What went wrong 🙁

Overall I was happy with the course, but several elements could be improved. Here are three that come to mind, but you can read more in our issues feed:

  1. I’ll probably get rid of the requirement that lab notebooks be graded. Honestly I forgot about that line in the grading rubric (oops!), and so everyone got 100/100. They probably deserved that grade, though. These students answered all of our lab questions, many of which didn’t have “right” answers and were not easy. For example, we asked students to hypothesize the function(s) of the elaborate surface sculpturing one can see in Tingidae (see photo below and question 9-13 in the handout). I don’t know the answer if there is one!
  2. Students need more guidance regarding how to take field notes – or at least what I expected from them for this aspect of the Discover Your Inner Darwin exercise – and iterative examination of their notes. Their field notebooks were quite inconsistent in their detail.
  3. I need to lecture (even) less and bring back required readings that are discussed as a group. I jettisoned this element in order to bring the work load more in line with what Penn State recommends for a 4-credit course (about 160–180 hours of work in a semester). Time to rework the load again. I missed the readings!
top of a lace bug, whose surface is elaborately sculptured like lattice work
Amazing photo of a lace bug (Hemiptera: Tingidae), by Gilles San Martin (CC BY-SA 2.0) Why is their cuticle so elaborately sculptured? I don’t know! And my students didn’t seem comfortable with that.

What went right 😀

I definitely feel like this course is morphing into one that is both effective and fun. Although imperfect, it was easily my best semester as an instructor. Highlights for me:

  1. The observation component of the natural history exercise was really fun for me to witness and read about, and most students found it incredibly rewarding. With more direction from me, and maybe multiple iterations per semester, it could emerge as a highlight for students and an avenue for future research.
  2. The collection is also heading in the right direction. Each one was relatively small but sufficiently diverse, and the specimen preps were almost immaculate. Clearly a lot of time and care was put into these collections!
  3. The blog post exercise was also good fun, and it was an opportunity for students to dig deeper into observations and subjects that inspired them.
  4. The collections resulted in real data that can be used for research! Each student submitted his/her data as Darwin Core Archives, which are basically ready to share through GBIF (I want to doublecheck them first!) With a little help from GBIF, I think we can make this element almost as compelling as the collection.

Changes and opportunities

I’ve discussed one possible change with three semesters of students now, and I feel confident now that it’s an idea worth pursuing: I’d love to partner with a likeminded professor at a university relatively close to ours, say within a 6-hour drive of Penn State, for a combined field trip. We mix our students into teams that collect, prep, cook, and learn together … It could be fun! Another possible change to my course could disrupt the potential for any partnerships – a move to the spring semester.

More on that later!

Originally posted on the Frost Curators’ blog: