Gall midges (Cecidomyiidae) in Sweden: towards completing the taxonomic inventory of a megadiverse family of Diptera

Mathias and Catrin Jaschhof (July 2021)

Now we are close to completing our project, which will definitely end in July 2021. The good news is that we have finally gained a fairly accurate idea of the number of gall midges living in Sweden. There is every indication that the Swedish fauna comprises 2000 different cecidomyiids, which renders this the most species-rich family of Diptera, and one of the most speciose insect families, in Sweden. About one half of these species are mycophages (our taxonomic speciality), of which 639 are scientifically named. Alone in the short period of 2018-2021 we described and named more than 90 species from Sweden that were previously unknown to science (see below the list of our recent publications). In spite of all our efforts made in the past 15 years we did not achieve the envisaged goal, to nearly complete the taxonomic inventory of non-herbivorous cecidomyiids in Sweden. This is mainly because of two reasons: our target group turned out to be much richer in species than anticipated, and the taxonomic issues pertaining to the involved species are more complex than initially thought. As a consequence, we have to leave 178 species found in Sweden unnamed for the time being. Also, for time reasons we have to stop screening fresh material for new discoveries that remained unrecognized so far. This is how taxonomic work unfolds in a megadiverse group of organisms! The final publication of our project, which summarizes our 15 years of gall midge survey funded by the Swedish Taxonomy Initiative, can be found in Entomologisk Tidskrift 142 (3): 105-184.

The Gall Midge Project continues working till July (and then takes a break)

Mathias and Catrin Jaschhof (February 2021)

Period: 2018–July 2021
Funding: SLU Artdatabanken Sweden (3,100,000 SEK)
Project leader: Dr Mathias Jaschhof
Other personnel: Catrin Jaschhof 

For all of us time is running, and we of The Gall Midge Project are wondering how fast the past three years have passed… It´s time again for an update! Most importantly, below is an update of the list of publications resulting from our project. Apart from all the details provided in our papers, the truth is that, once more, we have to acknowledge ourselves beaten by the taxonomic diversity of “our” midges. While our project is close to be finished, we have clear evidence that there is much left to be discovered in, and described from Sweden, meaning our job is not yet completed. When, in 2004, we began studying the Swedish fauna, there were 120 species of mycophagous gall midges known to occur in Sweden; now we know of more than 800 species, both named and unnamed. This is more species than are currently named in all of Europe, and half as many than are named from all over the Earth. And we are continuing discovering species in addition, wherever in Sweden we take samples of “our” midges. Most of the species we discover now, in the final phase of our project, are new to science, meaning they require scientific description according to the rules of zoological nomenclature; this is a laborious task. At the moment we know of 160 undescribed species in Sweden; to formally describe them is work for several more years… Nobody had expected mycophagous gall midges to be so rich in species. The last paper referred to in the list below,  currently in the stage of a manuscript in progress, is essentially a summary of our research in Sweden since 2004, with an outlook into the future. We think it is just a matter of time that we will be back with a project on Sweden’s gall midges… From August 2021 on, we will for two years work for “GBOL III – Dark Taxa”, a project initiated to gain a more accurate idea of the size of the German fauna of gall midges. Our expectation is that the German is more speciose than the Swedish fauna, meaning there is a lot of work ahead of us. In parallel to GBOL III, we will keep an active interest in Sweden´s fauna, mainly by continuing taking samples of specimens to study at some point in the future. We will particularly focus on Öland, where sampling is easiest for us, and the fauna is known to be quite different from that on the Swedish mainland. Needless to say, we will continue working from our lab in Station Linné ? we cannot imagine a better place for our research.

List of publications 2018-2021

Jaschhof, M. & Jaschhof, C. 2021. Mycophagous gall midges (Diptera: Cecidomyiidae excl. Cecidomyiinae) in Sweden: status report after 15 years of taxonomic inventory, taxonomic checklist and description of Camptomyia alstromi sp. nov. Entomologisk Tidskrift 142: 105–184.

Gagné, R.J. &. Jaschhof, M. 2021. A Catalog of the Cecidomyiidae (Diptera) of the World. Fifth Edition. Digital version 4. Jaschhof, M. 2021. A review of gall midges (Diptera: Cecidomyiidae) described from Mesozoic deposits. Palaeoentomology 4 (1): 1–14. 

Jaschhof, M. & Jaschhof, C. 2020. High time for omtanke: New mycophagous gall midges (Diptera: Cecidomyiidae) from a floodplain forest in central Sweden threatened by continuous clear-felling. Entomologisk Tidskrift 141 (4): 191–206.

Jaschhof, M. & Jaschhof, C. 2020. Wheeleriola perplexa gen. et sp. nov., the first member of Catotrichinae (Diptera: Cecidomyiidae) found in New Zealand. Zootaxa 4852 (5): 565–570.

Sikora, T., Jaschhof, M. & Kurina, O. 2020. Additions to the Estonian fauna of mycophagous Cecidomyiidae (Diptera), with a description of Unicornella estonensis gen. et sp. nov. Zootaxa 4851 (2): 349–363.

Jaschhof, M. & Jaschhof, C. 2020. Reevaluation of species richness in Winnertzia (Diptera, Cecidomyiidae, Winnertziinae), with descriptions of 37 new species from Sweden, Peru and Australia. Zootaxa 4829 (1): 1–72.

Ham, D., Jaschhof, M. & Bae, Y.J. 2020. Mycophagous Gall Midges (Diptera, Cecidomyiidae) in Korea: Newly Recorded Species with Discussion on Four Years of Taxonomic Inventory. Animal Systematics, Evolution and Diversity 36 (1): 60–77.

Jaschhof, M. & Jaschhof, C. 2020. An update of Micromyinae (Diptera: Cecidomyiidae) taxonomy, with descriptions of a new genus and 13 new species from Northern Europe. Zootaxa 4750 (3): 349–369.

Ronquist, F., Forshage, M., Häggqvist, S., Karlsson, D., Hovmöller, R.,, Bergsten, J., Holston, K., Britton, T., Abenius, J., Andersson, B., Buhl, P.N., Coulianos, C.-C., Fjellberg, A., Gertsson, C.-A., Hellqvist, S., Jaschhof, M., Kjærandsen, J., Klopfstein, S., Kobro, S., Liston, A., Meier, R., Pollet, M., Prous, M., Riedel, M., Rohácek, J., Schuppenhauer, M., Stigenberg, J., Struwe, I., Taeger, A., Ulefors, S.-O., Varga, O., Withers, P. & Gärdenfors, U. 2020. Completing Linnaeus’s inventory of the Swedish insect fauna: Only 5,000 species left? PLoS ONE 15 (3): e0228561.

Karlsson, D., Hartop, E, Forshage, M., Jaschhof, M. & Ronquist, F. 2020. The Swedish Malaise Trap Project: A 15 Year Retrospective on a Countrywide Insect Inventory. Biodiversity Data Journal 8: e47255.

Jaschhof, M. & Jaschhof, C. 2020. The Porricondylini (Diptera: Cecidomyiidae: Porricondylinae) of Sweden revisited: descriptions of nineteen new species of ten genera, including a new genus. Zootaxa 4728 (2): 151–182.

Kjaerandsen, J. & Jaschhof, M. 2019. New records and first DNA barcodes of the family Canthyloscelidae (Diptera) in Fennoscandia. Norwegian Journal of Entomology 66 (2): 81–93.

Jaschhof, M. & Jaschhof, C. 2019. New and rarely found species of asynaptine Porricondylinae (Diptera: Cecidomyiidae) in northern Europe. Zootaxa 4604 (2): 281–300.

Jaschhof, M. & Ševcik, J. 2019. Cabamofa orientalis sp. nov. from Thailand, congeneric with C. mira Jaschhof from Costa Rica (Diptera: Bibionomorpha: Sciaroidea incertae sedis). Zootaxa 4576 (2): 392–394.

Jaschhof, M. & Jaschhof, C. 2019. New Dicerurini from Europe, mostly Sweden (Diptera: Cecidomyiidae, Porricondylinae). Zootaxa 4559 (2): 245–264.

Sikora, T., Jaschhof, M., Mantic, M., Kasprák, D. & Ševcík, J. 2019. Considerable congruence, enlightening conflict: molecular analysis largely supports morphology-based hypotheses on Cecidomyiidae (Diptera) phylogeny. Zoological Journal of the Linnean Society 185: 98–110.

Jaschhof, M. 2018. Two million species of gall midges - who is surprised? Abstracts Volume of the 9th International Congress of Dipterology, Windhook, Namibia, November 25–30, 2018: 121.

Jaschhof, M. & Jaschhof, C. 2018. Descriptions of eighteen new species of Glemparon, a previously monotypic genus of Porricondylinae (Diptera, Cecidomyiidae). European Journal of Taxonomy 450: 1–38.

Jaschhof, M. & Spungis, V. 2018. Towards reliable identification of male Dicerura: descriptions of three new and seven poorly known species (Diptera: Cecidomyiidae, Porricondylinae). Zootaxa 4422 (1): 85–103.

Brown, B.V., Borkent, A., Adler, P.H., Amorim, D.S., Barber, K., Bickel, D., Boucher, S., Brooks, S.E., Burger, J., Burington, Z.L., Capellari, R.S., Costa, D.N.R., Cumming, J.M., Curler, G., Dick, C.W., Epler, J.H., Fisher, E., Gaimari, S.D., Gelhaus, J., Grimaldi, D.A., Hash, J., Hauser, M., Hippa, H., Ibáñez-Bernal, S., Jaschhof, M., Kameneva, E.P., Kerr, P.H., Korneyev, V., Korytkowski, C.A., Kung, G.-A., Kvifte, G.M., Lonsdale, O., Marshall, S.A., Mathis, W., Michelsen, V., Naglis, S., Norrbom, A.L., Paiero, S., Pape, T., Pereira-Colavite, A., Pollet, M., Rochefort, S., Rung, A., Runyon, J.B., Savage, J., Silva, V.C., Sinclair, B.J., Skevington, J.H., Stireman III, J.O., Swann, J., Thompson, F.C., Vilkamaa, P., Wheeler, T., Whitworth, T., Wong, M., Wood, D.M., Woodley, N., Yau, T., Zavortink, T.J. & Zumbado, M.A. 2018. Comprehensive inventory of true flies (Diptera) at a tropical site. Communications Biology 1 (21): 1–8.

Borkent, A., Brown, B.V., Adler, P.H., Amorim, D.S., Barber, K., Bickel, D., Boucher, S., Brooks, S.E., Burger, J., Burington, Z.L., Capellari, R.S., Costa, D.N.R., Cumming, J.M., Curler, G., Dick, C.W., Epler, J.H., Fisher, E., Gaimari, S.D., Gelhaus, J., Grimaldi, D.A., Hash, J., Hauser, M., Hippa, H., Ibáñez-Bernal, S., Jaschhof, M., Kameneva, E.P., Kerr, P.H., Korneyev, V., Korytkowski, C.A., Kung, G.-A., Kvifte, G.M., Lonsdale, O., Marshall, S.A., Mathis, W., Michelsen, V., Naglis, S., Norrbom, A.L., Paiero, S., Pape, T., Pereira-Colavite, A., Pollet, M., Rochefort, S., Rung, A., Runyon, J.B., Savage, J., Silva, V.C., Sinclair, B.J., Skevington, J.H., Stireman III, J.O., Swann, J., Thompson, F.C., Vilkamaa, P., Wheeler, T., Whitworth, T., Wong, M., Wood, D.M., Woodley, N., Yau, T., Zavortink, T.J. & Zumbado, M.A. 2018. Remarkable fly (Diptera) diversity in a patch of Costa Rican cloud forest: Why inventory is a vital science. Zootaxa 4402 (1): 53–90. 

Jaschhof, M., Ham, D. & Bae, Y.J. 2018. Loboplusia coreana sp. nov. from South Korea, only the second species of a genus originally described from Costa Rica (Diptera: Cecidomyiidae, Winnertziinae). Zootaxa 4399 (1): 131–133.

Jaschhof, M. & Jaschhof, C. 2018. A needle in a haystack: the new genus Bernadottea (Diptera: Cecidomyiidae, Winnertziinae) with four new species in the Old World. Zootaxa 4379 (4): 529–538.  

Gall Midge Project News: Again, it goes on!

Period: 2018–2020
Funding: SLU Artdatabanken Sweden (3,100,000 SEK)
Project leader: Dr Mathias Jaschhof
Other personnel: Catrin Jaschhof


November 2017 SLU Artdatabanken took the decision to continue funding our project on the taxonomy and diversity of gall midges (family Cecidomyiidae) in Sweden for another three years. Being aware of the fact that there are other groups of poorly known organisms in need of basic research in Sweden, we take that grant as an honour and privilege—and as an incentive for tackling new challenges. A part of the coming three years of research will address, for the first time in our career, the predators among the gall midges. This is another species-rich subgroup of Cecidomyiidae whose diversity, in Sweden and the world, is known only in fragments. Our project, designed to focus on Lestodiplosis and related genera, will contribute to the taxonomic revision of the world fauna, which is vitally needed (and demanded for a long time) for these tiny midges. It goes without saying that we will also continue doing research into the fungus-feeding gall midges of Sweden, which were recently shown to be more speciose than previously estimated by us—thanks to a new approach of combining morphology with molecular data (see below the outline of our project for the years 2015–2017).

Considering that SLU Artdatabanken has granted substantial amounts of taxpayer’s money in support of our research since 2004, some people might ask for the outcomes from this investment. Scientists’ currency is mostly publications, so below is a list of our publications on Swedish (and other Nordic) Cecidomyiidae produced during funding by SLU Artdatabanken. All these papers together contain descriptions of close to 200 species new to science. And: now it goes on!


The Gall Midge Project... goes on!

Period: 2015–2017
Funding: SLU Artdatabanken Sweden (3,180,000 SEK)
Project leader: Dr Mathias Jaschhof
Other personnel: Catrin Jaschhof 

Why Gall Midges? Cecidomyiidae (gall midges), one of the most biodiverse families of the order Diptera (two-winged insects), contain more than 6,200 described species—mostly plant-feeders, but also fungus-feeders and predators. Cecidomyiids are abundant and ubiquitous in terrestrial ecosystems worldwide. Even so, due to their small size and other difficulties in studying them, they have never attracted much research interest by dipterist-taxonomists and are thus ranked among the least known dipterans. At present only fragments of the world fauna are known and the number of world species is regarded as inestimable. To illustrate these facts, more than 800 species of gall midges, almost all unknown to science, have recently been found within just 4 hectares of tropical cloud forest!

To present knowledge, the Swedish fauna includes 240 plant-feeding species (classified in the subfamily Cecidomyiinae) and 409 fungus-feeding species (Lestremiinae, Micromyinae, Winnertziinae, and Porricondylinae). Only the fungus-feeders are well documented, thanks to two previous projects funded by SLU Artdatabanken. There are indications that the Swedish fauna actually comprises more than twice as many species as currently known, i.e. 500–600 plant-feeders, 500–600 fungus-feeders, and 100 predators. So there is still a long way to go before we may regard the Swedish Cecidomyiidae as adequately inventoried!

The Gall Midge Project, designed to closing this gap of faunistic and taxonomic knowledge, aims at the identification and documentation of 300+ species new to Sweden and 60+ species new to science. Specimens to study come from the rich Cecidomyiidae material generated by the Swedish Malaise Trap Project as well as from fresh collections (to be made in the course of the project) in Öland, Gotland, and several carefully selected sites on the Swedish mainland. The identification and classification of species will be based on both morphological and molecular evidence. Relevant results, such as new taxa, generic revisions, and identification keys, will be published in peer-reviewed journals, and voucher specimens will be deposited in the Naturhistoriska Riksmuseet Stockholm.

Why Station Linné? The Station is the ideal base for studying the biodiversity and taxonomy of Swedish Cecidomyiidae: it ensures closest distance to the Swedish Malaise Trap Project, which is a major source of study material; it provides an adequate environment for rearing gall midges from infested plant material, as the Station maintains a greenhouse facility; and it allows for short distances to the envisaged collecting sites. Moreover, Station Linné is adequately equipped for morpho-taxonomic research, and some of its employees are themselves actively involved in biodiversity research. Altogether, all the Station breathes the animating spirit of discovery.

Who is funding this research? The Gall Midge Project is a subproject of, and funded by The Swedish Taxonomy Initiative (STI), which strives to inventory all the multicellular fauna and flora of Sweden. Of 50,000–60,000 multicellular species estimated to occur in Sweden, up to 1,300 might be gall midges. Being an ambitious and unique endeavour in biodiversity research in general, STI has performed as one of the major supporters of particularly cecidomyiid research for many years now. In fact, STI means far more than a regional enterprise for the study of gall midges, especially fungus-feeders. Nowhere else on the planet are fungus-feeding cecidomyiids so intensively studied as in Sweden—thanks to STI.


List of publications relevant to Nordic Cecidomyiidae (and a few other, related Diptera) produced with funding from The Swedish Taxonomy Initiative

Jaschhof, M. 2017. A taxonomic review of Palaearctic Peromyia Kieffer (Diptera: Cecidomyiidae, Micromyinae), with descriptions of thirty-eight new species mostly from Sweden. Studia dipterologica  23 (1) 2016: 51–116.

Jaschhof, M. & Jaschhof, C. 2017. New species of Aprionus (Diptera, Cecidomyiidae, Micromyinae) from Sweden and other parts of the Palearctic region. European Journal of Taxonomy  378: 1–38.

Gagné, R.J. & Jaschhof, M. 2017. A Catalog of the Cecidomyiidae (Diptera) of the World. Forth edition. Digital version 3.

Jaschhof, M. 2017. Catochini, Strobliellini and Acoenoniini revisited: a taxonomic review of the small tribes of the Micromyinae (Diptera: Cecidomyiidae). Zootaxa 4250 (3): 275–295.

Sikora, T., Jaschhof, M. & Ševcik, J. 2017. Glossostyles perspicua gen. et sp. nov. and other fungivorous Cecidomyiidae (Diptera) new to the Czech and Slovak Republics.  European Journal of Taxonomy 303: 1–29.

Jaschhof, M. & Jaschhof, C. 2017. Mycophagous gall midges (Diptera, Cecidomyiidae: Lestremiinae, Micromyinae, Winnertziinae, Porricondylinae): first records in Sweden and descriptions of closely related new species from elsewhere. Zootaxa 4226 (4): 546–570.

Sevcik, J., Kasprak, D., Mantic, M., Fitzgerald, S., Sevcikova, T., Tothova, A. & Jaschhof, M. 2016. Molecular phylogeny of the megadiverse insect infraorder Bibionomorpha sensu lato (Diptera). PeerJ 4:e2563; DOI 10.7717/peerj.2563.

Wirta, H. K., Várkonyi, G., Rasmussen, C., Kaartinen, R. M., Schmidt, N. M., Hebert, P. D. N., Barták, M., Blagoev, G., Disney, H., Ertl, S., Gjelstrup, P., Gwiazdowicz, D. J., Huldén, L., Ilmonen, J., Jakovlev, J., Jaschhof, M., Kahanpää, J. V., Kankaanpää, T. T., Krogh, P. H., Labbee, C., Lettner, C., Michelsen, V., Nielsen, S. A., Nielsen, T. R., Paasivirta, L., Pedersen, S., Pohjoismäki, J., Salmela, J. A. M., Vilkamaa, P. A., Väre, H. U., von Tschirnhaus, M. & Roslin, T. V. 2016. Establishing a community-wide DNA barcode library as a new tool for arctic research. Molecular Ecology Resources 16 (3): 809–822.

Jaschhof, M. 2016. A review of world Diallactiini (Diptera, Cecidomyiidae, Winnertziinae), with the description of six new genera and seventeen new species. Zootaxa 4127 (2): 201–244.

Jaschhof, M. 2016. New species of Monepidosis Mamaev, 1966 and Antipodosis gen. nov., a closely related genus from New Zealand. European Journal of Taxonomy 192: 1–24.

Jaschhof, M. 2016. Taxonomic revision of some Micromyinae (Diptera: Cecidomyiidae) described by Zoya L. Berest. Zootaxa 4097 (2): 255–262.

Jaschhof, M. 2015. A review of the Aprionus flavidus group, with description of two new species close to Aprionus betulae Jaschhof (Diptera: Cecidomyiidae, Micromyinae).  Studia dipterologica 21 (2) (2014): 221–229.

Jaschhof, M. 2015. Morphological re-examination reveals that Campylomyza serrata Jaschhof, 1998 is a complex of five cryptic species (Diptera: Cecidomyiidae, Micromyinae). Contributions to Entomology 65 (2): 373–381.

Jaschhof, M. & Jaschhof, C. 2015. Linnaeomyia hortensis gen. et spec. nov. (Diptera: Cecidomyiidae, Porricondylinae) from a backyard site in Öland, Sweden. Zootaxa 4018 (2): 292–296.

Jaschhof, M. 2015. [Chapter] 17.12. Cecidomyiidae (Gall midges). – Pp. 572–574 in: Böcher, J., Kristensen, N.P., Pape, T. & Vilhelmsen, L. (eds), The Greenland Entomofauna. An Identification Manual of Insects, Spiders and their Allies. Fauna Entomologica Scandinavica 44, Leiden / Boston: Brill, 881 pp.

Jaschhof, M. & Jaschhof, C. 2015. New species and new distribution records of Lestremiinae, Micromyinae and Porricondylinae (Diptera: Cecidomyiidae) in Sweden. Zootaxa 3973 (1): 159–174.

Jaschhof, M., Skuhravá, M. & Penttinen, J. 2014. Checklist of the Cecidomyiidae (Diptera) of Finland. In: Kahanpää, J. & Salmela, J. (eds): Checklist of the Diptera of Finland. ZooKeys 441: 103–118.

Gagné, R.J. &.Jaschhof, M. 2014. A Catalog of the Cecidomyiidae (Diptera) of the World. Third edition. Digital version 2.


Jaschhof, M. & Jaschhof, C. 2013. The Porricondylinae (Diptera: Cecidomyiidae) of Sweden, with notes on extralimital species. Studia dipterologica Supplement 20: 1–392.

Jaschhof, M. & Jaschhof, C. 2009. The Wood Midges (Diptera: Cecidomyiidae: Lestremiinae) of Fennoscandia and Denmark. Studia dipterologica Supplement 18, 333 pp.

Penttinen, J. & Jaschhof, M. 2009. On the systematics of Sylvenomyia Mamaev & Zaitzev (Diptera, Cecidomyiidae, “Porricondylinae”), with the description of a new species from Finland. Zootaxa 2009: 48–54.

Jaschhof, M., Jaschhof, C., Viklund, B. & Kallweit, U. 2006. On the morphology and systematic position of Sciarosoma borealis Chandler, based on new material from Fennoscandia (Diptera: Sciaroidea). Studia dipterologica 12 (2005) (2): 231–241.

Figure 1: This is one of three Malaise traps that collected specimens for the Gall Midge Project in Grytsjön Nature Reserve near Bäckebo, Småland, in 2014. Note the abundance of dead aspen and birch wood on the forest floor: the perfect habitat for fungus-feeding gall midges.

Figure 2: One of the objects of desire: the gall midge Clinorhytis flavitarsis, described and named by the French reverend Kieffer, in 1896. This species was rediscovered only in the mid-1980s in Latvia and Ukraine, and first recorded in Sweden in 2010. With a body length of 5 mm it is a large member of the gall midge family.

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