My research focuses on the systematics, phylogeny and biogeography of the Chalcidoidea (Hymenoptera). Chalcidoid wasps is one of the largest groups of insects: there are about as 100,000 described species, but many are still undescribed and regional faunas are also poorly known. Some of chalcids are specialist, but many are also generalist parasitoids; lots are used in successful biological control programs against insect pests.
One area of my research is the family Eulophidae, the most numerous group of Chalcidoidea (ca. 4470 described species). The taxonomy of this group is relatively poorly understood. Eulophids attack wide variety of insect, and occasionally mite or spider hosts. Their larvae act as koino- or idiobionts, gregarious or solitary, ecto- or endoparasitoids; they attack eggs, larvae or pupae of their hosts. Some species are phytophagous (chiefly gall-formers on eucalypts).
A second area of research is the family Tetracampidae. This is a small family with about 50 described species classified in 15 genera. The extant groups of the family are represented by egg parasitoids of Hymenoptera and Coleoptera (Tetracampinae), larval parasitoids of Diptera (Tetracampinae, Platynocheilinae), and phytophagous species (Mongolocampinae). Unlike Eulophidae, the Tetracampidae look rather arteficial than natural group in its current concept.
Another area of my interests is fossil Chalcidoidea. Chalcids are relatively poorly known from the fossils. The amber inclusions are probably the only source for such fossil data because of minute size and weak body sclerotization of these insects.
There are many reasons for the comparatively modest progress in this area: poor condition of the amber samples, minute diagnostic characters not visible in amber specimens, poor representation of the chalcids in amber fossils (in comparison with other groups of insects), to list some.
The records of Eulophidae from ambers are rare. Boucek & Askew (1968) briefly mentioned Entedon sp. from Dominican amber, however, without exact identification of the amber specimen. Then Gumovsky (2001) recorded two eulophids also from Dominican amber (Chrysonotomyia dominicana and Achrysocharoides sp.).
Some amber insect inclusions are classified in Tetracampidae: namely this family is among the most represented ones in fossil resins. However, clarification of the correct family affiliation of fossil tetracampids requires additional studies. The placement of fossil groups in Tetracampidae was likely motivated by the conventional concept of this family as intermediate between Pteromalidae and Eulophidae, and thus serving as a “dumping ground” for the species with problematic status.
My current research in this field focuses on a survey of chalcid inclusions in the Canadian and Taymyr (Cretaceous, 80-90 million years ago), Baltic (Late Eocene, about 40 million years ago) and Dominican (Miocene, about 20 million years ago) ambers. Gumovsky et al. (2018) conducted a survey of the chalcidoid inclusions in Taimyr amber (84-100 Ma). As a result, and 11 new species, were described in Baeomorpha. Also, Baeomorphinae Yoshimoto, 1975, based on Baeomorpha Brues, 1937, is transferred from Tetracampidae Foerster, 1856 and recognized as a junior synonym of Rotoitidae Boucek and Noyes, 1987 (Hymenoptera: Chalcidoidea). One enigmatic rotoitid inclusion, which differs from Baeomorpha species in the possession of very short stigmal vein, is described as Taimyromorpha pusilla Gumovsky. Inclusions containing specimens identified as Baeomorpha and Taimyromorpha are found in amber from Taimyr and Canada that originated from Laurasia, not Gondwana. The newly described fossils indicate the southern hemisphere distribution of extant Rotoitidae is relictual with the pattern observed being formed at least in part by extinction events, though distributions of the only two extant rotoitid genera, Rotoita (New Zealand) and Chiloe (small area in the southern Chile) may have been more extensive in the past. Also, the Canadian amber genera Distylopus (Distylopinae) and Bouceklytus (Bouceklytinae) are excluded from Tetracampidae and regarded as Chalcidoidea incertae sedis.
Other my research interests include functional morphology of attachment and grooming structures in Hymenoptera. One of the structures is the pretarsus, that serves as the main attachment device. The pretarsus of Chalcidoidea is characterized by a distal elastic widening of the planta that spreads over the arcus, by a pair of folding plates at the dorsal side of the arolium (the dorsal plates), and by the absence of auxiliary sclerites. The surface of the fully spread arolium of chalcids has a spongiform structure. The peculiarities of the inverting/everting biomechanics of the pretarsus of chalcids involve: (1) interactions between the elastic part of the planta, the dorsal plates and the manubrium; (2) the functioning of the elastic part of the planta and the arcus together as a single unit.
A manubrium with a distinct proximal row of three setae characterizes almost all Eulophidae, Aphelinidae and Signiphoridae (‘eulophid lineage’) and Tetracampidae, whereas a row of two setae characterizes Mymaridae, Rotoitidae and Trichogrammatidae. Other studied families (Pteromalidae, Eurytomidae, Torymidae, Ormyridae, Eupelmidae, Encyrtidae, Perilampidae), which represent a ‘pteromalid lineage ’, are characterized mostly by five setae in a proximal row, which could represent a synapomorphy for these groups, or a symplesiomorphy in Chalcidoidea, depending on rooting. However, these characters may be correlated with differences in body size that characterize the different lineages rather than being phylogenetically important. Other characters that may be phylogenetically informative are: (1) shape of the manubrium (spindle-like in Mymaridae, Rotoitidae, Trichogrammatidae and the ‘eulophid lineage’, but mostly bottle-like in representatives of the ‘pteromalid lineage’); (2) pubescence of the proximal part of the planta (sparse, thick setae in Rotoitidae, Trichogrammatidae and the ‘eulophid lineage’, but dense, slender setae in representatives of the ‘pteromalid lineage’).
One more area of my research interests include morphology of immature stages of Chalcidoidea and all aspects of biology and evolution of egg-larval parasitism in Chalcidoidea, which are rather fragmentary up-to-date. Most discussions on larval morphology and biology of chalcid wasps concern ectoparasitoids, whereas endoparasitoid larvae traditionally attract less attention, mainly due to the difficulties with their preparation and identification.
The development of solitary and gregarious eulophid endoparasitoids (Entedon, in particular) differs in immature morphology and physiology. Generally, the first-instar larvae of solitary species have a beak-shaped head with sharp mandibles and get involved in siblicide (killing of rivals). The first-instar larvae of gregarious endoparasitoids are characterized by shorter mandibles and are tolerant to each other (Harvey et al. 2012), although the morphology of the larval gregarious parasitoids of the genus Entedon has not been described so far. Gumovsky & Ramadan (2011) described a unique biology of Afrotropical species Entedon erythrinae, a gregarious egg-larval endoparasitoid of the bruchid beetle Specularius impressithorax.
The early development of E. erythrinae differs from all other species of the genus and is peculiar among known life histories of chalcidoid wasps. Unlike other non-polyembryonic chalcidoids, the early development of the parasitoid egg is not associated with organogenesis, but simply results in multiplication of proliferating cells. The cells differentiate into the external layer (formed by the larger elongate cells) and the inner cell mass (formed by the smaller relatively rounded cells). The external layer gives rise to an extraembryonic membrane or serosa (so called ‘soccer ball’ chamber), while the internal cell aggregation develops further into the larva. This larva shares all the morphological characters with the second instar of other Entedon species, for which immature stages are known. Therefore, the first (aggressive and responsible for siblicide) instar larval stage is omitted, what may happen to be an adaptation to gregariousness (Gumovsky & Ramadan 2011). However, more data on other gregarious endoparasitoids are required to verify whether such embryonization is an ultimate or exceptional adaptation.
The origin and alteration of the egg-larval and larval parasitism within the same genus of parasitoids may shed light on the evolution of aggressiveness and tolerance in animal world as a whole (with possible implications for humans also: e.g., Gomez et al. 2016).
All these studies incorporate morphological, biological or molecular information into analyses that are used to produce hypotheses of phylogenetic relationships and the evolution of behavioral patterns.
The first signal of monophyly of Eulophidae was obtained by Campbell et al. (2000) based on 28S D2 rDNA gene of chalcidoids. Also, the genus Elasmus, the only genus of the former family Elasmidae, appeared a derived taxon within Eulophidae. Then Gauthier et al. (2000) used the same gene on broader sampling to clarify the phylogeny of Eulophidae. These authors supported the idea that Elasmidae are derived eulophids, the subfamilies Eulophinae and Tetrastichinae are closely related and provided some new characters to support monophyly of some groups of Eulophidae. Gumovsky (2002) also used the same gene for the subfamily Entedoninae and discussed distribution of some morphological characters in Entedoninae.
The families Eulophidae, Tetracampidae, Trichogrammatidae and also Aphelinidae are sometimes considered as the “eulophid lineage” (Gibson et al., 1997). This group is characterized by reduced number of antennal (generally 4-7, but occasionally more) and tarsal (3-4) segments. However, there are many overlaps in distribution of these characters as within the “eulophid lineage”, as with outgroups. Gladun & Gumovsky (2006) demonstrated that the representatives of the “eulophid lineage” share similar morphology of pretarsus. Gokhman (2004, 2009) and then Gokhman & Gumovsky (2009) suggested that Eulophidae belong to the so-called "low-numbered" chalcidoid families with the modal haploid number of chromosomes n=6 (but with occasional modifications to 5 or 7), whereas n=9-12 in many other families. However, such karyoptype occurs also outside Eulophidae and also the number of chromosomes reduced independently in various groups of Chalcidoidea (Gokhman & Gumovsky, 2009).
Our research (Gumovsky, 2011) based on the combined analysis of nuclear (28S D2 rDNA) and two mitochondrial (cytochrome oxidase subunit I, COI, and cytochrome b, Cyt b) sequences demonstrated that there is no signal of close relationships between the families Eulophidae, Tetracampidae and Trichogrammatidae, but at least Eulophidae and Trichogrammatidae are supported as monophyletic. Also, the Eulophidae-lineage appeared to consist of four internal lineages corresponding to the recognized subfamilies: Eulophinae, Tetrastichinae, Entedoninae and Entiinae (= Euderinae), what was also supported by morphological characters (including putative synapomorphies). Largely similar pattern was reported by other students foe Eulophidae (Burks et al., 2011) and Chalcidoidea as a whole (Munro et al., 2011; Heraty et al., 2013).
The most ancient records of fossil Chalcidoidea are dated as Cretaceous (78-115 Mya) and represented by the ‘basal’ group (e.g. Mymarommatidae and Mymaridae and the unspecified ‘aphelinid–trichogrammatid’ representatives: Heraty et al., 2013). The rest of the groups are common in Eocene deposits (ca. 40 Mya), and so the main diversification is presumed to have taken place in the Late Cretaceous or soon after. The phylogenetic analyses conducted so far (Burks et al., 2011; Munro et al., 2011; Heraty et al., 2013) suggest that the common ancestor of Eulophidae, Tetracampidae and Trichogrammatidae had branched off yet in Gondwana (ca. 120–60 Mya). In particular, the enigmatic genus Trisecodes (unplaced within Eulophidae) had diversified when the modern Africa and South America were not yet split (Gumovsky, 2014).
Research on functional morphology of Hymenoptera has been used for phylogenetic interpretation. Gibson (1985) and Heraty et al. (1997) discussed the phylogenetic implications of some pro- and mesothoracic structures. Basibuyuk et al. also inferred phylogenetic information from the morphology of the antennal cleaner (1995), hamuli (1997) and grooming behaviours (1999). Basibuyuk et al. (2000a) also discussed peculiarities of the structure and location of the sensilla on the manubrium in Hymenoptera in light of established hypotheses of relationships within Hymenoptera.
Gladun & Gumovsky (2006) demonstrated that character-state distribution of features of the pretarsus among studied taxa support some established groups or proposed associations of Chalcidoidea.
The evolution of host associations and peculiarities of host shifts during natural history of the group, are fascinating areas of research. Such studies are also of paramount importance to biocontrol programs: this demonstrates the utility and impact of systematics to almost every area of science.
Fellowships and research visits
2016 - visiting researcher, IZIKO, South African National Museum, Cape Town, South Africa.
2014, 2015, 2016 - visiting researcher, University of Witwatersrand, Johannesburg, South Africa.
2013 - University of Kisangani
(UNIKIS) - Centre de Surveillance de la Biodiversite, Democratic
Republic of Congo: collaborative research.
2012-2013 - Post-doctoral
fellow, the Museum of Central Africa (Tervuren) (MRAC-RMCA), BELSPO-Marie
Curie supported fellowship.
2010-2011 - Wageningen University and The Netherlands Institute of Ecology (NIOO-KNAW)
2009 - Bishop Museum, Honolulu, Hawaii, USA: a study of collection material.
2009 - University of Hawaii at Manoa, Department of Entomology, Honolulu, Hawaii, USA: a workshop with students and a study of collection material.
2009 - Hawaii Deapartment of Agriculture, Honolulu, USA, a collaborative project on invasive bruchid beetle pests and their parasitoids.
2009 - ICIPE, Nairobi, Kenya: a study of collection material
2008 - Mugla University, Mugla, Turkey: workshop with students, a study of collection material, collaborative research and field trips.
2008 - University of Dakar, Senegal: a study of collection material, collaborative research and field trips.
2007 - Mustafa Kemal University, Antakya, Turkey: a study of collection material, collaborative research and field trips
2006 - Paleontological Institute of Russian Academy of Sciences (Moscow, Russia): a study of fossil chalcidoids.
2005 - Entomology Research Museum, University of California (Riverside, USA): a study of collection material.
2005 - The Natural History Museum (London, UK): a study of collection material, molecular studies on eulophid wasps.
2004 - Naturkunde Museum (Stuttgart, Germany): a study of chalcidoid inclusions in fossil resins.
2004 - Zoologische Staatssammlung (Munchen, Germany): a study of collection material.
2004 - Max-Planck Institute for Metal Research (Stuttgart, Germany): a study on functional morphology of chalcids.
2003 - European Station of CABI Bioscience (Delemont, Switzerland): biocontrol project management.
2000 - CIRAD-Amis (Montpellier, France): a study of collection material.
2000 - Museum National d'Histoire Naturelle (Paris, France): a study of collection material.
2000 - Zoological Muzeum, University of Copenhagen (Denmark): a study of collection material.
2000 - The Natural History Museum (London, UK): a study of collection material.
2000 - Imperial College at Silwood Park (Ascot, UK): molecular studies on eulophid wasps.
1998 - Zoological Institute of Russian Academy of Sciences (St.-Petersburg, Russia): a study of collection material.
1996 - Regional Environmental Center for Eastern and Central Europe (Budapest, Hungary): nature conservation project management.
2014 - SYNTHESYS Project: the research project was conducted in The Natural History Museum (London).
2014 - The award of the State Fund of Fundamental Research (Ukraine), grant F50/029 for young holders of Dr. Sci. (Habilitation) degree: Parasitoid guilds of alien pests as a model for studying of the adaptive capacities of native and alien parasitoids.
2012-2013 - BELSPO-Marie Curie supported fellowship.
2011 - The award of the State Fund of Fundamental Research (Ukraine), grant F35/002 for young holders of Dr. Sci. (Habilitation) degree: Phylogenetic and faunistic research on Eulophidae (Hymenoptera).
2010-2011 - Wageningen University ‘Production Ecology and Resource Centre’ (PE&RC, Netherlands) Visiting Scientist Scholarship.
2011 - SFFR (The State Fund of Fundamental Research (Ukraine)), grant F 35/002.
2007- 2009 - Alexander von Humboldt Foundation, research fellowship award.
2006 - TUBITAK / NASU Academic Exchange grant.
2005 - SYNTHESYS Project (GB–TAF–535).
2004 - DAAD (German Academic Exchange Service, grant No. 322 – A/04/15867).
2001 - SFFR (The State Fund of Fundamental Research (Ukraine)) grant 05.07/00078.
2000 - European Commission research grant (under COBICE-programme).
1999 - Royal Society / NATO Grant (NATO/ 99A/bll).
2014 – I.I. Schmalhausen Award of the National Academy of Sciences of Ukraine (together with A. Radchenko and E.E. Perkovsky).
2002-2004 - President of Ukraine Scholarship for young scientists.
1997, 1998 - George Soros Foundation (International Foundation Vidrodgenia) grants.
1996 - The National Academy of sciences of Ukraine award for young scientists.
1997 - Travel grant of the International Association of Academies of Sciences.
1995 - Taras Shevchenko’s award of Kiev National University.
1995 - Special student award of Kiev National University.
Participation in international projects on phylogeny and taxonomy
2016-2018 - NSF An integrative approach to understanding the evolution and systematics of Chalcidoidea: A recent megaradiation of Hymenoptera (superviser Prof. J. Heraty, University of California, Riverside). Contribution to the project: phylogeny of Eulophidae, Tetracampidae.
2005-2009 - Tree of Life Project: Phylogeny of Chalcidoidea (superviser Prof. J. Heraty, University of California, Riverside). Contribution to the project: phylogeny of Eulophidae, Tetracampidae.
2006 - A project “Identifications of taxa of Entedoninae (Hym.: Eulophidae) from Turkey by using molecular DNA markers” supported by TUBITAK / NASU Academic Joint Research grant, in collaboration with Mustafa Kemal University, Antakya, Turkey. Contribution to the project: taxonomic and phylogenetic studies.
Nature conservation grants / Projects
1997 - Biodiversity Support Program (U.S.A.I.D. - funded consortium of World Wildlife Fund, The Nature Conservancy, and World Resources Institute): “Studying on and conservation of beneficial insects (Hymenoptera mainly) of the National Park “Podol’s’ki Tovtry” (South-Western Ukraine) [Project Director].
1994 - ISAR, a Clearinghouse on Grassroots Cooperation in Eurasia: “Biodiversity survey in the nature reserve "Lesniki" (Kiev vicinity)” [Project Director].
Participation in biocontrol projects
2009 - A project on biology of an invasive bruchid beetle pest Specularius impressithorax and its hymenopterous parasitoids (with Hawaii Deapartment of Agriculture, Honolulu, USA). Project contribution: studies biology, development and taxonomy of the beetle and its parasitoids.
2003-2005 - A project on approaches to control invasive moth pest Cameraria ohridella (curated by Schmalhausen Institute of Zoology, Kiev, Ukraine). Project contribution: studies on biology and taxonomy of the parasitoids of the moth.
2002-2004 - Institutional Partnership project ‘‘Emphasising Classical and Conservation Biological Control in Research and Teaching’’ (7 IP 65648). Project contribution: preparation of manuals on biocontrol; lectures and studies on biology of biocontrol agents (beetles and parasitoids).
2001-2002 - A project on approaches to control invasive moth pest Acrocercops brongniardella (curated by Schmalhausen Institute of Zoology, Kiev, Ukraine). Project contribution: studies on biology and taxonomy of the parasitoids of the moth.
England, France (2000)
Hungary (2001, 2010)
Turkey (2007, 2008)
The Democratic Republic of Congo (2013, 2014, 2015)
Uganda (2009, 2013, 2014, 2015, 2016)
Kenya (2009, 2013)
Namibia (2011, 2014-2016)
South Africa (2014-2016)
Anonymous referee in “Zoosystematica Rossica", "Entomotropica", "Zootaxa", “Journal of Natural history”.
Ukrainian (native), Russian (second native), English (fluent), German (functional).
Alex Gumovsky, I. I. Schmalhausen Institute of Zoology, 2017