The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps.

Chalcidoidea are mostly parasitoid wasps that include as many as 500 000 estimated species. Capturing phylogenetic signal from such a massive radiation can be daunting. Chalcidoidea is an excellent example of a hyperdiverse group that has remained recalcitrant to phylogenetic resolution. We combined 1007 exons obtained with Anchored Hybrid Enrichment with 1048 ultra-conserved elements (UCEs) for 433 taxa including all extant families, >95% of all subfamilies, and 356 genera chosen to represent the vast diversity of the superfamily. Going back and forth between the molecular results and our collective knowledge of morphology and biology, we detected bias in the analyses that was driven by the saturation of nucleotide data. Our final results are based on a concatenated analysis of the least saturated exons and UCE datasets (2054 loci, 284 106 sites). Our analyses support an expected sister relationship with Mymarommatoidea. Seven previously recognized families were not monophyletic, so support for a new classification is discussed. Natural history in some cases would appear to be more informative than morphology, as illustrated by the elucidation of a clade of plant gall associates and a clade of taxa with planidial first-instar larvae. The phylogeny suggests a transition from smaller soft-bodied wasps to larger and more heavily sclerotized wasps, with egg parasitism as potentially ancestral for the entire superfamily. Deep divergences in Chalcidoidea coincide with an increase in insect families in the fossil record, and an early shift to phytophagy corresponds with the beginning of the "Angiosperm Terrestrial Revolution". Our dating analyses suggest a middle Jurassic origin of 174 Ma (167.3-180.5 Ma) and a crown age of 162.2 Ma (153.9-169.8 Ma) for Chalcidoidea. During the Cretaceous, Chalcidoidea may have undergone a rapid radiation in southern Gondwana with subsequent dispersals to the Northern Hemisphere. This scenario is discussed with regard to knowledge about the host taxa of chalcid wasps, their fossil record and Earth's palaeogeographic history

On the identity of three little-known Microterys Thomson species (Hymenoptera: Encyrtidae)

Illustrated redescriptions are given for three species of encyrtid wasps first described in the early 1800s: Microterys cedrenus (Walker), M. cyanocephalus (Dalman) and M. interpunctus (Dalman), and four new synonyms are proposed: M. aldreyi Japoshvili (of M. cedrenus), M. dichrous (Mercet) (of M. cedrenus), M. steinbergi Sugonjaev (of M. cyanocephalus), and M. duplicatus (Nees) (of M. interpunctus)

From hell’s heart I stab at thee! A determined approach towards a monophyletic Pteromalidae and reclassification of Chalcidoidea (Hymenoptera)

We thank Daniel Alejandro Aquino (Museo de La Pla- ta) , Fernando H. A. Farache (Rio Verde) , Jeremy Frank (Bishop Museum) , and Ryan Perry (UC Santa Barbara) for helpful photographs of type specimens. We thank Nicole Fisher and Juanita Rodriguez Arrieta (ANIC, Canberra) as well as Christine Lambkin, Chris Burwell and Susan Wright (QM, Brisbane) for the loan of multiple Australian specimens. We acknowledge the Queensland government for collecting permits (WITK18248017-WITK18278817).International audienceThe family Pteromalidae (Hymenoptera: Chalcidoidea) is reviewed with the goal of providing nomenclatural changes and morphological diagnoses in preparation for a new molecular phylogeny and a book on world fauna that will contain keys to identification. Most subfamilies and some tribes of Pteromalidae are elevated to family level or transferred elsewhere in the superfamily. The resulting classification is a compromise, with the aim of preserving the validity and diagnosability of other, well-established families of Chalcidoidea. The following former subfamilies and tribes of Pteromalidae are elevated to family rank: Boucekiidae, Ceidae, Cerocephalidae, Chalcedectidae, Cleonymidae, Coelocybidae, Diparidae, Epichrysomallidae, Eunotidae, Herbertiidae, Hetreulophidae, Heydeniidae, Idioporidae, Lyciscidae, Macromesidae, Melanosomellidae, Moranilidae, Neodiparidae, Ooderidae, Pelecinellidae (senior synonym of Leptofoeninae), Pirenidae, Spalangiidae, and Systasidae. The following subfamilies are transferred from Pteromalidae: Chromeurytominae and Keiraninae to Megastigmidae, Elatoidinae to Neodiparidae, Nefoeninae to Pelecinellidae, and Erotolepsiinae to Spalangiidae. The subfamily Sycophaginae is transferred to Pteromalidae. The formerly incertae sedis tribe Lieparini is abolished and its single genus Liepara is transferred to Coelocybidae. The former tribe Tomocerodini is transferred to Moranilidae and elevated to subfamily status. The former synonym Tridyminae (Pirenidae) is treated as valid. The following former Pteromalidae are removed from the family and, due to phylogenetic uncertainty, placed as incertae sedis subfamilies or genera within Chalcidoidea: Austrosystasinae, Ditropinotellinae, Keryinae, Louriciinae, Micradelinae, Parasaphodinae, Rivasia, and Storeyinae. Within the remaining Pteromalidae, Miscogastrinae and Ormocerinae are confirmed as separate from Pteromalinae, the former tribe Trigonoderini is elevated to subfamily status, the former synonym Pachyneurinae is recognized as a distinct subfamily, and as the senior synonym of Austroterobiinae. The tribe Termolampini is synonymized under Pteromalini, and the tribe Uzkini is synonymized under Colotrechnini. Most former Otitesellinae, Sycoecinae, and Sycoryctinae are retained in the tribe Otitesellini, which is transferred to Pteromalinae, and all other genera of Pteromalinae are treated as Pteromalini. Eriaporidae is synonymized with Pirenidae, with Eriaporinae and Euryischiinae retained as subfamilies. Other nomenclatural acts performed here outside of Pteromalidae are as follows: Calesidae: elevation to family rank. Eulophidae: transfer of Boucekelimini and Platytetracampini to Opheliminae, and abolishment of the tribes Elasmini and Gyrolasomyiini. Baeomorphidae is recognized as the senior synonym of Rotoitidae. Khutelchalcididae is formally excluded from Chalcidoidea and placed as incertae sedis within Apocrita. Metapelmatidae and Neanastatidae are removed from Eupelmidae and treated as distinct families. Eopelma is removed from Eupelmidae and treated as an incertae sedis genus in Chalcidoidea. The following subfamilies and tribes are described as new: Cecidellinae (in Pirenidae), Enoggerinae (incertae sedis in Chalcidoidea), Erixestinae (in Pteromalidae), Eusandalinae (in Eupelmidae), Neapterolelapinae (incertae sedis in Chalcidoidea), Solenurinae (in Lyciscidae), Trisecodinae (in Systasidae), Diconocarini (in Pteromalidae: Miscogastrinae), and Trigonoderopsini (in Pteromalidae: Colotrechninae). A complete generic classification for discussed taxa is provided

From hell’s heart I stab at thee! A determined approach towards a monophyletic Pteromalidae and reclassification of Chalcidoidea (Hymenoptera)

We thank Daniel Alejandro Aquino (Museo de La Pla- ta) , Fernando H. A. Farache (Rio Verde) , Jeremy Frank (Bishop Museum) , and Ryan Perry (UC Santa Barbara) for helpful photographs of type specimens. We thank Nicole Fisher and Juanita Rodriguez Arrieta (ANIC, Canberra) as well as Christine Lambkin, Chris Burwell and Susan Wright (QM, Brisbane) for the loan of multiple Australian specimens. We acknowledge the Queensland government for collecting permits (WITK18248017-WITK18278817).International audienceThe family Pteromalidae (Hymenoptera: Chalcidoidea) is reviewed with the goal of providing nomenclatural changes and morphological diagnoses in preparation for a new molecular phylogeny and a book on world fauna that will contain keys to identification. Most subfamilies and some tribes of Pteromalidae are elevated to family level or transferred elsewhere in the superfamily. The resulting classification is a compromise, with the aim of preserving the validity and diagnosability of other, well-established families of Chalcidoidea. The following former subfamilies and tribes of Pteromalidae are elevated to family rank: Boucekiidae, Ceidae, Cerocephalidae, Chalcedectidae, Cleonymidae, Coelocybidae, Diparidae, Epichrysomallidae, Eunotidae, Herbertiidae, Hetreulophidae, Heydeniidae, Idioporidae, Lyciscidae, Macromesidae, Melanosomellidae, Moranilidae, Neodiparidae, Ooderidae, Pelecinellidae (senior synonym of Leptofoeninae), Pirenidae, Spalangiidae, and Systasidae. The following subfamilies are transferred from Pteromalidae: Chromeurytominae and Keiraninae to Megastigmidae, Elatoidinae to Neodiparidae, Nefoeninae to Pelecinellidae, and Erotolepsiinae to Spalangiidae. The subfamily Sycophaginae is transferred to Pteromalidae. The formerly incertae sedis tribe Lieparini is abolished and its single genus Liepara is transferred to Coelocybidae. The former tribe Tomocerodini is transferred to Moranilidae and elevated to subfamily status. The former synonym Tridyminae (Pirenidae) is treated as valid. The following former Pteromalidae are removed from the family and, due to phylogenetic uncertainty, placed as incertae sedis subfamilies or genera within Chalcidoidea: Austrosystasinae, Ditropinotellinae, Keryinae, Louriciinae, Micradelinae, Parasaphodinae, Rivasia, and Storeyinae. Within the remaining Pteromalidae, Miscogastrinae and Ormocerinae are confirmed as separate from Pteromalinae, the former tribe Trigonoderini is elevated to subfamily status, the former synonym Pachyneurinae is recognized as a distinct subfamily, and as the senior synonym of Austroterobiinae. The tribe Termolampini is synonymized under Pteromalini, and the tribe Uzkini is synonymized under Colotrechnini. Most former Otitesellinae, Sycoecinae, and Sycoryctinae are retained in the tribe Otitesellini, which is transferred to Pteromalinae, and all other genera of Pteromalinae are treated as Pteromalini. Eriaporidae is synonymized with Pirenidae, with Eriaporinae and Euryischiinae retained as subfamilies. Other nomenclatural acts performed here outside of Pteromalidae are as follows: Calesidae: elevation to family rank. Eulophidae: transfer of Boucekelimini and Platytetracampini to Opheliminae, and abolishment of the tribes Elasmini and Gyrolasomyiini. Baeomorphidae is recognized as the senior synonym of Rotoitidae. Khutelchalcididae is formally excluded from Chalcidoidea and placed as incertae sedis within Apocrita. Metapelmatidae and Neanastatidae are removed from Eupelmidae and treated as distinct families. Eopelma is removed from Eupelmidae and treated as an incertae sedis genus in Chalcidoidea. The following subfamilies and tribes are described as new: Cecidellinae (in Pirenidae), Enoggerinae (incertae sedis in Chalcidoidea), Erixestinae (in Pteromalidae), Eusandalinae (in Eupelmidae), Neapterolelapinae (incertae sedis in Chalcidoidea), Solenurinae (in Lyciscidae), Trisecodinae (in Systasidae), Diconocarini (in Pteromalidae: Miscogastrinae), and Trigonoderopsini (in Pteromalidae: Colotrechninae). A complete generic classification for discussed taxa is provided

Fig. 4 in The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps

Fig. 4. The Chalcidoidea bush of life. (a) IQ-TREE tree obtained from the combined exonsAA+UCEs90-25 datasets (see also Fig. S1). Monophyletic families are in grey, para- or polyphyletic families are in colour. Higher level groups/clades discussed in text are highlighted with boxes. Statistical support for backbone nodes are shown with single (SH-aLRT ≧80% or UFboot ≧95%) or double stars (SH-aLRT ≧80% and UFboot ≧95%). (b) Contribution of the exonsAA and UCEs90-25 datasets to the combined tree. Gene concordance factor (gCF); gene discordance factor due to polyphyly (gDFP); site concordance factor averaged over 100 quartets (sCF). Points: raw data (Table S2d). (c) Comparison of branch length for the backbone nodes and other ingroup nodes. Points: raw data (Table S2c). For (b) and (c), stars above box plots indicate statistical significance: ns, p> 0.05; ***, p ≤ 0.001; ****, p ≤ 0.0001. (d) Correlation between node age and sCF (outgroups excluded). Points: raw data (Table S2e); line: regression curve for the best-fit model (log linear model; p <2.2e—16).Published as part of <i>Cruaud, Astrid, Rasplus, Jean-Yves, Zha, Junxia, Burks, Roger, Delvare, Ǵerard, Fusu, Lucian, Gumovsky, Alex, Huber, John T., Jan̆sta, Petr, Mitroiu, Mircea-Dan, Noyes, John S., Noort, Simon van, Baker, Austin, Bohmova, Julie, Baur, Hannes, Blaimer, Bonnie B., Brady, Sean G., Bubeńıkova, Kristyna, Chartois, Marguerite, Copeland, Robert S., Papilloud, Natalie Dale-Skey, Molin, Ana Dal, Dominguez, Chrysalyn, Gebiola, Marco, Guerrieri, Emilio, Kresslein, Robert L., Krogmann, Lars, Lemmon, Emily, Murray, Elizabeth A., Nidelet, Sabine & Nieves-Aldrey, Jośe Luis, 2023, The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps, pp. 1-30 in Cladistics 2023</i> on page 13, DOI: 10.1111/cla.12561, <a href="http://zenodo.org/record/10115140">http://zenodo.org/record/10115140</a&gt

The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps

Cruaud, Astrid, Rasplus, Jean-Yves, Zha, Junxia, Burks, Roger, Delvare, Ǵerard, Fusu, Lucian, Gumovsky, Alex, Huber, John T., Jan̆sta, Petr, Mitroiu, Mircea-Dan, Noyes, John S., Noort, Simon van, Baker, Austin, Bohmova, Julie, Baur, Hannes, Blaimer, Bonnie B., Brady, Sean G., Bubeńıkova, Kristyna, Chartois, Marguerite, Copeland, Robert S., Papilloud, Natalie Dale-Skey, Molin, Ana Dal, Dominguez, Chrysalyn, Gebiola, Marco, Guerrieri, Emilio, Kresslein, Robert L., Krogmann, Lars, Lemmon, Emily, Murray, Elizabeth A., Nidelet, Sabine, Nieves-Aldrey, Jośe Luis (2023): The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps. Cladistics 2023: 1-30, DOI: 10.1111/cla.1256

Fig. 5 in The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps

Fig. 5. Global historical biogeography of Chalcidoidea and new classification. The chronogram obtained from the complete set of ingroup taxa is illustrated. The previous classification is used to annotate tips (four letter prefixes; see also Table S1 for complete information on sampling) with successive grey and white boxes grouping the tip labels. The new familial classification from Burks et al. (2022) is shown to the right. For clarity, ancestral ranges are given only up to family level and only for the BAYEAREALIKE + J model (which was selected by AICc). All inferences of ancestral ranges are provided in Fig. S5. Inferences of ancestral ranges were conducted with only one specimen per genus as shown with brackets that connect tips. Current distribution of genera is shown with coloured boxes at tips. Sampling area of specimens is indicated in tip labels. NEO = Neotropical; NEA = Nearctic; AFR = Afrotropical; PAL = Palaearctic; ORI = Oriental; AUS = Australasian. UKN = Unknown when collection data are unavailable. Stars indicate that specimens were sampled in areas where species was introduced or not yet cited. Sampling area for the specimen used for sequencing exons is listed first, sampling area for the specimen used for sequencing UCEs is listed second; n.a. is used when no specimen was sequenced and only one sampling area is reported when exons and UCEs were obtained from specimens sampled in the same areas (or from the same specimen). Unless specified, nodes are supported by SHaLRT ≥80%, UFBoot ≥95% and sCF ≥34.3 (minimum support for a family that is well defined morphologically, Trichogrammatidae). Nodes with a grey circle are supported by SHaLRT <80% or UFBoot <95%; nodes with a black circle are supported by SHaLRT <80% and UFBoot <95%; nodes with a black triangle are supported with sCF <34.3. Images on the left of tentative family names are all at the same scale. Images on the right of tentative family names have been magnified. Photos ©K. Bolte (Baeomorphidae); ©J.-Y. Rasplus (all others).Published as part of <i>Cruaud, Astrid, Rasplus, Jean-Yves, Zha, Junxia, Burks, Roger, Delvare, Ǵerard, Fusu, Lucian, Gumovsky, Alex, Huber, John T., Jan̆sta, Petr, Mitroiu, Mircea-Dan, Noyes, John S., Noort, Simon van, Baker, Austin, Bohmova, Julie, Baur, Hannes, Blaimer, Bonnie B., Brady, Sean G., Bubeńıkova, Kristyna, Chartois, Marguerite, Copeland, Robert S., Papilloud, Natalie Dale-Skey, Molin, Ana Dal, Dominguez, Chrysalyn, Gebiola, Marco, Guerrieri, Emilio, Kresslein, Robert L., Krogmann, Lars, Lemmon, Emily, Murray, Elizabeth A., Nidelet, Sabine & Nieves-Aldrey, Jośe Luis, 2023, The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps, pp. 1-30 in Cladistics 2023</i> on page 15, DOI: 10.1111/cla.12561, <a href="http://zenodo.org/record/10115140">http://zenodo.org/record/10115140</a&gt

The Chalcidoidea bush of life: evolutionary history of a massive radiation of minute wasps

Raw paired reads were uploaded as NCBI Sequence Read Archives (PRJNA884376 for AHE and PRJNA1017994 for UCEs).We dedicate this work to the memory of our dear friend and colleague John LaSalle, specialist of Eulophidae, who was an enthusiastic member of this project.International audienceChalcidoidea are mostly parasitoid wasps that include as many as 500 000 estimated species. Capturing phylogenetic signal from such a massive radiation can be daunting. Chalcidoidea is an excellent example of a hyperdiverse group that has remained recalcitrant to phylogenetic resolution. We combined 1007 exons obtained with Anchored Hybrid Enrichment with 1048 ultra-conserved elements (UCEs) for 433 taxa including all extant families, >95% of all subfamilies, and 356 genera chosen to represent the vast diversity of the superfamily. Going back and forth between the molecular results and our collective knowledge of morphology and biology, we detected bias in the analyses that was driven by the saturation of nucleotide data. Our final results are based on a concatenated analysis of the least saturated exons and UCE datasets (2054 loci, 284 106 sites). Our analyses support an expected sister relationship with Mymarommatoidea. Seven previously recognized families were not monophyletic, so support for a new classification is discussed. Natural history in some cases would appear to be more informative than morphology, as illustrated by the elucidation of a clade of plant gall associates and a clade of taxa with planidial first-instar larvae. The phylogeny suggests a transition from smaller soft-bodied wasps to larger and more heavily sclerotized wasps, with egg parasitism as potentially ancestral for the entire superfamily. Deep divergences in Chalcidoidea coincide with an increase in insect families in the fossil record, and an early shift to phytophagy corresponds with the beginning of the “Angiosperm Terrestrial Revolution”. Our dating analyses suggest a middle Jurassic origin of 174 Ma (167.3–180.5 Ma) and a crown age of 162.2 Ma (153.9–169.8 Ma) for Chalcidoidea. During the Cretaceous, Chalcidoidea may have undergone a rapid radiation in southern Gondwana with subsequent dispersals to the Northern Hemisphere. This scenario is discussed with regard to knowledge about the host taxa of chalcid wasps, their fossil record and Earth's palaeogeographic history