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

A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea)

Perry, Ryan K., Heraty, John M. (2019): A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea). Insect Systematics and Diversity (AIFB) 3 (5), No. 1: 1-23, DOI: 10.1093/isd/ixz012, URL: http://dx.doi.org/10.1093/isd/ixz01

Fig. 6 in A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea)

Fig. 6. Dorsal view of mesosomal color variation in Burkseus vittatus: (A) D4559, (B) D5320, (C) D5113, (D) D5101, (E) D4699, (F) D3640, (G) D4173, (H) D3665, (I) D3666, (J) D4177, (K) D3993, and (L) illustrated view. Scale bar = 0.2 mm.Published as part of Perry, Ryan K. & Heraty, John M., 2019, A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea), pp. 1-23 in Insect Systematics and Diversity (AIFB) 3 (5) on page 12, DOI: 10.1093/isd/ixz012, http://zenodo.org/record/716823

Fig. 2 in A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea)

Fig. 2. Fore wings, dorsal view: (A) Burkseus vittatus comb. n., (B) B. flavoviridis comb. n., (C) B. robustus n. sp., (D) B. sigillatus n. sp. The photographs of B. robustus and B. sigillatus were taken after DNA extraction, displaying how the dark colors on the submarginal vein setae (B. robustus and B. sigillatus), stigmal vein and uncus (B. robustus), and banding patterns (B. sigillatus) are resilient enough to remain visible after extraction. Scale bar = 0.5 mm. ams = admarginal setae, bsl = basal setal line, csl = cubital setal line, disc = fore wing disc, spc = speculum, smv = submarginal vein, stg = stigmal vein, unc = uncus.Published as part of Perry, Ryan K. & Heraty, John M., 2019, A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea), pp. 1-23 in Insect Systematics and Diversity (AIFB) 3 (5) on page 8, DOI: 10.1093/isd/ixz012, http://zenodo.org/record/716823

Fig. 3 in A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea)

Fig. 3. Combined molecular and morphological maximum likelihood tree.The different symbols correspond with specimens collected at the same location. Each specimen of these groups was collected during the same collecting event, with the exception of D3665, D3666, and D3791, collected at the same location but 2 wk apart.Published as part of Perry, Ryan K. & Heraty, John M., 2019, A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea), pp. 1-23 in Insect Systematics and Diversity (AIFB) 3 (5) on page 9, DOI: 10.1093/isd/ixz012, http://zenodo.org/record/716823

Fig. 5 in A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea)

Fig. 5. Mesosomal color variation. (A–H) Burkseus robustus: (A) D3674, (B) D3675, (C) D4591, (D) D4580, (E) D4581, (F) D4680, (G) D5326, (H) illustrated view. (I–L) B. sigillatus: (H) D5325, (I) D5324, (J) D4738, and (L) illustrated view. Scale bar = 0.2 mm.Published as part of Perry, Ryan K. & Heraty, John M., 2019, A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea), pp. 1-23 in Insect Systematics and Diversity (AIFB) 3 (5) on page 11, DOI: 10.1093/isd/ixz012, http://zenodo.org/record/716823

Fig. 8 in A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea)

Fig. 8. Gaster color variation within females (A–E) and males (F–J) of the four Nearctic Burkseus species.Published as part of Perry, Ryan K. & Heraty, John M., 2019, A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea), pp. 1-23 in Insect Systematics and Diversity (AIFB) 3 (5) on page 15, DOI: 10.1093/isd/ixz012, http://zenodo.org/record/716823

Fig. 9 in A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea)

Fig. 9. Habitus: (A) Burkseus elongatus (BMNH: NHMUK 10371836), (B) B. singa (MZH: UCRCENT 513243), (C) B. pinicolus (BMNH: NHMUK 10371840), (D) Cirrospilus curvineurus (MZH: UCRCENT 513242).Published as part of Perry, Ryan K. & Heraty, John M., 2019, A Tale of Two Setae: How Morphology and ITS2 Help Delimit a Cryptic Species Complex in Eulophidae (Hymenoptera: Chalcidoidea), pp. 1-23 in Insect Systematics and Diversity (AIFB) 3 (5) on page 16, DOI: 10.1093/isd/ixz012, http://zenodo.org/record/716823