Chalcidoidea (Hymenoptera) is extremely diverse with an estimated 500 000 species. We present the first phylogenetic analysis of the superfamily based on both morphological and molecular data. A web-based, systematics workbench mx was used to score 945 character states illustrated by 648 figures for 233 morphological characters for a total of 66 645 observations for 300 taxa. The matrix covers 22 chalcidoid families recognized herein and includes 268 genera within 78 of 83 subfamilies. Morphological data were analysed alone and in combination with molecular data from ribosomal 18S (2105 bp) and 28S D2-D5 expansion regions (1812 bp). Analyses were analysed alone and in combined datasets using implied-weights parsimony and likelihood. Proposed changes in higher classification resulting from the analyses include: (i) recognition of Eriaporidae, revised status; (ii) recognition of Cynipencyrtidae, revised status; (iii) recognition of Azotidae, revised status; (iv) inclusion of Sycophaginae in Agaonidae, revised status; (v) reclassification of Aphelinidae to include Aphelininae, Calesinae, Coccophaginae, Eretmocerinae and Eriaphytinae; (vi) inclusion of Cratominae and Panstenoninae within Pteromalinae (Pteromalidae), new synonymy; (vii) inclusion of Epichrysomallinae in Pteromalidae, revised status. At a higher level, Chalcidoidea was monophyletic, with Mymaridae the sister group of Rotoitidae plus the remaining Chalcidoidea. A eulophid lineage was recovered that included Aphelinidae, Azotidae, Eulophidae, Signiphoridae, Tetracampidae and Trichogrammatidae. Eucharitidae and Perilampidae were monophyletic if Eutrichosomatinae (Pteromalidae) was included, and Eupelmidae was monophyletic if Oodera (Pteromalidae: Cleonyminae) was included. Likelihood recovered a clade of Eupelmidae + (Tanaostigmatidae + (Cynipencyrtus + Encyrtidae). Support for other lineages and their impact on the classification of Chalcidoidea is discussed. Several life-history traits are mapped onto the new phylogeny.© The Willi Hennig Society 2013. Without question, Chalcidoidea is one of the most megadiverse groups of insects. Their morphological diversity is staggering (Fig. 1). They range in size from such veritable giants as females of Leptofoenus (Pteromalidae), which exceed 20 mm, to the minute and morphologically bizarre male of Dicopomorpha echmepterygis (Mymaridae), the smallest known specimen of which is 0.13 mm long. Males of D. echmepterygis have lost eyes, ocelli, mouthparts, antennal flagellum, wings, tarsi except for a highly modified arolium, and virtually any other feature that places them as parasitic wasps (Fig. 1a). Other bizarrities include male fig wasps, which can be reduced to turtle-like fighting machines that bear no resemblance to their corresponding females and are hardly recognizable as chalcidoids, or the grotesquely enlarged scutellum (Fig. 1h) of Galearia latreillei (Eucharitidae) and the dart-shaped ovipositor sheaths (Fig. 1j) of Cameronella (Pteromalidae). Convergent morphology is also rampant, and enlarged...
Chalcidoidea (Hymenoptera) are extremely diverse with more than 23,000 species described and over 500,000 species estimated to exist. This is the first comprehensive phylogenetic analysis of the superfamily based on a molecular analysis of 18S and 28S ribosomal gene regions for 19 families, 72 subfamilies, 343 genera and 649 species. The 56 outgroups are comprised of Ceraphronoidea and most proctotrupomorph families, including Mymarommatidae. Data alignment and the impact of ambiguous regions are explored using a secondary structure analysis and automated (MAFFT) alignments of the core and pairing regions and regions of ambiguous alignment. Both likelihood and parsimony approaches are used to analyze the data. Overall there is no impact of alignment method, and few but substantial differences between likelihood and parsimony approaches. Monophyly of Chalcidoidea and a sister group relationship between Mymaridae and the remaining Chalcidoidea is strongly supported in all analyses. Either Mymarommatoidea or Diaprioidea are the sister group of Chalcidoidea depending on the analysis. Likelihood analyses place Rotoitidae as the sister group of the remaining Chalcidoidea after Mymaridae, whereas parsimony nests them within Chalcidoidea. Some traditional family groups are supported as monophyletic (Agaonidae, Eucharitidae, Encyrtidae, Eulophidae, Leucospidae, Mymaridae, Ormyridae, Signiphoridae, Tanaostigmatidae and Trichogrammatidae). Several other families are paraphyletic (Perilampidae) or polyphyletic (Aphelinidae, Chalcididae, Eupelmidae, Eurytomidae, Pteromalidae, Tetracampidae and Torymidae). Evolutionary scenarios discussed for Chalcidoidea include the evolution of phytophagy, egg parasitism, sternorrhynchan parasitism, hypermetamorphic development and heteronomy.
Calesinae is a small group of Chalcidoidea (Hymenoptera) that are parasitoids of whiteflies (Hemiptera: Aleyrodidae). One species, Cales noacki Howard, has been introduced from South America into citrus‐growing regions of North America, the Mediterranean and Africa for biological control. The remaining species are found in Australia and New Zealand: a classic Gondwanan disjunction. The subfamily consists of a single genus, Cales, which is currently unplaced within Chalcidoidea. Its taxonomic position has historically been unstable, although most often Cales is associated with Aphelinidae. Here, we present a detailed morphological study of the group with an emphasis on Australian species. Although Cales shares many characteristics with Aphelinidae, especially Coccophaginae and Eretmocerus, more studies of character systems across Chalcidoidea are needed to determine which features may be synapomorphic. Consequently, we leave Cales incertae sedis within Chalcidoidea. We also describe a new species from New Zealand, Cales berryisp.n., reared from the whitefly Asterochiton pittospori on lemonwood, Pittosporum eugenioides, and we present a key and review the four known species of Cales.
The kudzu bug, Megacopta cribraria (F.) (Heteroptera: Plataspidae), is an invasive insect pest introduced from Asia in 2009 that poses a threat to soybeans (Glycine max [L.] Merr. [Fabales: Fabaceae]) and other legume crops in the United States. Initially discovered in Georgia, M. cribraria rapidly expanded across the southeast until 2014 when a significant decline in its population was observed across many locations. This notable decline in M. cribraria populations is attributed to the emergence of new parasitoids and pathogens in its new invasive range. So far, only a single egg parasitoid, Paratelenomus saccharalis (Dodd) (Hymenoptera: Platygastridae), is known to parasitize the eggs of M. cribraria in the United States. Here, we report a new egg parasitoid of M. cribraria identified as Ooencyrtus nezarae Ishii, 1928 (Hymenoptera: Encyrtidae), recovered from egg masses of M. cribraria collected from soybean in Alabama. O. nezarae is reported to parasitize eggs from a variety of heteropteran families and has been observed parasitizing M. cribraria in China. To the best of our knowledge, this is the first report of O. nezarae in North America. The potentials of O. nezarae for biological control of M. cribraria in the United States and the direction of future studies are discussed.
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