Matthew L. Buffington

Divergence estimates and early evolutionary history of Figitidae (Hymenoptera: Cynipoidea)

We examine the phylogenetic relationships of Figitidae and discuss host use within this group in light of our own and previously published divergence time data. Our results suggest Figitidae, as currently defined, is not monophyletic. Furthermore, Mikeiinae and Pycnostigminae are sister‐groups, nested adjacent to Thrasorinae, Plectocynipinae and Euceroptrinae. The recovery of Pycnostigminae as sister‐group to Mikeiinae suggests two major patterns of evolution: (i) early Figitidae lineages demonstrate a Gondawanan origin (Plectocynipinae: Neotropical; Mikeiinae and Thrasorinae: Australia; Pycnostigminae: Africa); and (ii) based on host records for Mikeiinae, Thrasorinae and Plectocynipinae, Pycnostigminae are predicted to be parasitic on gall‐inducing Hymenoptera. The phylogenetic position of Parnips (Parnipinae) was unstable, and various analyses were conducted to determine the impact of this uncertainty on both the recovery of other clades and inferred divergence times; when Parnips was excluded from the total evidence analysis, Cynipidae was found to be sister‐group to [Euceroptrinae + (Plectocynipinae (Thrasorinae + (Mikeiinae + Pycnostigminae)))], with low support. Divergence dating analyses using BEAST indicate the stem‐group node of Figitidae to be c. 126 Ma; the dipteran parasitoids (Eucoilinae and Figitinae), were estimated to have a median age of 80 and 88 Ma, respectively; the neuropteran parasitoids (Anacharitinae), were estimated to have a median age of 97 Ma; sternorrhynchan hyperparasitoids (Charipinae), were estimated to have a median age of 110 Ma; the Hymenoptera‐parasitic subfamilies (Euceroptinae, Plectocynipinae, Trasorinae, Mikeiinae, Pycnostigminae, and Parnipinae), ranged in median ages from 48 to 108 Ma. Rapid radiation of Eucoilinae subclades appears chronologically synchronized with the origin of their hosts, Schizophora (Diptera). Overall, the exclusion of Parnips from the BEAST analysis did not result in significant changes to divergence estimates. Finally, though sparsely represented in the analysis, our data suggest Cynipidae have a median age of 54 Ma, which is somewhat older than the age of Quercus spp (30–50 Ma), their most common host.

Phylogeny, Evolution and Classification of Gall Wasps: The Plot Thickens

Gall wasps (Cynipidae) represent the most spectacular radiation of gall-inducing insects. In addition to true gall formers, gall wasps also include phytophagous inquilines, which live inside the galls induced by gall wasps or other insects. Here we present the first comprehensive molecular and total-evidence analyses of higher-level gall wasp relationships. We studied more than 100 taxa representing a rich selection of outgroups and the majority of described cynipid genera outside the diverse oak gall wasps (Cynipini), which were more sparsely sampled. About 5 kb of nucleotide data from one mitochondrial (COI) and four nuclear (28S, LWRh, EF1alpha F1, and EF1alpha F2) markers were analyzed separately and in combination with morphological and life-history data. According to previous morphology-based studies, gall wasps evolved in the Northern Hemisphere and were initially herb gallers. Inquilines originated once from gall inducers that lost the ability to initiate galls. Our results, albeit not conclusive, suggest a different scenario. The first gall wasps were more likely associated with woody host plants, and there must have been multiple origins of gall inducers, inquilines or both. One possibility is that gall inducers arose independently from inquilines in several lineages. Except for these surprising results, our analyses are largely consistent with previous studies. They confirm that gall wasps are conservative in their host-plant preferences, and that herb-galling lineages have radiated repeatedly onto the same set of unrelated host plants. We propose a revised classification of the family into twelve tribes, which are strongly supported as monophyletic across independent datasets. Four are new: Aulacideini, Phanacidini, Diastrophini and Ceroptresini. We present a key to the tribes and discuss their morphological and biological diversity. Until the relationships among the tribes are resolved, the origin and early evolution of gall wasps will remain elusive.

Description, circumscription and phylogenetics of the new tribe Zaeucoilini (Hymenoptera: Figitidae: Eucoilinae), including a description of a new genus

Abstract Neotropical eucoiline genera that have been included in and allied with the Zaeucoila genus group are redescribed. Following character analysis and phylogenetic reconstruction (25 taxa, 96 morphological characters, 1452 ribosomal and mitochondrial characters), this informal genus group was found to be monophyletic, and hence raised to formal recognition as a tribe of eucoilines, namely Zaeucoilini new tribe. Through phylogenetic reconstruction, Aegeseucoela Buffington was determined to be polyphyletic; the type species of Aegeseucoela, A. grenadensis (Ashmead), is transferred to Agrostocynipscomb.n.; the orphaned species, A. flavotincta (Kieffer), is transferred to Marthiella Buffington gen.n., comb.n. Based on the examination of the holotype of Diranchis flavipes Ashmead, 1900 , this species is transferred to Rhabdeucoelacomb.n., where it is both a junior, subjective synonym of flavipes ( Ashmead, 1894 ) syn.n. and a junior, secondary homonym. Zaeucoilini contains the following genera: Agrostocynips Diaz, Dettmeria Borgmeier, Dicerataspis Ashmead, Lopheucoila Weld, Marthiella Buffington, Moneucoela Kieffer, Moritiella Buffington, Penteucoila Weld, Preseucoela Buffington, Rhabdeucoela Kieffer, Tropideucoila Ashmead and Zaeucoila Ashmead. Characters and character states applicable specifically to Zaeucoilini are defined and illustrated. Characters supporting the monophyly of each genus are discussed. The plesiomorphic host for members of the Zaeucoilini are postulated as agromyzid leaf‐mining Diptera, with one shift within the clade containing Dettmeria, Dicerataspis and Lopheucoila to fruit‐infesting Diptera. New host records are reported for species of Preseucoela, Rhabdeucoela and Zaeucoila. All known hosts for the species of each genus are reported, as are known distributions and locations of type specimens.

Biodiversity into your hands - A call for a virtual global natural history 'metacollection'

BackgroundMany scientific disciplines rely on correct taxon delineations and identifications. So does a great part of the general public as well as decision makers. Researchers, students and enthusiastic amateurs often feel frustrated because information about species remains scattered, difficult to access, or difficult to decipher. Together, this affects almost anyone who wishes to identify species or verify identifications. Many remedies have been proposed, but we argue that the role of natural history collections remains insufficiently appreciated. We suggest using state-of-the-art mass imaging technology and to join forces to create a global natural history metacollection on the internet, providing access to the morphology of tens of millions of specimens and making them available for automated digital image analysis.DiscussionRobotic high-resolution imaging technology and fast (high performance) computer-based image stitching make it now feasible to digitize entire collection drawers typically used for arthropod collections, or trays or containers used for other objects. Resolutions of 500 megapixels and much higher are already utilized to capture the contents of 40x50 cm collection drawers, providing amazing detail of specimens. Flanked by metadata entry, this helps to create access to tens of thousands of specimens in days. By setting priorities and combining the holdings of the most comprehensive collections for certain taxa, drawer digitizing offers the unique opportunity to create a global, virtual metacollection.The taxonomic and geographic coverage of such a collection could never be achieved by a single institution or individual. We argue that by joining forces, many new impulses will emerge for systematic biology, related fields and understanding of biodiversity in general.Digitizing drawers containing unidentified, little-curated specimens is a contribution towards the beginning of a new era of online curation. It also will help taxonomists and curators to discover and process the millions of “gems” of undescribed species hidden in museum accessions.SummaryOur proposal suggests creating virtual, high-resolution image resources that will, for the first time in history, provide access for expert scientists as well as students and the general public to the enormous wealth of the world’s natural history collections. We foresee that this will contribute to a better understanding, appreciation and increased use of biodiversity resources and the natural history collections serving this cause.

Key to Nearctic species of Trissolcus Ashmead (Hymenoptera, Scelionidae), natural enemies of native and invasive stink bugs (Hemiptera, Pentatomidae)

Trissolcus japonicus (Ashmead) and T. cultratus (Mayr), comb. rev. are under study as classical biological agents to control the brown marmorated stink bug Halyomorpha halys (Stål) in North America. Here we present diagnoses for all Nearctic species of Trissolcus, including T. japonicus and T. cultratus comb. rev., and identification keys to enable separation of these species from the existing fauna. Trissolcus cultratus comb. rev. is removed from synonymy with T. flavipes. Two new species are described, Trissolcus valkyria sp. n. and T. zakotos sp. n. A neotype is designated for T. brochymenae and a lectotype is designated for T. basalis.

Systematics of Australian Thrasorinae (Hymenoptera, Cynipoidea, Figitidae) with descriptions of Mikeiinae, new subfamily, two new genera, and three new species

Abstract The Australian Thrasorinae are revised and Mikeius is transferred to Mikeiinae Paretas-Martínez & Pujade-Villar, subfam. n., and Mikeius clavatus Pujade-Villar & Restrepo-Ortiz, sp. n., is described. Two new genera of Thrasorinae are erected: Cicatrix Paretas-Martínez, gen. n., including Cicatrix pilosiscutum(Girault), comb. n. from Amblynotus, Cicatrix schauffi (Buffington), comb. n. from Mikeius, and Cicatrix neumannoides Paretas-Martínez & Restrepo-Ortiz, sp. n.; and Palmiriella Pujade-Villar & Paretas-Martínez, gen. n., including Palmiriella neumanni (Buffington), comb. n. from Mikeius, Thrasorus rieki Paretas-Martínez & Pujade-Villar, sp. n., is also described. A phylogenetic analysis of 176 morphological and biological characters, including all these new taxa and all genera previously included in Thrasorinae, was conducted. All subfamilies were recovered as monophyletic, with the following relationships: Parnipinae (Euceroptrinae (Mikeiinae (Plectocynipinae (Thrasorinae)))). A worldwide key to the subfamilies of Figitidae is provided that includes the new subfamily, as well as a key to genera Thrasorinae.

Not All Oak Gall Wasps Gall Oaks: The Description of Dryocosmus rileypokei, A New, Apostate Species of Cynipini from California

Abstract Cynipini gall wasps (Hymenoptera: Cynipidae) are commonly known as oak gall wasps for their almost exclusive use of oak (Quercus spp.; Fagaceae) as their host plant. Previously, only three of the nearly 1,000 species of Cynipini have been recorded from hosts other than Quercus. These three are known from western chinquapin (Chrysolepis), chestnut (Castanea) and tan bark oak (Lithocarpus), all lineages of Fagaceae related to Quercus. Here we describe Dryocosmus rileypokei Morita & Buffington, new species, a second species of cynipine which attacks Chrysolepis. Unlike the previously known gall wasp D. castanopsidis, which produces a medium-sized spherical external gall near the base of the staminate (male) flowers of Chrysolepis sempervirens, D. rileypokei attacks the same host acting as a nut galler. Dryocosmus rileypokei creates a gall within the mesocarp wall of the nut and appears to draw nutrients away from the developing seed. Later instar larvae and teneral adults were found within these internal galls. It appears that the adult wasp eventually chews an exit hole from these galleries. The evolution of host use in the three, non-oak galling Dryocosmus species is discussed.

Genetic differentiation of Ganaspis brasiliensis (Hymenoptera: Figitidae) from East and Southeast Asia

Ganaspis brasiliensis (Ihering) (Hymenoptera: Figitidae: Eucoilinae) is a Drosophila parasitoid that has often been misidentified as G. xanthopoda (Ashmead) in recent studies. This study aims to clarify genetic differentiation of G. brasiliensis based on the nucleotide sequences of the mitochondrial cytochrome oxidase subunit 1 (CO1) gene and three nuclear DNA regions, the inter-transcribed spacers 1 and 2 (ITS1 and ITS2) and putative 60S ribosomal protein L37 (RpL37), as well as crossing experiments. Four lineages are recognized in individuals assigned as G. basiliensis by morphology, (1) individuals occurring in Japan and probably South Korea, (2) individuals from a small subtropical island of Japan, Iriomote-jima, (3) individuals from temperate lowlands of Japan and high altitude areas of Southeast Asia, and (4) individuals occurring widely in Asia, America, Hawaii and Africa. The first lineage is a specialist of Drosophila suzukii (Matsumura), a pest of fresh fruit, and also the fourth lineage has a capacity to parasitize this pest species. The first, third and fourth lineages occur sympatrically at least in Tokyo. The third and fourth lineages differed in mate choice and host use to some extent, but post-mating isolation between them was almost absent.

Revision of Plectocynipinae (Hymenoptera: Figitidae) with Descriptions of A New Genus and Three New Species from Chile.

Abstract. A revision of Plectocynipinae was initiated following the recent collections of specimens of Plectocynips Diaz and Araucocynips, new genus, from Chile. In addition to a redescription of the subfamily, we provide redescriptions, based on females, of the species of both genera. Descriptions are provided for Araucocynips ariasae, new species, Araucocynips queulensis, new species, and Plectocynips lago, new species, all from Chile. New host records are provided for Plectocynips pilosus Ros-Farré from galls of Aditrochus fagicolus Rübsaamen and Aditrochus coihuensis Ovruski (Hymenoptera: Chalcidoidea) on Nothofagus pumilio (Poepp. & Endl.) Krasser and Nothofagus dombeyi (Mirb.), respectively. In addition, males are described for Araucocynips ariasae, Araucocynips queulensis, and Plectocynips pilosus.

The occurrence and phylogenetic implications of wing interference patterns in Cynipoidea (Insecta:Hymenoptera)

Wing interference patterns (WIPs) are a potentially rich source of taxonomic data. We surveyed a broad range of Cynipoidea (Hymenoptera) for the presence of these patterns. Further, we used phylogeny to investigate the evolutionary patterns of WIPs throughout the superfamily. We also documented that WIPs occur across nearly all species sampled, in both the hindwing and forewing; in many instances, the forewing WIP is carried over to the hindwing. Only Austrocynips mirabilis Riek lacked WIPs. Among species that possess WIPs, we were able to recognise four distinct categories: radiform (radiating), striatiform (longitudinally striate), campiform (continuous field) and galactiform (amorphous mixture of colours). Wing interference patterns are not found in infuscate wings or areas of infuscation on otherwise hyaline wings, and are reduced on large wings. Within Figitidae, WIPs are more phylogenetically conservative, especially at the tribal level: within Cynipidae the phylogenetic conservatism of WIPs is somewhat less clear, though all Synergini sampled had a very consistent WIPs. For certain genera, such as Ganaspidium and Andricus, WIPs may prove useful for species-level discrimination.