Life cycle closure of Cerroneuroterus minutulus (Giraud, 1859) with C. aggregatus (Wachtl, 1880) (Hymenoptera: Cynipidae: Cynipini): experimental, taxonomic and molecular approaches.

The life cycle of Cerroneuroterus minutulus (Giraud, 1859), a species previously known only from its asexual generation, is closed. Our study demonstrates that C. minutulus exhibits the heterogony typical of Cynipini species, with alternating sexual and asexual generations. The identity of the sexual generation is demonstrated by laboratory experiments and confirmed by molecular data, with Cerroneuroterus aggregatus (Wachtl, 1880) determined to be the sexual generation, and junior synonym, of Cerroneuroterus minutulus (Giraud, 1859). Illustrations and diagnoses for adults and galls, observations on biology and distribution information are provided as is a detailed redescription of the sexual generation of C. minutulus.

Holocynips illinoiensis, sp. nov., a new species of oak gall wasp (Hymenoptera: Cynipidae) from the USA.

A new species, Holocynips illinoiensis Melika & Nicholls, sp. nov. is described from the Nearctic (America north of Mexico). Description, diagnoses, information on biology and host association is given for the new species, as well as preliminary discussion on the coherence of the genus Holocynips.

Precipitation is the main axis of tropical plant phylogenetic turnover across space and time.

Early natural historians-Comte de Buffon, von Humboldt, and De Candolle-established environment and geography as two principal axes determining the distribution of groups of organisms, laying the foundations for biogeography over the subsequent 200 years, yet the relative importance of these two axes remains unresolved. Leveraging phylogenomic and global species distribution data for Mimosoid legumes, a pantropical plant clade of c. 3500 species, we show that the water availability gradient from deserts to rain forests dictates turnover of lineages within continents across the tropics. We demonstrate that 95% of speciation occurs within a precipitation niche, showing profound phylogenetic niche conservatism, and that lineage turnover boundaries coincide with isohyets of precipitation. We reveal similar patterns on different continents, implying that evolution and dispersal follow universal processes.

Diversity and divergence: evolution of secondary metabolism in the tropical tree genus Inga.

Plants are widely recognized as chemical factories, with each species producing dozens to hundreds of unique secondary metabolites. These compounds shape the interactions between plants and their natural enemies. We explore the evolutionary patterns and processes by which plants generate chemical diversity, from evolving novel compounds to unique chemical profiles. We characterized the chemical profile of one-third of the species of tropical rainforest trees in the genus Inga (c. 100, Fabaceae) using ultraperformance liquid chromatography-mass spectrometry-based metabolomics and applied phylogenetic comparative methods to understand the mode of chemical evolution. We show: each Inga species contain structurally unrelated compounds and high levels of phytochemical diversity; closely related species have divergent chemical profiles, with individual compounds, compound classes, and chemical profiles showing little-to-no phylogenetic signal; at the evolutionary time scale, a species' chemical profile shows a signature of divergent adaptation. At the ecological time scale, sympatric species were the most divergent, implying it is also advantageous to maintain a unique chemical profile from community members; finally, we integrate these patterns with a model for how chemical diversity evolves. Taken together, these results show that phytochemical diversity and divergence are fundamental to the ecology and evolution of plants.

Revision of the Amphibolips niger group with the description of a new species (Hymenoptera: Cynipini).

Amphibolips is currently divided into two species-groups, clearly differentiated by adult and gall morphology. The niger group of Amphibolips species is revised. This complex includes eight species: A. gumia Kinsey, A. jubatus Kinsey, A. elatus Kinsey, A. maturus Kinsey, A. nebris Kinsey, A. niger Kinsey, A. pistrix Kinsey and A. ufo Cuesta-Porta & Pujade-Villar sp. nov. We provide descriptions of new species and re-descriptions of known species, diagnoses, keys to complexes and species. The possibility of the niger complex belonging to a new genus is also discussed.

Re-establishment of the Nearctic oak cynipid gall wasp genus Feron Kinsey, 1937 (Hymenoptera: Cynipidae: Cynipini), including the description of six new species.

The Nearctic cynipid oak gall wasp genus Feron Kinsey, comb. rev., is re-established with 34 species: F. albicomus (Weld, 1952), comb. nov., F. amphorus (Weld, 1926), comb. nov., F. apiarium (Weld, 1944), comb. nov., F. atrimentum (Kinsey, 1922), comb. nov., F. bakkeri (Lyon, 1984), comb. nov., F. caepula (Weld, 1926), comb. nov., F. californicum (Beutenmueller, 1911), comb. nov., F. clarkei (Bassett, 1890), comb. nov., F. comatum (Weld, 1952), comb. nov., F. crystallinum (Bassett, 1900), comb. nov., F. cylindratum (Kinsey, 1937), comb. nov., F. discale (Weld, 1926), comb. nov., F. discularis (Weld, 1926), comb. nov., F. dumosae (Weld, 1957), comb. nov., F. gigas (Kinsey, 1922), comb. nov., F. izabellae Melika, Nicholls & Stone, sp. nov., F. kingi (Bassett, 1900), comb. nov., F. parmula (Bassett, 1900), comb. nov., F. pattersonae (Fullaway, 1911), comb. nov., F. roberti Melika, Nicholls & Stone, sp. nov., F. rucklei Melika, Nicholls & Stone, sp. nov., F. scutellum (Weld, 1930), comb. nov., F. serranoae Pujade-Villar & Cuesta-Porta, sp. nov., F. splendens (Weld, 1919), comb. nov., F. stellare (Weld, 1926), comb. nov., F. stellulum (Burnett, 1974), comb. nov., F. sulfureum (Weld, 1926), comb. nov., F. syndicorum Pujade-Villar & Cuesta-Porta, sp. nov., F. tecturnarum (Kinsey, 1920), comb. nov., F. tetyanae Melika, sp. nov., F. tibiale Kinsey, 1937, comb. rev., F. tubifaciens (Weld, 1926), comb. nov., F. verutum Kinsey, 1937, comb. rev., and F. vitreum Kinsey, 1937, comb. rev. Most species are known only from the asexual generation; F. clarkei, F. comatum, and F. dumosae are known only from the sexual generation, while both generations are recognised for F. atrimentum, F. crystallinum, F. gigas, F. kingi, and F. pattersonae. Matching of alternate sexual and asexual generations is established for the first time for F. kingi and F. pattersonae (= Andricus pedicellatus (Kinsey, 1922), syn. nov.) based on molecular data (both cytb and ITS2 sequences). Morphological descriptions, re-descriptions, diagnoses, and a key to species are given, as well as data on DNA sequences, biology, phenology, and distribution.

Re-establishment of the Nearctic oak cynipid gall wasp genus Druon Kinsey, 1937 (Hymenoptera: Cynipidae: Cynipini), with description of five new species.

The Nearctic cynipid oak gall wasp genus Druon Kinsey comb. rev. is re-established, with 5 new species and 10 species previously placed in the genus Andricus Hartig 1840: D. alexandri Melika, Nicholls Stone, sp. nov., D. flocculentum (Lyon), comb. nov., D. fullawayi (Beutenmller), comb. nov., D. garciamartinonae Pujade-Villar, sp. nov., D. gregori Melika, Nicholls Stone, sp. nov., D. hansoni Cuesta-Porta, Melika Pujade-Villar, sp. nov., D. ignotum (Bassett), comb. nov., D. linaria Kinsey, comb. rev., D. pattoni (Bassett), comb. nov., D. protagion Kinsey, comb. rev. (D. malinum Kinsey, syn. nov., D. polymorphae Kinsey, syn. nov.), D. quercusflocci (Walsh), comb. nov., D. quercuslanigerum (Ashmead), comb. nov., D. receptum Kinsey, comb. rev., D. rusticum Kinsey, comb. rev. and D. serretae Pujade-Villar, Cuesta-Porta Melika, sp. nov.. All species are known only from their asexual generation except for D. ignotum and D. quercuslanigerum, for which alternating asexual and sexual generations are known. The sexual generation of D. ignotum and a new morphological variety of D. quercuslanigerum from Mexico are both described for the first time. We provide descriptions, redescriptions, diagnoses, keys to species, DNA sequence data and analyses, and information on biology, phenology, and distribution. Additionally, four Andricus species with woolly galls and striato-reticulated metasomas are transferred to Striatoandricus Pujade-Villar: S. furnessulus, comb. nov., S. furnessae, comb. nov., S. guatemalensis, comb. nov., and S. tenuicornis, comb. nov.

A catalogue, revision, and regional perspective of Eastern Palaearctic and Oriental oak gall wasps and their inquilines (Hymenoptera: Cynipidae: Cynipini, Synergini, Ceroptresini).

Recent years have seen rapid advances in the study of Fagaceae-associated gall wasps (Hymenoptera: Cynipidae) of the Eastern Palaearctic and the Oriental (EPO) regions, for both the gall inducing Cynipini (commonly termed oak gall wasps though many species gall non-oak Fagaceae) and the predominantly inquiline tribes Synergini and Ceroptresini. This process has propagated some taxonomic errors and involves many taxa whose taxonomic status is uncertain. To provide a stable foundation for further advance, here we review the taxonomic status of the 212 species (133 oak gall wasps and 79 oak cynipid inquilines) that have been described or recorded in these regions. Of this total, we treat 171 as valid species names (103 oak gall wasps and 68 oak cynipid inquilines) in 20 genera; 22 as synonym names, 13 as incertae sedis, three as nomen dubium, and three as species inquirenda. Callirhytis kunugicola Shinji, 1944 is proposed as syn. nov. of C. kunugicola Shinji, 1943, as Shinji described the same species twice. For all valid species names, we provide taxonomic references, synonyms, and geographical distributions. We summarize what is known of host plant associations and gall locations for gall inducers, and host associations for inquilines. We discuss geographic patterns in the known richness of currently valid species for both groups of organisms, and place this in the context of the biogeography of available Fagaceae host plants. We provide a brief historical review of the study of oak gall wasps and their inquilines in the EPO regions.

Pairing of sexual and asexual generations of Nearctic oak gallwasps, with new synonyms and new species names (Hymenoptera: Cynipidae, Cynipini).

Most oak gallwasps (Hymenoptera; Cynipidae, Cynipini) have lifecycles involving obligate alternation between a sexual and an asexual generation. Many species are currently known from only one of these generations, with the alternate generation either unknown or separately described with a different name. Here we describe previously unknown generations for 12 Nearctic oak cynipids. We describe the sexual generation for ten species previously known only from an asexual generation: Acraspis quercushirta (Bassett, 1864), Acraspis villosa Gillette, 1888, Amphibolips spinosa Ashmead, 1887, Andricus balanaspis (Weld, 1922), comb. nov., Antron quercusnubila (Bassett, 1881), Bassettia pallida Ashmead, 1896, Neuroterus floccosus (Bassett, 1881), comb. rev., Philonix fulvicollis Fitch, 1859, Phylloteras poculum (Osten Sacken, 1862) and Sphaeroteras carolina (Ashmead, 1887), comb. rev.. We describe the asexual generation of Bassettia flavipes (Gillette, 1889), comb. nov. and Melikaiella tumifica (Osten Sacken, 1865), both previously known only from a sexual generation. Three described taxa (Neuroterus vernus Gillette, 1889, syn. nov., Neuroterus papillosus Beutenmueller, 1910, syn. nov. and Neuroterus perminimus Bassett, 1900, syn. nov.) are all identified as the sexual generation of Neuroterus niger Gillette, 1888. We also comment on the presence of previously unknown non-functional males reared from galls of the asexual generation of Neuroterus niger. We identify two further synonyms: Acraspis macrocarpae Bassett, 1890 is syn. nov. of A. quercushirta (Bassett, 1864), and Bassettia archboldi Melika Abrahamson, 2007 is syn. nov. of Bassettia pallida Ashmead, 1896. In addition, we formally synonymise multiple varieties described by Kinsey within his Acraspis quercushirta, A. villosa, Antron quercusnubila, Neuroterus niger and Philonix fulvicollis species complexes. We provide descriptions and diagnoses for these taxa, and summarise what is known of their biology, including host associations. All taxa and relationships between alternate generations are supported by morphological and molecular data.

A new genus of Nearctic oak gall wasp, Grahamstoneia Melika amp; Nicholls, gen. nov. (Hymenoptera: Cynipidae, Cynipini).

A new genus, Grahamstoneia Melika Nicholls, gen. nov., with one new species, G. humboldti Melika Nicholls, sp. nov., asexual generation, is described. This new taxon occurs in the south-western Nearctic, inducing galls on two species within Quercus section Protobalanus (Q. vacciniifolia Kellogg and Q. chrysolepis Liebm.), an ecology and distribution shared with the closely related genus Heteroecus Kinsey. Descriptions, diagnoses, biology, and host associations for the new genus and species are given. The new taxon is supported by morphological and molecular data.

Phylogenomics reveals accelerated late Cretaceous diversification of bee flies (Diptera: Bombyliidae).

Bombyliidae is a very species-rich and widespread family of parasitoid flies with more than 250 genera classified into 17 extant subfamilies. However, little is known about their evolutionary history or how their present-day diversity was shaped. Transcriptomes of 15 species and anchored hybrid enrichment (AHE) sequence captures of 86 species, representing 94 bee fly species and 14 subfamilies, were used to reconstruct the phylogeny of Bombyliidae. We integrated data from transcriptomes across each of the main lineages in our AHE tree to build a data set with more genes (550 loci versus 216 loci) and higher support levels. Our overall results show strong congruence with the current classification of the family, with 11 out of 14 included subfamilies recovered as monophyletic. Heterotropinae and Mythicomyiinae are successive sister groups to the remainder of the family. We examined the evolution of key morphological characters through our phylogenetic hypotheses and show that neither the "sand chamber subfamilies" nor the "Tomophthalmae" are monophyletic in our phylogenomic analyses. Based on our results, we reinstate two tribes at the subfamily level (Phthiriinae stat. rev. and Ecliminae stat. rev.) and we include the genus Sericosoma Macquart (previously incertae sedis) in the subfamily Oniromyiinae, bringing the total number of bee fly subfamilies to 19. Our dating analyses indicate a Jurassic origin of the family (165-194 Ma), with the sand chamber evolving early in bee fly evolution, in the late Jurassic or mid-Cretaceous (100-165 Ma). We hypothesize that the angiosperm radiation and the hothouse climate established during the late Cretaceous accelerated the diversification of bee flies, by providing an expanded range of resources for the parasitoid larvae and nectarivorous adults.

Three new Nearctic genera of oak cynipid gall wasps (Hymenoptera: Cynipidae: Cynipini): Burnettweldia Pujade-Villar, Melika amp; Nicholls, Nichollsiella Melika, Pujade-Villar amp; Stone, Disholandricus Melika, Pujade-Villar amp; Nicholls; and re-establishment of the genus Paracraspis Weld.

We describe three new genera of cynipid oak gall wasps from the Nearctic: Burnettweldia Pujade-Villar, Melika Nicholls, gen. nov., Nichollsiella Melika, Pujade-Villar Stone, gen. nov., and Disholandricus Melika, Pujade-Villar Nicholls, gen. nov. (Hymenoptera: Cynipidae: Cynipini). Burnettweldia includes five species, B. californicordazi Cuesta-Porta, Melika Pujade-Villar, sp. nov., B. conalis (Weld), comb. nov., B. corallina (Bassett), comb. nov., B. plumbella (Kinsey), comb. nov., B. washingtonensis (Gillette), comb. nov.. Nichollsiella includes three species, N. arizonica (Cockerell), comb. nov., N. sulcata (Ashmead), comb. nov., and N. puigi Melika, Cuesta-Porta Pujade-Villar, sp. nov.. Disholandricus includes four species, D. chrysolepidis (Ashmead), comb. nov., D. lasius (Ashmead), comb. nov., D. reniformis (McCracken Egbert), comb. nov., D. truckeensis (Ashmead), comb. nov. The genus Paracraspis Weld, comb. rev. is re-established with three species, P. guadaloupensis (Fullaway), P. insolens (Weld), and P. patelloides (Trotter). Descriptions, re-descriptions, diagnoses, keys to genera and species are given, including data on DNA sequences, biology, phenology and distribution.

Faecal metabarcoding reveals pervasive long-distance impacts of garden bird feeding.

Supplementary feeding of wildlife is widespread, being undertaken by more than half of households in many countries. However, the impact that these supplemental resources have is unclear, with impacts largely considered to be restricted to urban ecosystems. We reveal the pervasiveness of supplementary foodstuffs in the diet of a wild bird using metabarcoding of blue tit (Cyanistes caeruleus) faeces collected in early spring from a 220 km transect in Scotland with a large urbanization gradient. Supplementary foodstuffs were present in the majority of samples, with peanut (Arachis hypogaea) the single commonest (either natural or supplementary) dietary item. Consumption rates exhibited a distance decay from human habitation but remained high at several hundred metres from the nearest household and continued to our study limit of 1.4 km distant. Supplementary food consumption was associated with a near quadrupling of blue tit breeding density and a 5-day advancement of breeding phenology. We show that woodland bird species using supplementary food have increasing UK population trends, while species that do not, and/or are outcompeted by blue tits, are likely to be declining. We suggest that the impacts of supplementary feeding are larger and more spatially extensive than currently appreciated and could be disrupting population and ecosystem dynamics.

Re-description and systematic re-appraisal of the genus Kokkocynips Pujade-Villar amp; Melika, (Hymenoptera: Cynipidae: Cynipini), including new combinations of Nearctic species and the description of a new species from Panama.

The monophyly and taxonomic validity of some currently accepted genera of gall wasps in the Cynipini (Hymenoptera: Cynipidae) are being challenged by recent systematic studies. Here we used morphological and molecular data to re-describe and revise the taxonomic limits of the monotypic genus Kokkocynips Pujade-Villar Melika, previously recorded only from Mexico.                We describe a new species from Panama, Kokkocynips panamensis Medianero Nieves-Aldrey, reared from galls on Quercus salicifolia Neé, and add new records from Mexico for the type species of the genus K. doctorrosae Pujade-Villar. Six Nearctic species, Dryocosmus rileyi (Ashmead, 1896), D. imbricariae (Ashmead, 1896), D. coxii (Basset, 1881), D. deciduus (Beutenmueller, 1913), Callirhytis difficilis (Ashmead, 1887) and C. attractans (Kinsey, 1922) are transferred to Kokkocynips. Species of Kokkocynips are associated only with red oaks (Quercus section Lobatae (Fagaceae)) and are distributed in the Nearctic and Neotropical regions, from Canada through Mexico and Panama. Taxonomic limits of Kokkocynips are discussed in light of the molecular phylogenetic relationships of the studied species. Diagnostic characters, gall description, distribution, and biological data of Kokkocynips species are given, including a key for the identification of the asexual generations of seven species.

New species of Nearctic oak gall wasps (Hymenoptera: Cynipidae, Cynipini).

Twenty nine new species of cynipid oak gall wasps from the Nearctic region (America north of Mexico) are described: Andricus archboldi Melika Abrahamson, sp. nov., A. catalinensis Melika, Nicholls Stone, sp. nov., A. chapmanii Melika Abrahamson, sp. nov., A. chiricahuensis Melika, Nicholls Stone, sp. nov., A. coconinoensis Melika, Nicholls Stone, sp. nov., A. columbiensis Melika, Nicholls Stone, sp. nov., A. cooki Melika, Nicholls Stone, sp. nov., A. fitzpatricki Melika Abrahamson, sp. nov., A. highlandensis Melika, Nicholls Stone, sp. nov., A. mellificus Nicholls, Stone Melika, sp. nov., A. menkei Melika Abrahamson, sp. nov., A. mogollonensis Melika, Nicholls Stone, sp. nov., A. nichollsi Melika Stone, sp. nov., A. schickae Nicholls, Melika Stone, sp. nov., A. torreyaensis Melika Abrahamson, sp. nov., A. williami Melika, Nicholls Stone, sp. nov., Antron lovellae Melika, Nicholls Stone, sp. nov., A.tomkursari Melika, Nicholls Stone, sp. nov., Dryocosmus archboldi Melika Abrahamson, sp. nov., Loxaulus virginianae Melika Buss, sp. nov., Neuroterus alexandrae Nicholls Melika, sp. nov., N. aliceae Melika, Nicholls Stone, sp. nov., N. bussae Melika Nicholls, sp. nov., N. oblongifoliae Nicholls, Stone Melika, sp. nov., N. quaili Melika, Nicholls Stone, sp. nov., N. rosieae Melika, Nicholls Stone, sp. nov., N. stonei Melika Nicholls, sp. nov., Zapatella abrahamsoni Melika, sp. nov., Z. brooksvillei Melika Abrahamson, sp. nov.. Alternate asexual and sexual generations are described for four species, Andricus archboldi Melika Abrahamson, sp. nov., A. fitzpatricki Melika Abrahamson, sp. nov., A. schickae Nicholls, Melika Stone, sp. nov., Neuroterus aliceae Melika, Nicholls Stone, sp. nov.. Descriptions, diagnoses, plus information on biology and host associations are given for all new species. All taxa are supported by morphological data; matching of generations is established using DNA sequence data. We also demonstrate that Neuroterus niger var. alimas Kinsey should be considered as a nomen dubium.

A new genus of Neotropical oak gall wasp, Prokius Nieves-Aldrey, Medianero amp; Nicholls, gen. nov. (Hymenoptera: Cynipidae: Cynipini), with description of two new species from Panama.

A new genus, Prokius Nieves Aldrey, Medianero Nicholls, gen. nov., and two new species of oak gall wasps (Hymenoptera: Cynipidae: Cynipini), Prokius cambrai Medianero Nieves-Aldrey sp. nov. and Prokius lisethiae Medianero Nieves-Aldrey sp. nov., are described from adults reared from galls on Quercus bumelioides Liebm (Fagaceae, sect. Quercus, white oaks) collected in Panama. The new genus is phylogenetically and morphologically close to Dros Kinsey and forms part of a large clade that includes species from several other genera that appear to require revision, including Andricus Hartig and Phylloteras Ashmead. Molecular and morphological data, diagnostic characters, gall descriptions, distribution and biological data of the new genus and the new species are given. This new genus represents the fourth recently described genus of Cynipidae endemic to the Neotropical region.

Andricus forni Pujade-Villar Nicholls n. sp., a new species of oak gallwasp from China (Hymenoptera: Cynipidae).

A new sexual form of a gallwasp, Andricus forni Pujade-Villar Nicholls n. sp., is described from China (Zhejiang province) based on males, females and galls collected on Quercus serrata. Data on the morphology, diagnosis, distribution and biology of the new species are provided. Molecular data are also provided to support this species. A key to Asian Andricus gallwasp species with similar integral leaf galls is presented. Finally we discuss the taxonomic status of Asian Andricus making these integral leaf galls, including uncertainty on the validity of both Andricus quercicola and A. marmoratus.

Hybrid capture of 964 nuclear genes resolves evolutionary relationships in the mimosoid legumes and reveals the polytomous origins of a large pantropical radiation.

PREMISE: Targeted enrichment methods facilitate sequencing of hundreds of nuclear loci to enhance phylogenetic resolution and elucidate why some parts of the "tree of life" are difficult (if not impossible) to resolve. The mimosoid legumes are a prominent pantropical clade of ~3300 species of woody angiosperms for which previous phylogenies have shown extensive lack of resolution, especially among the species-rich and taxonomically challenging ingoids. METHODS: We generated transcriptomes to select low-copy nuclear genes, enrich these via hybrid capture for representative species of most mimosoid genera, and analyze the resulting data using de novo assembly and various phylogenomic tools for species tree inference. We also evaluate gene tree support and conflict for key internodes and use phylogenetic network analysis to investigate phylogenetic signal across the ingoids. RESULTS: Our selection of 964 nuclear genes greatly improves phylogenetic resolution across the mimosoid phylogeny and shows that the ingoid clade can be resolved into several well-supported clades. However, nearly all loci show lack of phylogenetic signal for some of the deeper internodes within the ingoids. CONCLUSIONS: Lack of resolution in the ingoid clade is most likely the result of hyperfast diversification, potentially causing a hard polytomy of six or seven lineages. The gene set for targeted sequencing presented here offers great potential to further enhance the phylogeny of mimosoids and the wider Caesalpinioideae with denser taxon sampling, to provide a framework for taxonomic reclassification, and to study the ingoid radiation.

Description of the sexual generation of Dryocosmus destefanii (Hymenoptera: Cynipidae: Cynipini) and disclosure of its life cycle.

The sexual generation of Dryocosmus destefanii Cerasa Melika, 2018 that emerges from galls on Q. suber L. in Italy is described for the first time, establishing its heterogonic life cycle. We provide observations on its distribution, illustration of adults and galls and information on its biology as supported by morphological and molecular data. An illustrated identification key to Western Palaearctic Dryocosmus species is also given.