No full-text available
To read the full-text of this research,
you can request a copy directly from the authors.
A New Genus of Oak Gall Wasp, Striatoandricus Pujade-Villar (Hymenoptera: Cynipidae: Cynipini) from America with Descriptions of Two New Mexican Species
Abstract
A new genus of cynipid oak gall wasp, Striatoandricus Pujade-Villar (Hymenoptera: Cynipidae: Cynipini), is described. Striatoandricus gen. nov. includes four previously described species, Andricus nievesaldreyi n. comb., A. georgei n. comb., A. maesi n. comb., and A. barriosi n. comb., which induce pubescent leaves or twig galls on Quercus belonging to Quercus section. Two new species from México are also described: S. cuixarti Pujade-Villar n. sp. and S. sanchezi Pujade-Villar n. sp. in Quercus section. Descriptions of the genus and diagnostic characters, including DNA sequence data, are presented. This new genus is supported by both morphological and molecular data.
... As part of this ongoing revisionary process, some of the previously synonymized genera have been re-established: Erythres (Pujade-Villar & Melika 2014), Femuros (Pujade-Villar & Ferrer-Suay 2015), Dros (Pujade-Villar et al. 2017) and Trichoteras (Zimmermann 2018). New genera have also been established for species erroneously placed in Andricus: Protobalandricus Melika, Nicholls & Stone, 2018(Nicholls et al. 2018c, Striatoandricus Pujade-Villar, 2020 (Cuesta-Porta et al. 2020) and Disholandricus Melika, Pujade-Villar & Nicholls, 2021(Melika et al. 2021b). However, the status of many Nearctic and Neotropical Andricus species needs to be reviewed. ...
... (Bassett, 1881) Diagnosis. Morphologically most closely resembles S. barriosi (Medianero & Nieves Aldrey, 2019) known from Panama, both having the head and mesosoma dark brown to black, the metasoma reddish brown; the notaulus complete, entirely reticulate pronotum, second metasomal tergum entirely striate, not areolate-reticulate, without smooth dorsolateral area; the mesoscutum longer than broad; mesopleuron entirely and uniformly reticulate, without smooth areas; Rs+M inconspicuous, not reaching basalis, its projection reaching basalis in its lower half (Cuesta-Porta et al. 2020). But in S. barriosi the dorsal part of the mesopleuron is smooth, while in S. tenuicornis the entire mesopleuron is reticulate, except a small part of the speculum which is smooth, shining. ...
... Species in the genus Striatoandricus, which is sister to Druon, also induce woolly clustered galls, either on leaves or twigs. For example, S. tenuicornis from Arizona (transferred herein) and several Striatoandricus species from Mexico and Central America induce woolly leaf galls (Pujade-Villar et al. 2011, Medianero & Nieves-Aldrey 2019, Cuesta-Porta et al. 2020Fig. 242). ...
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 (Beutenmüller), 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.
... However, in recent years, significant new faunas of oak gall wasps have been discovered in the Neotropical and Oriental regions, leading to the description or re-establishment of 17 genera. Eight of the recently described genera belong to the American fauna; two are endemic to Central America, Coffeikokkos Pujade-Villar & Melika (Pujade-Villar et al. 2012b) and Barucynips Medianero & Nieves-Aldrey (Medianero & Nieves-Aldrey 2013); one, Protobalandricus Melika, Nicholls & Stone, is known only from California (Nicholls et al. 2018b); and five genera are more widely distributed in North America and Central America: Kinseyella Pujade-Villar & Melika , Zapatella Pujade-Villar & Melika (Pujade-Villar et al. 2012a), Kokkocynips Pujade-Villar & Melika (Pujade-Villar et al. 2013), Melikaiella Pujade-Villar (Pujade-Villar et al. 2014) and Striatoandricus Pujade-Villar (Cuesta-Porta et al. 2020). ...
... nievesaldreyi Pujade-Villar, 2011), leading to comparison of Kokkocynips with this group. However, as mentioned in Pujade-Villar et al. (2013), these species have toothed claws, are associated with white oaks and have recently been moved into their own new genus Striatoandricus (Cuesta-Porta et al. 2020). In fact, Kokkocynips is placed in a very different part of the phylogeny to Striatoandricus (see Fig. 1), implying that the distinct sculpturing on the metasomal tergites shared by these genera has been independently derived. ...
... Studies over the last decade have revealed new morphological diversity in Neotropical oak-associated cynipids; around 100 species of Nearctic and Neotropical Cynipidae have been described or revised, primarily associated with red oaks (Quercus section Lobatae) and white oaks (Quercus section Quercus; see for example Pujade-Villar et al. 2010, Pujade-Villar et al. 2012a, Pujade-Villar et al. 2012b, Pujade-Villar et al. 2013, Medianero & Nieves-Aldrey 2013, Pujade-Villar et al. 2014, Nicholls et al. 2018b, Cuesta-Porta et al. 2020. Incorporating this novel morphological diversity into the existing generic level classification has been a challenge because some taxa show combinations of characters otherwise diagnostic of different genera. ...
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.
... In addition, both Weld (1952) and Melika & Abrahamson (2002) have synonymized multiple genera into Andricus, rendering it polyphyletic and difficult to classify. In recent years, a significant effort was undertaken by various cynipid taxonomists to rectify this chaotic genus, which resulted in the resurrection of erroneous synonymizations (Pujade-Villar & Melika 2014, Pujade-Villar & Ferrer-Suay 2015, Pujade-Villar et al. 2017, Zimmermann 2018, and the establishment of new genera (Nicholls et al. 2018, Cuesta-Porta et al. 2020, Melika et al. 2021b. The majority of recent taxonomic studies of Cynipini in the US have focused on southern regions such as Arizona, California, Florida, and Texas, where there are higher diversities of oak species (Brandão-Dias et al. 2022, Cuesta-Porta et al. 2022, Cooke-McEwen & Gates 2020, Melika & Nicholls 2021, Melika et al. 2021a, Melika et al. 2021b, Nicholls et al. 2018, Zhang et al. 2021. ...
... Druon now contains a total of 15 species from USA, Mexico, and Costa Rica, forming galls on the leaves of oaks in the sections Quercus and Virentes (Kinsey 1937, Cuesta-Porta et al. 2022. Striatoandricus Pujade-Villar, 2020 is a newly described genus that includes nine species found in Mexico, Panama, and isolated mountain ranges in Arizona, USA (Cuesta-Porta et al. 2020. Druon can be distinguished from the morphologically similar Striatoandricus as the metasoma of the latter is matte, and tergites striate and/or reticulate instead of shiny (Cuesta-Porta et al. 2022). ...
A new species of the genus Druon Kinsey, 1937, D. laceyi Zhang, Sasan & O'Kennon sp. nov. is described on host plant Quercus laceyi Small from central Texas. We also re-establish Andricus lustrans Beutenmüller, 1913 comb.rev., and transfer Striatoandricus aciculatus (Beutenmüller, 1909) comb. nov. from Andricus. Finally, we report a new state and host record for Druon gregori Melika, Nicholls & Stone, 2022. All observations were first shared on the social platform iNaturalist, highlighting the potential of cybertaxonomy in uncovering overlooked biodiversity.
... The most recent classification divides the family Cynipidae into 12 tribes: Aylacini, Aulacideini, Ceroptresini, Cynipini, Diastrophini, Diplolepidini, Eschatocerini, Paraulacini, Pediaspidini, Phanacidini, Qwaqwaiini and Synergini (Ronquist et al. 2015). Oak gall wasps (Cynipini) are by far the most species-rich group of gall wasps, with more than 1,000 known species in 50 genera worldwide (Csóka et al. 2005, Ronquist et al. 2015, Nicholls et al 2018a, Cuesta-Porta et al. 2020, Fang et al 2020, Nieves-Aldrey et al. 2021. Between 90 and 95% of described Cynipini species induce galls on the genus Quercus L. (Fagaceae; Csóka et al. 2005), while the remainder induce galls on other genera in the family Fagaceae (Castanea, Castanopsis, Chrysolepis, Lithocarpus and Notholithocarpus) (Nieves-Aldrey et al. 2009, Tang et al. 2016, Nicholls et al. 2018b. ...
... In their review, Medianero & Nieves-Aldrey (2011) listed eight genera of native Cynipini from this region: Acraspis Mayr, 1881, Amphibolips Reinhard, 1865, Andricus Hartig, 1840, Atrusca Kinsey, 1930, Bassettia Ashmead, 1887, Disholcaspis Dalla Torre & Kieffer, 1910, Loxaulus Mayr, 1881and Odontocynips Kieffer, 1910. Since then, species from seven more genera have been recorded: Coffeikokkos Pujade-Villar & Melika, 2012(Pujade-Villar et al. 2012b, Medianero & Nieves-Aldrey 2013, Barucynips Medianero & Nieves-Aldrey, 2013 (Medianero & Nieves-Aldrey 2013), Striatoandricus Pujade-Villar, 2020 (Medianero & Nieves-Aldrey 2019, Cuesta-Porta et al. 2020, Melikaiella Pujade-Villar, 2014 (Medianero & Nieves-Aldrey 2014), Neuroterus Hartig,1840 (Medianero & Nieves-Aldrey 2017), Zapatella Pujade-Villar & Melika, 2012(Pujade-Villar et al. 2012a and Kokkocynips Pujade-Villar & Melika, 2013(Nieves-Aldrey et al. 2021; this brings the total number of genera of oak gall wasps currently known from the Neotropical region to 15. However, our sampling has revealed further taxa from this region that are morphologically and genetically distinct from known Cynipini genera and hence require formal description. ...
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.
... Synergus citriformis is considered a generalist species, even though it is mainly associated with galls of Amphibolips on oaks within the Lobatae section. Found also in association with woolly galls initiated by Andricus, Striatoandricus (arisen by Cuesta-Porta et al. (2020) to include some previously known species within Andricus) and Cynips, and with other gall morphotypes (never tuberous galls) initiated by Andricus, Atrusca, Cynips, Disholcaspis, Dros, and Loxaulus on oaks of both Quercus and Lobatae sections Lobato-Vila et al. 2020b, 2020c. ...
... Mexico is one of the richest regions of oak diversity along with Southeast Asia and harbors a great number of endemic Quercus species (Nixon 1998;Zavala 1998;Valencia 2004). A moderate assessment of the Mexican oak gall wasp fauna during the last decade revealed a great diversity of gall-makers and inquilines in this country, which were subsequently described or recorded within a number of publications (Melika et al. , 2011aPujade-Villar & Paretas-Martínez 2012;Pujade-Villar et al. 2009-b, 2012a, 2013a-b, 2014a-d, 2015b, 2019, 2019, 2020bCuesta-Porta et al. 2020). Therefore, more exhaustive fieldwork in the Nearctic region as well as in some Neotropical border countries, in combination with a detailed study of Kinsey's collection, will undoubtfully reveal further new species, especially within Synergus. ...
A complete revision of the genus Synergus Hartig (Hymenoptera: Cynipidae: Synergini) in the New World (Nearctic and Neotropical regions) is conducted for the first time in order to stabilize its taxonomy. A total of 11 new species are described from Mexico: S. ashmeadi Lobato-Vila & Pujade-Villar, sp. nov.; S. beutenmulleri Lobato-Vila & Pujade-Villar, sp. nov.; S. compressus Lobato-Vila & Pujade-Villar, sp. nov.; S. diversicolor Lobato-Vila & Pujade-Villar, sp. nov.; S. ebenus Lobato-Vila & Pujade-Villar, sp. nov.; S. linnei Lobato-Vila & Pujade-Villar, sp. nov.; S. macrackenae Lobato-Vila & Pujade-Villar, sp. nov.; S. oaxaquensis Lobato-Vila & Pujade-Villar, sp. nov.; S. personatus Lobato-Vila & Pujade-Villar, sp. nov.; S. ruficephalus Lobato-Vila & Pujade-Villar, sp. nov.; and S. weldi Lobato-Vila & Pujade-Villar, sp. nov. Synergus splendidus Fullaway, 1911 is proposed as a syn. nov. of S. dorsalis (Provacher, 1889); S. garryana Gillette, 1893 and S. oneratus coloradensis Gillette, 1896 as syn. nov. of S. oneratus (Harris, 1841); and S. magnificus Weld, 1957 as a syn. nov. of S. reniformis McCracken & Egbert, 1922. Redescriptions and illustrations are provided for poorly characterized species. A key to species and a summary table including all valid Synergus from the New World, their biology and distribution, are given. Distribution, morphology, and trophic associations are discussed and compared between New World and Palaearctic species.
... As a result, several species of in Neotropical oak gall wasp have been newly described belonging to the genera: Amphibolips Reinhard, Andricus Hartig, Bassettia Ashmead, Disholcaspis Dalla Torre Kieffer, Loxaulus Mayr, Melikaiella Pujade-Villar, Neuroterus Hartig, Odontocynips Kieffer and Striatoandricus Pujade-Villar, (e.g. Cuesta-Porta et al. 2020;Fernández-Garzón et al. 2017;Medianero and Nieves-Aldrey 2010, 2014Melika et al. 2009Melika et al. , 2011Nieves-Aldrey and Medianero 2010;Nieves-Aldrey et al. 2021;Pujade-Villar 2008;, 2017. New endemic genera have also been described: Barucynips Medianero and Nieves-Aldrey, Coffeikokkos Pujade-Villar and Melika, and Zapatella Pujade-Villar and Melika (Medianero and Nieves-Aldrey 2013;Pujade-Villar et al. 2012a, 2012b. ...
A new species of oak gallwasp, Andricus partali n. sp. (Hym., Cynipidae: Cynipini), is described from Costa Rica inducing galls on the leaves of Q. costaricensis Liebm. (Section Lobatae). We provide a description, diagnosis, and information about the distribution and biology of the new species.
... The first modern review of genera was provided by Melika & Abrahamson (2002), with some erroneous synonymizations of genera. Since then six Nearctic genera have been re-established: Sphaeroteras Ashmead, 1897, Trichoteras Ashmead, 1897, Dros Kinsey, 1937, Erythres Kinsey, 1937, Femuros Kinsey, 1937and Paracraspis Weld, 1952; while 12 new genera have been described from the Nearctic and Neotropics: Kinseyella Pujade-Villar & , Coffeikokkos Pujade-Villar & Melika, 2012, Zapatella Pujade-Villar & Melika, 2012, Barucynips Medianero & Nieves-Aldrey, 2013, Kokkocynips Pujade-Villar & Melika, 2013, Melikaiella Pujade-Villar, 2014, Protobalandricus Melika, Nicholls & Stone, 2018, Striatoandricus Pujade-Villar, 2020, Burnettweldia Pujade-Villar, , Nichollsiella Melika, Pujade-Villar & Stone, 2021, Disholandricus Melika, Pujade-Villar & Nicholls, 2021and Grahamstoneia Melika & Nicholls, 2021(Cuesta-Porta et al. 2020Medianero & Nieves-Aldrey 2013, Nicholls et al. 2018cPujade-Villar & Ferrer-Suay 2015, Pujade-Villar & Melika 2014, 2012a,b, 2013, 2017bZimmerman 2018). In addition, four new genera have been described from the Palaearctic and the Oriental regions: Cerroneuroterus , Cycloneuroterus Melika & Tang, 2011, Cyclocynips Melika, Tang & Sinclair, 2013and Heocynips Fang, Nieves-Aldrey & Melika, 2020(Fang et al. 2020Melika et al. 2010Melika et al. , 2013bTang et al. 2011). ...
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.
... The first modern review of genera was provided by Melika & Abrahamson (2002), with some erroneous synonymizations of genera. Since then, four Nearctic genera have been re-established (Sphaeroteras Ashmead, 1897, Dros Kinsey, 1937, Erythres Kinsey, 1937, and Femuros Kinsey, 1937, eight new genera with new species have been described or re-defined from the Nearctic and Neotropics (Kinseyella Pujade-Villar & Melika, 2010, Coffeikokkos Pujade-Villar & Melika, 2012, Zapatella Pujade-Villar & Melika, 2012, Barucynips Medianero & Nieves-Aldrey, 2013, Kokkocynips Pujade-Villar & Melika, 2013, Melikaiella Pujade-Villar, 2014, Protobalandricus Melika, Nicholls & Stone, 2018, Striatoandricus Pujade-Villar, 2020, and four new genera have been described from the Palaearctic and the Oriental regions (Cerroneuroterus , Cycloneuroterus Melika & Tang, 2011, Cyclocynips Melika, Tang & Sinclair, 2013and Heocynips Fang, Nieves-Aldrey & Melika, 2020 (Cuesta-Porta et al. 2020;Fang et al. 2020;Medianero & Nieves-Aldrey 2013;Melika et al. 2010Melika et al. , 2013Nicholls et al. 2018b;Nieves-Aldrey et al. 2021;Pujade-Villar & Ferrer-Suay 2015;Pujade-Villar & Melika 2014;Pujade-Villar et al. 2010, 2012a,b, 2013, 2018Tang et al. 2011). Further revision of genera and description of new species is ongoing, especially in relation to the Nearctic gall wasp fauna. ...
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.
... USA (states of Florida and California) (Ashmead 1885; Fullaway 1911; McCracken & Egbert 1922), Panama (Nieves-Aldrey & Medianero 2011) and Mexico (states of Aguascalientes, Ciudad de México, Guanajuato, Hidalgo, México, Michoacán, Morelos, Oaxaca, Puebla, Tlaxcala, Veracruz and Zacatecas) (Lobato-Vila & Pujade-Villar 2017; Lobato-Vila et al. 2020b; and in this work, see the material examined).Biology. Mainly associated with galls of Amphibolips on oaks of the Lobatae section, but also with woolly galls initiated by Andricus, Striatoandricus (a new Cynipini genus raised byCuesta-Porta et al. (2020) to include some previously known species within Andricus), and Cynips, and with other gall morphotypes (never tuberous galls) initiated by Andricus, Atrusca, Cynips, Disholcaspis, Dros and Loxaulus on oaks of both Quercus andLobatae sections (Ashmead 1885; McCracken & Egbert 1922; Nieves-Aldrey & Medianero 2011; Lobato-Vila & Pujade-Villar 2017; Lobato-Vila et al. 2020b; and in this work, see the material examined). Remarks. ...
Inquiline oak gall wasps from the genus Synergus Hartig (Hymenoptera: Cynipidae: Synergini) that occur in the New World should be classified into the following morphological groups: i) species with the radial cell of fore wings open; ii) species with the radial cell of fore wings closed and mesopleurae completely sculptured, always with transversal striae covering its surface; iii) species with the radial cell of fore wings closed and mesopleurae partially smooth, with the speculum always shiny and not sculptured. The latter group, which appears to be unique to the New World’s fauna, is the focus of this study. An exhaustive taxonomical revision of all the Synergus species within this morphological group is carried out for the first time after the initial treatment by Lobato-Vila & Pujade-Villar (2017). Redescriptions, images and new distribution and biological data are provided. A new species from Mexico, Synergus aurofacies Lobato-Vila & Pujade-Villar, sp. nov., is here formally described and illustrated. Synergus variegatus McCracken & Egbert, 1922 is a junior synonym of S. flavens McCracken & Egbert, 1922 syn. nov.; S. profusus McCracken & Egbert, 1922 is a junior synonym of S. pacificus McCracken & Egbert, 1922 syn. nov.; and S. flavus Kieffer, 1904 and S. varicolor Fullaway, 1911 are junior synonyms of S. pomiformis (Ashmead, 1885) syn. nov. A key to species of this group is provided. The morphological traits of the species with partially smooth mesopleurae are discussed.
Phylogenetically the genus Atrusca Kinsey forms a very distinct clade; the genus is most closely related to Cynips (= Besbicus), Philonix and Xanthoteras gen rev.. Morphologically, Atrusca most closely resembles Cynips (= Besbicus) and Antron. This study reviews the genus Atrusca including 42 species. Three new synonyms are proposed: Atrusca vanescens (Kinsey, 1930), syn. nov. of Atrusca bella (Bassett, 1881), Atrusca oriunda (Kinsey, 1936), syn. nov. of Atrusca oriens (Kinsey, 1936), and Atrusca vulgata (Kinsey, 1936), syn. nov. of Atrusca vasta (Kinsey, 1936). Two species are transferred to other genera, Atrusca sagata (Kinsey) is transferred to Sphaeroteras and A. clavuloides (Kinsey) is transferred back to the genus Xanthoteras gen. rev., which is here re-established. We provide the re-descriptions of all known Atrusca species, diagnoses, keys to species and information on biology, phenology, and distribution.
A new genus of oak gall wasps, Neuroandricus Pujade-Villar & Melika, gen. nov., with one new species, N. pustulatus Pujade-Villar & Melika, sp. nov. is described from Mexico. The new Cynipini species is known from its asexual generation which induces galls on leaves of Quercus (section Quercus). Also, a new species of gall-associated parasitoid is described, Sycophila baldufi Savall-Roig & Pujade-Villar, sp. nov. (Eurytomidae), emerged from N. pustulatus galls. Diagnosis, distribution and biological data of the new species are given.
A new genus of oak gall wasp, Reticulodermis Pujade-Villar, Cuesta-Porta & Melika gen. nov. and a new species R. lithogalla García-Martiñón & Pujade-Villar sp. nov. are described from Mexico. Reticulodermis is known only from the asexual generation that induces galls on leaves of Quercus crassipes Humb. & Bonpl. (section Lobatae). We provide the diagnosis, distribution and biological data on the new species. An identification key to the genera of oak gall wasps inducing galls on the section Lobatae is given.
Se resume el conocimiento actual de la taxonomía de las avispas gallícolas de encinos (Cynipini) en México. Se exponen todas las referencias de las especies descritas en México, así como todas las menciones posteriores y nuevos datos a partir del estudio de agallas de recolectas esporádicas realizadas en los últimos 10 años. Este conocimiento ha sido organizado en forma de catálogo estado por estado. En total se contabilizan 205 especies en 24 de los 30 estados mexicanos que presentan encinos, algunas de ellas se citan por primera vez en algunos estados, contabilizando 161 registros nuevos para un total de 16 estados. Para cada especie se proporcionan los datos de distribución y hospedador, incluyendo el trabajo donde se menciona por vez primera a la especie, la página de referencia y el lugar de recolección. De los 21 géneros de avispas gallícolas en encinos presentes en México, Andricus Hartig es el más ampliamente distribuido, con 37 especies. La Ciudad de México y Zacatecas son los territorios con más especies citadas, 48 y 40 respectivamente. En lo que se refiere a los hospedadores, todos pertenecen al género Quercus, contemplando 66 especies repartidas en 4 secciones: Quercus, Lobatae, Protobalanus y Virentes.
New information is provided on the galling fauna from oaks in Santa Fe (Cuajimalpa (Mexico City) obtained from collections on Quercus laeta Liebmann, 1854 and Q. rugosa Née, 1801 (section Quercus), Q. crassipes Humboldt and Bonpland, 1809 and Q. calophylla Schlechtendal and Chamisso, 1830 (section Lobatae). Seven species are cited for the first time: Andricus fusciformis Pujade-Villar, 2014 and A. guanajuatensis Pujade-Villar, 2013; Femuros lusum Kinsey, 1937 and F. repandae Kinsey, 1937; Kokkocynips doctorrosae Pujade-Villar and Melika, 2013; Loxaulus hyalinus Pujade-Villar and Melika, 2014 and Neuroterus fusifex Pujade-Villar, 2016. Of the 12 species originally described from Santa Fe, nine of them are only known in this studied area: Amphibolips cibriani Pujade-Villar, 2011, Andricus breviramuli Pujade-Villar, 2014, A. rochai Pujade-Villar, 2018, A. santafe Pujade-Villar, 2013, Disholcaspis crystalae Pujade-Villar, 2018, Loxaulus laeta Pujade-Villar, 2014, Neuroterus eugeros Pujade-Villar, 2018, N. verrucum Pujade Villar, 2014 and Zapatella polytryposa Pujade-Villar and Fernández-Garzón, 2020. New hosts are provided for some of the mentioned species. Some galls are shown that could constitute species yet to be described. Of all mentioned species, the gall is described and illustrated, and aspects of its biology, hosts and distribution are commented.
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.
The original description of Parandricus mairei Kieffer, 1906 included a misinterpretation of a relevant forewing trait. The species was subsequently transferred to the genus Andricus, despite presenting two very peculiar morphological characters, namely a simple tarsal claw and the absence of irradiating carinae on the lower face. Similarly, the original descriptions of Andricus deqingis Wang, Gui, Chen, 2013 and A. flavus Pujade-Villar, Wang, Guo & Chen, 2014 included some relevant mistakes. Here, we present the results of a molecular analysis that reveals that individuals of the three species are genetically very similar; supporting the proposal that A. mairei is a senior synonym of the other two species, A. deqingis n. syn. and A. flavus n. syn. In addition, our results indicate that Parandricus renders Andricus paraphyletic, which supports that Parandricus is a junior synonym of Andricus. We re-describe and illustrate the relevant characters of A. mairei and provide additional comments on the characters erroneously interpreted in the former descriptions and give notes about the biology and intraspecific variability.
We present the latest version of the Molecular Evolutionary Genetics Analysis (MEGA) software, which contains many sophisticated
methods and tools for phylogenomics and phylomedicine. In this major upgrade, MEGA has been optimized for use on 64-bit computing
systems for analyzing bigger datasets. Researchers can now explore and analyze tens of thousands of sequences in MEGA. The
new version also provides an advanced wizard for building timetrees and includes a new functionality to automatically predict
gene duplication events in gene family trees. The 64-bit MEGA is made available in two interfaces: graphical and command line.
The graphical user interface (GUI) is a native Microsoft Windows application that can also be used on Mac OSX. The command
line MEGA is available as native applications for Windows, Linux, and Mac OSX. They are intended for use in high-throughput
and scripted analysis. Both versions are available from www.megasoftware.net free of charge.
Lezak B, Varacallo M. Anatomy, Bony Pelvis and Lower Limb, Foot Veins. [Updated 2019 Jun 6]. In: StatPearls [Internet]. Treasure Island (FL): StatPearls Publishing; 2019 Jan-. Available from: https://www.ncbi.nlm.nih.gov/books/NBK542295/
A new species of Andricus Hartig 1840 (Hymenoptera: Cynipidae: Cynipini) is described from Panama: Andricus barriosi sp. nov. The new species induces galls on Quercus bumelioides Liebm. and Q. insignis M. Martens & Galeotti (Fagaceae, sect. Quercus) and is the first species of the genus Andricus 'sensu lato' recorded from Panama. Andricus barriosi is part of a species complex that includes A. nievesaldreyi Pujade-Villar, A. georgei Pujade-Villar and A. maesi Pujade-Villar, which differ from the other species of the genus Andricus by having a characteristic striate-reticulate sculpture on the second metasomal tergum that is typically smooth in the other Andricus species. The diagnostic characters, gall description, distribution and biological data of the new species are given, and a key for the differentiation of the new species from related species is also provided.
Mexican species of Biorhiza Westwood are of doubtful generic affiliation after Kinsey described 11 species from Mexico in 1937. Moreover, two of them were transferred to Trigonaspis Hartig (B. nitellina Kinsey, 1937 and B. zinzala Kinsey, 1937) and another one to Cynips Linnaeus (B. socia Kinsey, 1937). The type material of all these species has been examined, detailing the problems to keep considering B. innocens Kinsey, 1937, B. solita Kinsey, 1937, B. stelis Kinsey, 1937, B. tanos Kinsey, 1937, B. tarasco Kinsey, 1937, B. tricosa Kinsey, 1937, B. ulcus Kinsey, 1937 and B. urcea Kinsey, 1937 within Biorhiza or within any other valid genus with malar sulcus. The genus Sphaeroteras Ashmead is re-established to group most of the Biorhiza species described by Kinsey. Only three Mexican species are considered valid in Sphaeroteras: S. solita (Kinsey, 1937) n. comb. (= B. socia Kinsey, 1937 n. syn. = B. stelis Kinsey, 1937 n. syn. = B. tarasco Kinsey, 1937 n. syn. = B. tricosa Kinsey, 1937 n. syn. = B. ulcus Kinsey, 1937 n. syn. = B. urcea Kinsey, 1937 n. syn.), S. tanos (Kinsey, 1937) n. comb. and S. zinzala (Kinsey, 1937) n. comb. The Mexican species Biorhiza innocens and B. nitellina are considered ‘incertae sedis’, as well as other species in America north of Mexico included in Trigonaspis. The affiliation of the species previously included in Sphaeroteras in America north of Mexico is discussed. Atrusca pulchripennis (Ashmead, 1896) belongs to Sphaeroteras, S. pulchripennis (Ashmead, 1896) n. comb. The most important morphological characters are illustrated. The validity of Trigonaspis in America is commented as well as the possibility that Xanthoteras Ashmead could be a valid genus.
Bayesian inference of phylogeny using Markov chain Monte Carlo (MCMC) (Drummond et al., 2002; Mau et al., 1999; Rannala and Yang, 1996) flourishes as a popular approach to uncover the evolutionary relationships among taxa, such as genes, genomes, individuals or species. MCMC approaches generate samples of model parameter values - including the phylogenetic tree -drawn from their posterior distribution given molecular sequence data and a selection of evolutionary models. Visualising, tabulating and marginalising these samples is critical for approximating the posterior quantities of interest that one reports as the outcome of a Bayesian phylogenetic analysis. To facilitate this task, we have developed the Tracer (version 1.7) software package to process MCMC trace files containing parameter samples and to interactively explore the high-dimensional posterior distribution. Tracer works automatically with sample output from BEAST (Drummond et al., 2012), BEAST2 (Bouckaert et al., 2014), LAMARC (Kuhner, 2006), Migrate (Beerli, 2006), MrBayes (Ronquist et al., 2012), RevBayes (Höhna et al., 2016) and possibly other MCMC programs from other domains.
This study reviewed the aphid genus Mesocallis from the Korean peninsula. A total of five species are recognized using morphological and molecular evidence, including two new species, Mesocallis (Mesocallis) carpinicola sp. nov. and M. (Paratinocallis) occulta sp.
nov., one newly recorded species, M. (M.) pteleae Matsumura, 1919, and the two species M. (M.) sawashibae (Matsumura, 1917) and M. (P.) corylicola (Higuchi, 1972). Species description, illustrations and distributional and biological data were provided for all Korean Mesocallis species along with a pictorial key. Pairwise distances, a neighbor-joining tree and a median-joining network are also given.
The six Mexican species described by Kinsey in 1937 originally belonging to Dros have been studied: D. moreliensis, D. periscellus, D. perlentus, D. petasus, D. picatus and D. repicatus. Here is concluded that Dros is a valid genus and three of the six species should be considered valid: D. moreliense comb. rest. (= D. periscele n. syn.), D. picatum comb. rest. (= D. repicatum n. syn.) and D. perlentum comb. rest. (= D. petasum n. syn.). An identification key to separate adults and galls of these species is included; also, the most important characters that define them are illustrated. The taxonomic position of the Dros species north of Mexico is discussed, concluding than at least two species belong to the genus Dros: D. sessile (Weld, 1927) and D. viscidum (Weld, 1944).
Se describe un nuevo género de cinípido agallícola, Kokkocynips Pujade-Villar & Melika n. gen. (Hymenoptera: Cynipidae: Cynipini) de México. Se discuten en detalle los caracteres diagnósticos y los límites genéricos de este nuevo género. Las agallas se encuentran en las ramas de Quercus acutifolia Née. Se describen e ilustran los caracteres diagnósticos, su distribución y la biología de la nueva especie.
PartitionFinder 2 is a program for automatically selecting best-fit partitioning schemes and models of evolution for phylogenetic analyses. PartitionFinder 2 is substantially faster and more efficient than version 1, and incorporates many new methods and features. These include the ability to analyze morphological datasets, new methods to analyze genome-scale datasets, new output formats to facilitate interoperability with downstream software, and many new models of molecular evolution. PartitionFinder 2 is freely available under an open source license and works on Windows, OSX, and Linux operating systems. It can be downloaded from www.robertlanfear.com/partitionfinder The source code is available at https://github.com/brettc/partitionfinder.
Time spent in describing specimens or understanding descriptions is
onen substantially lengthened by tiresome searches for accurate termi·
nology or for interpretive insight where confusion exists in the descriptive
terminology. Consistent and correct use of descriptive terms is
essentÎal for efficient taxonomie work; synonymies abound where original
descriptions have inadequately defined a specimen. The fault does
not lie in a lack of suitable ferros sinee there are descriptive tenns for
almost any imaginable structure. The problem lies in the misapplication
of these terms when workers fail to acquaint themselves with the nomen·
clature or use terms too casually. Almost all the lenns have been avait·
able sinee Smith published his GlossaryofEn/omoJogyin 1906. but tbey
have never been available in a convenient fonnat. Smith weeded out a
lot ofsynonyms but more bave appeared through the years. Torre-Bueno
(1937) feh ail terms in current use should be included even ifthey were
synonymized, for the benefit of worken in restrictod are&S. Unfortunatf>
Iy, he did not indicate any of these restricted areas in his glossary. For
this reason many worken find it inconvenient to use his glossary. Further,
the tenns tbey seek are neither cross.referenced nor categorized.
A complete revision of the terminology is out of my grasp. Instead,
1 have revised one small area of tenninology particularly in need of it.
1 have collected and revised descriptive terminology ofsurface sculpturiDg
and presented it with SEM micrographs for reference. 1hope this will
standardize the use of specific terminology which will in tum increase
accurate information transfer in descriptive and diagnostic work. The
terms are indexed and synonymized and presented in related categories.
Most of the teems are cross-.referenced, though only to c10sely related
tenns since the categorica1 groupings serve a similar function at a broad·
er level. The photo-reference section should be helpful in attaining a
comprehensive undentanding of these tenns, and also provides a survey
of the diverse surface sculpturing in the Hymenoptera.
A new genus of a cynipid oak gallwasp, Kinseyella (Hymenoptera: Cynipidae: Cynipini) is described from Mexico. Diagnostic characters and generic limits of the new genus are discussed in details. A new species, Kinseyella quercusobtusata is described, which induces galls on leaves of Quercus obtusata, an endemic Mexican oak. One earlier known species from Mexico was transferred to the newly established genus, Kinseyella lapiei (Kieffer), comb. nova. Diagnostic characters for separation of two species are also given.
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.
The current classification of world genera of Cynipini follows Weld (1952a), who divided Cynipini into 39 genera. Later, Monzen (1954) described a new genus, Neoneuroterus from Japan; Maisuradze (1961) reported a new genus, Repentinia Belizin & Maisuradze from Ciscaucasus (Azerbaijan); Kovalev (1965) described two new genera, Belizinella and Ussuraspis from the Far East of Russia; Lyon (1993) synonymized Xystoteras to Phylloteras and described a new genus, Euxystoteras; and Melika & Abrahamson (1997b) described a new genus Eumayriella from Florida and synonymized Trisoleniella to Eumayria. Later, Melika, Ros-Farré & Pujade-Villar (2001) synonymized Fioriella to Plagiotrochus. Recently 41 genera of Cynipini were known to associate with oaks, generic division of which was based on the presence or absence of a basal lobe on the tarsal claws. A reclassification of world genera of oak gall-inducing cynipids of the tribus Cynipini (Hymenoptera: Cynipidae) is given in which 26 genera are proposed as valid, 15 are synonymized, one Neuroterus subgenus, Latuspina Monzen, 1954 has an uncertain status; 73 comb. nov. and 26 comb. rev. are made.
Barucynips panamensis Medianero & Nieves-Aldrey, a new genus and species of oak gallwasps (Hymenoptera: Cynipidae: Cynipini), is described from adults reared from galls on Quercus bumelioides in Panama. The new genus is taxonomically close to the recently described Coffeikokkos from Costa Rica, but differs from it and all of the described genera of Cynipini, by the shape and setation of the projecting part of the ventral spine of the hypopygium and by the sculpture of the propodeum. A new species of Coffeikokkos is also described from the same area, the Volcán Barú in Panama. Diagnostic characters, gall description, distribution, and biological data of the new genus and the two new species are given. The new genus is the first genus of oak gallwasps of the tribe Cynipini described in Panama.
According to the Host-tracking Hypothesis, species of higher trophic levels with a close relationship to their hosts, such as parasites or parasitoids, are expected to show spatio-temporal phylogeographic patterns similar to those of their host. Alternatively, with ecological sorting, a subset of the local species pools might shift to a related host species, thereby disengaging common phylogeographic patterns. Here, we compare the phylogeographic structures of the cynipid rose gall wasp Diplolepis rosae across Europe and of two of its most common parasitoids, the wasps Orthopelma mediator and Glyphomerus stigma, by analysing the sequences of two gene fragments (COI and ITS 2). The phylogeographic structures of the three species associated with roses were incongruent. D. rosae had the lowest genetic diversity with one major clade, O. mediator showed the classical phylogeographic structure for Europe with one eastern and one western clade, and G. stigma had the highest diversity but no geographical structuring. This discordance of geographical patterns may be explained by 1) the dispersal propensity of adult parasitoids or 2) the parasitoids having the ability to switch to another host, while the primary host becomes rare or is even not available. Furthermore there was no indication that phylogenetic patterns were affected by Wolbachia infections. Our results document that communities of closely interacting species may be the result of idiosyncratic biogeographic histories.
A new genus of cynipid oak gallwasp, Zapatella Pujade-Villar & Melika, gen. n. (Hymenoptera: Cynipidae: Cynipini), with two new species, Zapatella grahami Pujade-Villar & Melika, sp. n. and Zapatella nievesaldreyi Melika & Pujade-Villar, sp. n., is described from the Neotropics. Zapatella grahami,known only from the sexual generation,induces galls in acorns of Quercus costaricensis and is currently known only from Costa Rica. Zapatella nievesaldreyi, known only from the asexual generation, induces inconspicuous galls in twigs of Quercus humboldtii, and is known only from Colombia. Diagnostic characters for both new species are given in detail. Five Nearctic species are transferred from Callirhytis to Zapatella: Zapatella cryptica (Weld), comb. n., Zapatella herberti (Weld), comb. n., Zapatella oblata (Weld), comb. n., Zapatella quercusmedullae (Ashmead), comb. n.,
Zapatella quercusphellos (Osten Sacken), comb. n. (= Zapatella quercussimilis (Bassett), syn. n.). A key based on adults for the species belonging to Zapatella is also given. Generic limits and morphological characteristics of Zapatella and closely related genera are discussed.
We present the most comprehensive analysis of higher-level relationships in gall wasps conducted thus far. The analysis was based on detailed study of the skeletal morphology of adults, resulting in 164 phylogenetically informative characters, complemented with a few biological characters. Thirty-seven cynipid species from thirty-one genera, including four genera of the apparently monophyletic Cynipini and almost all of the genera in the other tribes, were examined. The outgroup included exemplar species from three successively more distant cynipoid families: Figitidae (the sister group of the Cynipidae), Liopteridae and Ibaliidae. There was considerable homoplasy in the data, but many groupings in the shortest tree were nonetheless well supported, as indicated by bootstrap proportions and decay indices. Partitioning of the data suggested that the high level of homoplasy is characteristic of the Cynipidae and not the result of the amount of available phylogenetically conservative characters being exhausted. The analysis supported the monophyly of the Cynipini (oak gall wasps) which, together with the Rhoditini (the rose gall wasps), Eschatocerini and Pediaspidini formed a larger monophyletic group of gall inducers restricted to woody representatives of the eudicot subclass Rosidae. The inquilines (Synergini) were indicated to be monophyletic, whereas the Aylacini, primarily herb gall inducers, appeared as a paraphyletic assemblage of basal cynipid groups. The shortest tree suggests that the Cynipidae can be divided into three major lineages: one including the inquilines, the Aylacini genera associated with Rosaceae, and Liposthenes; one consisting entirely of Aylacini genera, among them Aulacidea, Isocolus and Neaylax; and one comprising the woody rosid gallers (the oak and rose gall wasps and allies), the Phanacis-Timaspis complex and the Aylacini genera associated with Papaveraceae.
Understanding the evolutionary history of living organisms is a central problem in biology. Until recently the ability to infer evolutionary relationships was limited by the amount of DNA sequence data available, but new DNA sequencing technologies have largely removed this limitation. As a result, DNA sequence data are readily available or obtainable for a wide spectrum of organisms, thus creating an unprecedented opportunity to explore evolutionary relationships broadly and deeply across the Tree of Life. Unfortunately, the algorithms used to infer evolutionary relationships are NP-hard, so the dramatic increase in available DNA sequence data has created a commensurate increase in the need for access to powerful computational resources. Local laptop or desktop machines are no longer viable for analysis of the larger data sets available today, and progress in the field relies upon access to large, scalable high-performance computing resources. This paper describes development of the CIPRES Science Gateway, a web portal designed to provide researchers with transparent access to the fastest available community codes for inference of phylogenetic relationships, and implementation of these codes on scalable computational resources. Meeting the needs of the community has included developing infrastructure to provide access, working with the community to improve existing community codes, developing infrastructure to insure the portal is scalable to the entire systematics community, and adopting strategies that make the project sustainable by the community. The CIPRES Science Gateway has allowed more than 1800 unique users to run jobs that required 2.5 million Service Units since its release in December 2009. (A Service Unit is a CPU-hour at unit priority).
Since its introduction in 2001, MrBayes has grown in popularity as a software package for Bayesian phylogenetic inference using Markov chain Monte Carlo (MCMC) methods. With this note, we announce the release of version 3.2, a major upgrade to the latest official release presented in 2003. The new version provides convergence diagnostics and allows multiple analyses to be run in parallel with convergence progress monitored on the fly. The introduction of new proposals and automatic optimization of tuning parameters has improved convergence for many problems. The new version also sports significantly faster likelihood calculations through streaming single-instruction-multiple-data extensions (SSE) and support of the BEAGLE library, allowing likelihood calculations to be delegated to graphics processing units (GPUs) on compatible hardware. Speedup factors range from around 2 with SSE code to more than 50 with BEAGLE for codon problems. Checkpointing across all models allows long runs to be completed even when an analysis is prematurely terminated. New models include relaxed clocks, dating, model averaging across time-reversible substitution models, and support for hard, negative, and partial (backbone) tree constraints. Inference of species trees from gene trees is supported by full incorporation of the Bayesian estimation of species trees (BEST) algorithms. Marginal model likelihoods for Bayes factor tests can be estimated accurately across the entire model space using the stepping stone method. The new version provides more output options than previously, including samples of ancestral states, site rates, site d(N)/d(S) rations, branch rates, and node dates. A wide range of statistics on tree parameters can also be output for visualization in FigTree and compatible software.
A new genus of oak gallwasp, Coffeikokkos Pujade-Villar & Melika, gen. n., is described from Costa Rica. Diagnostic characters and generic limits of the new genus are discussed in detail. The new genus includes Coffeikokkos copeyensis Pujade-Villar & Melika, sp. n., which induces galls on stems of Quercus bumelioides, an endemic oak to Costa Rica, Honduras and Panama. The new species and galls are described and illustrated.
How geographically widespread biological communities assemble remains a major question in ecology. Do parallel population histories allow sustained interactions (such as host-parasite or plant-pollinator) among species, or do discordant histories necessarily interrupt them? Though few empirical data exist, these issues are central to our understanding of multispecies evolutionary dynamics. Here we use hierarchical approximate Bayesian analysis of DNA sequence data for 12 herbivores and 19 parasitoids to reconstruct the assembly of an insect community spanning the Western Palearctic and assess the support for alternative host tracking and ecological sorting hypotheses. We show that assembly occurred primarily by delayed host tracking from a shared eastern origin. Herbivores escaped their enemies for millennia before parasitoid pursuit restored initial associations, with generalist parasitoids no better able to track their hosts than specialists. In contrast, ecological sorting played only a minor role. Substantial turnover in host-parasitoid associations means that coevolution must have been diffuse, probably contributing to the parasitoid generalism seen in this and similar systems. Reintegration of parasitoids after host escape shows these communities to have been unsaturated throughout their history, arguing against major roles for parasitoid niche evolution or competition during community assembly.
Phylogenetic relationships among the holometabolous insect orders were inferred from cladistic analysis of nucleotide sequences of 18S ribosomal DNA (rDNA) (85 exemplars) and 28S rDNA (52 exemplars) and morphological characters. Exemplar outgroup taxa were Collembola (1 sequence), Archaeognatha (1), Ephemerida (1), Odonata (2), Plecoptera (2), Blattodea (1), Mantodea (1), Dermaptera (1), Orthoptera (1), Phasmatodea (1), Embioptera (1), Psocoptera (1), Phthiraptera (1), Hemiptera (4), and Thysanoptera (1). Exemplar ingroup taxa were Coleoptera: Archostemata (1), Adephaga (2), and Polyphaga (7); Megaloptera (1); Raphidioptera (1); Neuroptera (sensu stricto = Planipennia): Mantispoidea (2), Hemerobioidea (2), and Myrmeleontoidea (2); Hymenoptera: Symphyta (4) and Apocrita (19); Trichoptera: Hydropsychoidea (1) and Limnephiloidea (2); Lepidoptera: Ditrysia (3); Siphonaptera: Pulicoidea (1) and Ceratophylloidea (2); Mecoptera: Meropeidae (1), Boreidae (1), Panorpidae (1), and Bittacidae (2); Diptera: Nematocera (1), Brachycera (2), and Cyclorrhapha (1); and Strepsiptera: Corioxenidae (1), Myrmecolacidae (1), Elenchidae (1), and Stylopidae (3). We analyzed ∼1 kilobase of 18S rDNA, starting 398 nucleotides downstream of the 5′ end, and ∼400 bp of 28S rDNA in expansion segment D3. Multiple alignment of the 18S and 28S sequences resulted in 1,116 nucleotide positions with 24 insert regions and 398 positions with 14 insert regions, respectively. All Strepsiptera and Neuroptera have large insert regions in 18S and 28S. The secondary structure of 18S insert 23 is composed of long stems that are GC rich in the basal Strepsiptera and AT rich in the more derived Strepsiptera. A matrix of 176 morphological characters was analyzed for holometabolous orders. Incongruence length difference tests indicate that the 28S + morphological data sets are incongruent but that 28S + 18S, 18S + morphology, and 28S + 18S + morphology fail to reject the hypothesis of congruence. Phylogenetic trees were generated by parsimony analysis, and clade robustness was evaluated by branch length, Bremer support, percentage of extra steps required to force paraphyly, and sensitivity analysis using the following parameters: gap weights, morphological character weights, methods of data set combination, removal of key taxa, and alignment region. The following are monophyletic under most or all combinations of parameter values: Holometabola, Polyphaga, Megaloptera + Raphidioptera, Neuroptera, Hymenoptera, Trichoptera, Lepidoptera, Amphiesmenoptera (Trichoptera + Lepidoptera), Siphonaptera, Siphonaptera + Mecoptera, Strepsiptera, Diptera, and Strepsiptera + Diptera (Halteria). Antliophora (Mecoptera + Diptera + Siphonaptera + Strepsiptera), Mecopterida (Antliophora + Amphiesmenoptera), and Hymenoptera + Mecopterida are supported in the majority of total evidence analyses. Mecoptera may be paraphyletic because Boreus is often placed as sister group to the fleas; hence, Siphonaptera may be subordinate within Mecoptera. The 18S sequences for Priacma (Coleoptera: Archostemata), Colpocaccus (Coleoptera: Adephaga), Agulla (Raphidioptera), and Corydalus (Megaloptera) are nearly identical, and Neuropterida are monophyletic only when those two beetle sequences are removed from the analysis. Goleoptera are therefore paraphyletic under almost all combinations of parameter values. Halteria and Amphiesmenoptera have high Bremer support values and long branch lengths. The data do not support placement of Strepsiptera outside of Holometabola nor as sister group to Coleoptera. We reject the notion that the monophyly of Halteria is due to long branch attraction because Strepsiptera and Diptera do not have the longest branches and there is phylogenetic congruence between molecules, across the entire parameter space, and between morphological and molecular data.
Galls are highly specialized plant tissues whose development is induced by another organism. The most complex and diverse galls are those induced on oak trees by gallwasps (Hymenoptera: Cynipidae: Cynipini), each species inducing a characteristic gall structure. Debate continues over the possible adaptive significance of gall structural traits; some protect the gall inducer from attack by natural enemies, although the adaptive significance of others remains undemonstrated. Several gall traits are shared by groups of oak gallwasp species. It remains unknown whether shared traits represent (i) limited divergence from a shared ancestral gall form, or (ii) multiple cases of independent evolution. Here we map gall character states onto a molecular phylogeny of the oak cynipid genus Andricus, and demonstrate three features of the evolution of gall structure: (i) closely related species generally induce galls of similar structure; (ii) despite this general pattern, closely related species can induce markedly different galls; and (iii) several gall traits (the presence of many larval chambers in a single gall structure, surface resins, surface spines and internal air spaces) of demonstrated or suggested adaptive value to the gallwasp have evolved repeatedly. We discuss these results in the light of existing hypotheses on the adaptive significance of gall structure.
We describe "universal" DNA primers for polymerase chain reaction (PCR) amplification of a 710-bp fragment of the mitochondrial cytochrome c oxidase subunit I gene (COI) from 11 invertebrate phyla: Echinodermata, Mollusca, Annelida, Pogonophora, Arthropoda, Nemertinea, Echiura, Sipuncula, Platyhelminthes, Tardigrada, and Coelenterata, as well as the putative phylum Vestimentifera. Preliminary comparisons revealed that these COI primers generate informative sequences for phylogenetic analyses at the species and higher taxonomic levels.
Polymerase chain reaction (PCR) followed by sequencing of single-stranded DNA yielded sequence information from the cytochrome b (cyt b) region in mitochondrial DNA from the ant Tetraponera rufoniger. Compared with the cyt b genes from Apis mellifera, Drosophila melanogaster, and D. yakuba, the overall A+T content (A+T%) of that of T. rufoniger is lower (69.9% vs 80.7%, 74.2%, and 73.9%, respectively) than those of the other three. The codon usage in the cyt b gene of T. rufoniger is biased although not as much as in A. mellifera, D. melanogaster, and D. yakuba; T. rufoniger has eight unused codons whereas D. melanogaster, D. yakuba, and A. mellifera have 21, 20, and 23, respectively. The inferred cyt b polypeptide chain (PPC) of T. rufoniger has diverged at least as much from a common ancestor with D. yakuba as has that of A. mellifera (approximately 3.5 vs approximately 2.9). Despite the lower A+T%, the relative frequencies of amino acids in the cyt b PPC of T. rufoniger are significantly (P < 0.05) associated with the content of adenine and thymine (A+T%) and size of codon families. The mitochondrially located cytochrome oxidase subunit II genes (CO-II) of endopterygote insects have significantly higher average A+T% (approximately 75%) than those of exopterygous (approximately 69%) and paleopterous (approximately 69%) insects. The increase in A+T% of endopterygote insects occurred in Upper Carboniferous and coincided with a significant acceleration of PPC divergence. However, acceleration of PPC divergence is not significantly correlated with the increase of the A+T% (P > 0.1). The high A+T%, the biased codon usage, and the increased PPC divergence of Hymenoptera can in that respect most easily be explained by directional mutation pressure which began in the Upper Carboniferous and still occurs in most members of the order. Given the roughly identical A+T% of the cyt b and CO-II genes from the other insects whose DNA sequences are known (A. mellifera, D. melanogaster, and D. yakuba), it seems most likely that the A+T% of T. rufoniger declined secondarily within the last 100 Myr as a result of a reduced directional mutation pressure.
The Marble gallwasp Andricus kollari has a native range divided into two geographically separated lifecycles. In Eastern Europe and Turkey, the lifecycle involves a sexual generation on Turkey oak, Quercus cerris, while in Iberia and North Africa the sexual generation host is cork oak, Q. suber. Over the last 500 years, A. kollari has expanded its range into northern Europe, following human planting of Q. cerris from Italy and the Balkans. We ask: (i) what is the genetic relationship between eastern and western distributions of Andricus kollari? Can we determine which lifecycle is ancestral, and how long ago they diverged? (ii) To what extent have eastern and western native ranges contributed to northwards range expansion? (iii) Is there any evidence for hybridization between the two life cycle types? We present analyses of allozyme data for 13 polymorphic loci and of sequence variation for a 433 bp fragment of the mitochondrial cytochrome b gene. These show: (i) that four haplotype lineages (one in Spain, two in Hungary/Italy and one in Turkey) diverged more or less simultaneously between 1 and 2 million years ago, suggesting the existence of at least four refuges through recent ice age cycles. Our data cannot resolve which lifecycle type is ancestral. (ii) Populations north of putative refuges are divided into two sets. Populations in south-west France are allied to Spain, while all remaining populations in northern Europe have been colonized from Italy and the Balkans. (iii) The transition from one race to another in south-west France is marked by abrupt transitions in the frequency of refuge-specific private alleles and corresponds closely to the northern limit of the distribution of cork oak. Although hybrids were detected in north-west France, none were detected where the two lifecycles meet in south-western France. The biology of oak gallwasps predicts that any hybrid zone will be narrow, and limited to regions where Q. cerris and Q. suber meet. Our data suggest that eastern and western A. kollari are effectively separate species.
The recent development of Bayesian phylogenetic inference using Markov chain Monte Carlo (MCMC) techniques has facilitated the exploration of parameter-rich evolutionary models. At the same time, stochastic models have become more realistic (and complex) and have been extended to new types of data, such as morphology. Based on this foundation, we developed a Bayesian MCMC approach to the analysis of combined data sets and explored its utility in inferring relationships among gall wasps based on data from morphology and four genes (nuclear and mitochondrial, ribosomal and protein coding). Examined models range in complexity from those recognizing only a morphological and a molecular partition to those having complex substitution models with independent parameters for each gene. Bayesian MCMC analysis deals efficiently with complex models: convergence occurs faster and more predictably for complex models, mixing is adequate for all parameters even under very complex models, and the parameter update cycle is virtually unaffected by model partitioning across sites. Morphology contributed only 5% of the characters in the data set but nevertheless influenced the combined-data tree, supporting the utility of morphological data in multigene analyses. We used Bayesian criteria (Bayes factors) to show that process heterogeneity across data partitions is a significant model component, although not as important as among-site rate variation. More complex evolutionary models are associated with more topological uncertainty and less conflict between morphology and molecules. Bayes factors sometimes favor simpler models over considerably more parameter-rich models, but the best model overall is also the most complex and Bayes factors do not support exclusion of apparently weak parameters from this model. Thus, Bayes factors appear to be useful for selecting among complex models, but it is still unclear whether their use strikes a reasonable balance between model complexity and error in parameter estimates.
Unlabelled:
RAxML-VI-HPC (randomized axelerated maximum likelihood for high performance computing) is a sequential and parallel program for inference of large phylogenies with maximum likelihood (ML). Low-level technical optimizations, a modification of the search algorithm, and the use of the GTR+CAT approximation as replacement for GTR+Gamma yield a program that is between 2.7 and 52 times faster than the previous version of RAxML. A large-scale performance comparison with GARLI, PHYML, IQPNNI and MrBayes on real data containing 1000 up to 6722 taxa shows that RAxML requires at least 5.6 times less main memory and yields better trees in similar times than the best competing program (GARLI) on datasets up to 2500 taxa. On datasets > or =4000 taxa it also runs 2-3 times faster than GARLI. RAxML has been parallelized with MPI to conduct parallel multiple bootstraps and inferences on distinct starting trees. The program has been used to compute ML trees on two of the largest alignments to date containing 25,057 (1463 bp) and 2182 (51,089 bp) taxa, respectively.
Availability:
icwww.epfl.ch/~stamatak
Many studies have addressed the latitudinal gradients in intraspecific genetic diversity of European taxa generated during postglacial range expansion from southern refugia. Although Asia Minor is known to be a centre of diversity for many taxa, relatively few studies have considered its potential role as a Pleistocene refugium or a potential source for more ancient westward range expansion into Europe. Here we address these issues for an oak gallwasp, Andricus quercustozae (Hymenoptera: Cynipidae), whose distribution extends from Morocco along the northern coast of the Mediterranean through Turkey to Iran. We use sequence data for a fragment of the mitochondrial gene cytochrome b and allele frequency data for 12 polymorphic allozyme loci to answer the following questions: (1). which regions represent current centres of genetic diversity for A. quercustozae? Do eastern populations represent one refuge or several discrete glacial refugia? (2). Can we infer the timescale and sequence of the colonization processes linking current centres of diversity? Our results suggest that A. quercustozae was present in five distinct refugia (Iberia, Italy, the Balkans, southwestern Turkey and northeastern Turkey) with recent genetic exchange between Italy and Hungary. Genetic diversity is greatest in the Turkish refugia, suggesting that European populations are either (a). derived from Asia Minor, or (b). subject to more frequent population bottlenecks. Although Iberian populations show the lowest diversity for putatively selectively neutral markers, they have colonized a new oak host and represent a genetically and biologically discrete entity within the species.
A new genus of oak gallwasp, Melikaiella Pujade-Villar n. gen. is described, with four new species: M. amphibolensis Pujade-Villar n. sp., M. bicolor Pujade-Villar n. sp., M. reticulata Pujade-Villar n. sp. from Mexico and M. ostensackeni Pujade-Villar n. sp. from USA. Callirhytis sonorae Weld from Mexico, C. corrugis Bassett, C. flora Weld, C. fructicola Ashmead, C. fructuosa Weld, C. lupana Weld, C. tumifica (Osten-Sacken) and Cynips papula Bassett from USA, are transferred to Melikaiella. Callirhytis petrosa Weld is a new synonymy of Melikaiella corrugis n. comb. Cynips quercusmodesta (O. Sacken) has an uncertain status and it is considered here as a nomina dubia. Synonymy of Cynips papula and Cynips quercusmodesta is rejected. Data on biology and distribution of the new genus is commented. The diagnostic characters are illustrated. The variability of C. flora sexual and asexual forms is discussed. A key to twelve species of Melikaiella is given.
We describe a new genus of cynipid oak gallwasp, Protobalandricus Melika, Nicholls & Stone (Hymenoptera: Cynipidae: Cynipini). Protobalandricus gen. nov. includes one previously described species, Disholcaspis spectabilis (Kinsey), which induces stem swelling-like galls on golden cup oaks, Quercus section Protobalanus. Descriptions of the genus and diagnostic characters, including DNA sequence data, are presented.
Many herbivores employ reward-based mutualisms with ants to gain protection from natural enemies. We examine the evolutionary dynamics of a tetra-trophic interaction in which gall wasp herbivores induce their host oaks to produce nectar-secreting galls, which attract ants that provide protection from parasitoids. We show that, consistent with other gall defensive traits, nectar secretion has evolved repeatedly across the oak gall wasp tribe and also within a single genus (Disholcaspis) that includes many nectar-inducing species. Once evolved, nectar secretion is never lost in Disholcaspis, consistent with high defensive value of this trait. We also show that evolution of nectar secretion is correlated with a transition from solitary to aggregated oviposition, resulting in clustered nectar-secreting galls, which produce a resource that ants can more easily monopolize. Such clustering is commonly seen in ant guard mutualisms. We suggest that correlated evolution between maternal oviposition and larval nectar induction traits has enhanced the effectiveness of this gall defense strategy.
A multiple sequence alignment program, MAFFT, has been developed. The CPU time is drastically reduced as compared with existing
methods. MAFFT includes two novel techniques. (i) Homo logous regions are rapidly identified by the fast Fourier transform
(FFT), in which an amino acid sequence is converted to a sequence composed of volume and polarity values of each amino acid
residue. (ii) We propose a simplified scoring system that performs well for reducing CPU time and increasing the accuracy
of alignments even for sequences having large insertions or extensions as well as distantly related sequences of similar length.
Two different heuristics, the progressive method (FFT‐NS‐2) and the iterative refinement method (FFT‐NS‐i), are implemented
in MAFFT. The performances of FFT‐NS‐2 and FFT‐NS‐i were compared with other methods by computer simulations and benchmark
tests; the CPU time of FFT‐NS‐2 is drastically reduced as compared with CLUSTALW with comparable accuracy. FFT‐NS‐i is over
100 times faster than T‐COFFEE, when the number of input sequences exceeds 60, without sacrificing the accuracy.
Based on several structural and biological characteristics, the Cynipoidea can be divided into two groups, ‘macrocynipoids’ and ‘microcynipoids’. The macrocynipoids (i.e. the family Liopteridae and the genera Austrocynips, Eileenella, Heteribalia and Ibalia) are generally large insects that parasitize wood- or cone-boring insect larvae. The microcynipoids are smaller insects that are either phytophagous gall inducers and inquilines (Cynipidae) or parasitoids of larvae of Hymenoptera, Neuroptera or Diptera (Figitidae sensu lato, including the families Eucoilidae, Charipidae and Anacharitidae). The phylogenetic relationships among genera of macrocynipoids and between these and a sample of four genera representing the Figitidae and Cynipidae were examined by parsimony analysis of 110 external morphological characters of adults. Within the macrocynipoids, three monophyletic lineages emerged, classified here as different families: the Austrocynipidae, with a single species, Austrocynips mirabilis, the only cynipoid with a true pterostigma; the Ibaliidae, including the genera Eileenella, Ibalia and Heteribalia; and the Liopteridae, comprising the remaining genera of macrocynipoids. The analysis further supported the monophyly of the microcynipoids and indicated that the macrocynipoids form a paraphyletic group relative to the microcynipoids, with the shortest tree suggesting the relationship (Austrocynipidae, (Ibaliidae, (Liopteridae, microcynipoids))). These results imply that cynipoids were originally parasitoids of wood-boring insect larvae and that the other modes of life evolved secondarily within the group.
Ronquist, F. (1999). Phylogeny, classification and evolution of the Cynipoidea. Ð Zoologica Scripta 28, 139±164. I review cynipoid phylogeny and evolution and present an improved higher classification of the superfamily, including the proposal of some nomenclatural changes at the family and genus level. There is convincing morphological evidence for cynipoid monophyly. Relationships with other apocritan wasps are unclear although some characters suggest that diapriids may form their sister group. Cladistic analysis based on adult morphology indicates that cynipoids fall into five monophyletic lineages arranged in a Hennigian comb or ladder: (Austrocynipidae (Ibaliidae (Liopteridae (Figitidae sensu lato, Cynipidae)))). The three first families comprise the macrocynipoids, the two latter the microcynipoids. Relationships among macrocynipoids have been analysed down to the genus or species level and considerable attention has been paid to their historical biogeography. In the microcynipoids, comprising the bulk of cynipoid diversity, phylogenetic research has focused on the gall wasps (Cynipidae). Higher-level relationships are well studied and this allows reconstruction of the early evolution of the gall wasp-host plant association and the origin of the inquilines. The Figitidae are by far the least known cynipoid family and their classification is chaotic; here, I present a first attempt at a cladistic analysis of their higher-level relationships. Biogeographic and fossil evidence, as well as a morphological clock estimate, suggest that the cynipoid crown-group initially diverged in the Jurassic but the earliest cynipoid fossils are from the mid Cretaceous. Existing cynipoid fossils are here tentatively placed in a phylogenetic context but explicit cladistic analysis is likely to shed more light on their exact relationships. Current phylogenetic estimates suggest that cynipoids went through three successive phases in their early evolution, each leaving a set of surviving lineages: first in the community of wood-boring insects, then in the gall community, and finally in the aphid community. The parasitic cynipoids have apparently shifted only four times between hosts in different insect orders, demonstrating extreme conservatism in host association as might be expected of koinobiont endoparasites.
The Marble gallwasp Andricus kollari has a native range divided into two geographically separated lifecycles. In Eastern Europe and Turkey, the lifecycle involves a sexual generation on Turkey oak, Quercus cerris, while in Iberia and North Africa the sexual generation host is cork oak, Q. suber. Over the last 500 years, A. kollari has expanded its range into northern Europe, following human planting of Q. cerris from Italy and the Balkans. We ask: (i) what is the genetic relationship between eastern and western distributions of Andricus kollari? Can we determine which lifecycle is ancestral, and how long ago they diverged? (ii) To what extent have eastern and western native ranges contributed to northwards range expansion? (iii) Is there any evidence for hybridization between the two life cycle types? We present analyses of allozyme data for 13 polymorphic loci and of sequence variation for a 433 bp fragment of the mitochondrial cytochrome b gene. These show: (i) that four haplotype lineages (one in Spain, two in Hungary/Italy and one in Turkey) diverged more or less simultaneously between 1 and 2 million years ago, suggesting the existence of at least four refuges through recent ice age cycles. Our data cannot resolve which lifecycle type is ancestral. (ii) Populations north of putative refuges are divided into two sets. Populations in south-west France are allied to Spain, while all remaining populations in northern Europe have been colonized from Italy and the Balkans. (iii) The transition from one race to another in south-west France is marked by abrupt transitions in the frequency of refuge-specific private alleles and corresponds closely to the northern limit of the distribution of cork oak. Although hybrids were detected in north-west France, none were detected where the two lifecycles meet in south-western France. The biology of oak gallwasps predicts that any hybrid zone will be narrow, and limited to regions where Q. cerris and Q. suber meet. Our data suggest that eastern and western A. kollari are effectively separate species.
Diversification of insect herbivores is often associated with coevolution between plant toxins and insect countermeasures, resulting in a specificity that restricts host plant shifts. Gall inducers, however, bypass plant toxins and the factors influencing host plant associations in these specialized herbivores remain unclear. We reconstructed the evolution of host plant associations in Western Palaearctic oak gallwasps (Cynipidae: Cynipini), a species-rich lineage of specialist herbivores on oak (Quercus). (1) Bayesian analyses of sequence data for three genes revealed extreme host plant conservatism, with inferred shifts between major oak lineages (sections Cerris and Quercus) closely matching the minimum required to explain observed diversity. It thus appears that the coevolutionary demands of gall induction constrain host plant shifts, both in cases of mutualism (e.g., fig wasps, yucca moths) and parasitism (oak gallwasps). (2) Shifts between oak sections occurred independently in sexual and asexual generations of the gallwasp lifecycle, implying that these can evolve independently. (3) Western Palaearctic gallwasps associated with sections Cerris and Quercus diverged at least 20 million years ago (mya), prior to the arrival of oaks in the Western Palaearctic from Asia 5-7 mya. This implies an Asian origin for Western Palaearctic gallwasps, with independent westwards range expansion by multiple lineages.
The endosymbiont Wolbachia can be responsible for selective sweeps on mitochondrial DNA variability within species. Similar signals can also result from demographic processes, although crucially the latter affect nuclear as well as mitochondrial loci. Here we present data on Wolbachia infection status and phylogeographic patterning for a widely distributed insect host, the oak gallwasp Biorhiza pallida (Hymenoptera: Cynipidae). Two hundred and eighteen females from eight European countries were screened for Wolbachia. All individuals from Hungary, Italy, France, U.K., Ireland, Switzerland, Sweden, and northern and southern Spain were infected with a single group A strain of Wolbachia, while populations in central Spain were not infected. A mitochondrial marker (cytochrome b) shows low variation and departure from neutrality in infected populations, but greater variation and no deviation from neutrality in Wolbachia-free populations. This pattern is compatible with a Wolbachia-induced selective sweep. However, we also find parallel differences between infected and uninfected populations for nuclear markers (sequence data for ITS1 and ITS2). All markers support the existence of a deep split between populations in Spain (some free of Wolbachia), and those in the rest of Europe (all infected). Allelic variation for five allozyme loci is also consistent with the Spain-rest of Europe split. Concordant patterns for nuclear and mitochondrial markers suggest that differences in the nature and extent of genetic diversity between these two regions are best explained by differing demographic histories (perhaps associated with range expansion from Pleistocene glacial refugia), rather than a Wolbachia-associated selective sweep.
We assessed the utility of eight DNA sequence markers (5.8S rDNA, 18S rDNA, 28S rDNA, ITS regions, long-wavelength opsin, elongation factor 1-alpha, cytochrome b, and cytochrome oxidase I) in reconstructing phylogenetic relationships at various levels of divergence in gallwasps (Hymenoptera: Cynipidae), using a set of eight exemplar taxa. We report sequence divergence values and saturation levels and compare phylogenetic results of these sequences analyzed both separately and combined to a well-corroborated morphological phylogeny. Likelihood ratio tests were used to find the best evolutionary model fitting each of the markers. The likelihood model best explaining the data is, for most loci, parameter rich, with strong A-T bias for mitochondrial loci and strong rate heterogeneity for the majority of loci. Our data suggest that 28S rDNA, elongation factor 1-alpha, and long-wavelength opsin may be potentially useful markers for the resolution of cynipid and other insect within-family-level divergences (circa 50-100 mya old), whereas mitochondrial loci and ITS regions are most useful for lower-level phylogenetics. In contrast, the 18S rDNA marker is likely to be useful for the resolution of above-family-level relationships.
Oak gallwasps are cyclically parthenogenetic insects that induce a wide diversity of highly complex species- and generation-specific galls on oaks and other Fagaceae. Phylogenetic relationships within oak gallwasps remain to be established, while sexual and parthenogenetic generations of many species remain unpaired. Previous work on oak gallwasps has revealed substantial intra-specific variation, particularly between regions known to represent discrete Pleistocene glacial refuges. Here we use statistical phylogenetic inference methods on sequence data for a fragment of the mitochondrial cytochrome b gene to reconstruct the relationships among 62 oak gallwasp species. For 16 of these we also include 23 additional cytochrome b haplotype sequences from different Pleistocene refuge areas to test the effect of intra-specific variation on inter-specific phylogeny reconstruction. The reconstructed phylogenies show good intra-generic resolution and identify several conserved clades, but fail to reconstruct either very recent or very ancient divergences. Nine of the 16 species represented by multiple haplotypes are not monophyletic. The apparent discordance between the recovered gene tree and the current taxonomic classification can be explained through: (a) collapsing of some species currently known only from either a sexual or a parthenogenetic generation into a single cyclically parthenogenetic entity; (b) sorting of ancestral polymorphism in diverging lineages, and (c) horizontal transfer of haplotypes, perhaps due to hybridization within glacial refuges. Our conclusions emphasise the need for careful intra-specific sampling when reconstructing phylogenies for radiations of closely related species and imply that for certain taxonomic groups full phylogenetic resolution (using molecular markers) may not be attainable.
The phylogenetic relationships between Dryocosmus, Chilaspis and allied genera of oak gallwasps (Hymenoptera, Cynipidae: Cynipini)
- Z Ács
- G Melika
- Z Penzes
- J Pujade-Villar
- G N Stone
Ács Z, Melika G, Penzes Z, Pujade-Villar J, Stone GN. 2007. The
phylogenetic relationships between Dryocosmus, Chilaspis
and allied genera of oak gallwasps (Hymenoptera, Cynipidae:
Cynipini). Syst Entomol 32(1):70-80. doi:10.1111/j.1365-3113.2006.00351.x.
New species of North American Cynipidae
- H F Bassett
Bassett HF. 1890. New species of North American Cynipidae. T Am
Entomol Soc (1890-) 17(1):59-92.
- P Cameron
Cameron P. 1883. Hymenoptera Biology Centrali-America. Vol. 1.,
497 pp., 120 pl.
Molecular systematics of the Chalcidoidea using 28S-D2 rDNA. Hymenoptera-Evolution
- B Campbell
- J Heraty
- J Y Rasplus
- K Chan
- J Steffen-Campbell
- C Babcock
Campbell B, Heraty J, Rasplus JY, Chan K, Steffen-Campbell J,
Babcock C. 2000. Molecular systematics of the Chalcidoidea
using 28S-D2 rDNA. Hymenoptera-Evolution, Biodiversity and
Biological Control. CSIRO Publishing, Collingwood, pp. 59-73.
New species and synonymy of American Cynipidae
- A C Kinsey
Kinsey AC. 1920. New species and synonymy of American
Cynipidae. B Am Mus Nat Hist 42:293-317.
Description and biology of the sexual generation of Disholcaspis quercusmamma
- C Mcewen
- S Digweed
- J A Nicholls
- W Cranshaw
McEwen C, Digweed S, Nicholls JA, Cranshaw W. 2014. Description
and biology of the sexual generation of Disholcaspis
quercusmamma (Walsh and Riley) (Hymenoptera: Cynipidae),
with notes on associated parasitoids. P Entomol Soc Wash
116(3):294-310. doi:10.4289/0013-8797.116.3.294.
Andricus maesi n. sp. Primera especie de cinípido gallícola para Nicaragua (Hymenoptera: Cynipidae)
- J Pujade-Villar
Pujade-Villar J. 2015. Andricus maesi n. sp. Primera especie de
cinípido gallícola para Nicaragua (Hymenoptera: Cynipidae).