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Revision of world Ceroptresini (Hymenoptera: Cynipidae) with the description of a new genus and five new species
Abstract
A taxonomic revision of the tribe Ceroptresini (Hymenoptera: Cynipidae) is conducted for the first time. Prior to this study, the total number of valid species of Ceroptres, the only genus within Ceroptresini to date, was 23. As a result of this revision, 15 Ceroptres species are retained as valid and one species, Amblynotus ensiger Walsh, 1864, is desynonymized from Ceroptres petiolicola (Osten-Sacken, 1861), being considered here as a valid Ceroptres species: C. ensiger (Walsh, 1864) status verified and comb. nov. An additional five new species are described from Mexico: Ceroptres junquerasi Lobato-Vila & Pujade-Villar sp. nov.; C. lenis Lobato-Vila & Pujade-Villar sp. nov.; C. mexicanus Lobato-Vila & PujadeVillar sp. nov.; C. nigricrus Lobato-Vila & Pujade-Villar sp. nov.; C. quadratifacies Lobato-Vila & Pujade-Villar sp. nov., increasing the total number of valid Ceroptres species to 21. Ceroptres masudai Abe, 1997 is synonymized with C. kovalevi Belizin, 1973. Ceroptres niger Fullaway, 1911 is transferred to Andricus (Andricus confusus Lobato-Vila & Pujade-Villar comb. nov. and nom. nov.). Five species (Amblynotus inermis Walsh, 1864; Cynips quercusarbos Fitch, 1859; Cynips querficus Fitch, 1859; Cynips quercuspisum Fitch, 1859; and Cynips quercustuber Fitch, 1859) are not considered as valid Ceroptres. The status of Ceroptres quereicola (Shinji, 1938), previously classified as an unplaced species, is commented on. In addition, a Nearctic species from the USA, Ceroptres politus Ashmead, 1896, is here proposed as the type species of a new genus within Ceroptresini: Buffingtonella Lobato-Vila & Pujade-Villar gen. nov. Redescriptions, biological and distribution data, illustrations and keys to genera and species within Ceroptresini are provided. The diagnostic morphological traits of Ceroptresini, Ceroptres and the new genus are discussed.
We review the genera of Phanacidini and provide updated diagnoses for each genus and a new key to the four genera. We also redescribe the enigmatic herb gall wasp Diakontschukia saussureae Diakontschuk, 2001 (Hymenoptera: Cynipidae: Phanacidini), based on a specimen collected in Haopingsi Nature Reserve, China. This is the first record of the tribe Phanacidini in China. Lastly, we provide extensive commentary on the validity of diagnostic characters that have been used for Phanacidini relative to other herb gall wasp tribes. We carried out a survey of Aulacideini, Aylacini, and Phanacidini to identify whether the length of F1 relative to F2 was of diagnostic value at the tribal level, finding that F1 was always conspicuously longer than F2 in Phanacidini and Aylacini but at most subequal to F2 in Aulacideini, suggesting that this is indeed a valuable character for diagnosing herb gall wasp tribes. Other characters discussed are the closure of the marginal cell and the dimensions of the mesopectus.
Distributional checklists of the extant, described species of five superfamilies of Hymenoptera of Canada, Alaska and Greenland are presented. In total, 296 species in 79 genera in 12 families are recorded: 55 species of Ceraphronoidea, classified in 10 genera in 2 families, 205 species of Cynipoidea in 58 genera in 5 families, 30 species of Evanioidea in 5 genera in 3 families of Evanioidea, 2 species of Stephanoidea in 2 genera in 1 family and 4 species of Trigonalyoidea in 4 genera in 1 family. Of the reported species, 281 (in 79 genera in 12 families) are listed from Canada, 31 (in 16 genera in 6 families) from Alaska, and 7 (in 5 genera in 2 families) from Greenland. The list includes 8 new generic records for Canada (1 Ceraphronoidea, 6 Cynipoidea and 1 Evanioidea) and 43 new Canadian species records (13 Ceraphronoidea, 28 Cynipoidea and 2 Evanioidea). For each species in Canada, distribution is tabulated by province or territory, except the province of Newfoundland and Labrador is divided into the island of Newfoundland and the region of Labrador. These checklists are compared with previous Nearctic and Palaearctic surveys, checklists and catalogues. Kleidotoma minima Provancher, 1883 (Figitidae) is moved from this genus to Hexacola Förster, 1869 to form H. minimum (Provancher, 1883), comb. nov.Amblynotus slossonae Crawford, 1917 (Figitidae) is moved from Melanips Walker, 1835 to Amphithectus Hartig, 1840 forming A. slossonae (Crawford, 1917), comb. nov.
We describe new species in the genus Ceroptres Hartig, 1840 (Hymenoptera: Cynipidae: Ceroptresini) represented by voucher material sequenced by Ward et al. (2024). We describe 22 new species, all authored by Nastasi, Smith, & Davis: C. anansii sp. nov., C. anzui sp. nov., C. bruti sp. nov., C. curupira sp. nov., C. daleki sp. nov., C. dandoi sp. nov., C. demerzelae sp. nov., C. iktomii sp. nov., C. jabbai sp. nov., C. jarethi sp. nov., C. lokii sp. nov., C. lupini sp. nov., C. mallowi sp. nov., C. promethei sp. nov., C. sandiegoae sp. nov., C. selinae sp. nov., C. soloi sp. nov., C. songae sp. nov., C. swiperi sp. nov., C. thrymi sp. nov., C. tikoloshei sp. nov., and C. zorroi sp. nov. After our taxonomic treatment, the genus Ceroptres includes 43 species, all but three of which are known from North America. Among our new species are two reared from cecidomyiid midge galls, an association previously recorded but without valid taxonomic association. We provide new records for two additional previously described species; we record C. ensiger (Walsh, 1864) from Pennsylvania and confirm characters for the male, and we record C. lanigerae Ashmead, 1885 from Texas. We also examined several putative species corresponding to either C. cornigera Melika & Buss, 2002 and/or C. frondosae Ashmead, 1896, which we regard as a species complex that requires elucidation in future studies. To enable further studies on Ceroptres, we provide an updated key to North American females. Overall, we find that species of Ceroptres are host specialists associated with a single host gall species or several galls that are phylogenetically or ecologically related. We suggest that there are many North American species of Ceroptres, possibly hundreds, still awaiting collection and characterization.
Robust keys exist for the family-level groups of Cynipoidea. However, for most regions of the world, keys to genera are not available. To address this gap as it applies to North America, a fully illustrated key is provided to facilitate identification of the tribes and genera of rose gall, herb gall, and inquiline gall wasps known from the region. For each taxon covered, a preliminary diagnosis and an updated overview of taxonomy, biology, distribution, and natural history are provided.
Host shifts to new plant species can drive speciation for plant-feeding insects, but how commonly do host shifts also drive diversification for the parasites of those same insects? Oak gall wasps induce galls on oak trees and shifts to novel tree hosts and new tree organs have been implicated as drivers of oak gall wasp speciation. Gall wasps are themselves attacked by many insect parasites, which must find their hosts on the correct tree species and organ, but which also must navigate the morphologically variable galls with which they interact. Thus, we ask whether host shifts to new trees, organs, or gall morphologies correlate with gall parasite diversification. We delimit species and infer phylogenies for two genera of gall kleptoparasites, Synergus and Ceroptres, reared from a variety of North American oak galls. We find that most species were reared from galls induced by just one gall wasp species, and no parasite species was reared from galls of more than four species. Most kleptoparasite divergence events correlate with shifts to non-ancestral galls. These shifts often involved changes in tree habitat, gall location, and gall morphology. Host shifts are thus implicated in driving diversification for both oak gall wasps and their kleptoparasitic associates.
Host shifts to new plants can drive speciation for plant-feeding insects, but how commonly do host shifts also drive diversification for the parasites of those same insects? Oak gall wasps induce galls on oak trees and shifts to novel tree hosts and new tree organs have been implicated as drivers of oak gall wasp speciation. Gall wasps are themselves attacked by many insect parasites, which must find their hosts on the correct tree species and organ, but which also must navigate the morphologically variable galls with which they interact. Thus, we ask whether host shifts to new trees, organs, or gall morphologies correlate with gall parasite diversification. We delimit species and infer phylogenies for two genera of gall kleptoparasites, Synergus and Ceroptres, reared from a variety of North American oak galls. We find that most species were reared from galls induced by just one gall wasp species, and no parasite species was reared from galls of more than four species. Most kleptoparasite divergence events correlate with shifts to non-ancestral galls. These shifts often involved changes in tree habitat, gall location, and gall morphology. Host shifts are thus implicated in driving diversification for both oak gall wasps and their kleptoparasitic associates.
Field studies were conducted between September 2020 and June 2022 to assess the gall wasp (Hymenoptera: Cynipidae) fauna of Burdur province (Southwestern Türkiye). A total of 69 species in 18 genera were recorded. Gall samples were collected from 150 samples collected in 100 different localities in the surveyed area. The identified species include three new records (Andricus melikai Pujade-Villar & Kwast, 2002; A. pseudocecconii Melika, Tavakoli & Stone, 2022; Saphonecrus undulatus (Мауr, 1872)) for the Turkish cynipid fauna. As a result of this study, 44 species of the current Turkish cynipid fauna were recorded from Burdur province for the first time. A checklist of the species and their host plants and/or host galls recorded in the study area is provided. The gall wasp richness and geomorphic-topographic structure of Burdur province is discussed.
The identities of most arthropod associates of cynipid-induced oak galls in the western Palearctic are generally known. However, a comprehensive accounting of associates has been performed for only a small number of the galls induced by the estimated 700 species of cynipid gall wasps in the Nearctic. This gap in knowledge stymies many potential studies of diversity, coevolution, and community ecology,
for which oak gall systems are otherwise ideal models. We report rearing records of insects and other arthropods from more than 527,306 individual galls representing 201 different oak gall types collected from 32 oak tree species in North America. Of the 201 gall types collected, 155 produced one or more arthropods. A total of 151,075 arthropods were found in association with these 155 gall types, and of
these 61,044 (40.4%) were gall wasps while 90,031 (59.6%) were other arthropods. We identified all arthropods to superfamily, family, or, where possible, to genus. We provide raw numbers and summaries of collections, alongside notes on natural history, ecology, and previously published associations for each taxon. For eight common gall-associated genera (Synergus, Ceroptres, Euceroptres, Ormyrus, Torymus, Eurytoma, Sycophila, and Euderus), we also connect rearing records to gall wasp phylogeny, geography, and ecology - including host tree and gall location (host organ), and their co-occurrence with other insect genera. Though the diversity of gall wasps and the large size of these communities is such that many Nearctic oak gall-associated insects still remain undescribed, this large collection and identification effort should facilitate the testing of new and varied ecological and evolutionary hypotheses in Nearctic oak galls.
The galls induced by cynipids in host oaks have been proposed as biodiversity hotspots. They constitute support for the development of arthropod communities of great diversity and functional complexity that find in them nutrition and protection from external abiotic and biotic elements which change according to environmental gradients. We characterize the abundance, richness, and diversity of gall-inducing cynipids associated with the galls of the host oak Quercus rugosa, through an elevation gradient. This study was conducted on 140 oak trees belonging to seven populations (20/site) in central Mexico. We identified 39 gall-inducing cynipid species (Cinipini), three inquiline species of galls (Synergini), and seven parasitoids of gall-inducing cynipid (Chalcidoidea), and the secondary fauna was characterized by eleven arthropod orders. We detected a positive and significant effect exerted by the altitudinal gradient on abundance, species richness, and diversity of gall-inducing cynipids, parasitoids, inquilines, and secondary fauna (order richness level). We documented positive and significant relationships between the gall-inducing cynipids and their parasitoids, inquilines, and secondary fauna in terms of species richness, diversity, and abundance of individuals. Arthropod community composition associated with gall-inducing cynipids differed among localities. This study suggests that gall-inducing cynipids in Q. rugosa act as ecosystem engineers that modify the host plant to offer a new habitat to other arthropods. Also, our results supported that Cynipid gall acts to protect the galls-associated arthropod community from unfavorable abiotic conditions throughout the altitudinal gradient.
La introducción de Q. virginiana y Q. palustris en la Ciudad de Chihuahua (México) a partir de árboles provenientes de viveros localizados en los estados de Tamaulipas y Nuevo León, ha provocado la introducción de Andricus quercuslanigera (Ashmead, 1881) (= A. linaria (Kinsey, 1937) n. syn.). Esta especie agalladora de encinos, conocida solamente a partir de su forma agámica, ocasiona en la zona de origen agallas esféricas aterciopeladas en el envés de las hojas de Q. geminata, Q. minima y Q. virginiana. Su presencia en Chihuahua está ocasionando un daño estético de proporciones importantes en los encinos plantados provocando extrañamiento y preocupación en las autoridades estatales en materia sanitaria forestal y en los habitantes, no solo por el impacto de daño potencial que puede ocasionar este insecto sobre los Q. virginiana plantados, sino también por su posible efecto en otras especies de Quercus nativas de Chihuahua. En este estudio se abordan todas éstas problemáticas.
A review of the inquiline oak gall wasp species of the genus Synergus
Hartig that are associated with tuberous and other tumor-like galls on oaks (Quercus L.) from the New World is provided for the first time. Redescriptions, remarks, pictures and new location data for the species previously known from the New World are given. Three new species that emerged from tuberous galls collected in Mexico are formally described and illustrated: Synergus grahami Lobato-Vila and PujadeVillar, sp. nov.; S. shorthousei Lobato-Vila and Pujade-Villar, sp. nov.; and S. tenebrosus Lobato-Vila and Pujade-Villar, sp. nov. A morphological key to identify and separate all the species addressed in this work is given. In addition, Saphonecrus brevicornis (Ashmead, 1896) is returned to Synergus; a new name for Synergus luteus Nieves-Aldrey and Medianero, 2011 is given to avoid a primary homonymy with an European species; Synergus multiplicatus Fullaway, 1911 becomes a syn. nov. of S. mexicanus Gillette, 1896, and S. davisi (Beutenmu¨ller, 1907) a syn. nov. of S. lignicola (Osten-Sacken, 1862).
The inquiline oak gall wasp fauna emerged from agamic galls of
Andricus quercuslaurinus Melika and Pujade-Villar, 2009, a dangerous pest for
Quercus affinis Scheidw. and Q. laurina Humb. et Bonpl. forests in Mexico, was
studied for the first time. Three species in the genus Synergus Hartig, 1840
(Synergini) were obtained from these galls: Synergus davisi (Beutenmüller, 1907),
previously known only from the United States of America; Synergus estradae
Pujade-Villar and Lobato-Vila, 2016, described from the state of Morelos (Mexico);
and Synergus pseudofilicornis Lobato-Vila and Pujade-Villar n. sp., so far
determined as Synergus filicornis Cameron, 1883 (= Synergus furnessana Weld,
1913) and here considered as a new species. Synergus pseudofilicornis n. sp. is
described and illustrated. A redescription and photographs of S. davisi, as well as
new distributions and host associations of S. davisi and S. estradae, are given. The
value of the inquilines as biological control agents for A. quercuslaurinus is
discussed, and a preliminary list of parasitoids obtained from the galls is presented.
A new species, Synergus mongolicus Pujade-Villar & Wang n. sp., is described from galls collected on twigs of Quercus mongolica. The morphological characters useful for differentiating the new species are also discussed. Three inquilines species are recorded from China for the first time: Saphonecrus shirakashii, Synergus belizinellus and S. symbioticus. Saphonecrus tianmushanus is considered a n. syn. of S. shirakashii based on comparison of the types. Saphonecrus taiwanensis Pujade-Villar new name is proposed for Saphonecrus lithocarpii Schwéger & Melika, 2015 by homonymy with S. lithocarpi Pujade-Villar, Guo, Wang & Chen, 2015 and finally Saphonecrus chinensis Tang & Schwéger, 2015 is considered a n. syn. of S. lithocarpi Pujade-Villar, Guo, Wang & Chen, 2015.
A new term, agastoparasitism, is proposed for parasitism among closely related species. Cynipid inquilines are typical agastoparasites. They cannot induce galls; instead their larvae live inside the galls formed by other cynipids. As in many other groups of agastoparasites, there are two competing hypotheses for the evolutionary origin of cynipid inquilines: either they arose from one of their cynipid hosts, and later radiated to exploit other gall-inducing cynipids (monophyletic origin), or they arose repeatedly, each inquiline from its host (polyphyletic origin). These hypotheses for the origin of cynipid inquilines were tested by a phylogenetic analysis of representative species of cynipid gall inducers and inquilines based on adult morphological characters. The analysis supported the monophyly of the inquilines and indicated an origin from gall inducers related to the genus Diastrophus, one of the current host groups. To examine whether the result of the analysis was influenced by convergent similarities among inquilines because of their similar mode of life, all putative apomorphies shared by some or all of the inquilines but not occurring in any of the gall inducers were removed. Despite this, the phylogenetic conclusions essentially remained the same, that is, the support for inquiline monophyly was not caused by convergent evolution. Based on these results, adaptive aspects of the evolutionary origin and maintenance of cynipid inquilinism are discussed, as well as general patterns in the evolution of agastoparasitism.
Seven new cynipid species are described from Arizona and western Texas: Ceropteres snellingi, Xanthoteras pungens, X. tuckeri, Antron daileyi, A. franklinensis, A. madera, and Andricus flocculentus. Appropriate morphological features of the species are illustrated. Included is a key to the described United States species of Antron.