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Revision of the Amphibolips species of the ‘nassa’ complex from Mexico and central America (Hymenoptera: Cynipidae)
Abstract
The cynipid gall wasp species of Amphibolips (Cynipidae: Cynipini) belonging to the “nassa” complex are reviewed for Mexico and Central America. Five new species are described: A. bassae Cuesta-Porta & Pujade-Villar n. sp., A. bromus Pujade-Villar & Cuesta-Porta n. sp., A. kinseyi Cuesta-Porta & Pujade-Villar n. sp., A. rulli Pujade-Villar & Cuesta-Porta n. sp., and A. turulli Pujade-Villar & Cuesta-Porta n. sp. Amphibolips quercuspomiformis (Bassett) comb. nov. is redescribed and Amphibolips malinche Nieves-Aldrey & Maldonado is proposed as a new synonym of A. hidalgoensis Pujade-Villar & Melika. The validity of A. dampfi is discussed. Diagnostic characters, distribution, host-plant relationships, and biology are provided. Also new diagnostic characters and new distribution data for earlier described species are given. Keys to adults and galls for all known “nassa” complex species from Mexico and Central America are presented.
... Taxonomic studies have advanced the classification within the tribe Cynipini, including descriptions of new genera and species and the identification of synonymous species based on morphological traits (Castillejos-Lemus et al., 2020;Cuesta-Porta et al., 2020;Melika, Pujade-Villar, et al., 2021;Nicholls et al., 2018;Nieves-Aldrey et al., 2012;Nieves-Aldrey et al., 2021;Zhang et al., 2021). These efforts require the use of modern tools to clarify some of the remaining taxonomic controversies among closely related groups of wasps. ...
... In particular, Amphibolips induces some of the most charismatic and conspicuous galls, ranging from hollow galls with a central larval chamber suspended by radiating fibres, to juicy and extremely sour gall tissues that are thought to deter potential predators (Guiguet et al., 2023). Significant progress has been made in the description and species delimitation of the species within this genus over the past 15 years (Castillejos-Lemus et al., 2020;Cuesta-Porta et al., 2020, 2021Medianero & Nieves-Aldrey, 2010;Melika et al., 2011;Nicholls et al., 2022;Nieves-Aldrey et al., 2012;Pujade-Villar et al., 2018). ...
... While Cuesta-Porta et al. (2023) placed a species from each complex in a basal clade, this result should be interpreted with caution due to the limited taxon sampling. Some morphological characteristics of the galls and adults in the 'niger' complex have led to the possibility that they may belong to a different genus (Cuesta-Porta et al., 2020Nieves-Aldrey et al., 2012). We consider that it is necessary to include a larger number of species from the 'niger' complex to determine whether they belong to the genus Amphibolips or represent a sister genus. ...
In recent years, new wasp species and genera of Cynipidae have been described, and their species delimitation and evolutionary relationships have been supported using molecular markers. However, few studies have included comprehensive and extensive sampling of specimens across the complete distribution of a single genus. In this study, we analysed the phylogenetic relationships of the genus Amphibolips throughout its range using ultraconserved elements (UCEs). We collected 520 adult wasps from 401 sites in various geographic locations, predominantly in Mexico, and used 78 wasps for UCE analysis. Our results demonstrate that Amphibolips forms a monophyletic clade. Amphibolips can be further subdivided into four genetically well-differentiated clades. Each clade had morphological traits that could be collectively characterised. Species delimitation using molecular sequencing highlights the taxonomic complexity involved in separating and assigning species using exclusively morphological criteria. This approach offers an opportunity to refine current morphological taxonomic criteria or propose new criteria to establish more accurate classifications. The rich diversity of host species and geographic regions in Mexico represents a biodiversity hotspot for Amphibolips. The number of undescribed species remains high. This study facilitates the determination of appropriate boundaries for putative new Amphibolips species.
... The first thorough taxonomic studies of the cynipid fauna of Mexico were performed by Kinsey from the 1920s to 1940s where he described several new genera and broadened the distribution of other existing Nearctic genera ). After Kinsey's contributions, the study of Mexican fauna was not resumed until the first decade of the 2000s with a revisionary process to update the taxonomy of the Nearctic and Neotropical fauna (Melika & Abrahamson 2002;Pujade-Villar et al. 2009, 2013Pujade-Villar & Melika 2014;Pujade-Villar & Ferrer-Suay 2015;Pujade-Villar et al. 2017Cuesta-Porta et al. 2020;Nicholls et al. 2022). To this date, Amphibolips is one of the most actively studied genera in Mexico and the Neotropics, probably due to the large size of the galls and the adults. ...
... Firstly, this demonstrates the existence of previously unrecognised lineages within the 'nassa' species group. Additionally, this second lineage contains multiple morphologically divergent taxa including the 'niger' complex, A. quercusostensackenii and A. quercuspomiformis; the latter species has previously been reported as a morphologically and biologically divergent species within the 'nassa' group (Cuesta-Porta et al. 2020). Amphibolips quercuspomiformis, along with A. salicifoliae Medianero & Nieves-Aldrey, 2010, are the only Amphibolips species exhibiting simple tarsal claws (Medianero & Nieves-Aldrey 2010;Cuesta-Porta et al. 2020). ...
... Additionally, this second lineage contains multiple morphologically divergent taxa including the 'niger' complex, A. quercusostensackenii and A. quercuspomiformis; the latter species has previously been reported as a morphologically and biologically divergent species within the 'nassa' group (Cuesta-Porta et al. 2020). Amphibolips quercuspomiformis, along with A. salicifoliae Medianero & Nieves-Aldrey, 2010, are the only Amphibolips species exhibiting simple tarsal claws (Medianero & Nieves-Aldrey 2010;Cuesta-Porta et al. 2020). Amphibolips quercusostensackenii has strongly lobed tarsal claws but differs from most of the 'nassa' species-group along with A. quercuspomiformis and A. salicifoliae by the shiny smooth speculum (coarsely rugose in the rest of species, except A. aliciae which exhibits finely rugose speculum). ...
Amphibolips is currently divided into two species-groups, clearly differentiated by adult and gall morphology. The ‘niger’ group of Amphibolips species is revised. This complex includes eight species: A. gumia Kinsey, A. jubatus Kinsey, A. elatus Kinsey, A. maturus Kinsey, A. nebris Kinsey, A. niger Kinsey, A. pistrix Kinsey and A. ufo Cuesta-Porta & Pujade-Villar sp. nov. We provide descriptions of new species and re-descriptions of known species, diagnoses, keys to complexes and species. The possibility of the ‘niger’ complex belonging to a new genus is also discussed.
... From Mexico, a total of 205 species were reported until 2022 (Martínez-Romero et al. 2022), 59 of which are associated with the section Lobatae, and among them, five genera including 38 species induce galls exclusively in the section Lobatae: Amphibolips Reinhard, 1865 (29 species;Cuesta-Porta et al. 2020c, 2023a In this study, a new genus Reticulodermis Pujade-Villar, Cuesta-Porta & Melika gen. nov. ...
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.
... Distribución.. Citada de los estados de Ciudad de México (incluida Sta. Fe sobre Q. crassipes) y Zacatecas (descripción original), Cuesta-Porta et al. (2020b). ...
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.
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.
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.
Se describe de México una nueva especie de cinípido de encino, Amphibolips cibriani Pujade-Villar n. sp., conocida sólo a partir de su generación asexual que induce agallas en las yemas de Quercus mexicana Bonpl. (Sección Lobatae). Se dan datos referentes a la diagnosis, distribución y biología de esta nueva especie.
While much has been learned regarding the phylogeny and evolution of cynipoid wasps, clearly illustrated diagnostic tools and identification keys have remained stagnant. So too, where keys do exist, they are often to genus or species, and there are no user-friendly keys to groups such as tribes, subfamilies, or families. This state of affairs leaves a knowledge gap for non-specialists and slows future research on the group. To address this, we provide a fully illustrated key to the higher-level groups of world Cynipoidea. We also provide summaries of all higher-level taxa with updated generic lists, biological data, distribution, and literature resources. The dichotomous key presented here is complimented with a multi-entry matrix-based key, created in Lucid, and served on www.waspweb.org with online versions of the dichotomous keys also available.
The tree of life is highly reticulate, with the history of population divergence emerging from populations of gene phylogenies that reflect histories of introgression, lineage sorting and divergence. In this study, we investigate global patterns of oak diversity and test the hypothesis that there are regions of the oak genome that are broadly informative about phylogeny.
We utilize fossil data and restriction‐site associated DNA sequencing (RAD‐seq) for 632 individuals representing nearly 250 Quercus species to infer a time‐calibrated phylogeny of the world's oaks. We use a reversible‐jump Markov chain Monte Carlo method to reconstruct shifts in lineage diversification rates, accounting for among‐clade sampling biases. We then map the > 20 000 RAD‐seq loci back to an annotated oak genome and investigate genomic distribution of introgression and phylogenetic support across the phylogeny.
Oak lineages have diversified among geographic regions, followed by ecological divergence within regions, in the Americas and Eurasia. Roughly 60% of oak diversity traces back to four clades that experienced increases in net diversification, probably in response to climatic transitions or ecological opportunity.
The strong support for the phylogeny contrasts with high genomic heterogeneity in phylogenetic signal and introgression. Oaks are phylogenomic mosaics, and their diversity may in fact depend on the gene flow that shapes the oak genome.
In this chapter, we review major classification schemes proposed for oaks by John Claudius Loudon, Anders Sandøe Ørsted, William Trelease, Otto Karl Anton Schwarz, Aimée Antoinette Camus, Yuri Leonárdovich Menitsky, and Kevin C. Nixon. Classifications of oaks (Fig. 2.1) have thus far been based entirely on morphological characters. They differed profoundly from each other because each taxonomist gave a different weight to distinguishing characters; often characters that are homoplastic in oaks. With the advent of molecular phylogenetics our view has considerably changed. One of the most profound changes has been the realisation that the traditional split between the East Asian subtropical to tropical subgenus Cyclobalanopsis and the subgenus Quercus that includes all other oaks is artificial. The traditional concept has been replaced by that of two major clades, each comprising three infrageneric groups: a Palearctic-Indomalayan clade including Group Ilex (Ilex oaks), Group Cerris (Cerris oaks) and Group Cyclobalanopsis (cycle-cup oaks), and a predominantly Nearctic clade including Group Protobalanus (intermediate or golden cup oaks), Group Lobatae (red oaks) and Group Quercus (white oaks, with most species in America and some 30 species in Eurasia). In addition, recent phylogenetic studies identified two distinct clades within a wider group of white oaks: the Virentes oaks of North America and a clade with two disjunct endemic species in western Eurasia and western North America, Quercus pontica and Q. sadleriana. The main morphological feature characterising these phylogenetic lineages is pollen morphology, a character overlooked in traditional classifications. This realisation, along with the now available (molecular-)phylogenetic framework, opens new avenues for biogeographic, ecological and evolutionary studies and a re-appraisal of the fossil record. We provide an overview about recent advances in these fields and outline how the results of these studies contribute to the establishment of a unifying systematic scheme of oaks. Ultimately, we propose an updated classification of Quercus recognising two subgenera with eight sections. This classification considers morphological traits, molecular-phylogenetic relationships, and the evolutionary history of one of the most important temperate woody plant genera.
Oaks ( Quercus , Fagaceae) are the dominant tree genus of North America in species number and biomass, and Mexico is a global center of oak diversity. Understanding the origins of oak diversity is key to understanding biodiversity of northern temperate forests.
A phylogenetic study of biogeography, niche evolution and diversification patterns in Quercus was performed using 300 samples, 146 species. Next‐generation sequencing data were generated using the restriction‐site associated DNA ( RAD ‐seq) method. A time‐calibrated maximum likelihood phylogeny was inferred and analyzed with bioclimatic, soils, and leaf habit data to reconstruct the biogeographic and evolutionary history of the American oaks.
Our highly resolved phylogeny demonstrates sympatric parallel diversification in climatic niche, leaf habit, and diversification rates. The two major American oak clades arose in what is now the boreal zone and radiated, in parallel, from eastern North America into Mexico and Central America.
Oaks adapted rapidly to niche transitions. The Mexican oaks are particularly numerous, not because Mexico is a center of origin, but because of high rates of lineage diversification associated with high rates of evolution along moisture gradients and between the evergreen and deciduous leaf habits. Sympatric parallel diversification in the oaks has shaped the diversity of North American forests.
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.
Two new species of oak gallwasp, Amphibolips zacatecaensis Melika & Pujade-Villar and A. hidalgoensis Pujade-Villar & Melika, are described from Mexico, known to induce galls on Quercus eduardi, Q. crassipes, Q. candicans and Q. crassifolia (section Lobatae, red oaks). Only females are known. Data on the diagnosis, distribution and biology of the new species are 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.
Three new species of Amphibolips Reinhard 1865, Amphibolips castroviejoi, A. aliciae, and A. salicifoliae (Hymenoptera: Cynipidae: Cynipini) are described from Panama. The three new species induce galls on Quercus salicifolia Nee (Fagaceae, sect. Lobatae, Red Oaks). Diagnostic characters, gall descriptions, distribution, and biological data are given. The generic limits of Amphibolips are re-established with adding new generic morphological characters. The presence of the genus Amphibolips in the Neotropical region is discussed. The new species represent the first record of this genus from Central America.
Se ha efectuado el primer estudio faunístico y taxonómico de los cinípidos de Panamá, a la vez que se actualiza el conocimiento de las tribus, géneros y especies de esta familia de himenópteros en la Región Neotropical. Los resultados apoyan la
hipótesis de que las montañas de Centroamérica albergan una fauna rica de inductores e inquilinos de agallas de las tribus Cynipini y Synergini (Cynipidae). Como resultado de 20 meses de muestreo en los bosques de Quercus de las montañas de Panamá
se colectaron 65 diferentes agallas inducidas por cinípidos, se obtuvieron los adultos inductores de 45 de ellas, los cuales se clasificaron en 10 géneros de la tribu Cynipini. Se estima que un 94% de las especies aún no han sido descritas. Los géneros de ciní-
pidos inductores identificados son: Andricus, con 12 especies; Neuroterus, con 9; Dryocosmus, con 7 especies; Cynips, con 4 especies; Amphibolips, con 3 especies; Disholcaspis, Loxaulus y Odontocynips con 2 especies cada género; Callirhytis y Bassettia,
con una especie cada uno. Se han identificado 11 especies de inquilinos de la tribu Synergini, 10 del género Synergus, de las cuales 8 son nuevas para la ciencia, y una especie perteneciente al recientemente descrito género Agastoroxenia. Adicionalmente dos
nuevos géneros de cinípidos inductores, aún en periodo de estudio, han sido detectados en el material colectado. La fauna actual
en el Neotrópico de la familia Cynipidae queda actualizada en 6 tribus, 21 géneros y 45 especies, de las cuales 41 son nativas y 4
han sido introducidas en la región. Para Panamá se aporta una lista de 21 especies de la familia Cynipidae, de las cuales 10 son
inductores de agallas y 11 son inquilinas. Por último, se discute el estado actual del conocimiento de las especies de la familia Cynipidae en la República de Panamá.
The species of Amphibolips Reinhard 1865 (Hymenoptera: Cynipidae: Cynipini) of Mexico not included in the “niger complex” Kinsey 1937 are revised. The “nassa complex” recently created by Melika et al. (2011) for these species is questioned. As a result of faunistic surveys of oak gall wasps in Mexico, seven new species of Amphibolips are described: Amphibolips durangensis Nieves-Aldrey & Maldonado, A. malinche Nieves-Aldrey & Pascual, A. jaliscensis Nieves-Aldrey & Pascual, A. oaxacae Nieves-Aldrey & Pascual, A. nevadensis Nieves-Aldrey & Pascual, A. tarasco Nieves-Aldrey & Pascual and A. michoacaensis Nieves-Aldrey & Maldonado. The types of A. dampfi Kinsey 1937, A. nebris Kinsey 1937 and A. fusus Kinsey 1937 are revised, redescribed and compared with the new species described herein. A new key for the identification of adults of all species not included in the “niger complex” from Mexico, including the related species A. castroviejoi Medianero & Nieves-Aldrey and A. aliciae Medianero & Nieves-Aldrey from Panama, is presented. Diagnostic characters, distribution data, host plants and species biology are provided for of all of the studied
species. The richness and distribution of Amphibolips in Mexico and the Neotropical region, their known biological data, and the taxonomic limits of this genus are also discussed.
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.
The knowledge status of the gall-former cynipids in the Mexican oaks is commented herein. For each of the 157 cited species from 33 oak species, the most important biological and ecological information is included. Andricus championi (Cameron), a species described only from the gall, is considered as 'incertae sedis'. Andricus tumeralis Pujade-Villar nom. nov. is proposed to denominate the species Andricus ashmeadi Dalla Torre & Kieffer for being a homonymy of Andricus ashmeadi Basset. Atrusca dugesi (Mayr) n. comb. (formery in Cynips) and Antron sagata (Kinsey) n. comb. (formery in Sphaeroteras) are proposed. The reasons of these changes are commented as well as the need for further studies on Mexican cynipids.
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.
L'heterogònia o alternança generacional és característica de dos tribus de Cynipidae: Pediaspidini (amb 2 espècies conegudes) i Cynipini (amb més de 900 espècies descrites). Només uns 85 cicles són coneguts, per la qual cosa la majoria d'ells estan per descriure. En aquest treball es donen dades referides a la biologia dels Cynipini, es fa un repàs històric del coneixement de la heterogònia en aquest grup d'himenòpters, es dónen dades dels diferents models que els caracteritzen, es dicuteix l'origen del caràcter heterogònic i s'esmenten aquells cicles coneguts a la bibliografia. També es donen dades de les referències bibliogràfiques referents a aquesta problemàtica.
Se presenta una lista preliminar de 161 especies del género Quercus para México, ubicadas en tres secciones: 76 en la sección Lobatae (encinos rojos), 81 en la sección Quercus (encinos blancos) y cuatro especies en la sección Protobalanus (encinos intermedios). Se calcula que 109 especies son endémicas del país, de las cuales 47 pertenecen a la sección Quercus, 61 a la sección Lobatae y una a Protobalanus. México comparte con Estados Unidos 33 especies del género, mientras que con Centroamérica comparte 20. Los estados con mayor diversidad de especies son Oaxaca, Nuevo León, Jalisco, Chihuahua y Veracruz. Las especies con distribución más amplia en nuestro país son Q. candicans, Q. castanea, Q. crassifolia, Q. laeta, Q. microphylla, Q. obtusata y Q. rugosa. Altitudinalmente las especies de Quercus se desarrollan entre 0 y 3,500 m, pero son más frecuentes entre 1,000 y 3,000 m. El conocimiento del género Quercus en México es aún deficiente y se necesita realizar más estudios en torno a este importante género.
Oak gall wasps (Hymenoptera: Cynipidae, Cynipini) are characterized by possession of complex cyclically parthenogenetic life cycles and the ability to induce a wide diversity of highly complex species- and generation-specific galls on oaks and other Fagaceae. The galls support species-rich, closed communities of inquilines and parasitoids that have become a model system in community ecology. We review recent advances in the ecology of oak cynipids, with particular emphasis on life cycle characteristics and the dynamics of the interactions between host plants, gall wasps, and natural enemies. We assess the importance of gall traits in structuring oak cynipid communities and summarize the evidence for bottom-up and top-down effects across trophic levels. We identify major unanswered questions and suggest approaches for the future.
Cynipid gallwasps comprise 1364 species worldwide, predominantly in temperate regions of the Holarctic. The vast majority of recorded species are from the Nearctic and the Western Palaearctic, both of which are long-standing centers of research on the taxonomy and biology of this group. In contrast, the Eastern Palaearctic and the Oriental Region faunas are much less studied, but potentially extremely rich. Previously recorded species richness of cynipid groups in the Eastern Palaearctic and Oriental regions probably represents a significant underestimate. This is particularly marked for gallwasps associated with oaks (tribe Cynipini), as has been shown by recent works, which describe many new species galling oaks in the subgenera Quercus and Cyclobalanopsis and plants of the other Fagaceae genera (Castanopsis, Lithocarpus and Castanea). In the present paper, we summarise the current knowledge of oak gallwasps in the regions that extend from Pakistan, through Nepal and China to Japan and south to Indonesia, and list a total of 97 species of Cynipini from the area.
Mesoscutum uniformly rugate. Notauli traceable only in posterior third, furrow deep, broad, and smooth. Median propodeal area smooth and glabrous. Metasomal terga smooth
- . . . . Depressed Posteriorly
depressed posteriorly........................................................................ 3
2.
Mesoscutum uniformly rugate. Notauli traceable only in posterior third, furrow deep, broad, and smooth. Median propodeal
area smooth and glabrous. Metasomal terga smooth, without micropunctures. Forewing almost hyaline. Males and females..
8 7. Forewing with anal and basal half of discoidal cells hyaline (Figs 10A, 13D)
- . . . Propodeal Carinae Indistinct
Propodeal carinae indistinct............................................................................. 8
7.
Forewing with anal and basal half of discoidal cells hyaline (Figs 10A, 13D). Area aside of parapsidal lines rugose (Fig. 10C).
Forewing with discoidal and anal cells completely infuscate (Fig.13C)
- . . . . Only Females
Only females..............................................................................A. turulli n. sp.
7'.
Forewing with discoidal and anal cells completely infuscate (Fig.13C). Area aside of parapsidal lines coriaceous (Fig. 12B).
Scutellar foveae quadrangular, limited posteriorly by transverse carinae never by rugose sculpture, median carina as long as length of foveae; posterior depression reaching scutellar foveae, with transverse carinae, smooth interspaces (Figs 11D, 4C)
- Females . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Males
Males and females............................................................................. A. tarasco
20'.
Mesoscutum uniformly rugose, with similar cells, notauli and medial sulci inconspicuous (Fig. 11D). Lateral margin of eyes
rugose-coriaceous, not emarginate (Fig. 4A). Scutellar foveae quadrangular, limited posteriorly by transverse carinae never
by rugose sculpture, median carina as long as length of foveae; posterior depression reaching scutellar foveae, with transverse
carinae, smooth interspaces (Figs 11D, 4C). Metascutellum quadrate, less than 1.8x as broad as high (Fig. 4E). Only females.
....................................................................................... A. hidalgoensis
1.
Galls growing on the underside of the leaves................................................................ 2
1'.
Galls growing on buds................................................................................. 4
2.
Turgescent spherical galls (Figs 14F-G). Turning into a raisin-like wrinkled mass when dry...............A. turulli n. sp.
2'.
Solid galls, sometimes very fragile, never turgescent. Galls either spherical or toadstool shaped. Maintaining its shape when
dry................................................................................................ 3
3.
Toadstool-shaped galls. With an elongated stem. Light green coloured, sometimes tinged with rose or pink when young. Internal solid spongious parenchyma (Lyon 1959: Plate 12, Figs A-B).................A. quercuspomiformis n. comb. (sex).
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