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Avispas agalladoras de los encinos de Santa Fe (Ciudad de México, México) (Hymenoptera, Cynipidae: Cynipini; Fagaceae)
Abstract
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.
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.
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.
Background: Parasitoidism, a specialized life strategy in which a parasite eventually kills its host, is frequently found
within the insect order Hymenoptera (wasps, ants and bees). A parasitoid lifestyle is one of two dominant life strategies
within the hymenopteran superfamily Cynipoidea, with the other being an unusual plant-feeding behavior
known as galling. Less commonly, cynipoid wasps exhibit inquilinism, a strategy where some species have adapted to
usurp other species’ galls instead of inducing their own. Using a phylogenomic data set of ultraconserved elements
from nearly all lineages of Cynipoidea, we here generate a robust phylogenetic framework and timescale to understand
cynipoid systematics and the evolution of these life histories.
Results: Our reconstructed evolutionary history for Cynipoidea differs considerably from previous hypotheses.
Rooting our analyses with non-cynipoid outgroups, the Paraulacini, a group of inquilines, emerged as sister-group to
the rest of Cynipoidea, rendering the gall wasp family Cynipidae paraphyletic. The families Ibaliidae and Liopteridae,
long considered archaic and early-branching parasitoid lineages, were found nested well within the Cynipoidea as
sister-group to the parasitoid Figitidae. Cynipoidea originated in the early Jurassic around 190 Ma. Either inquilinism
or parasitoidism is suggested as the ancestral and dominant strategy throughout the early evolution of cynipoids,
depending on whether a simple (three states: parasitoidism, inquilinism and galling) or more complex (seven states:
parasitoidism, inquilinism and galling split by host use) model is employed.
Conclusions: Our study has significant impact on understanding cynipoid evolution and highlights the importance
of adequate outgroup sampling. We discuss the evolutionary timescale of the superfamily in relation to their insect
hosts and host plants, and outline how phytophagous galling behavior may have evolved from entomophagous,
parasitoid cynipoids. Our study has established the framework for further physiological and comparative genomic
work between gall-making, inquiline and parasitoid lineages, which could also have significant implications for the
evolution of diverse life histories in other Hymenoptera.
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 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.
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).
A new genus of oak gallwasp, Kokkocynips Pujade-Villar & Melika gen. n., is described from Mexico. Diagnostic characters and generic limits of the new genus are discussed in detail. Galls were found on branches of Quercus acutifolia Nee. Diagnostic characters, distribution and biology of the new species are described and illustrated.
A new species of oak gall wasp, Andricus sphaericus Pujade-Villar n. sp., is described from Mexico. The species is known only from sexual generation. It induces galls on Quercus rugosa Née y Q. obtusata Humb. & Bonpl. (Fagaceae) leaves. It represents the second Andricus Hartig sexual generation collect from Mexico. Data on the diagnosis, distribution, and biology of the new species are given.
Key words: Hymenoptera, Cynipidae, Andricus, new species, Q. rugosa, Mexico
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.
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.
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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.
A revision of tuberous oak galls induced by Andricus species from Mexico is given. Andricus guanajuatensis Pujade-Villar n. sp., A. santafe Pujade-Villar n. sp. and A. carrilloi Pujade-Villar n. sp. (Hymenoptera: Cynipidae: Cynipini) are described from Mexico. Diagnosis, biology and distribution data of all Andricus species are given. Some morphological characters are discussed, illustrated and compared between related species. A key for all Mexican species and two North American species collected next to Mexico (Andricus wheeleri and A. lebaue Pujade-Villar n. sp.) is included. The gall and host of Andricus dugesi Beutenmüller is described for the first time.
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.
A new species of Zapatella Pujade-Villar and Melika, Z. polytryposa Pujade-Villar and Fernández-Garzón n. sp., from Mexico is described here. It is an agamic form, potentially dangerous, killing branches of Quercus crassipes Humb. and Bonpl. Diagnostic data, distribution, and biology of the new species are provided. Principal morphological characters of the new species are described and illustrated. The damage also was described. Zapatella is mentioned for the first time for Mexico; previously, it was known from the USA (six species), Costa Rica (one species), and Colombia (seven species).
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.
A new species of oak gall wasp, Andricus breviramuli Pujade-Villar n. sp., is described from Mexico. The species is known only from asexual females. Data on the diagnosis, distribution, and biology of the new species are given. The wasp induces galls on twigs and young shoots of Quercus laeta Liebm. This gall wasp seems to be a serious pest of Q. laeta in Santa Fe (Delegación Cuajimalpa, DF), México City.
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