Research Article |
Corresponding author: Dohuglas Eliseo Castillejos-Lemus ( castillejos.lemus@gmail.com ) Academic editor: Andreas Köhler
© 2020 Dohuglas Eliseo Castillejos-Lemus, Ken Oyama, José Luis Nieves-Aldrey.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Castillejos-Lemus DE, Oyama K, Nieves-Aldrey JL (2020) Description of three new species of oak gallwasps of the genus Amphibolips Reinhard from Mexico (Hymenoptera, Cynipidae). ZooKeys 987: 81-114. https://doi.org/10.3897/zookeys.987.51366
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Three new species of oak gall wasps of the genus Amphibolips Reinhard, 1865 (Hymenoptera: Cynipidae: Cynipini) are described from Mexico: Amphibolips magnigalla Nieves-Aldrey & Castillejos-Lemus, Amphibolips kinseyi Nieves-Aldrey & Castillejos-Lemus and Amphibolips nigrialatus Nieves-Aldrey & Castillejos-Lemus. The specimens of the first two species were representative of sexual generations and come from the State of Oaxaca, while only a female, collected in the State of Veracruz, is described for A. nigrialatus. The new species induces galls on Quercus zempoaltepecana and Q. sapotifolia (Fagaceae, section Lobatae, red oaks). Descriptions of the diagnostic morphological characteristics of the three species and a key for their identification are provided. The taxonomic relationships of the new species with other species of Amphibolips are discussed; the three new species are closely allied amongst themselves and are related to A. dampfi Kinsey, 1937. With the three newly-described species, the number of Amphibolips in Mexico is increased to 23.
Amphibolips, Cynipini, Lobatae, Mexico, oak apple gall, oak gallwasps, Quercus
Oak gall wasps (Cynipidae: Cynipini) include approximately 41 genera with circa 1,000 species (
The Nearctic region, particularly Mexico, is one of the centres of diversity of the oak gall wasps, which have been estimated to include more than 700 species (
Amphibolips is exclusively associated with the Lobatae section of Quercus genus and is restricted to the American continent (
The morphological characteristics of adults and their galls are very uniform amongst the most well-known Amphibolips species. The galls induced by species of this genus develop mainly in buds, stems or leaves and are rarely found in acorns. They are usually globose or spindle-shaped and detachable, with a spongy parenchyma surrounding a central larval cell, sometimes supported by radiating filaments (
Before 1937, only two species had been described in Mexico (A. palmeri Basset, 1890 and A. nigra Beutenmüller, 1911) (
The objective of this study is to present a description of three new species of the Amphibolips genus in Mexico. One of these species represents the first record of Amphibolips for the State of Veracruz (A. nigrialatus Nieves-Aldrey & Castillejos-Lemus, new species) and the other two species were collected in the State of Oaxaca. One of the species from Oaxaca induces a strikingly-characteristic gall (A. magnigalla Nieves-Aldrey & Castillejos-Lemus, new species), while the other species from Oaxaca (A. kinseyi Nieves-Aldrey & Castillejos-Lemus, new species) shares characteristics with A. magnigalla and with another species previously described from Oaxaca (A. dampfi Kinsey, 1937). The species richness of this genus in Mexico is discussed, as well as the taxonomic problems existing within the group. An update to the identification key given in
Quercus species of the Lobatae section were sampled in Veracruz State in 2008 and in Oaxaca in 2018. The galls were collected directly from oak trees and stored in plastic containers with plastic or mesh lids until the emergence of the wasps. The emergence of the wasps occurred under laboratory conditions. The voucher specimens and their galls were deposited in the entomological collections of the Museo Nacional de Ciencias Naturales in Madrid, Spain and in the Colección Nacional de Insectos of the Instituto de Biología, Universidad Nacional Autónoma de México, Mexico City, Mexico. The Quercus species were identified by Dr Susana Valencia-Ávalos at the Facultad de Ciencias of the Universidad Nacional Autónoma de México (
The type specimen of Amphibolips dampfi Kinsey, 1937 was examined for comparison with the new species described. The male holotype of this species was borrowed from the American Museum of Natural History, New York (AMNH) (James Carpenter).
The images used for the morphological descriptions were taken with a FEI Quanta 200 (Oregon, EU) scanning electron microscope (SEM) in Madrid (Spain) and with a JEOL JSM-IT300 (Tokyo, Japan) SEM in Morelia (Mexico). For the SEM observations, two strategies were followed, depending on the number of individuals available for a given species. For the preservation of some unique specimens mounted in a conventional manner, a low vacuum technique was used without gold coating. When the number of specimens allowed it, some specimens were dissected in 99% alcohol and mounted in stubs to be coated with gold and observed with a high vacuum technique. The forewings were mounted on slides with euparal and examined with a Wild MZ8 and an Olympus SZX10 stereomicroscope. Images of the wings and adult habitus were acquired with a NIKON Coolpix 4500 digital camera attached to a Wild MZ8 light microscope, with the exception of some images taken with an Olympus SC100 camera with the help of CELLSENS STANDARD software. Measurements were made with a micrometric eyepiece calibrated to a Wild M5A stereomicroscope. Photographs of galls in the field and of gall dissections were taken with a Nikon D5300 camera.
The terminology of the morphological structures and abbreviations follow that of
Holotype
: 1♀ in the Museo Nacional de Ciencias Naturales, Madrid, Spain (
Named after the strikingly-large size of the galls of this species.
This new species belongs to the group of Amphibolips species that have a forewing with a transversal clear band that is variable in size and extends towards medial and cubital veins to the ventral margin of the wing (
Amphibolips magnigalla shares with Amphibolips dampfi, A. castroviejoi and the other two new species described herein, a mesoscutellum emarginate posteriorly. However, the emargination is comparatively less deep in A. magnigalla (Figs
Regarding the gall, the new species is easily distinguished by its large spindle-shaped gall (approximately 10 cm in length × 2.5 cm in diameter), which is at least 2× larger than any other spindle-shaped gall described from Mexico. Amphibolips fusus and A. durangensis induce galls morphologically similar to the gall of the new species. However, besides the differences in size, the inner structure of the gall induced by the referenced species is different, being filled with a dense soft tissue, while the inner structure of the gall induced by the new species is often almost empty, with visible radiating filaments from the central larval cell.
Body length: 5.8 mm (n = 1) for females; 5.2 mm (n = 3) for males.
Female (Fig.
Head, in dorsal view 2.3× wider than long. POL:OOL:DOL as 23:44:14. Head in anterior view (Fig.
Head posterior view (male) (Fig.
Mouthparts (male) (Fig.
Antenna (Fig.
Mesosoma in lateral view (Fig.
Mesonotum. Mesoscutum (Fig.
Metanotum. Metapectal-propodeal complex. Metapleural sulcus reaching posterior margin of mesopectus at about mid-height of metapectal-propodeal complex. Metascutellum rugose; metanotal trough smooth and pubescent. Median propodeal area (Fig.
Legs. Densely pubescent; femora and tibiae robust. Metatibia about as long as metatarsus; apical margin of metatarsomeres 1–4, with long strong erect setae. Metatarsal claws with strong triangular basal lobes or teeth.
Forewing (Fig.
Metasoma (Fig.
Male (Figs
Gall (Fig.
The galls develop on twigs of Quercus zempoaltepecana Trel. The gall closely resembles that of Amphibolips durangensis Nieves-Aldrey & Maldonado, 2012. However, the gall of A. magnigalla is distinguished by its larger size, which is at least 2× longer than that of A. durangensis and by its different internal structure, which is filled with less dense spongy tissue and radiant filaments (easily visible in the older galls).
A. magnigalla was found only in one site: Comaltepec (Oaxaca State, Mexico). The galls were relatively abundant on a single isolated tree, while we did not find galls on the nearby trees.
Sexual generation. The galls were collected at the end of April and the insects emerged shortly thereafter, in early May. It seems that it is normal for many insects to feed on the tissue of this species. A detached gall was observed in a field, relatively far from the tree, probably carried by a bird.
Holotype
: 1♀ in the Museo Nacional de Ciencias Naturales (
Named after Dr Alfred Kinsey, one of the most prominent cynipidologists and the pioneer of the study of Amphibolips in Mexico.
Amphibolips kinseyi is very similar to A. dampfi Kinsey, 1937. We collected the new species in sites near where collections by A. Dampf were made (near Ixtlán, Oaxaca), the material of which was later described by Kinsey (
Body length: 6.3 mm (n = 1) for females; 5.7 mm (n = 3) for males.
Female. Body predominantly black (Fig.
Head, in dorsal view 2.1× as wide as long; 0.8× as wide as mesosoma. POL 0.7× the OOL; lateral ocelli separated from inner margin of an eye for a distance of 3× the diameter of a lateral ocellus. Head in anterior view (Fig.
Mouthparts (Fig.
Antenna (Fig.
Mesosoma in lateral view 1.3× as long as high. Pronotum, moderately pubescent; lateral surface of pronotum with strong irregular reticulate rugose sculpture (Fig.
Mesonotum. Mesoscutum (Fig.
Metanotum. Metapectal-propodeal complex. Metapleural sulcus obscured by the strong sculpture. Metascutellum weakly rugose; metanotal trough deep, smooth and pubescent. Median propodeal area with strong and coarse reticulate rugae; densely pubescent; lateral propodeal carinae distinct, subparallel anteriorly and converging posteriorly.
Legs. Densely pubescent; femora and tibiae robust. Femur 4× as long as wide; metatibia 1.6× as long as metatarsus; apical margin of metatarsomeres 1–4, with long strong erect setae. Metatarsal claws with strong triangular basal lobes or teeth.
Forewing (Fig.
Metasoma (Fig.
Male (Figs
(Fig.
Known only from the type locality along the route from Ixtlán to Tepanzacoalcos (Oaxaca State, Mexico).
Sexual generation. The galls were collected in late April and the insects emerged shortly thereafter, in early May. It is normal to find galls deformed and/or attacked by inquilines and parasitoids; the deformed or attacked galls are usually relatively small.
Holotype: 1♀ in the Museo Nacional de Ciencias Naturales (
Named after the smoky black forewing.
Amphibolips nigrialatus is closely allied to A. dampfi Kinsey, 1937 and the new species Amphibolips kinseyi. Despite being based on a single female holotype, we found distinctive diagnostic characters that let us describe the specimen as belonging to a new species. The strongly-emarginate mesoscutellum relates the new species to A. kinseyi and A. dampfi, but in A. nigrialatus, the postero-lateral projections of the scutellum are pointed apically and curved upwards. Moreover, the scutellar foveae are very large in the new species, extending approximately one half of the length of the mesoscutellum, medially confluent and not separated by a carina or groove, while they are well separated by the mesoscutellar impression in the other two species. The forewing colour of A. nigrialatus is the darkest we have seen in females of Amphibolips species from Mexico and the black smoky colouration even extends to the costal and basal cells and below the cubital vein. In this last character, it resembles A. castroviejoi from Panama, but in this late species, the forewing area anterior to the transversal band is even darker (Fig.
Body length: 6.6 mm (n = 1) for the female.
Female. (Fig.
Head, in dorsal view (Fig.
Mouthparts (Fig.
Antenna (Fig.
Mesosoma in lateral view 1.1× as long as high. Pronotum, moderately pubescent; lateral surface of pronotum with strong irregular reticulate rugose sculpture (Fig.
Mesonotum. Mesoscutum (Fig.
Metanotum. Metapectal-propodeal complex. Metapleural sulcus distinct, reaching posterior margin of mesopectus at about mid-height of metapectal-propodeal complex. Metascutellum rugose; metanotal trough deep, smooth and pubescent. Median propodeal area with strong and coarse reticulate rugae; densely pubescent; lateral propodeal carinae distinct, subparallel anteriorly and converging posteriorly (Fig.
Legs. Densely pubescent; femora and tibiae robust. Metatibia about 1.7× as long as metatarsus. Metatarsal claws with strong triangular basal lobes or teeth (Fig.
Forewing (Fig.
Metasoma (Fig.
Male. Unknown.
(Fig.
On twigs of Quercus sapotifolia Liebm. Closely resembles that of Amphibolips oaxacae Nieves-Aldrey & Pascual, 2012, A. michoacaensis Nieves-Aldrey y Maldonado, 2012, A. trizonata Ashmead, 1896 and Amphibolips kinseyi sp. nov. However, the gall of A. nigrialatus differs in its size, which is approximately half that of the other species. The gall is similar to that of A. murata Weld, 1957, but not as rough when dry and to that of A. quercusfuliginosa Ashmead, 1885, from which it is impossible to differentiate according to the original description of the gall. Nonetheless, the adults are completely different.
Known only from the type locality in Veracruz State, Mexico.
Presumably, a sexual generation. The gall was collected in late April and the insect emerged shortly afterwards.
Details of mesoscutum and mesoscutellum (dorsal view) of Amphibolips species A Amphibolips castroviejoi Medianero & Nieves-Aldrey B Amphibolips durangensis Nieves-Aldrey & Maldonado C Amphibolips fusus Kinsey (type) D Amphibolips cibriani Pujade-Villar (last image taken from
Details of wings and habitus of Amphibolips species A habitus of Amphibolips near fusus from Zacatecas B detail of forewing of Amphibolips fusus Kinsey (type) C habitus of Amphibolips cibriani Pujade-Villar (image taken from
1 | Females. Antenna with 13–14 antennomeres; F1 not modified (Fig. |
2 |
– | Males. Antenna with 15 antennomeres; F1 modified, flattened ventrally (Fig. |
9 |
2 | Forewing with a heavily-infuscate spot on the basal area of radial cell; remainder of the forewing, hyaline to only slightly infuscate (Fig. |
aliciae Medianero & Nieves-Aldrey |
– | Forewing entirely infuscate, more heavily along a band on anterior margin of wing (Fig. |
3 |
3 | More heavily infuscate band along anterior margin of forewing with a clear transversal band on one-third apical part of radial cell which is more or less extended towards posterior margin of wing (Fig. |
4 |
– | More heavily infuscate band along the anterior margin of the forewing, without a clear transversal band on apical part of radial cell extended towards posterior margin of wing. If there is a clear colourless spot apically on the radial cell, it does not extend below the radial cell (Fig. |
|
4 | Basal and first cubital cells colourless or only weakly infuscate prior to the heavily-infuscate basal half of the radial cell (Fig. |
fusus Kinsey |
– | Basal and first cubital cells as heavily infuscate as basal half of radial cell (Figs |
5 |
5 | Mesoscutellum slightly or moderately emarginate posteriorly (Fig. |
6 |
– | Mesoscutellum strongly emarginate posteriorly (usually V-shaped in dorsal view) (Figs |
7 |
6 | Mesoscutellum only slightly emarginate posteriorly (Fig. |
durangensis Nieves-Aldrey & Maldonado |
– | Mesoscutellum more deeply emarginate posteriorly; postero-lateral projections of mesoscutellum moderately expanded (Fig. |
magnigalla sp. nov. |
7 | Scutellar foveae smooth. Notauli visible (Fig. |
castroviejoi Medianero & Nieves-Aldrey |
– | Scutellar foveae with carinate sculpture (Figs |
8 |
8 | Scutellar foveae large, extended approximately one-half of the length of mesoscutellum; medially confluent not separated by a carina or groove (Figs |
nigrialatus sp. nov. |
– | Scutellar foveae not as large, extended approximately one-third of the length of the mesoscutellum and medially separated by the mesoscutellar impression (Figs |
kinseyi sp. nov. |
Males
9 | Forewing with a heavily infuscate spot in the basal area of the radial cell; rest of the forewing only slightly infuscate | aliciae Medianero & Nieves-Aldrey |
– | Forewing entirely and heavily infuscate, with a transversal clear band or with a more infuscate longitudinal band on the anterior margin of the wing (Figs |
10 |
10 | Forewing with a clear transversal band on one-third of the apical part of the radial cell, which is more or less extended towards the posterior margin of the wing (Figs |
11 |
– | Forewing without a clear transversal band on one-third of the apical part of the radial cell; usually with a more infuscate longitudinal band along dorsal margin of forewing; if there is a clear colourless spot apically on the radial cell, it does not extend below the radial cell (Fig. |
|
11 | Transversal clear band large; dorsally extended on 2/3 of apical area of radial cell and extended posteriorly to reach margin of the wing (Fig. |
magnigalla sp. nov. |
– | Transversal clear band much more reduced in size; extended at most on one third of apical area of radial cell, and posteriorly not reaching the posterior margin of the wing (Figs |
12 |
12 | Scutellar foveae smooth. Transversal clear band relatively larger and more extended (Fig. |
castroviejoi Medianero & Nieves-Aldrey |
– | Scutellar foveae sculptured. Transversal clear band very reduced in size (Figs |
13 |
13 | Posterolateral projections of the mesoscutellum acute pointed (Fig. |
dampfi Kinsey |
– | Posterolateral projections of mesoscutellum flat and rounded apically (Fig. |
kinseyi sp. nov. |
The current study increases the number of Amphibolips in Mexico from 20 to 23. However, this number may rapidly increase since fieldwork is revealing new species that are still being studied and will eventually be published elsewhere.
The three species described herein present the typical diagnostic features of the Amphibolips species out of the “niger” group (
Considering that the complete life cycle of most species is unknown, it is difficult to propose a pattern of morphologies or phenologies for these species. In some cases, mistakes could be made, for example, some descriptions may pertain to different generations of currently-recognised species.
The new species described herein are very similar and share a set of similar morphological characteristics. The forewings are very dark and heavily black infuscate, but with a clear transversal band that is more or less extended in both sexes; the mesoscutellum is very deeply emarginate posteriorly, with the emargination reaching the scutellar foveae and almost dividing the mesoscutellum into two parts. Another shared feature of the three new species is that all are distributed in the States of Oaxaca and Veracruz in southern Mexico and none exceeds the Trans-Mexican volcanic belt. Their host oak species (Quercus zempoaltepecana and Q. sapotifolia) are similar in their affinity for slopes in humid climates near the Gulf of Mexico and their occurrence within tropical communities with many Quercus species. After studying large collections of Amphibolips collected across extensive areas of Mexico, we observed a distinct morphological pattern within the geographic distribution of the species, which is also confirmed in the species from the United States (unpublished observations). The more southern a species is distributed, the stronger and deeper its mesoscutellar emargination appears. These two patterns are consistent within a north-south distribution in Mexico.
The presence of four described species from the State of Oaxaca (Amphibolips oaxacae Nieves-Aldrey & Pascual, A. dampfi Kinsey, A. magnigalla sp. nov. and A. kinseyi sp. nov.) and three species from Panama (A. castroviejoi Medianero & Nieves-Aldrey, A. aliciae Medianero & Nieves-Aldrey and A. salicifoliae Medianero & Nieves-Aldrey) allows us to propose that Amphibolips must be present throughout Central America. Although they have not been cited from Chiapas, Mexico to Costa Rica, its presence in those geographic areas is likely, given that most host Quercus species recorded from south of Mexico and Panama are also present in Mesoamerica (
We thank George Melika, Matt Buffington and Zhiwei Liu their useful comments that contributed to improve the manuscript. JLNA was supported by the research project MINECO/FEDER, UE, CGL2015-66571-P. JLNA is grateful to Enrique Medianero (Panama) and Enrique Pascual (Mexico) for their help with sampling in Veracruz and DECL thanks Javier Piña and Antonio López for their help in sampling in Oaxaca. Thanks to the authorities of the communities of the Sierra Juárez, Oaxaca, for allowing us to carry out this work. We also thank Laura Tormo (
* To this couplet option run nine Amphibolips species from Mexico included in the above referred original key.
** To this couplet run also the recently described species Amphibolips cibriani Pujade-Villar. According the original description, we cannot find diagnostic morphological characters differentiating A. cibriani from A. fusus Kinsey (Figs
*** To this couplet option run other nine Amphibolips species from Mexico included in the above referred original key.