Una especie nueva de avispa agalladora (Hymenoptera: Cynipidae), Andricus protector Pujade-Villar and Cuesta-Porta sp. nova (Hymenoptera: Cynipidae: Cynipini) de México
Juli Pujade-Villar a, Ricardo Clark-Tapiab, Victor Aguirre-Hidalgo c, George Melikac, Víctor Cuesta-Portaa, *
a Universitat de Barcelona, Facultat de Biologia, Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals, Avda. Diagonal 645, 08028-Barcelona, Spain
b Universidad de la Sierra Juárez, Laboratorio de Estudios Ambientales, Cam. a la Universidad s/n, 68725 Ixtlán de Juárez, Oaxaca, Mexico
c National Food Chain Safety Office, Plant Health Diagnostic National Reference Laboratory, Keleti Károly u. 24, 1024, Budapest, Hungary
*Corresponding author: victorcp93@gmail.com (V. Cuesta-Porta)
Received: 20 January 2025; accepted: 20 August 2025
http://zoobank.org/urn:lsid:zoobank.org:pub:510B361C-ED79-48EB-A484-9100684E3D92
Abstract
A new species of oak gall wasp, Andricus protector Pujade-Villar and Cuesta-Porta sp. nova, known only from its asexual generation that induces deciduous galls on buds of Quercus crassifolia Humb. and Bonpl. (Lobatae section), is described from Mexico. Diagnosis, distribution and biological data of the new species are given. The validity of Andricus species from Mexico is commented. Four species are proposed here to have dubious affiliation: Andricus burnetti (Dailey and Sprenger, 1983), A. marmoreus Kinsey, 1920, A. setifer (Karsch, 1880) and A. strues (Kinsey, 1938).
Keywords: Hymenoptera; Cynipidae; Andricus; New species; Mexico; Quercus crassifolia
Resumen
Se describe una especie nueva de avispa agalladora, Andricus protector Pujade-Villar y Cuesta-Porta sp. nova de México, conocida solo por su generación asexual, que induce agallas deciduas en yemas de Quercus crassifolia Humb. y Bonpl. (sección Lobatae). Se proporcionan la diagnosis, los datos de distribución y biológicos de la nueva especie. Se comenta la validez de las especies de Andricus de México. Cuatro especies tienen afiliación dudosa: Andricus burnetti (Dailey y Sprenger, 1983), A. marmoreus Kinsey, 1920, A. setifer (Karsch, 1880) y A. strues (Kinsey, 1938).
Palabras clave: Hymenoptera; Cynipidae; Andricus; Especie nueva; México; Quercus crassifolia
Introduction
Oak gall wasps (Hymenoptera: Cynipidae: Cynipini) represent the most diverse group within the Cynipidae family, comprising over 1,000 described species in 59 genera (Buffington et al., 2020; Melika, Pujade-Villar et al., 2021; Ronquist et al., 2015; Stone et al., 2002). These wasps induce galls on Fagaceae, primarily on oaks (Quercus L.). In America, north of Mexico, nearly 500 cynipid species have been recorded (Burks, 1979) in association with 90 species of Quercus (Nixon, 2008). In contrast, Europe has approximately 150 described species of cynipids (Melika 2006), despite having only about 30 Quercus species (Uotila, 2011). More specifically, only in the Iberian Peninsula, about 70 cynipid species associated with 10 Quercus species have been documented (Nieves-Aldrey, 2001). Under the same rule-of-thumb, the Mexican oak gall wasp fauna is likely extraordinarily rich, with many species yet to be described, as Mexico is home to 135-161 Quercus species, 86 of which are endemic (Nixon, 1993a, b; Zavala, 1998; Valencia, 2004).
The recorded number of Mexican cynipid species has varied considerably over time. Pujade-Villar et al. (2009) listed 157 cynipid species, with 42 species attributed to the genus Andricus Hartig, 1840. In a subsequent review, Pujade-Villar and Ferrer-Suay (2015) increased the number of Mexican cynipid species to 183, and 43 species of Andricus. More recently, Martínez-Romero et al. (2022) listed 205 species, 30 of which were assigned to Andricus. The genus Andricus is the most diverse, with more than 400 species described worldwide (Melika, Nicholls et al., 2021; Stone et al.,2002), and one of the most taxonomically complex among oak gall wasps.
Since the review by Pujade-Villar et al. (2009), several species previously classified under Andricus have been reassigned to other genera, including Disholandricus Melika, Pujade-Villar and Nicholls, 2021, Dros Kinsey, 1937, Druon Kinsey, 1937, Erythres Kinsey, 1937, Femuros Kinsey, 1937, Feron Kinsey, 1937, Protobalandricus Melika, Nicholls and Stone, 2018, Striatoandricus Pujade-Villar, 2020, and Trichoteras Ashmead, 1897. In Mexico, a total of 31 species of Andricus are listed (Martínez-Romero et al., 2022), later also A. cylindratum (Kinsey, 1937), A. tecturnarum Kinsey, 1920, A. tibialis Kinsey, 1937 (= A. tostum Kinsey, 1937) (= A. uterinus Kinsey, 1937), A. vitreus Kinsey, 1937 (= A. validus Kinsey, 1937) and A. verutus Kinsey, 1937 were transferred to Feron by Cuesta-Porta et al. (2023). Furthermore, since Martínez-Romero et al. (2022), 2 additional Andricus species have been described from Mexico, A. coombesi Pujade-Villar and Pérez-Torres, 2024 and A. mazahua García-Martinón and Pujade-Villar, 2024 (Pujade-Villar et al., 2024, García-Martiñón et al., 2024), bringing the current number of Andricus species in Mexico to 25. Among these, 18 species induce galls on branches, 4 attack buds (rarely leaves or petioles), 1 is found exclusively on leaves, and 1 in acorns. The host of A. aztecus remains unknown, although it is suspected to induce tuberous galls (Martínez-Romero et al.,2022). The new species described here attacks the buds of Q. crassifolia (Lobatae section), producing a distinctive spherical gall with the larval chamber surrounded by longitudinal air chambers divided by thin partitions of spongious tissue.
Materials and methods
Asexual adult gall wasps (ŏ) of the new species described herein were extracted dead from galls on Quercus crassifolia in Oaxaca (Mexico) years after being collected. They were preserved in 100% ethanol in the laboratory by the second author.
The description of the new species follows the current terminology of morphological structures specific for Cynipidae (Liljeblad & Ronquist, 1998; Melika, 2006) and more general sources such as the Hymenoptera Anatomy Ontology Portal (Yoder et al., 2010). Abbreviations for the fore wing venation follow Ronquist and Nordlander (1989); cuticular surface terminology follows that of Harris (1979). Measurements and abbreviations used here include: F1-F11, 1st and subsequent flagellomeres; POL (postocellar distance), the distance between the inner margins of the posterior ocelli; OOL (ocellar-ocular distance), the distance from the outer edge of a posterior ocellus to the inner margin of the compound eye; LOL, the distance between lateral and frontal ocelli. The width of the fore wing radial cell is measured from the margin of the wing to the Rs vein.
The scanning electron microscope (SEM) pictures were taken by the first author at the University of Barcelona (UB) using a field-emission gun environmental scanning electron microscope (FEI Quanta 200 ESEM), with low-resolution imaging without gold-coating the specimens. The photographs of the galls and habitus were taken by the second author with a Canon PowerShot SX510 HS digital camera and a digital camera mounted on Carl Zeiss microscopy III followed by processing with GIMP 2.8 (GNU Image Manipulation Program), respectively.
The type material of the new species is deposited in the University of Barcelona (UB), Catalonia (J. Pujade-Villar coll.).
Andricus protector Pujade-Villar and Cuesta-Porta sp. nova
(Figs. 1-3)
http://zoobank.org/urn:lsid:zoobank.org:act:D806CCCD-2240-4F39-AF38-94D670D404A5
Diagnosis. The new species differs from all Mexican Andricus species by the following combination of characters: mesoscutum transversally rugose; median mesoscutal line present, with smooth bottom; margin of the fore wing shortly ciliated; second metasomal tergum smooth anteriorly, anterolaterally with dense white setae, and a band of punctures in the posterior third; subsequent terga and hypopygium punctuate, without setae. Andricus protector is closely related to Andricus formosalis Weld, 1944 (Fig. 3B), but differs from it by the following characters: antenna and legs brown to black (yellowish in A. formosalis); inner margins of eyes parallel (converging ventrally in A. formosalis); head quadrate and slightly broader than the mesosoma (triangular, narrower than the mesosoma in A. formosalis); F1 1.2 × as long as F2 (only slightly longer in A. formosalis); placodeal sensilla inconspicuous (present in A. formosalis); pronotum completely rugose (with parallel ridges in the posterior margin in A. formosalis); median mesoscutal line long and wide (short and becoming almost inconspicuous in A. formosalis); mesoscutellar foveae quadrate, separated by a carina, not delimited posteriorly (triangular, widely separated and well-delimited posteriorly in A. formosalis); 2nd metasomal tergum occupying most of the metasoma (very short in A. formosalis). Also, the galls of both species are different: with a velvety reddish surface when young and spherical when mature in A. proctector sp. nova, and with a velvety whitish surface when young and ovoid when mature in A. formosalis.
The new species can be morphologically similar to other Andricus inducingtuberous galls in branches. Despite the great divergence in gall morphology between A. protector and the tuberous gallers, Andricus guanajuatensis, A. montezumus, A. peredurus and A. protector have the mesoscutum transversally carinate, whilst the rest of Mexican species that induce tuberous galls have the mesoscutum strongly wrinkled. Andricus protector has a wide band of punctuation in the second metasomal tergum and a medial mesoscutal line (both characters absent in A. guanajuatensis, A. montezumus, and A. peredurus).
Description. Asexual female. Body black (Fig. 3A); mandibles brown, maxillary and labial palpi light brown; antenna brown, scape black; tegula dark brown to black; legs black, tibiae I and II and all tarsomeres brown; metasoma chestnut brown to black; fore wing veins light.
Head (Fig. 1A-D) 1.3 × as broad as high and very slightly broader than mesosoma in frontal view, 2.1 × as broad as long in dorsal view. Gena coriaceous, broadened behind eye, around 1.4 × as broad as transverse diameter of eye; malar space with striae radiating from clypeus and reaching eye; eye 3.1 × as high as length of malar space. Inner margins of eyes parallel. POL 2.0× as long as OOL, OOL 1.3 × as long as diameter of lateral ocellus, 1.3 × as long as LOL, ocelli rounded, all 3 equal in size. Transfacial distance as long as height of eye and 1.35 × as long as height of lower face (distance between antennal rim and ventral margin of clypeus); diameter of antennal torulus 3.3 × as long as distance between them, distance between torulus and eye as long as diameter of torulus. Lower face coriaceous, setose, with some irregular weak striae, with elevated median area coriaceous, with irregular weak striae, without setae. Clypeus quadrate, flat, carinated, ventrally slightly curved, not emarginate and slightly incised medially; with anterior tentorial pit, epistomal sulcus and clypeo-pleurostomal line distinct. Frons, interocellar area and vertex strongly coriaceous, with some weak rugae between ocelli and eye; occiput carinate, with few short setae; postgena punctuate, pubescent; postocciput and postocciput around occipital foramen weakly carinate, glabrous; posterior tentorial pit oval; hypostomal carina emarginate, present at the basis of postgenal sulci, gular sulci present; occipital foramen longer than height of postgenal bridge.
Antenna (Fig. 1E, F) as long as head + mesosoma, with 13 flagellomeres, sometimes with a partial suture towards the middle of F12, rarely F13 fused; pedicel 1.6 × as long as broad, F1 0.8 × as long as length of scape + pedicel and 1.25 × as long as F2, F3 0.8 × as long as F2, F3 as long as F4, F5 as long as F6, F7 slightly shorter than F6 and equal as F8, F9 slightly shorter than F8 and equal as F10, F11 slightly shorter than F10, F12 + 13 shorter than F10 + F11, F12 as long as F11 and slightly longer than F13; placodeal sensilla inconspicuous on F5 – F12, absent on F1 – F4. Antennal formula: 14:11 x 7: 20: 16: 13: 13: 12: 12. 11: 11: 10: 10: 9: 9: 8.
Mesosoma (Fig. 2A-C) slightly longer than high in lateral view. Pronotum rugose with coriaceous interspaces, glabrous, emarginate along lateral and dorsal edges. Propleuron alutaceous with weak irregular rugae and sparse setae. Mesoscutum rugose, with coriaceous interspaces, glabrous, broader than long (width measured across base of tegulae). Notaulus complete; median mesoscutal line present, extends 1/3 of mesoscutum length, with smooth bottom; anterior parallel line impressed, extends to half of mesoscutum length, coriaceous, glabrous; parapsidal line incised and surrounded by coriaceous surface, extends 2/3 of mesoscutum; parascutal carina narrow, coriaceous, reaching notaulus. Transscutal articulation deep, distinct. Mesoscutellum quadrate, shorter than mesoscutum, uniformly rugose, slightly overhanging metanotum; mesoscutellar foveae big, quadrate, with smooth to alutaceous bottom, open basally with some rugae, distinctly divided by a central carina with coriaceous sculpture. Mesopleuron including speculum rugose-carinate, with coriaceous interspaces, sparsely pubescent; mesopleural triangle coriaceous-rugose, with sparse white setae. Metapleural sulcus reaching mesopleuron above half its height, upper part of sulcus inconspicuous; dorsal axillar area rugose, glabrous; lateral axillar area and axillar carina weakly carinated, glabrous; subaxillular bar with parallel sides smooth, glabrous; metanotal trough smooth, setose. Metascutellum sub-rectangle coriaceous, glabrous, ventral impressed area smooth. Lateral propodeal carinae distinct, broad, parallel but basally curved towards the nucha; central propodeal area coriaceous, glabrous, without central longitudinal carina, with some rugae; lateral propodeal area coriaceous rugose, glabrous; nucha with delicate rugae.
![Figure 3. Lateral habitus of a) Andricus protector sp. nova., b) A. formosalis [Picture obtained from Type Search of Smithsonian Entomological Collections, http://n2t.net/ark:/65665/m36fc57b94-797e-4d41-b88c-58776f4ed1d3]](https://rev-mex-biodivers.mx/wp-content/uploads/5448-Fig_3-12cm.jpg)
Fore wing (Fig. 4A) slightly longer than body, hyaline, with the margin shortly ciliate; radial cell open, 3.0 × as long as broad, R1 and Rs not reaching the wing margin; areolet present, large; Rs + M visible in 2/3 of its length, its projection reaching basal vein slightly below half of its height.
Legs with a reticulate sculpture. Tarsal claws with a basal lobe (Fig. 1G).
Metasoma (Fig. 2D, E) shorter than head + mesosoma, longer than high in lateral view, 2nd metasomal tergum smooth anteriorly with dense white setae anterolaterally and a band of punctures in the posterior third; subsequent terga and hypopygium punctuate, without setae; prominent part of the ventral spine of the hypopygium 5.0 × as long as broad in ventral view, with short and sparse white setae, without apical tuft. Body length 3.7-3.8 mm (n = 3).
Gall (Fig. 4B, C). Unilocular, spherical gall located on the axillary buds of Q. crassifolia, in clusters of 2-4, with average size of 11.6 mm (SD ± 0.46 mm, N = 20). Light yellowish red with short pubescent surface when young, turning brown and glabrous when mature. Inside a single larval cell located in the center, surrounded by multiple radiating empty chambers. All chambers are surrounded by spongy tissue. The consistency of the gall is not very hard.
Taxonomic summary
Type material. Holotype: asexual female “MEX, La Resinera, Santa Catarina Ixtepeji (Ixtepec, Oaxaca), Q. crassifolia, (30.vii.2021) extr. 18.v.24”, R. Clark leg. (black label); “holotype Andricus protector Pujade-Villar and Cuesta-Porta n. sp., desig. JP-V 2023” (red label). Paratypes: 3 asexual females with same data as the holotype. Holotype and 3 paratypes are deposited in the collection of JP-V (University of Barcelona, UB), one of them dissected for SEM imaging.
Etymology. Named for the species’ gall model in which the larval chamber is protected (surrounded) by multiple empty chambers.
Host plant. Known only from Q. crassifolia (Lobatae section). This oak species is distributed in Mexico (Chiapas, Chihuahua, Guerrero, Guanajuato, Hidalgo, Jalisco, Estado de México, Michoacán, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tlaxcala, Veracruz, and Zacatecas) and Guatemala. Distribution. Mexico (Oaxaca).
Biology. Only the asexual generation is known. The galls appear in mid-June and fall to the ground between August and September, rarely in October. Once in the ground, the galls absorb a lot of soil moisture during the rainy season, which explains why they were very soft when they were collected. This might have affected the development of the adults and prevented them from emerging. The extracted adults were dead, but perfectly preserved.
Discussion
Of the total of 25 Andricus species known from Mexico (Table 1), Andricus burnetti (Dailey & Sprenger, 1983) was described in the genus Trichoteras and transferred to Andricus by Melika and Abrahamson (2002), it is associated with oaks of the Protobalanus section which matches with most Trichoteras species, however the gall differs from the rest of Trichoteras and is more similar to Feron. Andricus marmoreus Kinsey, 1920 has a gall similar to those of Disholcaspis, but the adult has simple tarsal claws and an unusual mesoscutal sculpture for a Mexican species of Disholcaspis. The gall of A. setifer (Karsch, 1880) may correspond to some of the genus Striatoandricus. Lastly, A. strues (Kinsey, 1938) was originally described in the genus Conobius Kinsey, 1938, which was later synonymized under Andricus by Melika and Abrahamson (2002), but unlike what is mentioned in the original description of Conobius, A. strues has the margin of its fore wings ciliated. Concerning Callirhytis quercusbatatoides, Pujade-Villar et al. (2014) mentioned that this species probably belongs to Andricus; later, in Pujade-Villar and Ferrer-Suay (2015), it is considered as an Andricus species but the formal change is not indicated; and finally, in Martínez-Romero et al. (2022) it is definitively named as Andricus quercusbatatoides (Ashmead, 1881).
Among the rest of Andricus species reported from Mexico, the gall morphology and host association is still remarkably diverse (Table 1): one species attacks acorns, 4 induce galls on leaves and/or buds, 3 attack branches without causing globular tuberous galls and 13 induce tuberous galls. The gall of A. aztecus (Cameron, 1888) is unknown, although it may induce tuberous galls.
The alternating life cycle is only known in one of the species (A. quercuslaurinus); the rest remain known by their asexual form except in A. protuberans and A. sphaericus, of which only the sexual generation is known. The Mexican Andricus induce galls in the sections Quercus, Lobatae, Virentes, and Protobalanus (Martínez-Romero et al., 2022; Table 1). The species described here corresponds to an asexual form inducing galls in Lobatae section.
Andricus species with linear elements in the mesoscutum (wrinkles or carinae) are restricted to tuberous gall specie (except A. mazahua). This trait is also present in the species here described, but A. protector does not induce tuberous galls. Among the Mexican Andricus species that produce galls on buds, A. burnetti is associated with Q. palmeri (Protobalanus section), while A. protuberans, A. rochai, and A. strues are associated with species in the Quercus section. Although A. protuberans has also been reported on oaks from the Lobatae section, Cuesta-Porta, Cibrián-Tovar et al. (2022) suggested that these records require further confirmation. Andricus protector has a very peculiar gall containing a set of empty chambers radially surrounding the central larval chamber. The gall of the newly described species is very similar to the gall of A. formosalis known from Arizona also associated with the Lobatae section, but the adults are morphologically different (see Diagnosis and Fig. 3).
Table 1
Andricus species reported in Mexico after this study, indicating the known generation of their lifecycle (Known generation), the plant organ they attack (Plant organ), and the host oak association (Host section). Species marked with an asterisk are considered of unclear affiliation in the Discussion section.
| Species | Authors | Known generation | Plant organ | Host section | |
| Andricus | aztecus | (Cameron, 1888) | Asexual | Unknown | Unknown |
| Andricus | bonanseai | Mayr, 1905 | Asexual | Branch (tuberous) | Lobatae, Quercus |
| Andricus | breviramuli | Pujade-Villar, 2014 | Asexual | Branch (non-tuberous) | Quercus |
| Andricus | burnetti * | (Dailey and Sprenger, 1983) | Asexual | Buds/leaf | Protobalanus |
| Andricus | carrilloi | Pujade-Villar, 2013 | Asexual | Branch (tuberous) | Quercus |
| Andricus | coombesi | Pujade-Villar and Pérez-Torres, 2024 | Asexual | Acorn | Lobatae |
| Andricus | dugesi | Beutenmüller, 1917 | Asexual | Branch (tuberous) | Lobatae, Quercus |
| Andricus | durangensis | Beutenmüller, 1911 | Asexual | Branch (tuberous) | Lobatae |
| Andricus | furnaceus | Kinsey, 1920 | Asexual | Branch (tuberous) | Quercus |
| Andricus | fusiformis | Pujade-Villar, 2014 | Asexual | Branch (non-tuberous) | Lobatae, Quercus |
| Andricus | guanajuatensis | Pujade-Villar, 2013 | Asexual | Branch (tuberous) | Lobatae, Quercus |
| Andricus | marmoreus * | Kinsey, 1920 | Asexual | Branch (non-tuberous) | Quercus |
| Andricus | mazahua | García-Martinón and Pujade-Villar, 2024 | Asexual | Branch (tuberous) | Lobatae |
| Andricus | montezumus | Beutenmüller, 1913 | Asexual | Branch (tuberous) | Quercus |
| Andricus | peredurus | Kinsey, 1920 | Asexual | Branch (tuberous) | Quercus |
| Andricus | protector | Pujade-Villar & Cuesta-Porta, sp. nova. | Asexual | Buds/leaf | Lobatae |
| Andricus | protuberans | Pujade-Villar and Ferrer-Suay, 2015 | Sexual | Buds/leaf | Lobatae |
| Andricus | quercusbatatoides | (Ashmead, 1881) | Asexual | Branch (tuberous) | Virentes |
| Andricus | quercuslaurinus | Melika and Pujade-Villar, 2009 | Sex. and asex. | Branch (non-tuberous) | Lobatae |
| Andricus | rochai | Pujade-Villar, 2018 | Asexual | Buds/leaf | Quercus |
| Andricus | santafe | Pujade-Villar, 2013 | Asexual | Branch (tuberous) | Quercus |
| Andricus | setifer * | (Karsch, 1880) | Asexual | Branch (non-tuberous) | Unknown |
| Andricus | sphaericus | Pujade-Villar, 2016 | Sexual | Buds/leaf | Quercus |
| Andricus | strues * | (Kinsey, 1938) | Asexual | Buds/leaf | Quercus |
| Andricus | tumefaciens | Pujade-Villar and Paretas-Martínez, 2012 | Asexual | Branch (tuberous) | Lobatae, Quercus |
| Andricus | tumeralis | Pujade-Villar, 2009 | Asexual | Branch (tuberous) | Quercus |
Air chambers in galls are hypothesized to act as a defensive mechanism against parasitoids by hindering oviposition or confusing parasitoids simulating larval chambers and (Pujade-Villar et al., 2025; Stone & Cook 1998; Stone et al., 2002). Several independent cynipid lineages have converged into gall morphologies with hollow internal spaces in the parenchyma, such as the galls of Atrusca, Kinseyella, and some Amphibolips in the Nearctic and Trichagalma in Eastern Palearctic, usually exhibit radiating filaments that connect the larval chamber with the outer shell of the gall; or less common cases of free rolling larval chambers like some Disholcaspis or Belizinella (Cuesta-Porta et al.,2020, 2024, 2025; Ide & Koyama, 2023; Pujade-Villar et al., 2010). In the case of A. protector and A. formosalis, galls have hollow longitudinal chambers divided by longitudinal parenchymatous partitions. Despite the gall similarity, the adults of A. protector and A. formosalis exhibit significant differences (see Diagnosis). Further research is needed to determine the phylogenetic relationships between A. protector and the rest of Andricus species. The generic limits of the American Andricus species are still under revision and more data is needed to confirm that the similarities between A. protector and A. formosalis are phylogenetically informative or the result of morphological convergence.
Acknowledgements
We thank Denis Brothers (School of Biological and Conservation Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa) for his help with a ZNC question. This research was funded by the project “PID2021-128146NBI00/MCIN/AEI/10.13039/501100011033/” and “FEDER una manera de hacer Europa” from the Ministry of Science and Innovation of Spain and the European Region al Development Fund (ERDF). Also is supported by UNSIJ 2-EA-2203.
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