Introduction
The taxonomy of Charipinae (Hymenoptera: Cynipoidea: Figitidae) has always been difficult. Too many species have been described in too little detail, sometimes focusing on organisms from particular biogeographical regions with no comparison of material from other regions. Comparison of the available type material has revealed numerous synonymies. The taxonomy of Charipinae is further complicated because they are small wasps characterised mainly by a smooth and shiny body so there are very few diagnostic features for use in distinguishing species and establishing species limits.
Biologically, Charipinae are all secondary parasitoids of Aphididae via Aphidiinae (Hymenoptera: Braconidae) and Aphelininae (Hymenoptera: Aphelinidae) and secondary parasitoids of Psyllidae via Encyrtidae (Hymenoptera: Chalcidoidea) (Menke and Evenhuis Reference Menke and Evenhuis1991). For this reason, they are economically important. The first step to correctly determine which species are valid is to study type material. Charipinae includes eight genera: Alloxysta Förster, 1869 (cosmopolitan), Apocharips Fergusson, Reference Fergusson1986 (eastern Palaearctic and Neotropical), Dilapothor Paretas-Martínez and Pujade-Villar, 2006 (Australia), Dilyta Förster, 1869 (cosmopolitan except Australia), Lobopterocharips Paretas-Martínez and Pujade-Villar, 2007 (Nepal), Lytoxysta Kieffer, Reference Kieffer1909 (North America), Phaenoglyphis Förster, 1869 (cosmopolitan), and Thoreauana Girault, 1930 (Australia). Alloxysta is the most widespread genus and includes the largest number of species: 180 have been described, 168 of which are considered valid (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
In the Nearctic Region 34 species have been described. Andrews (Reference Andrews1976) described Alloxysta nothofagi and Andrews (Reference Andrews1978) described 11 Alloxysta species: A. anthracina, A. commensuratus, A. conniferensis, A.dicksoni, A. filimentosus, A. halli, A. minuscula, A. quebeci, A. rauchi, A. schlingeri, and A. vandenboschi. Ashmead described seven species: Allotria lachni in 1885, A. brassicae and A. megourae in 1887, A. xanthopsis in 1896, A. australiae in 1900, Alloxysta alaskensis in 1902, and Xystus japonicus in 1904. Baker (Reference Baker1896) described 12 species: Alloxysta apicalis, A. abdominalis, A. gracilis, A. longiventris, A. magna, A. rufipleura, A. robusta, Dilyta affinis, D. bicolor, D. coloradensis, D. ruficeps, and D. similis. The few species described by three other authors are: Charips leguminosa Weld, Reference Weld1920, Allotria tritici Fitch, Reference Fitch1861, and Charips areolata Kieffer, Reference Kieffer1909.
Here, we revise the Alloxysta type material deposited in the National Museum of Natural History, Washington, District of Columbia, United States of America (USNM), and the Canadian National Collection of Insects, Ottawa, Ontario, Canada (CNC) described by Andrews, Ashmead, and Baker, a total of 34 species, only 19 of which were previously considered valid.
The species we consider to be valid here are redescribed and illustrated to show their main diagnostic features. Notes on the type material of each studied species, valid or not, are given.
Materials and methods
Material studied is deposited in the following institutions:
USNM: United States National Museum of Natural History (Smithsonian Institution) (Washington, District of Columbia, United States of America).
Alloxysta abdominalis, A. alaskensis, A. apicalis, A. commensuratus, A. coniferensis, A. dicksoni, A. filimentosus, A. gracilis, A. halli, A. longiventris, A. magna, A. rauchi, A. robusta, A. rufipleura, A. schlingeri, A. vandenboschi, Allotria australiae, A. brassicae, A. lachni, A. megourae, A. tritici, A. xanthopsis, Charips leguminosa, C. areolata, Dilyta affinis, D. bicolor, D. coloradensis, D. ruficeps, D. similis, Xystus japonicus
CNCI: Canadian National Collection of Insects (Ottawa, Ontario, Canada).
Alloxysta anthracina, A. minuscula, A. quebeci
Type specimens were studied with a Leica M10 microscope (Solms, Germany) and photographed using an Entovision imaging system (Hagerstown, Maryland, United States of America), comprised of a Leica Z16 lens fitted to a JVC KY-75 3-CCD digital camera (Yokohama, Kanagawa, Japan). Specimens were illuminated using LED dome lighting. Images were stacked using Cartograph 6.1 and Archimed 6.2 (Microvision Instruments, Evry, France) at the USNM. At the CNC photos were taken with a ProgRes C14+ a 14 mpix CCD camera and Image-Pro Plus was used to automatically run the motor drive (ProScan III, Prior. (Toronto, Ontario, Canada)) and Zerene Stacker stack layers and obtain one final, entirely focused image.
Morphological terms used follow Paretas-Martínez et al (Reference Paretas-Martínez, Arnedo, Melika, Selfa, Seco-Fernández and Fülöp2007). Abbreviations used are F1–F12 for the first and subsequent flagellomeres. Width of the forewing radial cell is measured from the wing margin to the beginning of vein Rs. The transfacial line ratio was calculated by dividing the length of the transfacial line (measured from the inner margins of compound eyes, across the face through the antennal sockets) by the compound eye height. The malar space ratio was calculated by dividing the length of the lower part of the gena (from the mouthparts to the ventral margin of the compound eye), by the compound eye height. Females and males of the species described have the same characters except where indicated in the redescriptions.
Results and discussion
Alloxysta australiae (Ashmead, Reference Ashmead1900)
(Fig. 1)
Alloxysta australiae (Ashmead, Reference Ashmead1900): (A) forewing; (B) radial cell; (C) pronotum; (D) antenna; (E) propodeum; (F) body.
Allotria australiae Ashmead, Reference Ashmead1900: 330.
Type material of Allotria australiae Ashmead. Holotype ♀ (USNM), with the following labels: “Australia, Koebele”, “Type no 4876, USNM”, “Allotria australiae ♀ Ashm” (handwritten). As far as we can determine Alloxysta australiae is known only from the holotype.
Diagnosis.Alloxysta australiae is mainly characterised by a closed radial cell and the presence of pronotal and propodeal carinae. It is similar to A. fuscicornis (Hartig, 1841) but the latter lacks propodeal carinae. The species is also very similar to A. japonicus (Ashmead Reference Ashmead1904) but as only the female of A. australiae and male of A. japonicus are known a synonymy cannot be established.
Redescription. Female.
Colour. Head, mesosoma and metasoma yellowish brown. Scape, pedicel, F1, and F2 dark yellow, F3–F12 yellowish brown. Legs yellow and veins yellowish brown.
Head. Transversally ovate, smooth and shiny, slightly wider than high in front view. Setae present below and between toruli, scattered setae above toruli. Few scattered setae on vertex and many setae on face. Transfacial line 1.1 times compound eye height. Malar space 0.4 times compound eye height.
Antenna. F1–F3 thinner and smoother than remaining flagellomeres, F4–F11 with rhinaria and club-shaped. Pedicel 1.5 times as long as wide; F1 4.0 times as long as wide; F2 3.3 times as long as wide; F3 2.9 times as long as wide; F4 2.9 times as long as wide. F1 2.0 times as long as pedicel; F2–F11 subequal in length, width and shape (Fig. 1D).
Mesosoma. Mesosoma covered with sparse setae, the setae nearly absent at anterior margin of pronotum and denser on apical scutellum and propodeum. Pronotum with two thick carinae (Fig. 1C). Mesoscutum smooth and shiny, round in dorsal view. Scutellum smooth and shiny. Propodeum with two carinae straight and well defined, joining ventrally (Fig. 1E).
Forewing. Longer than body, 1.5 times as long as mesosoma and metasoma together (Fig. 1A). Closed radial cell, 2.4 times as long as wide. R1 short and curved; Rs long and also curved (Fig. 1B).
Metasoma. T1 with incomplete ring of setae on anterior part, glabrous at centre, wider laterally. Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Distribution. Australia (Ashmead Reference Ashmead1900; Carver Reference Carver1992, Reference Carver2004).
Natural history. According to Carver (Reference Carver1992, Reference Carver2004) this species usually attacks Aphidius colemani Viereck, 1912 (Hymenoptera: Braconidae). The complete list of hosts is given in Ferrer-Suay et al. (Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
Alloxysta brevis (Thomson, 1862)
Charips leguminosa Weld, Reference Weld1920: 15 new synonym.
Allotria megourae Ashmead, Reference Ashmead1887: 19 new synonym.
Alloxysta rauchi Andrews, Reference Andrews1978new synonym.
Type materialof Charips leguminosa Weld.Holotype ♀ (USNM), with the following labels: “Twin Falls, Id., 3700ft. Att, RHR114” (handwritten), “Type No. 22589, USNM” (red label), “Charips (Charips) leguminosa Weld ♀” (handwritten). Paratypes 1♂ and 5♀ (USNM), with the following labels: “Twin Falls, Id., 3700ft. Att, 7-1919”, “RHD 114” (handwritten), “Paratype No. 22589, USNM” (red label), “Charips (Charips) leguminosa Weld” (handwritten). Andrews (Reference Andrews1978) mentioned that two more specimens are deposited in the Weld's collection, probably in the USNM.
After examining non-type specimens from several countries and hosts (Ferrer-Suay et al. Reference Ferrer-Suay, Selfa and Pujade-Villar2012b) we conclude that the little variation in shape of F1 (Andrews Reference Andrews1978, Fig. 103), represents intraspecific variability, not a diagnostic character to differentiate A. leguminosa from A. brevis, so we place A. leguminosa in synonymy.
Type materialof Allotria megourae Ashmead. Holotype ♀ (USNM), with the following labels: “Jacksonville Fla”, “Collection Ashmead”, “Type No. 2849, USNM” (red label), “Allotria megourae ♀ Ashm.” (handwritten).
The main diagnostic features of the holotype correspond to those of A. brevis, so A. megourae is here placed in synonymy. Andrews (Reference Andrews1978) mentions a “megourae” complex from the New World but it is the same to “brevis” complex from the Palaearctic area, as explained in Ferrer-Suay et al. (Reference Ferrer-Suay, Selfa and Pujade-Villar2012b).
Type materialof Alloxysta rauchi Andrews. Holotype ♀ (USNM), with the following labels: “8 mi, S.E. Spences, Bridge, B.C. CANADA, vii-4-66, F.G. Andrews”, “Host Plant Populus trichocarpa”, “Host Aphid Chaitophorus popilicolis”, “♀”, “Holotype Alloxysta rauchi Andrews” (red label), “Type Number 103513, USNM” (red label).
Paratype 1♀ (USNM), with the following labels: “8 mi, S.E. Spences, Bridge, B.C. Canada, VII-4-66, F.G. Andrews”, “Host Plant Populus trichocarpo”, “♀”, “Paratype Alloxysta rauchi Andrews” (yellow label).
According to Andrews the remaining paratypes (25 females and 13 males) are deposited in the author's collection. They were not examined. After examining the two USNM types and non-type specimens from several countries and hosts (Ferrer-Suay et al. Reference Ferrer-Suay, Selfa and Pujade-Villar2012b) we conclude that the curved shape of F1 (Andrews Reference Andrews1978, Fig. 102) represents intraspecific variability and does not differentiate A. rauchi from A. brevis so A. rauchi is placed here in synonymy.
Comments. Three valid species belong to the “brevis” complex: A. brevis (=A. leguminosa, A. rauchi, and A. megourae), A. arcuata (=A. ligustri) and A. mullensis.
Distribution. Holarctic and Neotropical (Ferrer-Suay et al. Reference Ferrer-Suay, Selfa and Pujade-Villar2012b).
Natural history. This species attacks a widely range of hosts in both Europe and the New World (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
Alloxysta castanea (Hartig, 1841)
Dilyta affinis Baker, Reference Baker1896: 132 new synonym.
Alloxysta abdominalis Baker, Reference Baker1896. Synonymised by Andrews (Reference Andrews1978: 51).
Alloxysta apicalis Baker, Reference Baker1896. Synonymised by Andrews (Reference Andrews1978: 51).
Dilyta coloradensis Baker, Reference Baker1896. Synonymised by Andrews (Reference Andrews1978: 51).
Alloxysta gracilis Baker, Reference Baker1896. Synonymised by Andrews (Reference Andrews1978: 51).
Alloxysta magna Baker, Reference Baker1896. Synonymised by Andrews (Reference Andrews1978: 51).
Alloxysta rufipleura Baker, Reference Baker1896. Synonymised by Andrews (Reference Andrews1978: 51).
Dilyta ruficeps Baker, Reference Baker1896. Synonymised by Andrews (Reference Andrews1978: 51).
Dilyta similis Baker, Reference Baker1896. Synonymised by Andrews (Reference Andrews1978: 51).
Alloxysta quebeci Andrews, Reference Andrews1978: 66 new synonym.
The synonymies proposed by Andrews (Reference Andrews1978) were confirmed and we propose two new synonymies, as indicated above and discussed further below.
Type material of Dilyta affinis Baker.Holotype ♀ (USNM), with the following labels: “Colo 1632”, “Ft Collins Col”, “9-7-95, misc. sweeping, Carex Rhus, willow weeds, etc.” (handwritten), “CF Baker Collector”, “Dilyta affinis Baker (red label, handwritten), “Type number 103532, USNM” (red label).
Type material of Alloxysta abdominalis Baker. Holotype ♀ (USNM), with the following labels: “Colo 1547”, “Ft Collins Col”, “6-7-95 misc. sweeping” (handwritten), “CF Baker Collector”, “Alloxysta abdominalis Baker” (red label, handwritten), “Type Number 103525, USNM” (red label).
Type material of Alloxysta apicalis Baker. Holotype ♀ (USNM), with the following labels: “Colo 1632”, “Ft. Collins Col”, “9-7-95, misc. sweeping, Carex Rhus, Willow weeds, etc.” (handwritten), “CF Baker Collector”, “Dilyta affinis Baker (red label, handwritten), “Type Number 103532, USNM” (red label).
Type material of Dilyta coloradensis Baker. Holotype ♀ (USNM), with the following labels: “Colo 1563”, “Ft Collins Col”, “6-13-95, misc. sweeping” (handwritten), “CF Baker Collector”, “Dilyta coloradensis Baker” (red label, handwritten), “Type Number 103534, USNM” (red label).
Type material of Alloxysta gracilis Baker. Holotype ♀ (USNM), with the following labels: “Colo 1632”, “Ft Collins Col”, “9-7-95, misc. sweeping, Carex Rhus” (handwritten), “Willow needs, etc.” (handwritten), “CF Baker Collection”, “Alloxysta gracilis Baker” (red label, handwritten), “Type Number 103527, USNM” (red label).
Type material of Alloxysta magna Baker. Holotype ♀ (USNM), with the following labels: “Colo 1563”, “Ft Collins Col”, “6-13-95, misc. sweeping” (handwritten), “CF Baker Collector”, “Alloxysta magna Baker” (red label, handwritten), “Type Number 103529, USNM” (red label).
Type material of Alloxysta rufipleura Baker. Lectotype ♂ (USNM), with the following labels: “Colo1563”, “Ft Collins Col”, “6-13-95, misc. sweeping” (handwritten), “CF Baker Collector”, “Type Number 103531, USNM” (red label), “Alloxysta rufipleura Baker” (red label, handwritten), “Lectotype Alloxysta rufipleura Baker, Design.: A.S. Menke” (handwritten). Paralectotype ♀ (USNM), with the following labels: “Colo1563”, “Ft Collins Col”, “6-13-95, misc. sweeping” (handwritten), “CF Baker Collector”, “Paralectotype Alloxysta rufipleura Baker, Design.: A.S. Menke”, “Alloxysta affinis Baker Det. Menke, 198”.
Type material of Dilyta ruficeps Baker. Holotype ♀ (USNM), with the following labels: “Colo 1563”, “Ft Collins Col”, “6-13-95, misc. sweeping” (handwritten), “CF Baker Collector”, “Dylita ruficeps Baker” (red label, handwritten), “Type Number, 103535, USNM” (red label).
Type material of Dilyta similis Baker. Holotype ♂ (USNM), with the following labels: “Colo 1632”, “Ft Collins Col”, “9-7-95, misc. sweeping, Carex Rhus, Willow Weeds, etc.”, (handwritten), “Dylita similis Baker” (red label, handwritten), “Type Number 103536, USNM” (red label).
Type material of Alloxysta quebeci Andrews. Holotype ♀ (CNC), with the following labels: “Chimo, Quebec, 17-18.viii.59, W.P.M. Mason”, “♀”, “Holotype Alloxysta quebeci Andrews” (red label), “Holotype Alloxysta quebeci Andrews, CNC No 19030” (red label). Paratypes Four ♀ (CNC) and two ♀ (Andrews collection), with the same data as holotype. According to Andrews this species is similar to A. minuscula but the two species can be easily differentiated by the presence of propodeal carinae in A. quebeci and absence in A. minuscula and the features are those of A. castanea, so we propose the new synonymy here.
Distribution. Palaearctic and Neotropical (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a, Reference Ferrer-Suay, Selfa and Pujade-Villar2012c).
Natural history.Alloxysta castanea has been found mainly in Europe attacking Hyalopterus Koch, 1854 (Hemiptera: Aphididae) species through Praon Haliday, 1833 (Hymenoptera: Braconidae) species on Prunus Linnaeus (Rosaceae) species and Phragmites Adanson (Poaceae) species (Evenhuis Reference Evenhuis1982; Hübner et al. Reference Hübner, Völkl and Francke2002). The list of European hosts is presented in Ferrer-Suay et al. (Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a). The hosts of specimens collected in the United States of America are unknown.
Alloxysta commensuratus (Andrews, 1978)
(Fig. 2)
Alloxysta commensuratus Andrews, Reference Andrews1978: (A) forewing; (B) radial cell; (C) antenna male; (D) pronotum; (E) propodeum.
Alloxysta commensuratus Andrews, Reference Andrews1978: 58.
Type materialof Alloxysta commensuratus Andrews. Holotype ♂ (USNM), with the following labels: “Bowie Md, vi-7-‘45, DDT Expt.”, “5” (handwritten), “Holotype Alloxysta commensuratus Andrews” (red label), “Type Number 103570, USNM” (red label). Paratypes 2♂ and 1♀ (USNM), with the following labels: “Bowie Md, vi-7-‘45, DDT Expt.”, “♂”, “Paratype Alloxysta commensuratus Andrews” (yellow label): 1♂; “Bowie Md, vi-7-‘45, DDT Expt.”, “♀”, “Paratype Alloxysta commensuratus Andrews” (yellow label), “Slide USNM, 47-6-1” (handwritten): 1♀; “Bowie Md, vi-2-‘45, DDT Expt.”, “♂”, “Paratype Alloxysta commensuratus Andrews” (yellow label), “Slide USNM, 47-6-1” (handwritten): 1♂. Two other paratypes are deposited in Andrews's collection.
Diagnosis.Alloxysta commensuratus is mainly characterised as having a partially open radial cell, pronotal carinae present and propodeal carinae absent. Based on these features it is similar to A. macrophadna (Hartig, 1841) and A. obscurata (Hartig, 1840), but they can be differentiated by the presence of rhinaria on F4 in A. commensuratus (Fig. 2C) rather than F3 in A. macrophadna and A. obscurata; radial cell 2.2 times as long as wide in A. commensuratus (Fig. 2B) but 3.0 times as long as wide in A. macrophadna and 2.7 times as long as wide in A. obscurata.
Redescription.
Colour. Head, mesosoma and metasoma dark brown. Scape, pedicel, F1, and F2 dark yellow, F3–F12 yellowish brown. Legs yellow and veins yellowish brown.
Head. Transversally ovate, smooth and shiny, slightly wider than high in front view. Setae below and between toruli but none above toruli. Face with many setae. Vertex with few scattered setae. Transfacial line 1.2 times compound eye height. Malar space 0.4 times compound eye height.
Antenna. Female: F1–F2 thinner and smoother than remaining flagellomeres, F3–F11 with rhinaria and club-shaped. F1–F3 subequal in length, ratio 25:24:25. Male: F1–F3 thinner and smoother than remaining flagellomeres, F4–F12 with rhinaria and club-shaped. Pedicel 2.2 times as long as wide; F1 3.3 times as long as wide; F2 2.9 times as long as wide; F3 2.9 times as long as wide; F4 3.0 times as long as wide. F1 1.1 times as long as pedicel; F1 1.2 times as long as F2; F2 subequal to F3; F4 1.1 times as long as F3; F4–F12 subequal in length, width and shape (Fig. 2C).
Mesosoma. Mesosoma covered with sparse setae, the setae are denser on apical scutellum and propodeum. Pronotum with two thick carinae clearly visible (Fig. 2D). Mesoscutum smooth and shiny, round in dorsal view. Scutellum also smooth and shiny. Propodeum without carinae (Fig. 2E).
Forewing. Longer than body, 1.6 times as long as mesosoma and metasoma together (Fig. 2A). Partially open radial cell, 2.2 times as long as wide. R1 short and curved; Rs long and also curved (Fig. 2B).
Metasoma. T1 with incomplete ring of setae on anterior part, glabrous at centre, wider laterally. Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Distribution. United States of America (Maryland) (Andrews Reference Andrews1978).
Natural history. Unknown.
Alloxysta fuscicornis (Hartig, 1841)
Allotria brassicae Ashmead, Reference Ashmead1887: 14. Synonymised by Evenhuis (Reference Evenhuis1982).
Type material studiedof Allotria brassicae Ashmead.Holotype ♀ (USNM), with the following labels: “Jacksonville Fla”, “Collection Ashmead”, “Type No. 2845, USNM” (red label), “Lectotype H.H. Evenhuis” (orange label), “Allotria brassicae Ashm.” (handwritten), “Allotria brassicae Ashmead, Det. H.H. Evenhuis, 1974”.
According to Andrews (Reference Andrews1978) there are no paratypes. This species was synonymised by Evenhuis (Reference Evenhuis1982) with A. fuscicornis. We confirm this synonymy here.
Distribution. Cosmopolitan (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
Natural history. This species has been found on a wide range of hosts (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a). The most common are Brevicoryne brassicae (Linnaeus, 1758) (Hemiptera: Aphididae), and Myzus persicae (Sulzer, 1776) (Hemiptera: Aphididae) through Diaretiella rapae (M'Intosh, 1855) (Hymenoptera: Braconidae) (Horn Reference Horn1988; Carver Reference Carver2004).
Alloxysta halliAndrews 1978species dubia
Alloxysta halli Andrews, Reference Andrews1978: 62.
Type material of Alloxysta halli Andrews. According to Andrews (Reference Andrews1978) there are 9♂ and 3♀ (USNM). None can be located (M. Buffington personal communication (United States National Museum of Natural History (Smithsonian Institution), Washington, District of Columbia, United States of America), P. Hanson personal communication (Universidad de Costa Rica, San Pedro, Costa Rica), and S. Triapitsyn personal communication (University of California, Riverside, California).
Additional material: (4♂ and 2♀). “mebrose, mass” (handwritten), “Ex nutarophora” (handwritten), “HSSmith Collector”, “Alloxysta halli Andrews Det. Fred G. Andrews” (3♂ and 1♀); “Host: Aphid on Solidago” (handwritten), “Marmora Ont., 4-vii-1952, J. R. Vockeroth”, “Alloxysta halli Andrews Det. Fred G. Andrews”: 1♂; “Vineland Station, Ont. 29.v.1937, W.L. Putman”, “Host. Aphidius” (handwritten), “Alloxysta halli Andrews Det. Fred G. Andrews 1977”, “Alloxysta halli Andrews”: 1♀.
Comments. According to Andrews (Reference Andrews1978) the holotype ♀ and 1 paratype ♀ were deposited in the USNM collection. This material is probably lost (M. Buffington, USNM, personal communication). The rest of the type material is deposited in the collection of the University of California, Riverside, California, United States of America. According to S. Triapitsyn the paratypes there could not be found.
Some specimens determined by Andrews (Reference Andrews1978) as A. halli were found in the USNM. These specimens belong, without doubt, to A. castanea (Hartig, 1841). Additionally, two more specimens (1♂ and 1♀) determined as A. halli by Andrews and deposited at the University of Barcelona, Spain were examined; these two specimens also correspond to the male and female of A. castanea. In the original description of A. halli some important characters are not mentioned, namely, the presence/absence of pronotal and propodeal carinae. Moreover, the male antennae and radial cell are not conspecific with A. castanea. For these reasons, we consider A. halli as a species dubia until the types can be found and examined.
Alloxysta japonicus (Ashmead, Reference Ashmead1904)
(Fig. 3)
Alloxysta japonicus (Ashmead, Reference Ashmead1904): (A) forewing; (B) pronotum; (C) head; (D) antenna; (E) body; (F) propodeum.
Xystus japonicus Ashmead, Reference Ashmead1904: 77.
Type material of Xystus japonicus Ashmead. Holotype ♂ (USNM), with the following labels: “8921” (handwritten), “Japan, Koebele”, “Type No. 7137 USNM” (red label), “Xystus japonicus ♂ Ashm.” (handwritten). Alloxysta japonicus is represented only by the holotype ♂, received from Mr. A. Koebele from Japan (Ashmead Reference Ashmead1904).
Diagnosis.Alloxysta japonicus is mainly characterised as having a closed radial cell, pronotal and propodeal carinae present. Based on these features it is similar to A. xanthopsis (Ashmead Reference Ashmead1896), but differentiated by the length/width ratio of the flagellomeres (longer in A. japonicus than in A. xanthopsis) and dimensions of the radial cell (2.4 times as long as wide in A. japonisus but 2.1 times as long as wide in A. xanthopsis). This species is very similar to A. australiae (Ashmead Reference Ashmead1900), but as only the female of A. australiae and male of A. japonicus are known this synonymy cannot be established with certainty.
Redescription. Male.
Colour. Head and mesosoma yellowish brown, metasoma dark yellow. Scape yellowish brown, pedicel, F1–F3 dark yellow, F4–F12 yellowish brown. Legs yellow and veins yellowish brown.
Head. Transversally ovate, smooth and shiny, slightly wider than high in front view. Setae below and between toruli, few scattered setae above toruli. Without or with few scattered setae on vertex and many setae on face. Transfacial line 1.3 times compound eye height. Malar space 0.5 times compound eye height (Fig. 3C).
Antenna. F1–F12 with rhinaria and club-shaped. Pedicel 1.8 times as long as wide; F1 3.6 times as long as wide; F2 3.5 times as long as wide; F3 3.9 times as long as wide; F4 3.4 times as long as wide. F1–F12 subequal in length, width and shape (Fig. 3D).
Mesosoma. Mesosoma covered with sparse setae, abundant at anterior margin of pronotum and, denser on apical scutellum and propodeum. Pronotum with two long and thick carinae clearly visible (Fig. 3B). Mesoscutum smooth and shiny, round in dorsal view. Scutellum also smooth and shiny. Propodeum with two carinae thick and straight, reaching the base independently (Fig. 3F).
Forewing. Longer than body, 1.6 times as long as mesosoma and metasoma together (Fig. 3A). Closed radial cell, 2.4 times as long as wide. R1 short and straight; Rs long and slightly curved (Fig. 3A).
Metasoma. T1 with incomplete ring of setae on anterior part, glabrous at centre, wider laterally. Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Distribution. Japan (Ashmead Reference Ashmead1904).
Natural history. Only found on Aphis Linnaeus, 1758 (Hemiptera: Aphididae) species according to Dalla Torre and Kieffer (Reference Dalla Torre and Kieffer1910).
Alloxysta lachni (Ashmead, Reference Ashmead1885)
(Fig. 4)
Alloxysta lachni (Ashmead, Reference Ashmead1885): (A) forewing; (B) pronotum; (C) antenna male; (D) antenna female; (E) propodeum.
Allotria lachni Ashmead, Reference Ashmead1885: 302.
Type materialof Allotria lachni Ashmead.Holotype ♀ (USNM), with the following labels: “Jacksonville Fla, “Allotria lachni Ashm.” (handwritten), “not understandable”, “Type No, USNM” (red label).
Additional material (1♂ and 1♀): “Asheville, N.C., (Bent creek), 4-7, 1944”, “Parasite of Pine aphis”, “C.H. Hoffmann Collector”, “52.”, “Alloxysta lachni (Ashmead), Det. Fred G. Andrews, 1977”: 1♀; “Gainesville Fla. Ahr. 10’ 48”, “Alloxysta lachni (Ashmead), Det. Fred G. Andrews, 1977”: 1♂.
The holotype has both antennae missed. The identification has been made on the basis of specimens identified by Andrews.
Diagnosis.Alloxysta lachni is mainly characterised as having a closed radial cell, pronotal carinae absent and propodeal carinae present. It is similar to A. fracticornis (Thomson, 1862) but can be differentiated by the relative sizes of flagellomeres: F1 longer than F2, F2 subequal to F3 in A. lachni females (Fig. 4D), while F1–F3 subequal in length in A. fracticornis females; with rhinaria and club-shaped from F1 in A. lachni males, without any flagellomere curved (Fig. 4C), but with rhinaria and club-shaped from F3 and F3 curved in A. fracticornis males.
Redescription.
Colour. Head, mesosoma, and metasoma dark brown. Antennae yellowish brown. Legs yellow and veins yellowish brown.
Head. Transversally rounded, smooth and shiny, slightly wider than high in front view. Long setae below, between and above toruli. Scattered long setae on vertex and on face. Transfacial line 1.2 times compound eye height. Malar space 0.5 times compound eye height.
Antenna. Female: F1–F2 thinner and smoother than remaining flagellomeres, F3–F11 with rhinaria and club-shaped. Pedicel 1.8 times as long as wide; F1 5.6 times as long as wide; F2 2.3 times as long as wide; F3 1.8 times as long as wide; F4 1.6 times as long as wide. F1 2.0 times as long as pedicel; F1 2.0 times as long as F2; F2–F11 subequal in length, width and shape (Fig. 4D). Male: F1–F12 with rhinaria and club-shaped. Pedicel 1.5 times as long as wide; F1 4.6 times as long as wide; F2 2.9 times as long as wide; F3 2.4 times as long as wide; F4 2.2 times as long as wide. F1 2.6 times as long as pedicel; F1 4.6 times as long as F2; F2 1.2 times as long as F3; F3–F12 subequal in length, width and shape (Fig. 4C).
Mesosoma. Mesosoma covered with sparse setae, the setae are abundant at anterior margin of pronotum and, denser on apical scutellum and propodeum. Pronotum without carinae (Fig. 4B). Mesoscutum smooth and shiny, round in dorsal view. Scutellum also smooth and shiny. Propodeum with two carinae thin and well defined on top widening at base forming a plate (Fig. 4E).
Forewing. Longer than body, 1.3 times as long as mesosoma and metasoma together (Fig. 4A). Closed radial cell, 2.4 times as long as wide in female; 2.2 in male. R1 short and slightly curved; Rs long and curved.
Metasoma. T1 with incomplete ring of setae on anterior part, glabrous at centre, wider laterally. Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Distribution. Nearctic (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
Natural history. Only found on Cinara Curtis, 1835 (Hemiptera: Aphididae) species on conifers through Aphidiinae (Hymenoptera: Braconidae) and Pauesia Quilis, 1931 (Hymenoptera: Braconidae) species (Andrews Reference Andrews1978).
Alloxysta longiventris Baker, Reference Baker1896
(Fig. 5)
Alloxysta longiventris Baker, Reference Baker1896: (A) forewing; (B) pronotum; (C) antenna; (D) propodeum.
Alloxysta longiventris Baker, Reference Baker1896: 134.
Type materialof Alloxysta longiventris Baker.Holotype, ♀ (USNM), with the following labels: “Colo 1229”, “Ft Collins Col”, “5-26-94, sweeping alfalfa” (handwritten), “CF Baker Collector”, “Alloxysta longiventris Baker” (red label, handwritten), “Type Number 103528, USNM” (red label).
Diagnosis.Alloxysta longiventris is mainly characterised as having a partially open radial cell, pronotal carinae absent and propodeal carinae present, with rhinaria and club-shaped from F4. It is similar to A. patens Hellén, 1963 but can be differentiated by the relative sizes of flagellomeres: F1–F3 subequal in length in A. longiventris (Fig. 5C) but F1 longer than F2 and F2 subequal to F3 in A. patens; and dimensions of radial cell, which is 2.3 times as long as wide in A. commensuratus (Fig. 5A) but 2.8 times in A. patens.
Redescription. Female.
Colour. Head, mesosoma and metasoma dark brown. Scape, pedicel, F1–F3 dark yellow; F4–F11 yellowish brown. Legs yellow and veins yellowish brown.
Head. Transversally ovate, smooth and shiny, slightly wider than high in front view. Setae below and between toruli, few scattered setae above toruli. Scattered setae on vertex and on face. Transfacial line 1.2 times compound eye height. Malar space 0.5 times compound eye height.
Antenna. F1–F3 thinner and smoother than remaining flagellomeres, F4–F11 with rhinaria and club-shaped. Pedicel 2.0 times as long as wide; F1 2.5 times as long as wide; F2 2.0 times as long as wide; F3 2.2 times as long as wide; F4 2.5 times as long as wide. Pedicel 1.2 times as long as F1; F1–F3 subequal in length; F4 1.3 times as long as F3; F4–F11 subequal in length, width and shape (Fig. 5C).
Mesosoma. Mesosoma covered with sparse setae, the setae nearly absent at anterior and distolateral margins of pronotum and, denser apical scutellum and propodeum. Pronotum without carinae present (Fig. 5B). Mesoscutum smooth and shiny, round in dorsal view. Scutellum also smooth and shiny. Propodeum with two carinae forming a plate, with few setae on bottom and sides straight (Fig. 5D).
Forewing. Longer than body, 1.5 times as long as mesosoma and metasoma together (Fig. 4A). Partially open radial cell, 2.3 times as long as wide (Fig. 5A). R1 short and curved; Rs long and also curved.
Metasoma. T1 with incomplete ring of setae on anterior part, glabrous at centre, wider laterally. Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Distribution. United States of America (Colorado) (Baker Reference Baker1896).
Natural history. Unknown.
Alloxysta macrophadna (Hartig, 1841)
Alloxysta alaskensis Ashmead, Reference Ashmead1902: 142 new synonym.
Alloxysta coniferensis Andrews, Reference Andrews1978: 59 new synonym.
Type material of Alloxysta coniferensis Andrews. Holotype ♀ (USNM), with the following labels: “1Hs5, 118”, “LH Source-fir, Gatlinburg, Tenn, GSMNP, 5900, NE sweeps 15 Jun ‘47”, “R.H. Whittaker”, “♂”, “Holotype Alloxysta coniferensis Andrews” (red label), “Type Number 103511, USNM” (red label). Paratypes 2♂ and 3♀ (USNM), with the following labels: “3 Hh, 97”, “3H Source-fir, Gatlinburg, Tenn, GSMNP, 5900, NE sweeps 11 Jul ‘47”, “R.H. Whittaker”, “♂”, “Paratype Alloxysta coniferensis Andrews” (yellow label): 1♂; “1Hh5, 137”, “3H Source-fir, Gatlinburg, Tenn, GSMNP, 5900, NE sweeps 18 Jun ‘47”, “R.H. Whittaker”, “♂”, “Paratype Alloxysta coniferensis Andrews” (yellow label): 1♂; “2ES4, 67”, “2E Hemlock For., Gatlinburg, Tenn, GSMNP, 4000, NE sweeps 29 Jun ‘47”, “R.H. Whittaker”, “US National Museum Collection” (yellow label), “♀”, “Paratype Alloxysta coniferensis Andrews” (yellow label): 1♀; “2Eh, 1, 32, 67”, “2E Hemlock For., Gatlinburg, Tenn, GSMNP, 4000, NE sweeps 29 Jun ‘47”, “R.H. Whittaker”, “US National Museum Collection” (yellow label), “♀”, “Paratype Alloxysta coniferensis Andrews” (yellow label): 1♀; “3Eh4, 42”, “2E Hemlock For., Gatlinburg, Tenn, GSMNP, 4000, NE sweeps 13 Jul ‘47”, “R.H. Whittaker”, “US National Museum Collection” (yellow label), “♀”, “Paratype Alloxysta coniferensis Andrews” (yellow label): 1♀.
The female of A. coniferensis has a slightly longer F2 than F1 according to Andrews (Reference Andrews1978), but these two flagellomeres in the paratypes look very similar under the microscope. Taking into account the radial cell dimensions, presence of pronotal carinae, absence of propodeal carinae and curved flagellomeres in the male, we establish this new synonymy here.
Type material of Alloxysta alaskensis Ashmead. Lectotype ♀ (USNM), with the following labels: “8, 6, 85” (handwritten), “St. Paul Isl. Alaska”, “Collector T. Kincald”, “♀ Type No 5525, USNM” (red label), “Alloxysta alaskensis ♀Ashm.” (handwritten), “Lectotype alaskensis Ashmead, Design.:A.S. Menke”. Paralectotype ♂ (USNM), with the following labels: “unclear, 6-12-99” (handwritten), “Harriman Expedition ‘99, T. Kincald Collector”, “Type No. 5525, ♂, USNM” (red label), “Alloxysta alaskensis ♂, Ashm.” (handwritten).
The type series of A. alaskensis consists of five specimens (3♂ and 2♀). However, the species is represented by only two specimens in the USNM: the lectotype ♀ designated by Andrews (Reference Andrews1978) and one paralectotype ♂. Andrews regarded this specimen as the holotype, but this designation is incorrect because Ashmead based his description on more than one specimen. According to Menke and Evenhuis (Reference Menke and Evenhuis1991) “Andrew's recognition of a holotype constituted a lectotype designation”. These authors placed a lectotype label on the specimen.
Distribution. Holarctic, previously only known from the Palaearctic region (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
Natural history. Usually found on Medicago sativa Linnaues (Fabaceae) through Aphidius ervi Haliday, 1843 (Hymenoptera: Aphidiidae) (Evenhuis Reference Evenhuis1982; Fergusson Reference Fergusson1986), Macrosiphum ptericolens (Patch) (Hemiptera: Aphididae) through Aphidius Nees, 1819 (Hymenoptera: Braconidae) species and Praon species (Müller et al. Reference Müller, Adriaanse, Belshaw and Godfray1999). The complete list of hosts is presented on Ferrer-Suay et al. (Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a). The hosts of the species collected in the United States of America are still unknown.
Alloxysta minuscula Andrews, Reference Andrews1978
(Fig. 6)
Alloxysta minuscula Andrews, Reference Andrews1978: (A) forewing; (B) antenna; (C) radial cell; (D) pronotum; (E) propodeum; (F) body.
Alloxysta minuscula Andrews, Reference Andrews1978: 66.
Type materialof Alloxysta minuscula Andrews.Holotype ♂ (CNC), with the following labels: “Ottawa, Ont. in lab 1937, G.A. Hobbs” (handwritten), “Holotype Alloxysta minuscula Andrews” (red label), “Holotype Alloxysta minuscula Andrews CNC No. 19023” (red label). Paratypes 6♂ (CNC) and 2♂ in author's collection with the same data as holotype. According to Andrews (Reference Andrews1978) the holotype is a female but the specimen studied that has the holotype label is a male.
Diagnosis.Alloxysta minuscula is mainly characterised having partially open radial cell, pronotal carinae present, propodeal carinae absent, rhinaria and club-shaped begin in F4 in female and F2 in male; F1 subequal to pedicel. According with these features it is easily differentiated from the other Alloxysta species.
Redescription.
Colour. Head and metasoma brown, mesosoma yellowish brown. Antenna yellowish brown. Legs yellow and veins yellowish brown.
Head. Transversally ovate, smooth and shiny, slightly wider than high in front view. Setae below and between toruli, no setae above toruli. No setae on vertex and many setae on face. Transfacial line 1.2 times compound eye height. Malar space 0.4 times compound eye height.
Antenna. Female: F1–F3 thinner and smoother than remaining flagellomeres, F4–F11 with rhinaria and club-shaped. F1 subequal to pedicel and F2, F2 slightly longer than F3. Male: F1 thinner and smoother than remaining flagellomeres, F2–F12 with rhinaria and club-shaped. Pedicel 1.5 times as long as wide; F1 2.8 times as long as wide; F2 2.1 times as long as wide; F3 2.1 times as long as wide; F4 2.1 times as long as wide. Pedicel-F12 subequal in length (Fig. 6B).
Mesosoma. Mesosoma covered with sparse setae, the setae nearly absent at distolateral margins of pronotum and, denser on apical scutellum. Pronotum with two thick carinae present (Fig. 6D). Mesoscutum smooth and shiny, round in dorsal view. Scutellum also smooth and shiny. Propodeum without carinae present (Fig. 6E).
Forewing. Longer than body, 1.8 times as long as mesosoma and metasoma together (Fig. 6A). Partially open radial cell, 2.5 times as long as wide (Fig. 6C). R1 short and curved; Rs long and also curved.
Metasoma. T1 with incomplete ring of setae on anterior part, glabrous at centre, wider laterally. Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Distribution. Canada (Ottawa) (Andrews Reference Andrews1978).
Natural history. Unknown.
Alloxysta nothofagi Andrews, Reference Andrews1976
(Fig. 7)
Alloxysta nothofagi Andrews, Reference Andrews1976: (A) forewing; (B) radial cell male; (C) radial cell female; (D) antenna female; (E) pronotum; (F) antenna male; (G) propodeum.
Alloxysta nothofagi Andrews, Reference Andrews1976: 286.
Type material of Alloxysta nothofagi Andrews.Paratypes 2♀ and ♂ (USNM), with the following labels: “18 Km, W. Angol, Malleco Prov. Chile, ii-10.1967, E.I. Schlinger, E67-2-10b” (handwritten), “Host Plant: Nothofagus obliqua” (handwritten), “Host aphid: Neuquenaphis schlingeri” (handwritten), “Primary parasite: Pseudephedrus sp.” (handwritten), “Paratype: Alloxysta nothofagae Andrews ♀” (handwritten, yellow label): 2♀; “3.7 Km, S. Puerto Moreno, Rio Negro Prov., xi-17-1966 Argentina, E.I. Schlinger, E66-11-17a”, “Host plant: Nothofagus antarctica” (handwritten), “Host aphid: Neuquenaphis edwardsi” (handwritten), “Paratype: Alloxysta nothofagae Andrews” (handwritten, yellow label): 1♂. The holotype is deposited in the University of Chile (Santiago, Chile) (Andrews Reference Andrews1976). Two more paratypes are deposited in author's collection.
Diagnosis.Alloxysta nothofagi is very similar to A. postica (Hartig, 1841) because of its partially open radial cell, pronotal and propodeal carinae and F1 subequal to pedicel. However, they can be differentiated by the coloration of the body (bicoloured in A. nothofagi and brown in A. postica); grade of pubescence at the pronotum (abundant in A. nothofagi and with scattered setae in A. postica); length/width ratio of the flagellomeres (larger in A. postica than in A. nothofagi) and the host (Neuquenaphis Blanchard, 1939 (Hemiptera: Aphididae) species in A. nothofagi but Aphis species in A. postica).
Redescription.
Colour. Head and metasoma brown, mesosoma yellowish brown. Antenna yellowish brown. Legs yellow and veins yellowish brown.
Head. Transversally ovate, smooth and shiny, slightly wider than high in front view. Setae below and between toruli, no setae above toruli. Scattered setae on vertex and many setae on face. Transfacial line 1.1 times compound eye height. Malar space 0.4 times compound eye height.
Antenna. Female: F1–F3 smoother than remaining flagellomeres, F4–F11 with rhinaria; club-shaped not visible. Pedicel 2.0 times as long as wide; F1 3.3 times as long as wide; F2 2.9 times as long as wide; F3 2.8 times as long as wide; F4 3.0 times as long as wide. F1 subequal to F1, F1 1.1 times as long as F2, F3 1.1 times as long as F2, F4 1.1 times as long as F3, F4–F11 subequal in length, width and shape (Fig. 7D). Male: F1 thinner and smoother than remaining flagellomeres, F2–F12 with rhinaria and club-shaped. Pedicel 2.0 times as long as wide; F1 4.0 times as long as wide; F2 3.0 times as long as wide; F3 2.4 times as long as wide; F4 2.0 times as long as wide. F1 1.2 times as long as pedicel, F1–F3 subequal in length, F3 1.2 times as long as F4, F4–F12 subequal in length, width and shape (Fig. 7F).
Mesosoma. Mesosoma covered with sparse setae, the setae nearly absent at anterior margin of pronotum and, denser on apical scutellum and propodeum. Pronotum with two thick carinae present clearly visible (Fig. 7E). Mesoscutum smooth and shiny, round in dorsal. Scutellum also smooth and shiny. Propodeum with two carina forming a plate (Fig. 7G).
Forewing. Longer than body, 1.7 times as long as mesosoma and metasoma together (Fig. 7A). Partially open radial cell, 2.4 times as long as wide in both male and female (Figs 7B, 7C). R1 short and curved; Rs long and also curved.
Metasoma. T1 with incomplete ring of setae on anterior part, glabrous at centre, wider laterally. Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Distribution. Argentina and Chile (Andrews Reference Andrews1976).
Natural history. Only found on Neuquenaphis edwardsi (Laing) on Nothofagus antarctica (Forster) Oersted (Nothofagaceae) through “Pseudephedreus” species; Neuquenaphis schlingeri Hille Ris Lambers, 1968 (Hemiptera: Aphididae) on Nothofagus obliqua (Mirbel) Oersted through “Pseudephedreus” species (Andrews Reference Andrews1976).
Alloxysta obscurata (Hartig, 1840)
Dilyta bicolor Baker, Reference Baker1896: 132 new synonym.
Alloxysta robusta Baker, Reference Baker1896: 133. Synonymised with A. bicolor by Andrews (Reference Andrews1978: 55).
A. anthracina Andrews, Reference Andrews1978: 54 new synonym.
Type material of Alloxysta anthracina Andrews. Holotype ♂ (CNC), with the following labels: “Mile 236 Alaska, Richard. Hwy, 16-vi-1951, W.R.M. Mason”, “Holotype Alloxysta anthracina Andres” (red label), “Holotype Alloxysta anthracina Andrews CNC No. 19077” (red label).
Alloxysta anthracina was synonymised with A. pallidicornis (Curtis, 1838) by Evenhuis and Kiriak (Reference Evenhuis and Kiriak1985). However, examination of the lectotype showed that A. anthracina can be easily differentiated from A. pallidicornis by the absence of propodeal carinae in the latter and presence in the former (independent, thin on top and thick on bottom), and shape of the radial cell (partially open in A. anthracina, completely open in A. pallidicornis). The features of A. anthracina correspond to those of A. obscurata, considered a valid species following revision of type material (see above).
Type material of Dilyta bicolor Baker. Holotype ♀ (USNM), with the following labels: “Colo 1563”, “Ft Collins Col”, “6-13-95, misc. sweeping” (handwritten), “CF Baker Collector”, “Dilyta bicolor Baker” (red label, handwritten), “Type Number 103533, USNM” (red label). According to Andrews (Reference Andrews1978) there is also a paratype but it has not been found in the USNM collection. Another specimen determined as A. bicolor by Weld is present but the locality and date of capture are different from the type material so it cannot be considered as paratype.
Type material of Alloxysta robusta Baker. Holotype ♀ (USNM), with the following labels: “Colo 1547”, “Ft Collins Col”, “6-7-95, misc. sweeping” (handwritten), “CF Baker Collector”, “Alloxysta robusta Baker” (red label, handwritten), “Type Number 103530, USNM” (red label).
Alloxysta robusta was synonymised with A. bicolor by Andrews (Reference Andrews1978). This synonymy is here confirmed.
Distribution. Holarctic and Neotropical (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a, Reference Ferrer-Suay, Selfa and Pujade-Villar2012c).
Natural history. It has been found on different combinations of hosts. The complete list of hosts is presented on Ferrer-Suay et al. (Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
Alloxysta pilipennis(Hartig, 1840)
Alloxysta dicksoni Andrews, Reference Andrews1978: 60 new synonym.
Type materialof Alloxysta dicksoni Andrews.Holotype ♀ (USNM), with the following labels: “Bowie Md, vii-4-'45, DDT Expt”, “♀”, “Holotype Alloxysta dicksoni Andrews” (red label), “Type Number 103512, USNM” (red label). Paratypes 6♂ (USNM) with the following labels: “Bowie Md, vii-4-'45, DDT Expt”, “♂”, “Paratype Alloxysta dicksoni Andrews” (red label), “Type Number 103512, USNM” (yellow label): 3♂; “Bowie Md, vii-7-'45, DDT Expt”, “♂”, “Paratype Alloxysta dicksoni Andrews” (red label), “Type Number 103512, USNM” (yellow label): 1♂; “Bowie Md, vii-2-'45, DDT Expt”, “♂”, “Paratype Alloxysta dicksoni Andrews” (red label), “Type Number 103512, USNM” (yellow label): 2♂. Other two paratypes are deposited in the author's collection. The antennae of the holotype are missing, but we have checked them with the original description and figures (Andrews Reference Andrews1978).
Distribution. Palaearctic and Neotropical (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a, Reference Ferrer-Suay, Selfa and Pujade-Villar2012c).
Natural history. Only found on Aphis species through Pontania proxima (Audinet-Serville, 1823) (Hymenoptera: Tenthredinidae) (Dalla Torre and Kieffer Reference Dalla Torre and Kieffer1910).
Alloxysta schlingeri Andrews, Reference Andrews1978 species dubia
Alloxysta schlingeri Andrews, Reference Andrews1978: 68.
Type material of Alloxysta schlingeriAndrews.Holotype ♂ and 9 Paratypes (4♂ and 5♀). They have not been found (M. Buffington, USNM, personal communication).
Comments. According to Andrews (Reference Andrews1978) the holotype and one female paratype were deposited in the USNM, but we have been unable to locate this material in the USNM collections. The deposition of the rest of the type material was not mentioned. Thus, we conclude that this type material has been lost.
Some important characters of A. schlingeri are not mentioned in the original description, including the presence/absence of pronotal and propodeal carinae, and the radial cell shape. Moreover, pictures of the first antennomeres (Andrews Reference Andrews1978, Figs. 67–69) cannot validate this species. For all these reasons, we conclude that A. schlingeri should be considered a species dubia until the types can be found and examined.
Alloxysta vandenboschi Andrews, Reference Andrews1978 revised status
(Fig. 8)
Alloxysta vandenboschi Andrews, Reference Andrews1978: (A) forewing; (B) pronotum; (C) antenna; (D) propodeum.
Alloxysta vandenboschi Andrews, Reference Andrews1978: 69.
Type materialof Alloxysta vandenboschi Andrews.Holotype ♂ (USNM), with the following labels: “2 mi. S. Kelsey Bay, Vancouver Is., Canada, vii-1-66, F.G. Andrews”, “F66-7-1a, Alnus sp., Euceraphis gillettie, Davidson”, “♂”, “Primary parasite Praon sp”, “Holotype Alloxysta vandenvoschi Andrews” (red label), “Type Number 103514, USNM” (red label). Paratype ♂ (CNC), with the following labels: “Mile 140, Steese Bay, Alaska, VI-21-1951 (Mason and McGillis).
Diagnosis.Alloxysta vandenboschi is mainly characterised having completely open radial cell, pronotal carinae present, propodeal carinae absent without setae where the carinae are usually present in Charipinae, F2 and F3 curved. There is no other known Alloxysta species closely related.
Redescription. Male.
Colour. Head yellowish brown, mesosoma and metasoma dark brown. Scape, pedicel, F1–F3 yellow; F4–F12 yellowish brown. Legs yellow and veins yellowish brown.
Head. Transversally ovate, smooth and shiny, slightly wider than high in front view. Setae below and between toruli, no setae above toruli. Scattered setae on vertex and on face. Transfacial line 0.8 times compound eye height. Malar space 0.4 times compound eye height.
Antenna. F1 thinner and smoother than remaining flagellomeres, F2–F12 with rhinaria and club-shaped. F2 and F3 strongly curved. Pedicel 1.7 times as long as wide; F1 6.3 times as long as wide; F2 5.0 times as long as wide; F3 4.8 times as long as wide; F4 3.7 times as long as wide. F1 1.9 times as long as pedicel; F1–F3 subequal in length; F4 1.1 times as long as F3; F4–F11 subequal in length, width and shape (Fig. 8C).
Mesosoma. Mesosoma covered with sparse setae, the setae nearly absent at anterior margin of pronotum and denser on apical scutellum, pronotum, and propodeum. Pronotum covered with two thick carinae (Fig. 8B). Mesoscutum smooth and shiny, round in dorsal view. Scutellum also smooth and shiny with few scattered setae. Propodeum without carinae and without setae where carina usually placed (Fig. 8D).
Forewing. Longer than body, 1.7 times as long as mesosoma and metasoma together (Fig. 8A). Completely open radial cell, 2.9 times as long as wide. R1 short and curved; Rs long and also curved.
Metasoma. T1 with incomplete ring of setae on anterior part, glabrous at centre, wider laterally. Metasoma smooth and shiny, T3 and T4 clearly distinguished.
After studying the holotype, Menke and Evenhuis (Reference Menke and Evenhuis1991) synonymised this species with A. obscurata. However, these two species are clearly different, they can be differentiated by the radial cell shape (completely open in A. vandenboschi, partially open in A. obscurata) and relative sizes of flagellomeres (F1–F3 subequal in length in A. vandenboschi but not in A. obscurata, and F2 and F3 clearly curved in A. vandenboschi but only F2 slightly curved in A. obscurata).
Distribution. United States of America (Alaska) (Andrews Reference Andrews1978).
Natural history. Only found on Euceraphis gillettei Davidson, 1915 (Hemiptera: Aphididae) on Alnus Miller (Betulaceae) species through Praon species (Andrews Reference Andrews1978).
Alloxysta victrix (Westwood, 1833)
Charips areolata Kieffer, Reference Kieffer1909: 481. Synonymised by Menke and Evenhuis (Reference Menke and Evenhuis1991: 145).
Allotria tritici Fitch, Reference Fitch1861: 841. Synonymised by Menke and Evenhuis (Reference Menke and Evenhuis1991: 148).
Type material of Charips areolata Kieffer. Lectotype ♀ (USNM), with the following labels: “Forest Hills, Masa, 29.v.1909”, “384” (handwritten), “Ex. same to as types. Exped. with Fayetteville ail 1924”, “US National Museum Collection”, “Alloxysta victrix (Westwood) Det. Fred G. Andrews, 1977”, “Lectotype Charips areolata Kieffer Desig. A.S. Menke”. Paralectotypes 1♂ and 2♀ (USNM), with the following labels: “384”, “US National Museum collection” (yellow label), “Alloxysta victrix (Westwood) Det. Fred G. Andrews, 1977”, “Paralectotype Charips areolata Kieffer, Design.: A.S. Menke”.
Kieffer had eight syntypes of this species of which one of them was deposited in the University of Arkansas (Menke and Evenhuis Reference Menke and Evenhuis1991). The validity of this synonymy is here confirmed.
Type material of Allotria tritici Fitch. Lectotype ♀ (USNM), with the following labels: “15, 341” (handwritten), “Fitch's coll”, “Allotria tritici Fch, a. nigricephs” (handwritten), “lectotype Allotria tritici Desig.: A.S. Menke”.
Complete notes about this type material were given by Menke and Evenhuis (Reference Menke and Evenhuis1991). The validity of this synonymy is here confirmed.
Distribution. Cosmopolitan, present in all continents except Africa and Antarctica (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
Natural history. This species has been found on a wide range of hosts (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a). The most common species where it has been found are Medicago sativa and Acyrthosiphon pisum (Harris, 1776) (Hemiptera: Aphididae) (Andrews Reference Andrews1978; Evenhuis and Barbotin Reference Evenhuis and Barbotin1987) and Aphidius species (Andrews Reference Andrews1978; Fergusson Reference Fergusson1986).
Alloxysta xanthopsis (Ashmead, Reference Ashmead1896)
(Fig. 9)
Alloxysta xanthopsis (Ashmead, Reference Ashmead1896): (A) forewing; (B) radial cell; (C) antenna; (D) pronotum; (E) propodeum.
Allotria xanthopsis Ashmead, Reference Ashmead1896: 185.
Type material of Allotria xanthopsis Ashmead.Holotype ♂ (USNM), with the following labels: “orange aphis” (handwritten), “5”, “Type No. 3282, USNM” (red label), “Allotria xanthopsis Ashm.” (handwritten).
Diagnosis.Alloxysta xanthopsis is mainly characterised by a closed radial cell, pronotal carinae present, propodeal carinae present being thin and well defined, joining at the base, F2 and F3 curved in males. It is similar to A. victrix, because of the colouration of the head, closed radial cell and curved F2–F3 in male specimens, but they can be easily differentiated by the presence of propodeal carinae in A. xanthopsis and absence in A. victrix.
Redescription. Male.
Colour. Head yellowish brown, mesosoma and metasoma dark brown. Scape, pedicel, F1–F3 yellow; F4–F12 yellowish brown. Legs yellow and veins yellowish brown.
Head. Transversally ovate, smooth and shiny, slightly wider than high in front view. Setae below and between toruli, no setae above toruli. Scattered setae on vertex and on face. Transfacial line 1.2 times compound eye height. Malar space 0.5 times of compound height.
Antenna. F1 thinner and smoother than remaining flagellomeres, F2–F12 with rhinaria and club-shaped. F2 and F3 curved. Pedicel 2.5 times as long as wide; F1 2.9 times as long as wide; F2 2.9 times as long as wide; F3 2.7 times as long as wide; F4 2.5 times as long as wide. F1 1.9 times as long as pedicel; F1–F3 subequal in length; F4 1.1 times as long as F3; F4–F11 subequal in length, width and shape (Fig. 9C).
Mesosoma. Mesosoma covered with sparse setae, the setae nearly absent at anterior margin of pronotum and denser on apical scutellum and propodeum. Pronotum with two carinae present (Fig. 9D). Mesoscutum smooth and shiny, round in dorsal view. Scutellum also smooth and shiny. Propodeum with two carinae straight and well defined, joining at base (Fig. 9E).
Forewing. Longer than body (Fig. 9A). Closed radial cell, 2.1 times as long as wide (Fig. 9B). R1 short and curved; Rs long and also curved.
Metasoma. T1 with incomplete ring of setae on anterior part, glabrous at centre, wider laterally. Metasoma smooth and shiny, T3 and T4 clearly distinguished.
Distribution. Nearctic (Ferrer-Suay et al. Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
Natural history.Alloxysta xanthopsis has been found on different combinations of hosts. The complete list of hosts is presented on Ferrer-Suay et al. (Reference Ferrer-Suay, Paretas-Martínez, Selfa and Pujade-Villar2012a).
Acknowledgements
The authors are grateful to Dr. Matthew Buffington and Dr. John Huber for their hospitality during our stay in the USNM and CNC, respectively. We also want to thank to Jennifer Read for taking the photos of the type material deposited in the CNC and her kind attention during our stay. This research was supported by the project CGL2011-22889 of the Ministerio de Ciencia e Innovación (Spain) and the grant AP2009-4833 of the Ministerio de Educación (Spain).
