6urn:lsid:arphahub.com:pub:DE333E49-E4EE-5693-9BEF-D446E6301C98urn:lsid:zoobank.org:pub:55528528-5C97-4D79-A718-8B3D153B37CCJournal of Hymenoptera ResearchJHR1070-94281314-2607Pensoft Publishers10.3897/jhr.54.1045710457Research ArticleAphelinidaeChalcidoideaSystematicsTaxonomyCenozoicNeogeneWorldRevision of the asychis species group of Aphelinus (Hymenoptera: Aphelinidae)ShirleyXanthe A.xanthe.shirley@gmail.com1WoolleyJames B.1HopperKeith R.2Department of Entomology, Texas A&M University, College Station, TX, 77843 USATexas A&M UniversityCollege StationUnited States of AmericaUSDA-ARS-BIIRL, Newark, DE, 19713 USAUSDA-ARS-BIIRLNewarkUnited States of America
Corresponding author: Xanthe A. Shirley (xanthe.shirley@gmail.com)
Academic editor: Hannes Baur
20172702201754132FFE4FFD9-484C-AB0C-2932-1C29FF94FFE16F3F5A35-3EBD-48FE-9DFC-9B350F129DE33228720909201612122016Xanthe A. Shirley, James B. Woolley, Keith R. HopperThis 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.http://zoobank.org/6F3F5A35-3EBD-48FE-9DFC-9B350F129DE3
Aphelinus (Hymenoptera: Aphelinidae) is a genus of parasitoid wasps that has a long history of use in biological control programs against aphids. Past research shows that species delimitation within Aphelinus is greatly complicated by lack of comprehensive literature and the existence of cryptic species complexes. One of these complexes is the Aphelinusasychis species group. Through the development of a morphological character set, a revision of the Aphelinusasychis species group was conducted. Two new species, Aphelinussinensissp. n., and Aphelinuskazakhstanensissp. n., are described, and the two existing valid species within the asychis group, Aphelinusasychis and Aphelinussemiflavus are redescribed and lectotypes are designated for Aphelinussemiflavus and Aphelinusbrevipennis (a junior synonym of A.semiflavus). We also provide a key for identifying species in the asychis group.
Shirley XA, Woolley JB, Hopper KR (2017) Revision of the asychis species group of Aphelinus (Hymenoptera: Aphelinidae). Journal of Hymenoptera Research 54: 1–32. https://doi.org/10.3897/jhr.54.10457
Introduction
Aphelinus Dalman, 1820 is a genus of chalcidoid wasps in the family Aphelinidae, subfamily Aphelininae (see Hayat 1972). All species of Aphelinus are endoparasitoids of aphids, and the genus has a long association with biological control research against aphid pests (Arce Gomez and Rumiatto 1989, Cate et al. 1973, Clausen 1978, Elliott et al. 1995, Raney 1971, Sell and Kuo-Sell 1989, Starks et al. 1976). The focus of this paper is the Aphelinusasychis group, which differs from other Aphelinus species groups in two traits: submarginal vein with only 2 [not 3 or more] setae (Hayat 1998) and oviposition probing/penetration site on host dorsal [not ventral] surface (De Farias and Hopper 1999). There are currently two valid species in the asychis group: A.semiflavus and A.asychis. Aphelinussemiflavus Howard, 1908 was originally collected near Fort Collins, Colorado, parasitizing Myzuspersicae (Sulzer, 1776), green peach aphid. Aphelinusasychis Walker was described in 1839 from British and Irish material.
Aphelinusasychis is common in both the Old World and New World, and it is an important parasitoid of aphids (ca. 60 documented aphid hosts). It has been used in biological control of at least six aphid species (Kalina and Stary 1976, Noyes 2016). Prominent biological control targets of A.asychis include Therioaphistrifolii (Monell, 1882), the spotted alfalfa aphid, A.asychis was also the focus of work on biological control programs of Schizaphisgraminum (Rondani, 1852), the greenbug, during the 1970s and 1980s (Cate et al. 1973, Johnson et al. 1979, Summy et al. 1979) and Diuraphisnoxia (Mordvilko, 1921), the Russian wheat aphid, during the 1990s (Brewer et al. 2001, Hopper et al. 1998, Prokrym et al. 1998). During foreign exploration for natural enemies of the Russian wheat aphid in the 1990s, A.asychis was found in Europe, Asia, northern Africa, and South America, where it parasitizes Diuraphis spp. (Gonzalez et al. 1994, Hopper et al. 1998).
A major factor that impedes the success of biological control programs is delayed recognition of cryptic species. Testing for reproductive compatibility is one way to discover cryptic species. Kazmer et al. (1996) tested for interculture reproductive compatibility among seven lab cultures of A.asychis collected during foreign exploration. They examined all possible crosses (49), and found three completely and reciprocally reproductively incompatible groups: (1) Mediterranean basin, (2) Kazakhstan, and (3) China. These were also reflected in clusters from phenetic analysis of 61 RAPD loci banding patterns. As explained in the descriptions below, we were able to examine voucher material from the original collections referred to in Kazmer et al. (1996), or material from cultures established from these collections.
A second issue in this species group is considerable confusion over whether A.asychis and A.semiflavus are different species. Ferrière (1965) and Nikol’skaya and Yasnosh (1966) synonymized A.semiflavus with A.asychis. Mackauer and Finlayson (1967) argued that they should remain separate because they differ in host ranges, and could be cryptic species. Aphelinussemiflavus has since been treated as a valid species by several researchers (Raney 1971, Raney et al. 1973, Ro and Long 1997). However, distinguishing between these species has been difficult. For example, during the biological control program against T.trifolii, in the 1950’s, A.asychis was initially recorded as A.semiflavus (Clausen 1978, Hagen and van den Bosch 1968).
Further work in the asychis species group is needed to (1) understand the nature and relationships among the three groups found by Kazmer et al. (1996) and among other species in the asychis species complex and (2) determine whether A.asychis and A.semiflavus are races of one species with different host ranges or are two distinct cryptic species.
Diagnosis of asychis species group
Following Hayat (1998), we consider a submarginal vein with only two setae to be diagnostic for the A.asychis group. In females, F1 and F2 are subquadrate, and F3 is 1.2-2.0× longer than wide. In males, F1 and F2 are wider than long, and F3 is >3.0× longer than wide. Brachyptery is also common in this group, particularly in males. There is marked sexual dimorphism, particularly in antennal proportions and coloration, which is not found in other Aphelinus species.
Material and methods
Specimens used in this study were killed in 95% ethanol and stored in freezers. Most were then critical-point-dried using a Samdri 790 CPD unit. Critical-point-dried specimens were card mounted with Franklin International’s water soluble Titebond Liquid Hide Glue. Selected specimens were slide-mounted following Noyes’ (1982) protocol. All card-mounted and slide-mounted specimens were assigned individual accession numbers (e.g., TAMU-ENTO X0852885, USNM ENT 4532898, etc.). Label data for type specimens are reported verbatim, where | signifies a new line on a label and || separates different labels. The following acronyms for museum collections are used, followed in some cases by the actual acronym used with specimen accession numbers: ANIC, Australian National Insect Collection (Canberra); BMNH [BMNH(E)], Natural History Museum (London); CNC (CNC HYMEN), Canadian National Collection of Insects and Nematodes (Ottawa); EMEC, Essig Museum Entomology, University of California (Berkeley); NHW, Naturhistorisches Museum (Vienna); TAMU (TAMU-ENTO), Texas A&M University Insect Collection (College Station, TX); UCR (UCRC ENT), University of California (Riverside, CA); USNM (USNM ENT) National Museum of Natural History (Washington, D.C.). Complete label data for any or all material examined are available upon request to the authors.
Images for figures were acquired using digital imaging and image-stacking. Specimens photographed for coloration were removed from alcohol storage, placed on a layer of water-based, water-soluble jelly in a small watch glass, submerged in alcohol, and photographed using a Leica M205 FA stereomicroscope and Leica Applications Suite software (ver. 4.5) as were card-mounted or point-mounted specimens. Slide-mounted specimens were photographed using an Olympus BH2 microscope with DIC illumination and Image-Pro Plus software (ver. 7.0). Zerene Stacker (http://zerenesystems.com) was used for all image stacking. Adobe Photoshop CS6, Adobe Lightroom 5, and Adobe InDesign CS6 were used for final modifications to images and layout of plates. All images were deposited in mx, a web-based content management database system. Morphological codings were conducted in mx. The mx system is open source, with further documentation available at http://mx.phenomix.org.
Morphology
See Appendix Table 1 for a list of the morphological terms used in this study, followed by a definition and a URI (uniform resource identifier) that links to more information on that character in the Hymenoptera Anatomy Ontology (Yoder et al. 2010) database.
Morphology measurements
Measurements from slide mounts were taken using an eyepiece reticle in a Zeiss standard 16 microscope. Measurements from card mounts were taken using an eyepiece reticle in a Leica MZ16 microscope. Raw measurements are only reported for body length, which is followed by the range and the number of specimens measured. The remaining measurements are reported as ratios.
Head. The length of the head was measured from the anterior to the posterior margin in dorsal view (Fig. 1a, hl). The frontovertex length was measured in dorsal view from the dorsal margin of the scrobal impression to the occiput (Fig. 1a, fvl). Head and frontovertex widths were both measured at their widest points. The posterior ocellar diameter (Fig. 1a, ow), distance from posterior ocelli to eye margin, (Fig. 1a, ool), distance between ocelli (Fig. 1a, pol) and distance from posterior ocelli to occipital margin (Fig. 1a, ocp) were measured as illustrated. The widths of each antennal segment (scape, pedicel, F1, F2, F3, and club) were measured at their widest points. The lengths of each antennal segment were measured from proximal to distal end. The scape of males in the asychis group bears a longitudinal row of pores on a convex ridge on the anterior (ventral) surface (Fig. 2), that appear to be openings for internal glands.
Meso/Metasoma length. Meso/metasoma length of specimens was measured from the anterior margin of the pronotum to the apex of the epiproct using slide-mounted specimens. The lengths of the mesosoma, the mid lobe of mesoscutum, and the scutellum were measured from their anterior to posterior margins along the mid line and widths were measured at their widest points.
Wings. Fore wing measurements are shown in Figure 1b; hind wing measurements follow those of the fore wing.
Male Genitalia and Ovipositor. The length of the phallobase was measured from the anterior margin of the genital capsule to the posterior end of the digiti (Fig. 1c, phl). The width of the phallobase was measured at its widest point (Fig. 1c, phl). The length of the digitus was measured between its most anterior to most posterior points, and its width was measured at its widest point (Fig. 1c, dig). Ovipositor length was measured as illustrated in Figure 1d.
Aphelinussinensis n. sp., male, scape, ventral view. Note the five linearly arranged exocrine gland pores.
https://binary.pensoft.net/fig/124060ResultsKey to species in the asychis group of Aphelinus, male or female specimens.
1
Procoxa yellow (Fig. 9c–f)
sinensis Shirley & Woolley, sp. n.
–
Procoxa brown (Figs 3c, 3d, 5c–5f, 7c, 7d)
2
2
Profemur entirely yellow or pale (Figs 7c, 7d)
semiflavus Howard
3
Profemur brown at base (Figs 3c, 3d, 5c–5f,)
3
4
Mesotibia entirely yellow (Figs 5c–5f)
kazakhstanensis Shirley & Woolley, sp. n.
5
Mesotibia brown at base or dark with apex pale (Figs 3c, f)
asychis Walker
AnimaliaHymenopteraAphelinidae76AA7B51-1413-52D7-93AD-F6AC4A25C48EAphelinusasychisWalker, 1839Figs 3, 4AphelinusasychisWalker, 1839, lectotype designation by Graham (1976).AphelinuseuthriaWalker, 1839, synonymy and lectotype designation by Graham (1976).MyinaaffinisFörster, 1841, synonymy and lectotype designation by Graham (1976).Aphelinusaffinis(Förster, 1841): Dalla Torre (1898).AphelinusbrevicalcarThomson, 1876, synonymy and lectotype designation by Graham (1976).AphelinusbrachypteraKurdjumov, 1913, synonymy and lectotype designation by Graham (1976).AphelinusdubiaKurdjumov, 1913, synonymy and lectotype designation by Graham (1976).Diagnosis.
Females and males. Legs with procoxa brown (Fig. 3d) [not yellow (Fig. 9d)], profemur and mesofemur dark brown with apex yellow or pale (Fig. 3d) [not entirely yellow (Figs 7d, 9d) or light brown with apex yellow or pale (Fig. 5d)], mesotibia dark brown with apex yellow (Figs 3c, 3d) [not entirely yellow (Figs 5d, 7d, 9d)] and metatibia dark brown with apex yellow (Figs 3c, 3d) [not entirely yellow (Fig. 9d) or light brown with apex yellow or pale (Figs 5d, 7d)].
Color (Fig. 3b, 3d, 3f). Head and mesosoma dark brown; radicle and basal portion of scape yellow/white, apical portion of scape and pedicel brown, and F1, F2, F3, and club yellow, tip of club dusky; legs with mesocoxa and metacoxa brown, metafemur yellow, protibia yellow; metasoma yellow from base to apex, lateral margins of metasoma darker than mesal area except in basal quarter.
Body length. 0.9 mm (n=2; slide mounts).
Head (Figs 3b and 4b). Width 0.8–1.1 head length in anterior view; frontovertex width 0.5× head width and 1.4–2.6× frontovertex length; posterior ocelli diameter 0.5× posterior ocelli to eye margin distance and 1.5× posterior ocelli to occipital margin distance; antenna as in Figure 4b with scape length 6.5–6.6× scape width, pedicel length 1.9–2.1× pedicel width, F1 and F2 subquadrate, length of both 1.1× width, F3 length 1.3–1.5× F3 width, and club length 2.9–3.2× club width and 2.8–3.4× F3 width, club with 8 longitudinal sensilla.
Mesosoma (Figs 3d, 3f and 4g). Mid lobe of mesoscutum length 0.7–0.8× mid lobe width with two pairs of long setae (one pair lateral and one pair posterior) and 29 short setae; side lobes of mesoscutum each with one pair of long setae and one pair of short setae; scutellum with two pairs of long setae (one pair anterior and one pair posterior); mesotibial spur length 0.6× mesobasitarsus length, metatibial spur length 0.6–0.8× metabasitarsus length.
Fore wing (Fig. 4d). Length 2.8× fore wing width, longest marginal seta 0.2× fore wing width; costal cell length 0.7× length of marginal vein, with one line of 5 setae on ventral surface and 1 dorsal setae in apical quarter; marginal vein with two rows of 10 large dorsal setae, one row of 5 small dorsal setae, and one row of 7 ventral setae; interspace between basal cell and linea calva with 25 setae arranged in three complete line and one incomplete line; linea calva closed with three setae at its posterior end, setae bordering linea calva proximally are arranged uniformly and evenly to posterior margin of wing.
Hind wing (Fig. 4e). Length 4.4× hind wing width, longest marginal seta 0.5× hind wing width.
Metasoma (Figs 3d, 3f and 4f). Length 1.8–2.0× mesosoma length; ovipositor length 1.2–1.3× mesotibia length and 1.0–1.2× metatibia length; third valvula length 0.2–0.3× ovipositor length.
Description. Male (Figs 3a, 3c, 3e and 4a, 4c, 4h). Similar to female except:
Color (Fig. 3a, 3c, 3e). All antennal segments yellow/brown; metasoma yellow at base darkening gradually to light brown at apex.
Head (Figs 3a and 4a). Antenna with scape length 4.7–4.8× scape width with five pores along mid line of single continuous convex ridge on ventral surface, pores small, approximately same diameter as base of adjacent seta, pedicel length 1.8–1.9× pedicel width, F1 length 0.6× F1 width, F2 length 0.5–0.6× F2 width, F3 length 3.5–3.7× F3 width, club length 3.3–6.5× club width and 1.5–1.6× F3 length.
Aphelinusbrevicalcar Thomson, 1876, lectotype female (LUZN, examined). Card-mounted. Label data: “Aphelinusbrevicalcar | Lectotype ♀. Thomas. | M. de V. Graham || Lectotype || Type No. 1574:1”.
Aphelinusbrachyptera Kurdjumov, 1913, lectotype female (NHW, examined). Card-mounted. Label data: “Aphelinusbrachyptera | (Först. MS). | Lectotype | M. de V. Graham || A.brachyptera | Förster Type || Collect. G. Mayr || 182”.
Aphelinusdubia Kurdjumov, 1913, lectotype female (NHW, examined). Mounted on minuten pin on block. Label data: “Aphelinus | (Föst. MS.) K | Lectotype || M.dubia | Förster, Type || Collect. | G. Mayr”. Paralectotypes, five females. Mounted on minuten pins on block. Label data as lectotype.
Old World and some New World populations, discussed below.
Discussion.
The collections from France, Italy, Morocco, and Spain correspond to those discussed in Kazmer et al. (1996). The leg coloration of of A.asychis specimens as described in the diagnosis is unique among species in the asychis group. We are treating collections from Spain, Morocco, and Italy as conspecific, noting that the metafemora are dark brown [not yellow]. We are also treating the collection from England, Sussex, as conspecific, noting that the profemora and mesofemora are yellow [not dark brown with apex yellow or pale]. We are treating the population from Hoda, Hawaii, and Bangalore, India, as conspecific, noting that the protibia are brown [not yellow]. Although the male antennal club in asychis is usually longer than in other species, we note that it is of comparable length (4.0–4.5 L/W) in the series from Montpellier and Antibes, France.
Numerous populations of A.asychis are present in North America. The population from Randall Co., Texas, is presumed to have originated from biological control program releases of A.asychis against the Russian wheat aphid in that area. The population from Wayne Co., Missouri, was collected from a Malaise trap, and one specimen from Florida was collected by a flight interception trap, both in the late 1980’s. The populations from Stillwater, Oklahoma, originated from a lab culture of A.asychis for Schizaphisgraminum biological control. The two specimens collected from Manhattan, Kansas, from spotted alfalfa aphid in a greenhouse probably result from agricultural research at Kansas State University. The series from Alameda Co., California, 1962, is from the former UC Insectary, Oxford Tract, and was possibly being researched as a candidate for biological control of yellow clover aphid or other pest aphids.
However, in a few cases, A.asychis was found in North America before any documented biological control releases. An A.asychis specimen was collected in Florida in 1952, which does not have any associated host data. Two specimens from Maryland, one from Myzuspersicae and one from strawberry aphids were collected in 1962 and 1950, respectively. One specimen from Maine in 1958 was collected from a Capitophorus aphid mummy. Two specimens from Colorado were collected from Myzuspersicae mummies, one in 1940 and the other in 1988. These records are especially interesting because they are geographically close to the type locality and host of A.semiflavus. The specimens from Minnesota, Ohio, and California (UCRC ENT 75429 and 75431) did not have collecting date or host information.
The lectotype and paralectotype females of affinis Förster were examined (NHW, Vienna). They are point-mounted specimens in good condition. We concur with Graham (1976) that affinis Förster is a junior synonym of asychis Walker. The lectotype of brachyptera Kurdjumov (NHW, Vienna) was examined. It is a brachypterous female mounted on a minuten pin through the mesosoma. We concur with Graham (1976) that brachyptera Kurdjumov is a junior synonym of asychis Walker. The lectotype female of brevicalcar Thomson (LUZN, Sweden) was examined. It is a card-mounted female in reasonably good condition. We concur with Graham (1976) that brevicalcar Thomson is a junior synonym of asychis Walker.
The type material of dubia Kurdjumov (NHW, Vienna) which consists of a female lectotype and five female paralectotypes, mounted together on a small wooden block on minuten pins, and a second pin with a wooden block bearing five additional female paralectotypes, was examined. A red dot next to one pin identifies the lectotype. The specimens are dirty and in poor condition and the minuten pins are rusting. Although we note some color variation in the metasoma of these specimens, we concur with Graham (1976) that dubia Kurdjumov is a junior synonym of asychis Walker. The lectotype female and paralectotype females of euthria Walker (BMNH, London) were examined. The lectotype and one paralectotype are mounted on the same card and are in poor condition (lectotype is missing all of the metasoma). Three additional paralectotypes individually card mounted are in reasonably good condition. We concur with Graham (1976) that euthria Walker is a junior synonym of asychis Walker.
Females and males. Legs with procoxa brown (Fig. 5d) [not yellow (Fig. 9d)], profemur and mesofemur light brown with apex yellow or pale (Fig. 5d) [not entirely yellow (Figs 7d, 9d) or dark brown with apex yellow or pale (Fig. 3d)], mesotibia yellow (Fig. 5d) [not dark brown with apex yellow or pale (Fig. 3d)], metatibia light brown with apex yellow (Fig. 5d) [not entirely yellow (Fig. 9d), or dark brown with apex yellow or pale (Fig. 3d)].
Color (Fig. 5b, 5d, 5f). Head and mesosoma dark brown; radicle and basal portion of scape yellow/white, apical portion of scape and pedicel brown, and F1, F2, F3, and club yellow, tip of club dusky; legs with mesocoxa and metacoxa brown, metafemur yellow, protibia yellow; metasoma yellow from base to apex, lateral margins of metasoma darker than mesal area except in basal quarter.
Body length. 0.7–0.9 mm (n=3; slide mounts) (Holotype 0.7 mm).
Head (Figs 5b and 6b). Width 1.1–1.2× head length in anterior view; frontovertex width 0.5–0.6× head width and 1.9–2.6× frontovertex length; posterior ocelli diameter 0.6–0.8× posterior ocelli to eye margin distance and 1.5× the posterior ocelli to occipital margin distance; antenna as in Figure 6b with scape length 5.8–7.3× scape width, pedicel length 1.8–2.2× pedicel width, F1 and F2 subquadrate, length of both 1.0× width, F3 length 1.4–1.6× F3 width, and club length 2.9–3.7× club width and 2.8–3.1× F3 width, club with 8 longitudinal sensilla.
Mesosoma (Figs 5d, 5f and 6g). Mid lobe of mesoscutum length 0.8× mid lobe width, and with two pairs of long setae (one pair lateral and one pair posterior) and 34–35 short setae; side lobes of mesoscutum each with one pair of long setae and one pair of short setae; scutellum with two pairs of long setae (one pair anterior and one pair posterior); mesotibial spur length 0.7–0.8× mesobasitarsus length, metatibial spur length 0.4–0.6× metabasitarsus length.
Fore wing (Fig. 6d). Length 2.3–2.4× fore wing width, longest marginal seta 0.1–0.2× fore wing width; costal cell 0.6–0.7× length of marginal vein, with one line of 6–7 setae on ventral surface and 1–2 dorsal setae in apical quarter; marginal vein with two rows of 14–15 large dorsal setae, one row of 7–8 small dorsal setae, and one row of 8–10 ventral setae; interspace between basal cell and linea calva with 30–33 setae arranged in three complete lines and two incomplete lines; linea calva closed with 2–3 setae at its posterior end, setae bordering linea calva proximally arranged uniformly and evenly to posterior margin of wing.
Hind wing (Fig. 6e). Length 4.6× hind wing width, longest marginal seta 0.7× hind wing width.
Metasoma (Figs 5d, 5f and 6f). Length 1.4× mesosoma length; ovipositor length 1.6× mesotibia length and 1.3× metatibia length; third valvula length 0.3–0.4× ovipositor length.
Description: Male (Figs 5a, 5c, 5e and 6a, 6c, 6h). Similar to female except:
Color (Fig. 5a, 5c, 5e). All antennal segments yellow/brown; metasoma yellow at base darkening gradually to light brown at apex.
Head (Figs 5a and 6a). Antenna with scape length 5.2–7.5× scape width with five pores along mid line of single continuous convex ridge on ventral surface, pores small, approximately same diameter as base of adjacent seta, pedicel length 2.0–2.2× pedicel width, F1 length and F2 length both 0.6× width, F3 length 2.8–3.3× F3 width, club length 3.5–4.7× club width and 1.6–1.7× F3 length.
https://binary.pensoft.net/fig/124064Other material examined.
None.
Hosts.
The original material was collected from Diuraphisnoxia in the field in Dmitrievka, Kazakhstan. In lab culture, Diuraphisnoxia on wheat was used as the host.
Distribution.
The species is known only from type material from Dmitrievka, Kazakhstan.
Discussion. The collections from Kazakhstan, Dmitrievka correspond exactly to those discussed in Kazmer et al. (1996). The most notable distinction between A.kazakhstanensis and all other A.asychis group species is the yellow mesotibia, combined with the profemur, mesofemur, and metatibia light brown at base and yellow at apex. Aphelinusasychis specimens have mesotibia, metatibia, profemur, and mesofemur dark brown at base and yellow at apex. Aphelinussemiflavus has mesotibia, profemur, and mesofemur yellow, and metatibia brown at base and yellow at apex. Aphelinussinensis specimens have mesotibia, metatibia, profemur, and mesofemur all yellow.
AnimaliaHymenopteraAphelinidae69DA474E-B332-583B-AD85-B7A841DEA5ACAphelinussemiflavusHoward, 1908AphelinussemiflavusHoward, 1908.AphelinusbrevipennisGirault, 1917, synonymy by Gahan (1924).Diagnosis.
Females and males. Legs with procoxa brown (Fig. 7d) [not yellow (Fig. 9d)], profemur, mesofemur, mesotibia entirely yellow (Fig. 7d) [not dark or light brown at base and yellow or pale at apex (Figs 3d, 5d)] and metatibia light brown with apex yellow (Fig. 7d) [not entirely yellow (Fig. 9d) or dark brown with apex yellow or pale (Fig. 3d)].
Aphelinussemiflavus, point-mounted specimens 7a male, antennae and face, lateral view (CNC 00122807) 7b female, antennae and face, anterior view (CNC 00122818) 9c male, habitus, lateral view (USNM ENT 2076436) 7d female, habitus, lateral view (paralectotype) 7e male, habitus, dorsal view (USNM ENT 2076436) 7f female, habitus, dorsal view (paralectotype).
https://binary.pensoft.net/fig/124065Description.
Female (Figs 7b, 7d, 7f and 8b, 8d–8g).
Color (Fig. 7b, 7d, 7f). Head and mesosoma dark brown; radicle and basal portion of scape yellow/white, apical portion of scape and pedicel brown, and F1, F2, F3, and club yellow, tip of club dusky; legs with mesocoxa and metacoxa brown, metafemur and protibia, entirely yellow; metasoma yellow from base to apex, lateral margins of metasoma darker than mesal area except in basal quarter.
Body length. 0.5–1.0 mm (n=3; slide mounts) (Paralectotypes 0.5–0.6 mm).
Head (Figs 7b and 8b). Width 1.2–1.3× head length in anterior view; frontovertex width 0.4–0.5× head width and 3.1–3.7× frontovertex length; posterior ocelli diameter 0.5× posterior ocelli to eye margin distance and 1.5× posterior ocelli to occipital margin distance; antenna as in Figure 8b with scape length 6.8–7.0× scape width, pedicel length 2.0–2.3× pedicel width, F1 and F2 subquadrate, length of both 0.9– 1.1× width, F3 length 1.5–1.9× F3 width, and club length 3.2–4.4× club width and 2.6–2.3× F3 width, club with 8 longitudinal sensilla.
Mesosoma (Figs 7d, 7f and 8g). Mid lobe of mesoscutum length 0.7–0.8× mid lobe width with two pairs of long setae (one pair lateral and one pair posterior) and 30–31 short setae; side lobes of mesoscutum each with one pair of long setae and one pair of short setae; scutellum with two pairs of long setae (one pair anterior and one pair posterior); mesotibial spur length 0.6–0.7× mesobasitarsus length, metatibial spur length 0.4–0.5× metabasitarsus length.
Fore wing (Fig. 8d). Length 2.3–2.8× fore wing width, longest marginal seta 0.1× fore wing width; costal cell length 0.6–0.7× length of marginal vein, with one line of 10–12 setae on ventral surface and 1–2 dorsal setae in apical quarter; submarginal vein with two setae; marginal vein with two rows of 14–17 large dorsal setae, one row of 7–13 small dorsal setae, and one row of 7–10 ventral setae; interspace between basal cell and linea calva with 31–40 setae arranged in three complete line and one to two incomplete lines; linea calva closed with 3 setae at its posterior end, setae bordering linea calva proximally are arranged uniformly and evenly to posterior margin of wing.
Hind wing (Fig. 8e). Length 3.5–3.7× hind wing width, longest marginal seta 0.2–0.3× hind wing width.
Metasoma (Figs 7d, 7f and 8f). Length 2.0× mesosoma length; ovipositor length 0.9× mesotibia length and 1.2× metatibia length; third valvula length 0.3× ovipositor length.
Description. Male (Figs 7a, 7c, 7e and 8a, 8c, 8h). Similar to female except:
Color (Fig. 7a, 7c, 7e). All antennal segments yellow/brown; metasoma yellow at base darkening gradually to light brown at apex.
Head (Figs 7a and 8a). Antenna with scape length 6.4× scape width with five pores along mid line of single continuous convex ridge on ventral surface, pores small, approximately same diameter as base of adjacent seta, pedicel length 1.7–2.2× pedicel width, F1 length 0.6× F1 width, F2 length 0.6× F2 width, F3 length 4.2–4.4× F3 width, club length 6.2× club width and 1.5× F3 length.
https://binary.pensoft.net/fig/124066Type material examined.
AphelinussemiflavusHoward 1908, lectotype female (USNM, examined). Label data: “Type | No. 12031 | USNM || Aphelinus n.sp. | near mali || Myzuspersicae | Ft. Collins, Colo. | C.B. Gillette | det. July 15, 1908”. The slide containing the lectotype has 3 female and 2 male specimens under one cover slip. The female specimen in the lowest middle portion of the slide is herein designated lectotype, and the slide has been labeled accordingly. Paralectotypes. (USNM, examined). Two females and two males on same slide, data as lectotype. Two females, one male, one sex unknown, on cardmounts with label data reading as “Myzuspersicae | Fort Collins, CO || C. B. Gillette | Det. July, 15, 08 || Type | No. 12031| U.S.N.M.”.
AphelinusbrevipennisGirault 1917, lectotype female. (USNM, examined). Slide-mounted parts: “Aphelinus | brevipennis | Girault | female type || 19801”. Point mount: “Ohio || 1693 || 19801”. The female lectotype designated herein is mounted on a point, and parts consisting of an antenna, a fore wing, a hind wing, a mid leg and a hind leg were dissected by Girault and mounted on a slide. Paralectotype male (USNM). Point-mounted. Label data: “Ohio || 1704 || 19801”.
New World, with few populations in Old World (from France, Korea and Spain).
Discussion.
There has been confusion in the past about whether or not A.asychis and A.semiflavus are separate species. Based on the material examined, leg coloration patterns clearly differ between them. Regarding the New World populations examined, we are treating most North American populations as semiflavus, noting that in the Mexico and Winnipeg, Canada, populations, the metatibia are yellow/brown [not dark brown at base with apex pale].
We consider the Bangalore, India, and Dezful, Iran populations as sp. nr.semiflavus, noting that the India population has mid tibia and mid femora with brown [not yellow] and the Dezful, Iran, population has legs like semiflavus except one female with brown [not yellow] on mid tibia and mid femora. There is one specimen from Virginia that we are treating as possibly semiflavus, noting that the fore femora and fore tibia are brown [not yellow].
We have examined the lectotype female and paralectotype male of brevipennis Girault and agree with the conclusion of Gahan (1924) that it is a junior synonym of semiflavus Howard. Although Ferrière (1965), Nikol’skaya and Yasnosh (1966), and Yasnosh (1978) treated semiflavus as a junior synonym of asychis Walker, for reasons discussed above we consider it to be a distinct and valid species.
Future work should use reciprocal crosses and perhaps molecular data to determine whether populations of A.semiflavus in Old World vs. New World are in fact one species, are races of one species with different host ranges, or are two distinct cryptic species.
Females and males. Legs with procoxa yellow (Fig. 9d) [not brown (Figs 3d, 5d, 7d)], profemur, mesofemur, mesotibia, and metatibia yellow (Fig. 9d) [not brown at base with apex yellow or pale (Figs 3d, 5d, 7d)].
Color (Figs 9b, 9d, 9f). Head and mesosoma dark brown; radicle and basal portion of scape yellow/white; apical portion of scape and pedicel brown; F1, F2, F3, and club yellow; tip of club dusky; legs with mesocoxa and metacoxa brown, metafemur and protibia yellow; metasoma yellow from base to apex, lateral margins of metasoma darker than mesal area except in basal quarter.
Body length. 0.6–0.9 mm (n=3; slide mounts) (Holotype 0.8 mm).
Head (Figs 9b and 10b). Width 1.2–1.3× head length in anterior view; frontovertex width 0.4× head width and 1.0–1.4× frontovertex length; posterior ocelli diameter 0.6× posterior ocelli to eye margin distance and 1.5× posterior ocelli to occipital margin distance; antenna as in Fig. 4b with scape length 5.0–5.9× scape width; pedicel length 1.5–1.9× pedicel width; F1 and F2 subquadrate, length of both 0.9–1.0× width; F3 length 1.4–1.8× F3 width; club length 3.1–3.3× club width and 2.5–3.0× F3 width, club with 8 longitudinal sensilla.
Mesosoma (Figs 9d, 9f and 10g). Mid lobe of mesoscutum length 0.7× mid lobe width, with two pairs of long setae (one pair lateral and one pair posterior) and 31–33 short setae; side lobes of mesoscutum each with one pair of long setae and one pair of short setae; scutellum with two pairs of long setae (one pair anterior and one pair posterior); mesotibial spur length 0.7–0.80× mesobasitarsus length; metatibial spur length 0.5× metabasitarsus length.
Fore wing (Fig. 10d). Length 2.5–2.8× fore wing width, longest marginal seta 0.1–0.2× fore wing width; costal cell 0.8× length of marginal vein, with one line of 6–7 setae on ventral surface and 1–2 dorsal setae in apical quarter; submarginal vein with two setae; marginal vein with two rows of 12–18 large dorsal setae, one row of 7–11 small dorsal setae, and one row of 7–10 ventral setae; interspace between basal cell and linea calva with 18–33 setae arranged in two complete lines and two incomplete lines; linea calva closed with 2–3 setae at its posterior end, setae bordering linea calva proximally are arranged uniformly and evenly to posterior margin of wing.
Hind wing (Fig. 10e). Length 3.7–4.5× hind wing width; longest marginal seta 0.3–0.5× hind wing width.
Metasoma (Figs 9d, 9f, and 10f). Length 1.1× mesosoma length; ovipositor length 1.3× mesotibia length and 1.2× metatibia length; third valvula length 0.4× ovipositor length.
Description. Male (Figs 9a, 9c, 9e and 10a, 10c, 10h). Similar to female except:
Color (Fig. 9a, 9c, 9e). All antennal segments yellow/brown; metasoma yellow at base darkening gradually to light brown at apex.
Head (Figs 9a and 10a). Antenna with scape length 4.9× scape width, with five pores along mid line of single continuous convex ridge on ventral surface (Fig. 2); pores small, approximately same diameter as base of adjacent setae, pedicel length 1.7× pedicel width; length of F1 and F2 both 0.5× their width; F3 length 4.4× F3 width; club length 4.6× club width and 1.4× F3 length.
Holotype (deposited in USNM). Female, card mounted. Label data: “Texas: Brazos Co. | College Station | TAMU Lab Culture | 15.xii.1992 T92/051 | ex: Diuraphisnoxia | on wheat || T92/051 orig. collection | P.R. of China | Ningxia | 21.vi.1992 | Keith Hopper coll. | ex. Diuraphis | agropyronophaga || TAMU-ENTO X0852864”.
Paratypes (deposited in USNM, TAMU, BMNH, EMEC, CNC). 30 card mounts (9 female, 21 male). 10 card mounts (2 female, 8 male) from original material with label data reading “P.R. China: Pingluo | Ningxia 19.xi.1992 | T92/051 orig. mat. | Keith Hopper | ex. Diuraphis” (TAMU-ENTO accession numbers: females: X0852882, X0852886; males: X0852878 to –79, X0852883, X0852887, X0852889 to –92). 10 card mounts (7 female, 3 male) from F1 progeny with the same label data as holotype (TAMU-ENTO accession numbers: females: X0852864 to –67, X0852870, X0852873, X0852876; males: X0852874, X0852871, X0852863). 10 (all male) card mounts, voucher specimens from non-destructive DNA extraction, with same label data as original material (TAMU-ENTO accession numbers: X0856046 to –55). 7 slide mounts (5 female, 2 male). 3 slide mounts from original material with same label data as above (females: X0852880, X0852888; male: X0852885). 4 slide mounts from F1 progeny, same label data as above (females: X0852869, X0852875, X0852877; male: X0852868).
The original material was collected from Diuraphistritici (Gillette 1911) (the junior synonym Diuraphisagropyronophaga is given on the holotype label) in the field in China. In lab culture, Diuraphisnoxia on wheat was used as the host.
Distribution.
Northern China and Japan.
Discussion.
The collections from China, Pingluo, Ningxia correspond exactly to those discussed in Kazmer et al. (1996). The most notable traits of the type specimens of A.sinensis from Pingluo, China, that distinguish them from other A.asychis group specimens examined, are the presence of yellow procoxae, yellow femora, and yellow tibiae. In all other asychis group species, all coxae are brown, and femora and tibiae are patterned with brown. The series from Harbin, China, exhibits the same leg-pattern coloration as the type series. We are treating the Harbin series as conspecific, noting that the head and metasoma are much darker, almost black, and the antennal club is darker at the apex than in material from Pingluo. There is one specimen from the Honshu, Japan, series that has the same leg coloration patterns as the type series from Pingluo, however, the other specimens in this series resemble A.asychis.
Other Potential New Species
There are two additional potential new species, but there is not enough material to describe them. One potential new species is from Harrow, Canada, and has leg coloration like asychis, but includes a brown metafemur [not yellow]. There are two other Canadian series with one specimen in each series that also has this leg coloration. The other potential new species is from Dorking, England, and is represented by a single specimen that has leg coloration with all segments very dark brown.
Acknowledgements
We would like to thank Jewel Coffey, Bryant McDowell and Itzel Cetina, Texas A&M lab technicians, for digital imaging/processing, plate preparation, database management, and specimen preparation. Thanks to Courtney Hendler, Bethany Lefner and Ada Morales, Texas A&M undergraduate student workers, for digital imaging, and thanks to Kathryn Lanier, USDA-ARS-BIIRL, for maintaining cultures of Aphelinus and shipping specimens. So many museum curators and colleagues have helped with this research that we cannot list them all, but we would like to recognize Serguei Belokobylski (ZIS), Natalie Dale-Skey (BMNH), John Heraty (UCR), John Huber (UCR), Mike Gates (NMNH), Gary Gibson (CNC), Jason Mottern (NMNH), John Noyes (BMNH), Andy Polaszek (BMNH), Konstantin Samartsev (ZIS), Serguei Triapitsyn (UCR), Ekaterina Tselikh (ZIS), Zoltan Vás (HNHM), Doug Yanega (UCR), Dominique Zimmerman (NMW), Bob Zuparko (EMEC and CASC). This research has been supported by two awards from the NSF, USA: DEB 1257601 and DEB 1555790.
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Morphological terms used, their definitions, and URI locations on the Hymenoptera Anatomy Ontology web site (Yoder et al. 2010).
Term
Definition
URI
antenna
The anatomical cluster that is composed of the scape, pedicel and flagellum.
http://purl.obolibrary.org/obo/HAO_0000101
base
The tergum that is located on abdominal segment 2 AND The tergum that is located on the abdominal segment 3.
http://purl.obolibrary.org/obo/HAO_0000053 and http://purl.obolibrary.org/obo/HAO_0000056
body
The anatomical cluster that is composed of the whole organism but which excludes the antennae, legs and wings.
http://purl.obolibrary.org/obo/HAO_0000182
club
The anatomical cluster composed of the apical flagellomeres that are differentiated by size from the basal flagellomeres.
http://purl.obolibrary.org/obo/HAO_0001185
compound eye
The compound organ that is composed of ommatidia.
http://purl.obolibrary.org/obo/HAO_0000217
costal cell
The membranous region of the fore wing anterior to the submarginal vein, measured from the basal constriction that delimits the apex of the humeral plate of the wing to the point at which the submarginal vein touches the leading edge of the wing.
http://purl.obolibrary.org/obo/HAO_0000226
coxa
The leg segment that is connected to the body and to the trochanter via conjunctivae and muscles.
http://purl.obolibrary.org/obo/HAO_0000228
digitus
The sclerite that is located distally on the parossiculus.
http://purl.obolibrary.org/obo/HAO_0000385
edge
The margin that extends along the border of two areas that are oriented differently.
http://purl.obolibrary.org/obo/HAO_0000285
eye margin
The margin of the compound eye.
http://purl.obolibrary.org/obo/HAO_0000672
F1
The flagellomere that is proximally attached to the pedicel.
http://purl.obolibrary.org/obo/HAO_0001148
F2
The flagellomere that is located distal to the first flagellomere.
http://purl.obolibrary.org/obo/HAO_0001883
F3
The flagellomere that is located immediately distal to the second flagellomere.
http://purl.obolibrary.org/obo/HAO_0001895
femur
The leg segment that is distal to the trochanter and proximal to the tibia.
http://purl.obolibrary.org/obo/HAO_0000327
Fore wing
The wing that is located on the mesothorax.
http://purl.obolibrary.org/obo/HAO_0000351
frontovertex
The anatomical cluster that is composed of the vertex and the dorsal area of the upper face dorsal to the frontofacial ridge.
http://purl.obolibrary.org/obo/HAO_0001823
genitalia
The anatomical cluster that is composed of the cupula, gonostyle, volsella and the aedeagus
http://purl.obolibrary.org/obo/HAO_0000312
head
The tagma that is located anterior to the thorax.
http://purl.obolibrary.org/obo/HAO_0000397
hind wing
The wing that is located on the metathorax.
http://purl.obolibrary.org/obo/HAO_0000400
leg
The anatomical cluster that is composed of the coxa and all distal leg segments and is connected to the pectus.
http://purl.obolibrary.org/obo/HAO_0000494
longitudinal sensillum
The multiporous plate sensillum that is elongate.
http://purl.obolibrary.org/obo/HAO_0001936
margin
The line that delimits the periphery of an area.
http://purl.obolibrary.org/obo/HAO_0001133
marginal vein
The abscissa that is located along the anterior margin of the fore wing and is thought to correspond to the anterior abscissa of the radius (R1).
http://purl.obolibrary.org/obo/HAO_0000635
mesobasitarsus
The basitarsus that is located in the mid leg.
http://purl.obolibrary.org/obo/HAO_0001131
mesocoxa
The coxa that is located on the mid leg.
http://purl.obolibrary.org/obo/HAO_0001490
mesofemur
The femur that is located on the mid leg.
http://purl.obolibrary.org/obo/HAO_0000576
mesoscutum
The area that is located anterior to the transscutal articulation.
http://purl.obolibrary.org/obo/HAO_0001351
mesosoma
The anatomical cluster that is composed of the prothorax, mesothorax and the metapectal-propodeal complex.
http://purl.obolibrary.org/obo/HAO_0001120
mesotibia
The tibia that is located on the mid leg.
http://purl.obolibrary.org/obo/HAO_0001142
mesotibial spur
The tibial spur that is located on the mesotibia.
http://purl.obolibrary.org/obo/HAO_0001142
metabasitarsus
The basitarsus that is located on the hind leg.
http://purl.obolibrary.org/obo/HAO_0000587
metacoxa
The coxa that is located on the hind leg.
http://purl.obolibrary.org/obo/HAO_0000631
metasoma
The tagma that is connected anteriorly to the metapectal-propodeal complex at the pro- podeal foramen and consists of abdominal segments.
http://purl.obolibrary.org/obo/HAO_0001121
metatibia
The tibia that is located on the hind leg.
http://purl.obolibrary.org/obo/HAO_0000679
metatibial spur
The tibial spur that is located on the metatibia.
http://purl.obolibrary.org/obo/HAO_0000706
mid lobe of mesoscutum
The area that is located between the notauli.
http://purl.obolibrary.org/obo/HAO_0000520
occipital margin
The edge that separates the occiput from the vertex.
http://purl.obolibrary.org/obo/HAO_0001963
ocellus
The multi-tissue structure that is located on the top of the head, composed of the corneal lens, pigment cell, rhabdoms and synaptic plexus.
http://purl.obolibrary.org/obo/HAO_0000661
ovipositor
The anatomical cluster that is composed of the first valvulae, second valvulae, third valvu- lae, first valvifers and second valvifers .
http://purl.obolibrary.org/obo/HAO_0000510
pedicel
The antennal segment that is the second segment of the antenna and is connected proxi- mally with the scape and distally with the flagellum.
http://purl.obolibrary.org/obo/HAO_0000512
phallobase
The anatomical cluster that is composed of the cupulae, gonostipites and volsellae.
http://purl.obolibrary.org/obo/HAO_0000713
posterior ocellus
The ocellus that is paired.
http://purl.obolibrary.org/obo/HAO_0000481
procoxa
The coxa that is located on the fore leg.
http://purl.obolibrary.org/obo/HAO_0001122
profemur
The femur that is located on the fore leg.
http://purl.obolibrary.org/obo/HAO_0001124
protibia
The tibia that is located on the fore leg.
http://purl.obolibrary.org/obo/HAO_0000350
radicle
The area that is located proximally on the scape, is limited distally by a constriction and bears proximally the basal knob.
http://purl.obolibrary.org/obo/HAO_0000889
row
The anatomical cluster that is composed of repeated units of anatomical structures.
http://purl.obolibrary.org/obo/HAO_0000901
scape
The antennal segment that is proximal to the pedicel and is connected with the head via the radicle.
http://purl.obolibrary.org/obo/HAO_0000908
scutellum
The area that is located posteriorly of the transscutal line and is composed of the axillae and the mesoscutellum.
http://purl.obolibrary.org/obo/HAO_0000572
secretory pore
The anatomical space that corresponds to the distal end of an exocrine gland.
http://purl.obolibrary.org/obo/HAO_0001966
seta
The sensillum that is multicellular and consists of trichogen, tormogen, and sense cells. The area that is located between the notaulus and the parascutal carina.
http://purl.obolibrary.org/obo/HAO_0000935
side lobe of mesoscutum
The area that is located between the notaulus and the parascutal carina.
http://purl.obolibrary.org/obo/HAO_0000466
stigma
The patch on the wing that is sclerotized and is located on the anterior margin of the fore wing.
http://purl.obolibrary.org/obo/HAO_0000957
submarginal vein
Basal-most portion of the fore wing vein complex that occurs behind the costal cell; measured from the constriction that delimits the humeral plate to the point at which the vein touches the leading edge of the wing apically.
http://purl.obolibrary.org/obo/HAO_0000972
tarsus
The leg segment that is apical to the tibia.
http://purl.obolibrary.org/obo/HAO_0000992
third valvula
The sclerite that is located posterior to the second valvifer and is connected to the second valvifer via conjuntiva.
http://purl.obolibrary.org/obo/HAO_0001012
wing
The appendage that is between the notum and the pectus and is connected to the body by the axillary sclerite muscles.